BIOGEOGRAPHY

Space, Time, and Life

GLEN M. MAcDONALD
Departments of Geography and Organismic Biology,
Ecology and Evolution
University of California-Los Angeles

JOHN WILEY & SONS, INC.

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PREFACE

This book was written to provide an introductory text that would acquaint under-
graduate students with the field of biogeography. A special emphasis is placed on
instructing geography students in North America, although the topics and exam-
ples are global in scope. After a number of years teaching biogeography to stu-
dents in both the United States and Canada, it seemed to me that three things
were required in such an introductory text.
First, because introductory biogeography students come to the subject from
a wide array of backgrounds, ranging from the humanities, to the social sciences,
to the natural sciences, the text has to presuppose little in the way of background
knowledge in biology or physical geography. Therefore, the book contains ele-
mentary discussions of important concepts and terms from biology and physical
geography with which every biogeographer should be familiar. In addition, as so
much of biogeography depends upon an understanding of how the physical and
biological environments actually affect organisms and influence their geographic
distributions, ample space is provided to examine how physical and biological
factors such as heat, moisture stress, or predation impact individuals, populations,
and species.
Second, biogeographers today work on a diverse range of topics, ranging
from genetic analyses focused on unraveling evolutionary history, to disturbance
studies, to the design of reserves for the conservation of biodiversity. In many
cases, the research of modern biogeographers overlaps considerably with the
work of evolutionary scientists, ecologists, and conservation biologists. The text
was designed to present the scope of biogeographical work as it is defined by the
activities of the practitioners, rather than sticking to narrow academic defini-
tions of the discipline. Therefore the book contains chapters on issues such as
human evolution, disturbance, and conservation as well as discussions of tradi-
tional topics such as biomes, cladistics, and island biogeography. In addition, the
text uses examples from, and provides references to, the recent work of a very
wide variety of biogeographers. Some teachers will focus on those chapters that
most closely follow their own work and the text is designed to be used in this
selective manner or in a broader more encompassing examination of the field.
Third, biogeography is a fascinating subject in large part because of the wonder-
ful variety of organisms and ecosystems we are studying and the engrossing and
powerful scientific concepts that have been formulated and developed by bio-
geographers and scientists in closely aligned disciplines. The text contains many
examples of the natural histories of different species of plants and animals to
help convey this wonderful variety of life. To help relate the human history of
biogeography the text contains short biographical sketches of some of the major
figures in the field. It is hoped that students will see that the ideas that underlie
the science of biogeography were developed by people who were also once
young and curious like themselves. Photographs of both interesting organisms
v
 Vi PREFACE

and seminal figures in biogeography are provided to help establish the tie
between the classroom and the 'real world'.
In the formulation and writing of this text a number of biogeographers
provided excellent critical readings that resulted in insightful comments, help-
CON]
ful suggestions, and important corrections. These readers included: Karen
Arabas-Willamette University; C. John Burk-Smith College; William P. Coff-
man-University of Pittsburgh; Laura E. Conkey-Dartmouth College;
Anthony Davis-University of Toronto; James P. Doerner-University of
Northern Colorado; Melanie DeVore-Georgia College and State U niversity; OtoAPTER 1 AN
Holly Freifeld; Thomas W. Gillespie- University of South Florida; N. C. Hey-
wood-UW Stevens Point; Sally Horn-University of Tennessee; Kam-Biu
Liu-Louisiana State University; Robert Feet-University of North Carolina;
George Robinson-SUNY Albany; Gary L. Walker-Appalachian State Uni- PART I Sl
versity; Barry B. Warner-University of Waterloo; Susy S. Ziegler-U niversity
of Minnesota. Any mistakes that remain in this edition are solely the responsi-
bility of the author. ~R 2 SOJ
Many biogeographers generously provided copies of their publications for Bioi<
my use in preparation of this text. I owe these colleagues a debt of gratitude for
introducing me to much fine science and enlarging my view of our discipline. It is
a shame that due to the constraints of space, I could use only a fraction of this
research for detailed and specific examples in this book. Many of the concepts Phys
and ideas from these publications appear in more general ways throughout the
book and are accordingly referenced in the appropriate chapters.
The preparation of this text would not have been possible without the won-
derful guidance, assistance, support, and encouragement of the present and past
employees of John Wiley and Sons and Hermitage Publishing Services. I cannot
name all of the fine people who have aided me but I must acknowledge Karen
OWY'TER 3 TH
Ayoub, Ryan Flahive, Jennifer MacMillan, Larry Meyer, Hilary Newman, Joan
Petrokofsky, Denise Powell, Sandra Rigby, and Anne Smith. Last but not least I Ligh
must give a special acknowledgment to Nanette Kauffman who got me into all of Tern]
this in the first place and remains a good friend despite that (Breathe, breathe the
air. .. ).
Moi~
I am afraid that writing this book took much time and energy away from
some very deserving people, namely my graduate students and my family. I
thank those students who were affected: Alexis Aguilar, Roslyn Case, Steve Otht
Esselman, Bruce Gervais, Judy King, Amanda Pete!, Sarah Peugh, David Por- Intel
inchu, Aaron Potito, and Sigrid Rian for their understanding and unflagging En vi
enthusiasm-and for keeping it real. Most importantly, I owe a huge debt of
gratitude to my wife Joanne and my children Alex and Hilary. There were far ::lloAPTl:R 4 BI(
too many weekends, evenings, and vacation periods spent with me hunched
over the computer or engrossed in reading to be fair to a young family. The Pred
Com
patience and encouragement they showed when I was confronted by problems
Sym
in writing or meeting deadlines is deeply appreciated. Joanne also contributed Con:
much in the way of critical reading, text preparation, and generally keeping me Bioi
under control. So, to my family I say-thank you and now let's go out and have
some fun! ::x.A.PTeR s DL
Frre
\Vm
Floc
\
to help establish the tie

Imber of biogeographers
tsightful comments, help- CONTENTS
readers included: Karen
1 College; William P. Coff-

y-Dartmouth College;
Doerner-University of
~ge and State University;
CHAPTER 1 AN INTRODUCTION
:outh Florida; N. C. Hey-
of Tennessee; Kam-Biu
ersity of North Carolina;
-Appalachian State Uni- PART I SPACE AND LIFE
sy S. Ziegler-University
n are solely the responsi-
CHAPTER 2 SOME BASICS 9
s of their publications for Biology and the Hierarchies of Life 9
1es a debt of gratitude for Taxonomic Hierarchy 9
1iew of our discipline. It is Ecological Hierarchy 15
tse only a fraction of this Trophic Hierarchy 16
)k. Many of the concepts Physical Geography and the Functioning of the Earth 22
eral ways throughout the Global Climate 23
: chapters. Microclimate 31
possible without the won- World Soils 32
It of the present and past The Physical Environment of Lakes 35
1lishing Services. I cannot The Physical Environment of Oceans 38
nust acknowledge Karen
CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE 43
er, Hilary Newman, Joan
mith. Last but not least I Light 45
tan who got me into all of Temperature 47
hat (Breathe, breathe the Plants 48
Animals 54
:e and energy away from Moisture 60
udents and my family. I Plants 60
Animals 65
ilar, Roslyn Case, Steve
Other Physical Factors 66
)arab Peugh, David Par- Interacting Physical Controls on Geographic Distributions 68
standing and unflagging Environmental Gradients and Species' Niches 70
ly, I owe a huge debt of
d Hilary. There were far CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE 77
spent with me hunched
· to a young family. The Predation 77
confronted by problems Competition 83
Symbiosis: Mutualism, Commensalism, Parasitism, and Mimicry 87
Joanne also contributed
Combined Physical and Biological Controls on Distribution 90
rrd generally keeping me Biological Interactions, Gradients, and Niches 91
ow le t's go out and have
CHAPTER 5 DISTURBANCE 97

Fire 104
Wind 113
Flooding 116

vii
viii CONTENTS

Other Physical Disturbances 122 ~10 Rl

Pathogens 126 SL
Marine Disturbances 126
Defu
Dete
CHAPTER 6 COMMUNITIES, FORMATIONS AND BIOMES 132 Factc

Communities 133
Plant Physiognomy, Vegetation Structure, and Formations 139 The1
Ecological Equivalents, Life Zones, and the Biomes 141
Tropical Rainforest 145
Tropical Seasonal Forest 153
ltopical Savanna 154
Desert 159
The Mediterranean Biome 164
Temperate Grassland 167 ~11 BI
Temperate Forests 171
Temperate Rainforest 176 The I
Coniferous Boreal (Taiga) and Montane Forests 178 Earl)
Tundra 181 The I
The I
The I
The <
PART II TIME AND LIFE ::J1oU0TER 12 Hl
Hum
CHAPTER 7 CHANGING CONTINENTS AND CLIMATES 191

Life and the Geologic Time Scale 191

Hum
Shifting Continents 196
Quaternary Climatic Change 205
Future Changes in Continents and Climate 217

CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION 221

PART Ill 1
Dispersal 228
Colonization, Seasonal Migrations, and Irruptions 234
::.r.PTER 13 D1
Diffusion Versus Jump Dispersal 241
Dl
Barriers, Corridors, Filters, Stepping Stones, and Sweepstakes 246
Recent Invasions by Exotic Species 252 Map]
Biog
Com
CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION 262

Evolution and Speciation 262

Some Basic Genetics 263
Historical Development of Evolutionary Theory 267
Isolation and Speciation 272
The Temporal Pattern of Evolution 275
Direction in Evolution 277
Perfection in Evolution 278
Increasing Species Diversity 279 Biog
Geography and Evolution: Founder Effects, Bottlenecks, Vicariance Events, Adaptive
Radiation, and Evolutionary Convergence 280
Extinction 287
The Relationship Between Evolution and Extinction 295 Bey<
 CONTENTS ix

CHAPTER 10 REALMS, REGIONS, AND PROVINCES: THE BIOGEOGRAPHIC

SUBDIVISIONS OF THE EARTH 299
Defining Biogeographic Realms, Regions, and Provinces 299
Determining the Boundaries Between Regions 305
Factors Behind the Modern Biogeographical Regions 307
Evolution of the Mammals 308
Evolution of the Flowering Plants 311
139
The Modern Biogeographic Regions 315
Nearctic and Palearctic Regions-The Holarctic 315
Neotropical Region 317
Ethiopian (African) Region 319
Oriental Region 320
Australian Region 321

CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION 324

The Primate Linkage 324

Early Primates 327
The Hominids: Australopithecus 330
The Hominids: Early Homo 336
The Hominids: Homo Sapiens 340
The Geographic Expansion of Modern Humans 344

CHAPTER 12 HUMANS ASA FORCE IN EVOLUTION AND EXTINCTION 350

Humans as an Evolutionary Force 350

Animal and Plant Domestication 351
Questions of the Origin and Spread of Agriculture 357
Humans as a Force of Extinction 362
Prehistoric Extinctions 362
Historic Extinctions 368

PART Ill THEORY AND PRACTICE

CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC

DISTRIBUTIONS 377
' 246
Mapping Biogeographic Distributions 378
Biogeography of Range Size and Range Shape 380
Common Biogeographical Distributional Patterns 386
Endemic and Cosmopolitan Distributions Revisited 387
Continuous Zonal Biogeographic Distributions 388
Amphiregional Disjunct Distributions 388
Dispersal Disjunctions 389
Climatic Disjunctions 390
Geologic Disjunctions 391
Evolutionary Disjunctions 391
Biogeographic Relicts 392
Biogeographic Distributions and the Reconstruction of Evolutionary History 394
1riance Events, Adaptive Centers of Origin and the Dispersalist Model 395
Cladistic Biogeography 396
Panbiogeography and Vicariance Models 398
Beyond Vicariance Biogeography: The Phylogeographic Revolution 402
X CONTENTS

CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY 406

What is Biological Diversity? 406 CHA P T ER

How Many Different Species are there on the Earth? 409
Latitudinal and Altitudinal Diversity Gradients 411
Controls on Geographic Gradients of Species Diversity 415
Historical Theories 416 N I"
Equilibrium Theories 418
Island Biogeography 428
Geographic Patterns of Island Biodiversity 429
The Equilibrium Theory of Island Biogeography 431
lno
The Theory of Island Biogeography Today 439
geo1
CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE 451
stan
Bio1
The Value of Conservation 452 plaiJ
Endangered and Threatened Species 454 that
Biogeography and Endangered Species 459 pres
Biogeography and Conservation Planning 464
General Strategies for Species Conservation and Biodiversity Conservation 466
Habitat Restoration and Conservation 473 ate
The Global Warming Challenge 477 help
Some Reflections 480 enct
dot
tion
GLOSSARY 485
dire'
that
PHOTO CREDITS 495 hav(
bird
ILLUSTRATION CREDITS 496 you
you
IND EX 499 this
plan

prm
and
ple 1
and
tras1
mah
Stat
in$(

that
Flor
absc::
Sou1
will
sud
ever
anin

lJbr
terrm
ry 406

~09
CHAPTER 1
415
AN INTRODUCTION

431
In opening this text, you probably have three basic questions: What exactly is bio-
geography? Why should I bother studying it? How will this book help me under-
fALLENGE 45 1 stand biogeography? Let's start with a simple answer to the first question.
Biogeography is the study of the past and present geographic distributions of
plants and animals and other organisms. 1 Of course, there is much more to it than
that. We will explore this question further and expand on this definition
presently, but first let us consider why you might want to study biogeography.
Jdiversity Conservation 466
More than most sciences, biogeography helps us to understand and appreci-
ate the living environment that we experience every single day. Biogeography
helps us answer questions such as how the great diversity of life that we experi-
ence today arose, where the modern human species came from, and what we can
do to preserve the natural environment in the face of increasing human popula-
tion growth and environmental change. How does answering such questions
directly affect you? You cannot set foot outside your door without seeing plants
that are native to your area. Many of the plants you see are exotic species that
have been introduced by humans. You cannot escape hearing the calls of wild
birds, some of which are native and some of which have been introduced. Even if
you do not know the scientific names of the plants and animals near your home,
you are familiar with the way they look and sound. You must have wondered how
this diversity of life around you arose and specifically where all of these different
plants and animals that live near you originated.
If you have ventured far from your home, perhaps visiting another state,
province, or country, you have undoubtedly noticed differences in the vegetation
and animal life you encountered. For example, during the winter millions of peo-
ple travel from the northeastern United States and Canada to enjoy the sunshine
and palm-lined beaches of southern Florida. The green vegetation of Florida con-
trasts greatly with the cold and leafless winter forests of the Northeast. Many ani-
mals found in Florida, such as alligators, are not found in the northeastern United
States or in Canada. Why are alligators and most other plants and animals found
in southern Florida not found in the Northeast? The obvious answer might be
that these plants and animals require the warm and humid environment of
Florida to survive. However, many plant and animal species from Florida are also
absent from the other warm tropical and subtropical areas of the earth. Travel to
South America, West Africa, Southeast Asia, or northeastern Australia and you
will see palm trees and many interesting animals, including relatives of alligators
such as crocodiles, but not a single native alligator (Fig. 1.1). Surprisingly, how-
ever, you will find native alligators in southern China. Why are certain plant and
animals species limited to relatively small areas of the earth? Why are other types

1 Throughout the text key terms and concepts will appear in bold lettering. Short definitions for these

terms and concepts can be found at the end of the book.

1
2 CHAPTER 1 AN INTRODUCTION

us un
wher'

plant
sible
envir
peop
tion
lesseJ
a reas
proVJ
anirn
alter'
mals,
tant :
take
FIGURE 1.1 The world distribution of alligator species (Alligator mississipiensis and
Alligator sinensis). Understanding why alligators are found only in North America and
China is a classic biogeographical question. Alligators are found in warm subtropical past
regions because they cannot tolerate prolonged exposure to cold and freezing tempera- expl<
tures. In the geologic past, Europe, Asia, and North America were joined together in one est p
large continent that experienced relatively warm climates. Therefore the geographic range mals
of the ancestors of modern alligators was much larger than it is today. Over time, as conti-
nents shifted, climate changed, and alligators faced competition from crocodiles and orga
caimans, the geographic range of alligators became fragmented and small. lion
struc
that
of organisms widely distributed? Why can't you grow many of the plants found in ent '
Florida in a garden in New York? men
The role of biogeography is to record such geographic differences in vegeta- chal
tion and wildlife and seek to explain them. Biogeography can tell us why North cant
American alligators (Alligator mississipiensis) are found only in warm, moist areas fam
such as Florida and adjacent states. Biogeography can also help us to understand (the
phy.
why alligators are also found in China (Alligator sinensis), and not in tropical
regions of Africa and Australia. In studying biogeography, you will come to under- geo:
stand that a combination of geographical, environmental, and historical factors led bioi
to the great diversity of plant and animal life. In studying biogeography, you can cipl
develop a much greater awareness, understanding, and appreciation of the won- and
derful diversity of plant and animal life that you encounter every single day. pal1
The study of biogeography also helps us to appreciate the development bet
and history of our own species. We do not stand as far apart from the natural tha
world as we might think. Our development, both biologically and culturally, is a gee
product of geography, the earth's physical environment, and interactions with gra
other organisms. The evolution and spread of humans around the world is one of inc
the great stories of biogeography. At present, humans play an increasingly rar:
greater role in changing the world's environments and have a significant impact me
on the lives of plants and animals with which they share the earth ..We have
helped some species spread throughout the world while driving others from Ou
much of their former habitat. Some plants and animals owe their existence to bn
an
human activity. Sadly, many more species owe their extinction to us. Since our
existence depends on our relationships with the other species ,of the earth, our dis
future will ultimately be determined by how we treat them. Biogeography helps be

'gator mississipiensis and
liy in North America and
nd in warm subtropical
old and freezing tempera-
lere joined together in one
1refore the geographic range
is today. Over time, as conti-
m from crocodiles and
~d and small.
1any of the plants found in
phic differences in vegeta-
phy can tell us why North
I only in warm, moist areas
1lso help us to understand
~nsis), and not in tropical
1y, you will come to under-
1, and historical factors led
ng biogeography, you can
appreciation of the won-
ter every single day.
1reciate the development
tr apart from the natural
gically and culturally, is a
nt, and interactions with
round the world is one of
ms play an increasingly
have a significant impact
hare the earth ..We have
hile driving others from
Is owe their existence to
'tinction to us. Since our
species ·of the earth, our
1em. Biogeography helps
AN INTRODUCTION 3
us understand where living organisms, including humans, have come from and
where we and earth's other inhabitants are potentially headed.
The ability of humans to alter the environment and affect the survival of
plants and animals bestows a great deal of responsibility upon us. We are respon-
sible for making sure our actions cause as little damage as possible to the natural
environment. In this effort we will always face tradeoffs between the needs of
people for food, resources, living space, and recreational areas, and the preserva-
tion and conservation of nature. In some cases, we can alter our activities to
lessen the damage we cause. In other instances, we might try to restore damaged
areas to a semblance of their natural state. In these enterprises, biogeography can
provide important guidance. By examining the long-term histories of plant and
animal communities, biogeographers provide information on how humans have
altered the environment. In studying the natural distributions of plants and ani-
mals, biogeographers can help preserve nature. Biogeography provides impor-
tant guidance on how we are affecting the environment and what actions we can
take to conserve resources and preserve natural environments.
Let us now return to definirlg biogeography. In studying the present and
past distributions of life, biogeographers have two basic tasks: description and
explanation. Describing where a plant or animal species occurs today is the easi-
est part, and yet this task remains incomplete for vast numbers of plants and ani-
mals. So far, a little less than 2 million living species of plants, animals, and other
organisms have been identified. It is thought that an additional 4 to over 20 mil-
lion remain undiscovered by modern science. It is even more difficult to recon-
struct the past distributions of organisms because we must rely on fossil records
that are often incomplete and difficult to interpret. Explaining exactly how pres-
ent and past distributions are controlled by the complex geographic, environ-
mental, and historical factors that affect all organisms presents the greatest
challenge. In order to understand how the physical and biological environment
controls the distribution of plants and animals today, biogeographers must be
familiar with concepts and techniques in physiology, anatomy, ecology, pedology
(the study of soils), climatology, limnology (the study of lakes), and oceanogra-
phy. In order to study how plants and animals were distributed in the past, bio-
geographers must know some basic geology, paleontology, and evolutionary
biology. Since biogeography requires knowledge from a wide variety of other dis-
ciplines, it is called a synthetic science. The interests of the biogeographer blend
and overlap with the interests of the ecologist, the evolutionary biologist, and the
paleontologist to such an extent as to make the identification of firm boundaries
between biogeography and these disciplines difficult. Indeed, it might be argued
that from a life sciences perspective, all of these disciplines are subfields of bio-
geography. Given the synthetic nature of the field, it is not surprising that biogeo-
graphers can be found working in many different university departments,
including geography, biology, geology, paleontology, and anthropology. Biogeog-
raphers can also be found working in park services, forestry services, environ-
mental services, conservation groups, and private consulting firms.
Clearly, biogeography encompasses a huge area of the natural sciences.
One way in which biogeographers tackle the complexity of their subject is to
break it down into different subdisciplines. Phytogeographers study the present
and past distributions of plants. Zoogeographers examine the present and past
distributions of animals. The biogeographic study of the modern relationships
between organisms and the environment is called ecological biogeography.
4 CHAPTER 1 AN INTRODUCTION

Some biogeographers concentrate on studying past distributions and the evolu- bu

tion of life. This line of inquiry is called historical biogeography. The field of ana- USI
lytical biogeography is concerned with developing general rules that explain te1
how geography affects the evolution and distribution of plants and animals and di1
how past distributions and evolutionary history are reflected in modern distribu- the
tions. The application of the lessons learned from ecological, historical, and ana- ap
lytical biogeography, including the protection or restoration of the natural
environment, is called conservation biogeography. va
So, what are the aims of this textbook, and how can it help you understand r01
this fascinating and diverse field? The book is written to provide you with a solid Sa
introduction to the field of biogeography. It does not assume extensive back- liv
ground knowledge of biology, geography, or geology. There are, however, funda- we
mental terms and concepts from these three fields that all biogeographers must r01
know, and so they are outlined in this book. In Chapter 2 you will be introduced to ce1
some important basics of biology and physical geography including climatology, bi<
oceanography, and limnology. Later, in Chapter 6, we will explore some important oq
concepts from geology regarding the history of the earth. Throughout the text, yo
you will also be introduced to widely applied concepts from ecology and evolu- wh
tionary biology. These concepts will be explained in detail, and their importance in
to biogeography will be highlighted. enc
In recognition of the ecological, historical, and analytical facets of biogeog-
raphy, the book is divided into three major sections. The first section, called Space
and Life, is concerned with ecological biogeography. In these chapters we will
examine how present physical and biological conditions affect organisms and their KEYWOR
distributions. Chapter 3 describes how physical conditions affect organisms and
control their geographic distributions. In Chapter 4 we will look at the interactions Analytical biog
that occur between different organisms and discuss how these biological factors 3 geography
can control geographic distributions. In Chapter 5 we will examine how distur- ::-onservation b
bances such as fires or floods impact ecosystems and influence the geographic dis-
tribution of organisms. The final chapter of this section, Chapter 6, discusses how
plants and animals often occur together in communities, and how the earth can be
subdivided into biogeographic zones based on vegetation communities.
The second section of the book concerns historical biogeography and is
entitled Time and Life. In Chapter 7 we will learn a little basic geology and dis-
cover how the physical geography of the earth has changed over the last 500 mil-
lion years. We will also explore how the earth's climate has undergone periodic
changes. We will pay special attention to the shifts between glacial and nonglacial
conditions that have occurred over the past 2 million years. We will conclude the
chapter with a consideration of how human activity is now changing the climate
of the planet. Chapter 8 will delve into how organisms change their distributions
in response to geographic and environmental conditions. In Chapter 9 we will
explore how geography impacts processes and patterns of evolution and extinc-
tion. In Chapter 10, we will consider how the earth can be divided into different
biogeographic regions based on the species that occur together and the evolu-
tionary history of plants and animals. We will focus on biogeography and the
evolution of humans in Chapter 11 and examine our impact on other species in
Chapter 12.
The final section of the book is called Biogeography: Theory and Practice.
In these chapters, we examine the analytical and conservation aspect of biogeog-
raphy. In Chapter 13, we will focus on some different common geographic distri-
 KEY WORDS AND TERMS 5

listributions and the evolu- butions and their causes. We will explore how biogeographical patterns can be
~eography. The field of ana- used to trace the evolutionary history of modern plant and animal species. Chap-
general rules that explain ter 14 describes biogeographic theories that explain global variations in the
i of plants and animals and diversity of plant and animal species on continents, in oceans, and on islands. In
:fleeted in modern distribu- the concluding chapter of the book, we will explore how biogeography is being
•logical, historical, and ana- applied to problems of conservation.
·estoration of the natural One of the greatest attractions of biogeography comes from the wonderful
variety of life and the fantastic adaptations that allow organisms to live in envi-
can it help you understand ronments ranging from the coldest depths of the ocean to the hot sands of the
to provide you with a solid Sahara Desert. Biogeography is a science concerned with life, and it should be
ot assume extensive back- lively! In this book you will be introduced to many examples of fantastic and
There are, however, funda- wonderful plants and animals. These organisms, and the many interesting envi-
at all biogeographers must ronments and communities in which they live, will be used to illustrate the con-
2 you will be introduced to cepts and theories of biogeography. We will also examine many examples of how
tphy including climatology, biogeography has impacted people and how human activity has affected other
rill explore some important organisms. These examples should make your studies more enjoyable and allow
arth. Throughout the text, you to translate your knowledge of biogeography to the real world. In the end,
s from ecology and evolu- when you travel across the world, walk down your own street, or look at yourself
etail, and their importance in the mirror, you will realize that every single plant, animal, and person you
encounter is a result of, and contributor to, the biogeographical story of life.
1alytical facets of biogeog-
e first section, called Space
In these chapters we will
: affect organisms and their KEY WORDS AND TERMS
:ions affect organisms and
¥ill look at the interactions Analytical biogeography Ecological biogeography Phytogeography
>w these biological factors Biogeography Historical biogeography Zoogeography
will examine how distur- Conservation biogeography
luence the geographic dis-
' Chapter 6, discusses how
, and how the earth can be
m communities.
"ical biogeography and is
tie basic geology and dis-
tged over the last 500 mil-
e has undergone periodic
een glacial and nonglacial
~ars. We will conclude the
1ow changing the climate
;hange their distributions
ms. In Chapter 9 we will
: of evolution and extinc-
be divided into different
together and the evolu-
n biogeography and the
1pact on other species in

1hy: Theory and Practice.

mtion aspect of biogeog-
mmon geographic distri-
PART I
SPACE AND LIFE
CHAPTER 2
SOME BASICS

Biogeography is a very broad science, and biogeographers are a highly diverse

group of scientists. This multidisciplinary scope is one of the most satisfying
aspects of biogeography-and one of the most challenging. Biology, geography,
and geology all contribute concepts, specialized vocabularies, and ways of classi-
fying information that are important to biogeography. In order to understand and
appreciate the science of biogeography, we must therefore have some knowledge
of its foundations in these other sciences. Before we get down to the business of
biogeography, we will review some key concepts from these associated disci-
plines. In this chapter, we will concentrate on aspects of biology and physical
geography that are fundamental to biogeography. Later in this book we will dis-
cuss some concepts from geology that are also important to biogeography. Much
of the material that we cover here will be familiar to anyone who has taken intro-
ductory courses in biology and physical geography. However, even for those with
such a background, a little review won't hurt.

BIOLOGY AND THE HIERARCHIES OF LIFE

Biology can be defined as the science of life and all of its phenomena. Life on
earth includes millions of different types of organisms, ranging from viruses to
whales. The study of any of these organisms could include examination of myriad
phenomena, from biochemical reactions to social behavior. It is clear that biolo-
gists face a daunting challenge! The science of biology tackles this immense task
by breaking it down into smaller components that can be studied individually.
One way to categorize and organize the constituent units of a large entity is to
develop a hierarchy. A hierarchy is a system of organization in which components
are ordered by rank. Most corporations are good examples of hierarchies. A large
group of office staff is under the direction of one office manager, who, along with
several other managers, is under the direction of the vice president of operations.
The vice president, along with two or three other vice presidents, reports to the
president of the firm. The president is at the top of the hierarchy, and the office
staff is its base. Every biogeographer should be familiar with three important bio-
logical hierarchies: the taxonomic hierarchy, the ecological hierarchy, and the
trophic hierarchy. We will examine each of these.

Taxonomic Hierarchy
Taxonomy is the subdiscipline of biology concerned with the classification and
naming of organisms. Taxonomy is also known as systematics when the main goal
is to determine the evolutionary relationship between groups of organisms. In
this case, taxonomists are also called systematisists. The evolutionary histories of
9
10 CHAPTER 2 SOME BASICS

organisms that are reconstructed by sytematisists are referred to as phylogenies. clost

Taxonomists use observable traits, referred to as characters or character states to encc
group similar individual organisms together and to separate groups of different ever
organisms. The groups that taxonomists develop are called taxa (the singular Araz
form is taxon). The characters that taxonomists use to classify organisms usually diffe
start with physical differences, such as color variations in flowers or in the nam
plumage of different birds. Taxonomists may also consider differences that are can;
apparent through chemical analysis of the tissue or fluids of the organism. Such
studies are referred to as chemotaxonomy. Finally, cytotaxonomists examine the that
chromosome structure of organisms to detect genetic similarities and differences siste
in order to classify organisms. thes
The classification and naming of organisms by individual taxonomists ent 1
would not be very useful if everyone had their own system and terminology. For- ica '
tunately, there is a single system of taxonomic classification that is generally the 1
accepted by all biologists and biogeographers. The development of our present Pim
taxonomic system is at the foundation of biology and biogeography. It is a history cone
that can be traced back to ancient Greece. The roots of the modern taxonomic betv
system began with Aristotle (384-322 B.C.). Aristotle, a student of Plato, is best mos
known as a philosopher but was also active in biology, physics, astronomy, and dles
psychology. He developed an early scientific system for classifying animals into The
groups that shared similar features. Aristotle believed that the specific form and
behavior of individual plants and animals were inherited and immutable. He con-
sidered that all individuals belonged to groups, or species (from the Greek word
"eidos"), of taxonomically similar individuals. Aristotle believed that the form
and behavior of these species did not change from generation to generation. He
taught that dogs form one species and cats another. Aristotle also argued that
plant and animal species formed a hierarchy ranging from simple organisms, such
as worms, to the most complex organism, which he considered to be humans. In
the Middle Ages, European scientists, influenced by Aristotelian logic, grouped
organisms that appeared to be generally, but not exactly, similar to each other
into taxonomic units called genera (the singular form is genus). A genus would
include, for example, the Scots pine trees of Scandinavia and the Mediterranean
pines of southern Europe. Both the Scots pines and the Mediterranean pines
belong to the genus called Pinus.
Before we proceed, let's examine the words "genus" and "species" and con-
sider the continuing role of Latin in biology and biogeography. Genus is the
Latinized form of the Greek word "genos." In the Middle Ages the language of
scholarship in Europe was Latin, and the names of the genera came from that
language or are latinized versions of words from other languages. For example, in
Latin the genus for pines is called Pinus. In comparison, the genus for cats is Felis.
Although Latin is no longer widely spoken or understood, the formal names of
organisms are still written in that language. The benefit of this convention is that
no matter which language the scientist is working in, the names of the organisms
are always presented in Latin using the Latin alphabet. Even papers and books
written in Russian Cyrillic or Chinese characters will always present the scientific a
names of organisms in the Latin alphabet. Biologists and biogeographers always
FIG
know which organisms are being discussed, even if they can read nothing else in Poe
the document. The use of the proper scientific names for organisms is also impor- spe
tant for avoiding confusion among scientists who speak the same language. Take bot
pines, for example. In North America we have many different trees that are oft
 BIOLOGY AND THE HIERARCHIES OF LIFE 11

eferred to as phylogenies.
:ters or character states to
Jarate groups of different
called taxa (the singular
classify organisms usually
ems in flowers or in the
sider differences that are
ids of the organism. Such
>taxonomists examine the
milarities and differences
if individual taxonomists
:em and terminology. For-
lication that is generally
telopment of our present
ogeography. It is a history
)f the modern taxonomic
1 student of Plato, is best
1, physics, astronomy, and
or classifying animals into
that the specific form and
:i and immutable. He con-
ies (from the Greek word
e believed that the form
eration to generation. He
Uistotle also argued that
om simple organisms, such
tsidered to be humans. In
ristotelian logic, grouped
:tly, similar to each other
is genus). A genus would
_a and the Mediterranean
the Mediterranean pines
closely related and belong to the genus Pinus. In the South Pacific, you might
encounter a tree that is commonly called the Norfolk Island pine. This plant, how-
ever, is unrelated to our North American pines and belongs instead to the genus
Araucaria (Fig. 2.1). In many instances the same plant or animal will have several
different common names, but every organism has only one accepted scientific
name. In addition, the Latin names of organisms often contain descriptions that
can be understood by people with only a limited knowledge of Latin.
So, what is the difference between a species and a genus? It is recognized
that genera of plants and animals contain organisms that are related but are con-
sistently distinguishable on the basis of their morphology. In addition, many of
these taxa, though members of the same genus, cannot interbreed. These differ-
ent members of a genus are species. For example, people in eastern North Amer-
ica will be familiar with the eastern white pine, and those in the western part of
the continent might know the lodgepole pine. Both species belong to the genus
Pinus and share certain broad similarities, such as possessing long thin needles,
cones, and upright growth (Fig. 2.2). There are, however, clear differences
between the trees. White pines can grow to well over 30m in height, whereas
most lodgepole pines only achieve heights of 20m. The white pine carries its nee-
dles in bundles of five. In contrast, the lodgepole pine has bundles of two needles.
The white pine has large cones that open immediately upon ripening and release

_s" and "species" and con-

•geography. Genus is the
dle Ages the language of
e genera came from that
anguages. For example, in
the genus for cats is Felis.
;)Od, the formal names of
of this convention is that
e names of the organisms
.. Even papers and books
JVays present the scientific a b
td biogeographers always
FIGURE 2.1 White pine trees of the genus and species Pinus strobus growing in the
r can read nothing else in
Pocono Mountains of Pennsylvania (a) and so-called Norfolk Island pines of the genus and
organisms is also impor- species Araucaria heterophylla growing on Norfolk Island in the South Pacific (b). Although
the same language. Take both species are called pines, they are unrelated, and only white pine is actually a member
different trees that are of the pine genus.
12 CHAPTER 2 SOME BASICS

mon
chan

orga
ral e<
Eastern Alth
white pine defir
beha

~
duci1
COmJ

Lodgepole
pine desc
grea
FIGURE 2.2 The needles, cones, and shapes of a mature eastern white pine (Pinus era~

strobus) and a western lodgepole pine (Pinus contorta). Notice that both pine species share spec
a general resemblance but possess clear differences in terms of their needles, cones, and won
mature form. isms
Swe1
knm
their seeds. Lodgepole pines have small cones, many of which remain closed and ited
retain their seeds until a fire causes the cone scales to open. In addition, eastern spec
white pine and lodgepole pine cannot interbreed.The two trees are recognized as naet
different species within the genus Pinus. Following the concepts of Aristotle, the by a
English scholar John Ray (1627-1705) attempted to systematically define a exar
species. He concluded that if seeds came from the same plant, the seedlings must Pim.
be related and similar in character. He recognized that variations might occur tiate
between seedlings from the same parent, but he considered these to be "acci- nam
dents." Ray's conception of species can be paraphrased as "Like begets like." also
Since the time of Ray, species have formed the basic unit of the modern tax- nam
onomic system, but there remains much debate among scientists as to just how a nam
species should be defined. Three main definitions are in use today. The phyloge- kno
netic species concept identifies a species as a group of sexually reproducing organ- the
isms that share at least one diagnostic character that is present in all members of
the species but absent in other organisms. Unique behavioral traits are accepted spe<
as defining characters. This concept does not adequately take into account natu- catf
ral variations within species, such as differences in human hair color. If this con- the
cept were widely applied by biologists, far more species would be identified than reff
is now the case. In addition, the phylogenetic species concept does not consider ital
reproductive interactions between members of the species or their evolutionary stru
history. The biological species concept defines a species as a group of organisms imi
that can interbreed freely under natural conditions. This definition was articu- mo
lated by the great evolutionary biologist Ernst Mayr in 1942, although its roots nar
extend into the early part of the twentieth century. It remains the most widely ent
used definition of a species in biology. Finally, paleontologists who study the fos- spe
sil record often use the evolutionary species concept. Evolutionary species are
organisms that have a direct ancestor-descendant relationship that is traceable in
the fossil record. Usually, such a relationship is inferred from morphological sim-
ilarities such as the size or shape of the fossils. This is because morphological fea-
tures are generally all that is available from the fossil record. New species are
differentiated when there are clear divisions of one evolutionary line into two or

,,
Eastern
white pine
\I;I'
~
Ji
- Lodgepole
pine
!astern white pine (Pinus
~e that both pine species share
1 of their needles, cones, and
of which remain closed and
J open. In addition, eastern
two trees are recognized as
e concepts of Aristotle, the
to systematically define a
1e plant, the seedlings must
hat variations might occur
1sidered these to be "acci-
d as "Like begets like."
1sic unit of the modern tax-
g scientists as to just how a
in use today. The phyloge-
~xually reproducing organ-
; present in all members of
tavioral traits are accepted
!ly take into account natu-
man hair color. If this con-
:s would be identified than
concept does not consider
~cies or their evolutionary
:s as a group of organisms
rhis definition was articu-
in 1942, although its roots
: remains the most widely
)logists who study the fos-
Evolutionary species are
ionship that is traceable in
l from morphological sim-
~cause morphological fea-
1 record. New species are
)lutionary line into two or
BIOLOGY AND THE HIERARCHIES OF LIFE 13
more new lineages. The evolutionary species concept allows for morphological
changes in the species over time as evolution occurs.
In practice, biogeographers combine these concepts and view species as
organisms that are morphologically similar and can interbreed freely under natu-
ral conditions. It is assumed that such organisms must share a common ancestor.
Although this sounds pretty straightforward, we still encounter problems in
defining species due to natural variability in the morphology and reproductive
behavior. In addition, species are generally thought of in terms of sexually repro-
ducing organisms. Organisms that reproduce asexually raise difficulties when the
common phylogenetic and biological species definitions are used.
Until the eighteenth century, scientists named species simply by adding some
descriptive words to the name of the genus. The eighteenth century was a period of
great geographical and biological exploration and discovery when many new gen-
era and species were found and described. To differentiate these newly discovered
species, their names often became polynomials containing a dozen or more Latin
words. Clearly, this system was unwieldy. The simplified system of naming organ-
isms that we use today was invented in the mid-eighteenth century by the great
Swedish scientist Carolus Linnaeus (1707-1778). He attempted to catalog all of the
known species of the world. Linnaeus was trained as a medical doctor but inher-
ited a keen interest in plants and natural history from his father. Many of the
species he identified and described are still accepted by taxonomists today. Lin-
naeus introduced the binomial system whereby every organism could be identified
by a unique combination of a generic (genus) and specific (species) name. For
example, white pine is known as Pinus strobus, while lodgepole pine is known as
Pinus contorta. These related species share the same genus, Pinus, but are differen-
tiated by their species names, strobus and contorta. Species with the same generic
names are assumed to be related. With the binomial system, different species can
also share the same specific name. This occurs frequently because the specific
names often describe some feature of the organism. As an example, the scientific
name for white oak is Quercus alba, whereas the scientific name for a seabird
known as the snowy sheathbill is Chionis alba. In this case, the word alba refers to
the light coloring of the organisms and in no way implies a genetic relationship.
Because different, unrelated species can share the same specific name, a
species is never referred to by its specific name alone- the genus must be indi-
cated. If it is clear which genus is being discussed, you can use the first initial of
the generic name instead of spelling the complete name (i.e., Quercus alba can be
referred to as Q. alba). By convention, the generic and specific names are always
italicized, and the generic name starts with a capital letter. The specific name
starts with a lower-case letter. In this book you will find the scientific names of
important genera and species in parentheses following the first usage of the com-
mon name. Sometimes the scientific name of a species is followed by the full
name, abbreviated name, or initials of the taxonomist who first described it. A sci-
entific name followed by the letter L, such as Quercus alba L., means that the
species was first described by Linnaeus himself.
So what does the modern taxonomic hierarchy consist of? As we have seen,
species form the lowest level of the hierarchy, although some species may be fur-
ther subdivided into distinctive subspecies and races. Collections of species that
are similar to one another, and presumably related, are grouped together in the
next level, the genera. Genera that are morphologically similar and likely possess
evolutionary linkages are grouped together as families. Oaks, for example,
14 CHAPTER 2 SOME BASICS

belong to the family of trees called Fagaceae. This family also includes the genus ider
beech (Fagus) and several other genera of trees. How families and higher taxo- ass:
nomic groups are classified remains a subject of revision and debate. According
to one classic system, families that are related are grouped together in taxonomic Ecc
orders. The order for oaks is Fagales. In turn, orders are grouped into classes. All
flowering plants (sometimes called angiosperms or the Magnoliophyta) are The
grouped into one of two classes. The oaks are placed in the class Dicotyledons mor
(also called Magnoliopsida). Among other traits, plants in this class have seeds man
that produce two initial leaves, or cotyledons, when they germinate. Many of the fort
familiar trees of the deciduous forest, such as maples (Acer), birch (Betula), elm isms
(Ulmus), willows (Salix), and garden flowers such as roses (Rosa) and daisies wou
(Aster) are dicotyledons. The other class for flowering plants is Monocotyledons orga
(also called Liliopsida). Some examples of monocotyledons include orchids biog
(Orchidaceae) and grasses (Poaceae). When the seeds of these plants germinate, The~
they produce one cotyledon. men
Classes are grouped together into phyla (the singular is phylum and in
plants the term division is often used). The phylum for the dicotyledons and vidu
monocotyledons is Anthophyta. The Anthophyta are also referred to as the flow- arcti
ering plants. Conifers (cone-bearing plants) such as the pines, the spruces (Picea), migl
and the firs (Abies) are neither dicotyledons nor monocotyledons. They do not indi'
belong to the Anthophyta but are instead members of the division Coniferophyta popt
and are also known as gymnosperms. Finally, at the highest level, the phyla are inter
grouped into five kingdoms or sometimes a lesser number of domains. Debate faun
continues on how to classify organisms at many levels, including this highest and
level. According to one system, the Kingdoms include Monera, Protista, Fungi, may
Plantae, and Animalia. The Monera are simple prokaryotic organisms such as are r
bacteria. The Protista are single-celled eukaryotic organisms such as amoeba. reco:
You can probably guess who the members are of the Fungi, Plantae, and Ani- that
malia kingdoms. Thus, we can trace the affinity of every organism, including quer
humans (Table 2.1), up through the taxonomic hierarchy to a kingdom. However, travt
the exact number of kingdoms continues to be debated. are<
The taxonomic hierarchy provides a convenient and accepted way to clas- freq1
sify life. It should be remembered, however, that this system is not static. New arcti
species are continuously being discovered and added. At times, new genera and latio
families are added. Even the status of different kingdoms is debated. In addition, agri<
taxonomists may change the genus or family of older recognized species when
new information comes to light. It sometimes happens that several taxonomists all tl
ecoi
TABLE 2.1 A Systematics (Taxonomic Hierarchy) of orga
Eastern White Pine (Pinus strobus) and focu
Humans (Homo sapiens sapiens) focu
colo
White Pines Humans su~

Species Pinus strobus Homo sapiens sapiens

stud
Genus Pinus Homo flo"
Family Pinaceae Homonidae
the
ec(Y.
Order Coniferales Primates
refe
Class Coniferopsida Mammalia
guil
Division or Phylum Coniferophyta Chordata
forr
Kingdom Plantae Animalia
guil

nily also includes the genus
w families and higher taxa-
sian and debate. According
uped together in taxonomic
l.re grouped into classes. All
1r the Magnoliophyta) are
d in the class Dicotyledons
mts in this class have seeds
hey germinate. Many of the
; (Acer), birch (Betula), elm
as roses (Rosa) and daisies
tg plants is Monocotyledons
cotyledons include orchids
is of these plants germinate,
, singular is phylum and in
m for the dicotyledons and
· also referred to as the flow-
h.e pines, the spruces (Picea),
onocotyledons. They do not
rf the division Coniferophyta
: highest level, the phyla are
aumber of domains. Debate
evels, including this highest
tde Monera, Protista, Fungi,
)karyotic organisms such as
organisms such as amoeba.
he Fungi, Plantae, and Ani-
f every organism, including
rchy to a kingdom. However,
ted.
nt and accepted way to clas-
llis system is not static. New
:d. At times, new genera and
dams is debated. In addition,
ier recognized species when
ens that several taxonomists
~ Hierarchy) of
strobus} and
rpiens}
Humans
-lama sapiens sapiens
-lama
~omonidae
'rimates
-.1ammalia
:hordata
~nimalia
BIOLOGY AND THE HIERARCHIES OF LIFE 15
identify the same species and give it different names. These names are referred to
as synonyms, and the earliest published name generally takes precedence.
Ecological Hierarchy
The taxonomic hierarchy provides for a division and ranking of life based on the
morphology and the evolutionary relationship of organisms. It is the foundation for
many aspects of biological research. It does not, however, provide much guidance
for the ecologist or biogeographer who goes out into the field to study how organ-
isms are affected by the environment or how they interact with other species. It
would be an intractable task to set out to study the relationship between every
organism with the environment and all other organisms. Therefore, ecologists and
biogeographers often concentrate on specific spatial and taxonomic scales of study.
These levels of ecological study can be organized into a hierarchy that reflects the
increasing geographic and taxonomic scale being examined.
The lowest scale of study that is of interest to the biogeographer is the indi-
vidual organism. For example, scientists might put a radio tracking collar on one
arctic fox (Alopex lagopus) and monitor its movements. At the next level, they
might consider studying a population of arctic foxes. A population is defined as all
individuals of a given species in a prescribed area. Usually, members of the same
population are assumed to be in close enough proximity to be able to interact and
interbreed frequently. Of course, in many cases, members of the same species are
found in different locations and do not interact on a regular basis. Interbreeding
and other interactions between these separated populations of the same species
may only occur relatively frequently or extremely infrequently. Such populations
are referred to as metapopulations. Two broad types of metapopulations can be
recognized. Loose metapopulations consist of subpopulations of the same species
that live in different locations and interact with other subpopulations very infre-
quently. The distance between subpopulations is farther than most individuals
travel during their life spans. Tight metapopulations consist of subpopulations that
are close enough for individuals to travel between them and thus interact more
frequently. The arctic fox populations living on different islands in the Canadian
arctic separated by large expanses of water are an example of a loose metapopu-
lation. Birds that live and nest in different woodlots that are separated only by
agricultural fields are an example of a tight metapopulation.
We might also consider examining the interaction of our fox population with
all the other species of organisms in its environment. In this case, we are studying an
ecological community. A community can be broadly defined as all populations of
organisms that live and interact within a prescribed area. Ecological research that
focuses on one species is sometimes referred to as autecology, while research that
focuses on the interactions between species in communities is referred to as syne-
cology. Some ecologists and biogeographers limit their studies of communities to
subsets of the total community. For example, phytogeographers might restrict their
study to flowering plant species only. Some biogeographers would refer to this as the
flowering plant community. However, other ecologists and biogeographers feel that
the term community should only be used to refer to all species of organisms in an
ecosystem. Subsets of a community, such as the flowering plant species, are often
referred to as assemblages. One important subset of the complete community is the
guild. Guilds are groups of animal species within a community that have similar
forms, habitat, and resource requirements. The insectivorous bird species form one
guild in a community, while the seed-eating bird species form another.
16 CHAPTER 2 SOME BASICS

If we were to consider the relationship between the species of our commu-

nity and the physical factors of the environment, particularly when we examine
flows of energy and matter through this biophysical system, we would be con-
ducting research at the scale of the ecosystem. It could be argued that the spatial
IIi
boundaries of any ecosystem extend over the whole earth. Even the smallest area
is linked to the world at large by physical processes such as rainfall and biological
processes such as the input of fine airborne organisms. In practice, the boundaries
of ecosystems are often defined by the researcher and can vary from very small
areas such as the trunk of a decaying tree to large areas such as the tundra of
Baffin Island in arctic Canada. Very large areas of the earth's surface that have a
similar climate and vegetation are referred to as biomes. The biomes are an
important area of research in biogeography, and we will discuss them in detail
later. Finally, all of life on the planet is collectively referred to as the biosphere-
the highest and broadest level of ecological research. The other realms of the
earth are physical ones of the atmosphere, hydrosphere, and lithosphere. The
atmosphere includes all the components of the air; the hydrosphere includes all
the water in the oceans, lakes, streams, and ground; and the lithosphere is the
solid earth of rock and sediment. Although the research of biogeographers may
focus on smaller levels such as ecosystems, the end goal of biogeography is to
combine the results from all of these more specific studies and draw general con-
clusions about how the biosphere functions today, how it developed, and where it
might be headed in the future. Remember, our future and that of the biosphere
are really one and the same.

Trophic Hierarchy c
A third way of ordering the biosphere is to examine the flow of energy through Ir
ecosystems. The various levels through which energy flows from its initial capture 0<

by the biosphere until its dissipation as waste heat are called trophic levels. The ti
biosphere functions through the acquisition of energy by organisms and the flow A
21
of energy from one organism to another. With very few exceptions, all life is
cJ
dependent on the sun to provide the energy that is consumed. Notable excep-
a
tions are the amazing ecosystems that have developed around undersea volcanic v
vents (Technical Box 2.1). v
Most of the solar energy that reaches the earth is visible light, which is a p
form of short-wave radiation. The electromagnetic waves that make up visible r
light range in wavelength from -0.4 to 0.6 microns. Of this incoming energy, f
approximately 32% is reflected back into space by the earth's atmosphere. a
Roughly 18% is absorbed by the earth's atmosphere. The surface of the earth c
absorbs about 50% of the incoming radiation, and about 20% of this is used in r
evaporation. Visible light energy from the sun is absorbed by the atmosphere and t

the earth's surface and then radiated as long-wave (3- 25 micron wavelength)
thermal energy. This long-wave radiation is what we sense as heat. It is clear that
almost 100% of the incoming solar radiation is reflected back into space or used
in heating and evaporation. What then powers the biosphere? The entire bios-
phere, including the human race, is supported by a tiny fraction of the incoming
solar radiation. The amount of solar energy captured for direct use by the bios-
phere is only 0.1- 0.3% of the total input!
The solar energy used to power the biosphere is captured through the
process of photosynthesis. For most plants, the radiation used for photosynthesis
 BIOLOGY AND THE HIERARCHIES OF LIFE 17

the species of our commu-

ticularly when we examine
system, we would be con- TECHNICAL BOX 2.1
d be argued that the spatial
arth. Even the smallest area
.ch as rainfall and biological
. In practice, the boundaries
td can vary from very small
ueas such as the tundra of
! earth's surface that have a
tiomes. The biomes are an
will discuss them in detail
'erred to as the biosphere-
:h. The other realms of the
there, and lithosphere. The
he hydrosphere includes all
and the lithosphere is the
trch of biogeographers may
goal of biogeography is to
1dies and draw general con-
wit developed, and where it Deep-sea crabs and other marine life near a "black smoker" hydrothermal vent
e and that of the biosphere in the deep ocean. Illumination for the picture is provided by artificial lighting
from a deep-sea submersible vehicle.

Chemosynthesis and the Ecosystems of Oceanic Hydrothermal Vents

the flow of energy through
!lows from its initial capture
re called trophic levels. The
y by organisms and the flow
y few exceptions, all life is
consumed. Notable excep-
d around undersea volcanic
h is visible light, which is a
waves that make up visible
s. Of this incoming energy,
>y the earth's atmosphere.
·e. The surface of the earth
.bout 20% of this is used in
rbed by the atmosphere and
(3-25 micron wavelength)
sense as heat. It is clear that
:ted back into space or used
biosphere? The entire bios-
.ny fraction of the incoming
j for direct use by the bios-
In 1977 scientists discovered that hydrothermal vents located 2500 m below the
ocean surface near the Galapagos Islands supported unexpectedly dense concentra-
tions of organisms. Similar vents have been discovered in both the Pacific and
Atlantic oceans. The biomass of these vents is astounding and can range as high as
2(}-.30 kg/m2. The vent organisms include giant red worms (Riftia pachyptila), large
clams (Calyptogena magnifica), and mussels (Bathymodiolus thermophilus). In
addition, various species of crabs, shrimps, and sea anemones are found at some
vents. In all, some 300 new species of animals have been discovered near undersea
vents. How can such fauna be supported so far from the upper ocean waters where
photosynthesis occurs? These vents emit mineral-laden waters at temperatures
ranging up to 450° C. When mixed with the cold ocean waters, this produces tem-
peratures in the vent area of 8° to 23° C. Hydrogen sulfide (H2S) is particularly
abundant in these plumes. Studies have revealed that these vents support a food
chain based entirely on geothermal energy. At the base of the food chain are bacte-
ria that oxidize the sulfur from the vents to form carbohydrates. In contrast to pho-
tosynthesis, this process is called chemosynthesis and proceeds as follows:
C02 + HzS + 02 + HzO ~ CHzO + H2S04
The C02 and 02 in the process comes from the sea water. The bacteria are then
eaten by primary consumers such as limpets, mussels, and clams. It is suspected that
some other types of bacteria may utilize methane or ammonia from the vents in
energy synthesis.

re is captured through the

:ion used for photosynthesis
18 CHAPTER 2 SOME BASICS

falls in the red through blue visible light ( -0.4-0.6 micron wavelength) portion of
the electromagnetic spectrum. This light is referred to as photosynthetically
active radiation, or PhAR. Most of the solar energy that the earth receives is
PhAR, so it is no coincidence that plants evolved to use this portion of the elec-
tromagnetic spectrum in photosynthesis. It is also not surprising that our eyes are
adapted to detecting radiation in these wavelengths. We can see visible light spec-
trum energy but not ultraviolet or infrared energy.
During photosynthesis, atmospheric carbon and water vapor are trans-
formed into sugar, water, and oxygen. The energy for photosynthesis comes from
PhAR. The process takes place in chloroplasts, which are small, green bodies
within plant cells. The chloroplasts hold chlorophyll, which is the primary light
capturing the pigment of plants. The overall process can be summarized as:
Light
6COz + 12H20 --7 C6H1206 + 6H20 + 602
Carbon Water Sugar Water Oxygen
Dioxide

The C02 enters the leaves of plants through openings created by specialized sets
of cells called stomata. The stomata also allow the release of oxygen and water
vapor from the interior of the leaf.
There are three different biochemical pathways that green plants use in
photosynthesis. Most plants capture energy using the C 3 pathway described by
Melvin Calvin of the University of California at Berkeley. In C3 plants, the C02
from the atmosphere is converted into a 3-carbon molecule called 3-phospho-
glyceric acid. In the 1960s it was discovered that sugar cane converts C02 into
two 4-carbon molecules: malic and aspartic acid. This process became known as
the C 4 pathway. Finally, some plants, such as the prickly pear cactus (Opuntia),
use a modified form of photosynthesis called crassulacean acid metabolism
(CAM). In CAM plants, C02 is absorbed at night and stored as malic acid. Dur-
ing the light of day, photosynthesis is conducted by the C3 pathway. In general, C4
plants have the highest rates of photosynthesis, while CAM plants display the
lowest rates. Interestingly, however, all plants are relatively inefficient in terms of
energy fixation through photosynthesis. Only about 1% to 3% of the light hitting
a leaf is transformed into chemical energy in the form of simple carbohydrates FIG I
such as the sugars glucose and fructose. simr
La vi
Photosynthetic plants are the foundation of the trophic hierarchy (Figure 2.3).
and
Plants are referred to as primary producers because they fix the energy of the sun
into chemical energy used to power the biosphere. This term is slightly mislead-
ing, for plants cannot actually produce energy but merely transform it from one
state to another. Plants are also referred to as autotrophs or phototrophs because
of their ability to fix energy through photosynthesis rather than derive it from the con
consumption of other organisms. Organisms that eat plants to obtain energy are in a
called primary consumers. Species that eat the primary consumers are referred to ally
as secondary consumers, while species that eat secondary consumers are referred levc
to as tertiary consumers. Plant-eating species are also called herbivores, and eco
meat-eating species are called carnivores. Animals that eat both meat and veg- one
etable matter, such as most humans, are omnivores. When plants and animals die, ace•
decomposers consume them. Decomposition is the ultimate fate of all trophic con
levels. Herbivores, carnivores, omnivores, and decomposers are collectively attf
known as heterotrophs because they rely on other organisms to provide energy. wei
 BIOLOGY AND THE HIERARCHIES OF LIFE 19

cron wavelength) portion of

~d to as photosynthetically
;y that the earth receives is
use this portion of the elec-
; surprising that our eyes are
Ne can see visible light spec-

md water vapor are trans-

, photosynthesis comes from
ich are small, green bodies
I, which is the primary light
~an be summarized as:

Water Oxygen

:s created by specialized sets

·elease of oxygen and water

tys that green plants use in

le c3 pathway described by
·keley. In C3 plants, the C02
molecule called 3-phospho-
gar cane converts C02 into
[s process became known as
[ckly pear cactus (Opuntia),
tssulacean acid metabolism
td stored as malic acid. Dur-
te C3 pathway. In general, C4
ile CAM plants display the
1tively inefficient in terms of
l% to 3% of the light hitting
FIGURE 2.3 The complexity of energy flow within an ecosystem illustrated by a
rm of simple carbohydrates
simplified and small part of the food web for the northwestern Atlantic Ocean (after
Lavinge, 1992). Despite its complexity, a trophic hierarchy with planktonic plants at the base
rophic hierarchy (Figure 2.3). and sharks, mammals, and birds at the top is still apparent.
hey fix the energy of the sun
:Ois term is slightly mislead-
terely transform it from one
,phs or phototrophs because The trophic levels extending from primary producers to the highest level of
ather than derive it from the consumers are sometimes called food chains. However, the idea that energy flows
: plants to obtain energy are in a linear fashion up the trophic hierarchy is simplistic and misleading. Gener-
ry consumers are referred to ally, there is a skipping of levels and a back and forth exchange between different
dary consumers are referred levels of consumers and decomposers. As a result, the actual flow of energy in an
also called herbivores, and ecosystem is more like a food web than a linear chain (Fig. 2.3). Elton (1927) was
:hat eat both meat and veg- one of the earliest scientists to identify the importance of ordering ecosystems
Vhen plants and animals die, according to energy flows. He also recognized that the flow of energy was more
ultimate fate of all trophic complex than a simple linear chain. The work of Lindeman in 1942 was the first
composers are collectively attempt to formally conceptualize and quantify the flow of energy within food
rganisms to provide energy. webs. The study of food webs is the basis of modern systems ecology. It has been
20 CHAPTER 2 SOME BASICS

suggested that understanding food web structure is fundamental to understand- Ia

ing basic ecosystem structure, function, and response to disturbance and change. Sf
Relatively complex trophic hierarchies have been identified from both ter- 01
restrial and marine ecosystems. However, tracking and quantifying the flow of
energy in such systems are not easy. The three approaches commonly used to b)
investigate and define food webs are direct observation of feeding habits, inspec- fo
tion of the stomach contents of dissected organisms, and use of radioactive trac- ve
ers such as phosphorus-32 (32P), which are introduced through injection into the ity
vegetation and can be easily traced in tissue samples from animals. en
The average length of time during which energy captured through photo- en
synthesis is held by living plants in food webs varies from a few days in the open pi<
ocean to 3 years in grasslands and 22 to 25 years in some forests. Energy captured an
by photosynthesis can also be held in plant litter for significant periods of time tht
before being used by decomposers. In the tropics, the energy held in plant litter
may be released within three months, while it can be held for over 100 years in rar
the litter of temperate and northern forests. However, studies using radioactive vid
tracers show that, despite the potential long residence time of energy in plants wo
and litter, much of the energy captured by photosynthesis moves very quickly san
through the food web. In both terrestrial and freshwater ecosystems, some of the In
energy captured by photosynthesis is passed up to the highest trophic levels bel
within a matter of weeks. rna
Elton noted that trophic hierarchies appeared to be limited to only four or ma1
five levels. More recent examinations of the number of trophic levels in many
food webs ranging from invertebrate communities living in dung to birds in forest
stands to fish, birds, and mammals of the Antarctic pack-ice have found that most wht
ecosystems support only three to six trophic levels. Why is this? It has been dB
argued that since the transfer of energy between different trophic levels is rela-
tively inefficient, the energy available to higher trophic levels will rapidly B1'
decrease. All organisms use energy to function through the process of respiration, B2'
which is the oxidative reaction that breaks the high-energy bounds of carbohy- Ho·
drates to release energy for the organism's metabolism. Thus, respiration is the
pla1
reverse of the process of photosynthesis, with metabolic energy being produced. dB
Metabolic Energy gra
Energy + 6C02 + 12H20 ~ C6H1206 + 6H20 + 60z bio
Carbon Water Sugar Water Oxygen
squ
Dioxide ecc
era
On average, only about 10% of the energy of any trophic level is passed on to the lCS
next trophic level. This generalization is known as the 10% rule. As a result of the pr<
10% rule, each trophic level is expected to have about 90% less energy available me
to support it than the preceding trophic level. There is a high degree of variation 90
in the actual energy efficiencies of different organisms in food webs. Birds and OC(
mammals are the least efficient because they use energy to maintain constant tril
body heat. They assimilate only about 3% of the energy they receive in food.
Insects have an energy efficiency of around 39% , while fish have efficiencies of trc
around 10%. The decrease in available energy with each trophic level is bu
extremely important in ordering ecosystems. When you consider how little m<
energy is passed up through each trophic level, it is easy to understand why the ca
bulk of all living things on earth are plants (perhaps >90% by mass) and very ap

'undamental to understand-
to disturbance and change.
en identified from both ter-
md quantifying the flow of
roaches commonly used to
on of feeding habits, inspec-
and use of radioactive trac-
d through injection into the
from animals.
~y captured through photo-
from a few days in the open
me forests. Energy captured
~ significant periods of time
e energy held in plant litter
~ held for over 100 years in
~r, studies using radioactive
ce time of energy in plants
nthesis moves very quickly
tter ecosystems, some of the
• the highest trophic levels
o be limited to only four or
r of trophic levels in many
ing in dung to birds in forest
ck-ice have found that most
;. Why is this? It has been
ferent trophic levels is rela-
trophic levels will rapidly
h the process of respiration,
-energy bounds of carbohy-
ism. Thus, respiration is the
•lie energy being produced.
r Water Oxygen
>hie level is passed on to the
10% rule. As a result of the
.t 90% less energy available
s a high degree of variation
ns in food webs. Birds and
nergy to maintain constant
1ergy they receive in food.
1ile fish have efficiencies of
vith each trophic level is
n you consider how little
~asy to understand why the
s >90% by mass) and very
BIOLOGY AND THE HIERARCHIES OF LIFE 21
large carnivores are rare, occurring at much lower numbers than their prey
species. In theory, it takes a minimum average of 10 prey organisms to support
one bird or mammal predator of equal mass.
The total number of consumers that any ecosystem can support is limited
by the productivity of the plants. The productivity of vegetation is therefore a key
focus of study for understanding the structure of ecosystems. The productivity of
vegetation in fixing energy is known as primary productivity. Primary productiv-
ity can be viewed in two different ways. Gross primary productivity is the total
energy fixed in an ecosystem by photosynthesis. However, since some of this
energy is used through respiration to support the metabolic activities of the
plants, not all of the photosynthetic energy is available to produce plant matter
and be passed onto higher trophic levels. The gross primary productivity minus
the energy lost by respiration is referred to as net primary productivity. So, how is
net primary productivity measured? Many different techniques may be used,
ranging from the measurement of carbon fixation and respiration losses of indi-
vidual plants to satellite-based estimates of the photosynthetic activity of the
world's oceans. A relatively direct way is to harvest and weigh a representative
sample of the plant matter produced during a given time period in a given area.
In terrestrial environments, the plants can be harvested from both above and
below the ground to produce an estimate of aerial and root production. The plant
material is dried and weighed. The mass of living material is referred to as bio-
mass. The plant biomass is used to calculate primary productivity as follows:
dB=B1-B2
where
dB = the change in biomass between time period 1 (t1) and time period 2 (t2)
B1 =biomass at time t 1
B 2 = biomass at time tz
However, we must also account for losses in biomass due to death and decay of
plants (L) and losses to consumers (G), so that the true net primary productivity=
dB+ L + G. Net primary productivity can be measured in terms of biomass as
grams per square meter per year (g/m2/yr). Alternatively, the caloric content of the
biomass can be measured and net primary productivity reported as calories per
square meter per year. Since the rate of vegetation growth varies in different
ecosystems, net primary productivity is highly variable from place to place. In gen-
eral, plant biomass and primary productivity are highest in the warm and wet trop-
ics and lowest in cold polar ecosystems and dry deserts. The mean net primary
productivity of a tropical rainforest ecosystem is some 2200 grams per square
meter per year. In desert environments, the net primary productivity averages from
90 to 3 grams per square meter per year. The net primary productivity of the open
ocean averages 125 grams per square meter per year. We will discuss the global dis-
tribution of net primary production further when we examine the world's biomes.
When we look at the distribution of the standing biomass of different
trophic levels, the impact of the 10% rule quickly becomes apparent. These distri-
butions often have a pyramidal shape in which biomass can decrease by 90% or
more with each increase in trophic level. However, the shape of such pyramids
can vary widely. In Panamanian rainforests, it has been shown that it takes
approximately 40,000 grams per square meter of vegetation biomass to support
22 CHAPTER 2 SOME BASICS

14 grams per square meter biomass of primary consumers, secondary consumers, Gl<
and decomposers. In coral reef systems, 703 grams per square meter of primary
producer biomass can support 132 grams per square meter of herbivores and 11 Bef1
grams per square meter of carnivores. fere
It would therefore seem that we should be able to predict the number of diff(
trophic levels that a given ecosystem will support. The food webs of insects, which phy~
have high-energy efficiencies, living in ecosystems with high primary productivity char
should contain more trophic levels than other types of communities. Interest- of tl
ingly, this has not been found to be the case. Most terrestrial food webs, despite we a·
differences in primary productivity and the thermal efficiency of the organisms, desc
tend to have three to four trophic levels. This means that energy flow cannot be Ang
the only factor limiting the number of trophic levels. It is also interesting to note such
that despite the low primary productivity of the oceans, many marine communi-
ties have five trophic levels, and carnivores are much more abundant compared pher
to the number of carnivores found in terrestrial systems. It is possible that the eral
transfer of energy between the marine primary producers, which are mostly (Fig.
microscopic plants that float in the water, and primary consumers, which are trop1
mainly microscopic crustaceans called copepods, is much more efficient than the tains
transfer of energy between land plants and terrestrial herbivores. Indeed, in some air ir
oceans the standing biomass of photosynthetic plankton may be low relative to 0.03f
the biomass of other trophic levels because of the efficiency with which the cope- sphe1
pods harvest the phytoplankton and incorporate the energy captured by them. In humi
the English Channel, 21 grams per square meter of herbivore standing biomass posp:
are supported by only 4 grams per square meter of primary producer standing 3.3%
biomass. 1000-
As a final thought on trophic levels, we might reflect on the place of humans coole
in the food webs of the world's ecosystems. Since we are omnivores, we function
as both primary consumers and higher level consumers. It is easy to see that when recei·
we eat the meat of herbivores such as cows or sheep, we are secondary con- trum
sumers. When, however, do we function as tertiary or quaternary consumers and
eat other carnivores? Most people eat fish. Important food species such as tuna
and salmon are carnivorous tertiary and quaternary consumers. Thus, in terms of
10
marine food webs, we function as quaternary consumers or even higher in the
trophic hierarchy!
8

PHYSICAL GEOGRAPHY AND THE E'

~ 6
FUNCTIONING OF THE EARTH Q)
-o
::J
E
The science of physical geography is concerned with the physical patterns and <( 4
processes that occur at the earth's surface. Physical geographers conduct research
on phenomena ranging from local landforms caused by glacial activity to large- 2
scale variations in the world climate. The most important areas of physical geog-
raphy for biogeographers are climatology, which is the study of the earth's
atmosphere and climate; pedology, which is the study of soils; limnology, which is
the study of freshwater; and oceanography, which is the study of the world's
oceans. It is worthwhile to review a few important concepts from the areas above
in order to investigate how features of the earth's physical environment interact FIGU
with the biosphere to influence the distribution of life. and c:

tmers, secondary consumers,
>er square meter of primary
meter of herbivores and 11
•le to predict the number of
e food webs of insects, which
th high primary productivity
~s of communities. Interest-
errestrial food webs, despite
efficiency of the organisms,
s that energy flow cannot be
. It is also interesting to note
ans, many marine communi-
;h more abundant compared
stems. It is possible that the
>reducers, which are mostly
imary consumers, which are
much more efficient than the
tl herbivores. Indeed, in some
1kton may be low relative to
'ficiency with which the cope-
~ energy captured by them. In
f herbivore standing biomass
,f primary producer standing
eflect on the place of humans
e are omnivores, we function
ers. It is easy to see that when
1eep, we are secondary con-
or quaternary consumers and
ant food species such as tuna
r consumers. Thus, in terms of
;umers or even higher in the
'ith the physical patterns and
geographers conduct research
~d by glacial activity to large-
ortant areas of physical geog-
l is the study of the earth's
dy of soils; limnology, which is
11 is the study of the world's
;oncepts from the areas above
physical environment interact
ife.
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 23
Global Climate
Before we begin our examination of the earth's climate, we should note the dif-
ference between the terms weather, climate, and atmosphere. These are three very
different concepts and are sometimes confused. Weather is the condition, or
physical state, of the atmosphere at a given place at a given time. The weather
changes from day to day or even hour to hour. Climate is the average condition
of the atmosphere at a given place based on statistics from a long period of
weather observations. When someone says it is raining in Boston today, he is
describing the weather. When you read that the average high temperature in Los
Angeles in July is 28° C, you are reading a description of climate. By convention,
such climatic averages are usually based on 30 years of observation.
The atmosphere is the layer of gas that surrounds the earth. The atmos-
phere extends over 100 km above the surface of the earth and is divided into sev-
eral different layers based on temperature, pressure, and chemical composition
(Fig. 2.4). However, all life exists in the lowest level of the atmosphere, called the
troposphere. The troposphere extends some 9 to 17 km above the earth and con-
t~i~s over 90% of the total mass of the atmosphere. The composition of pure dry
au m the troposphere is roughly 78% nitrogen, 21 % oxygen, 0.93% argon, and
0.036% carbon dioxide and a small amount of other gases. The air in the tropo-
sphere can be relatively dry or contain up to 5% water vapor in places such as the
humid tropics. Both air pressure and temperature decrease with height in the tro-
posphere. For every 275-m rise in elevation, the air pressure decreases by about
3.3 %, while temperature decreases at a rate of approximately 6.5° C for every
1000-m increase in altitude. This is why the air is thinner and temperatures are
cooler at the tops of high mountains.
As you recall from our discussion of photosynthesis, most of the sunlight
received by the earth is in the visible light portion of the electromagnetic spec-
trum ( ~0.4-0.6 micron wavelength). This energy is absorbed at the earth's surface
100
Thermosphere
80 - - - - - - - - - - - - - - - - - - - - Mesopause - -
Mesosphere
'E 60
6
(j)
"C
- - - - - - - Stratopause- -
.g
40
< Stratosphere
20
0 50
Air pressure relative to Air temperature
sea level value(%) (OC)
FIGURE 2.4 The atmosph eric stratification of the earth and th e variation in temperature
and air pressure within the troposphere (after Strahler and Strahler, 1997).
24 CHAPTER 2 SOME BASICS

and then emitted as heat energy at longer wavelengths of -5.0 to 20.0 microns.
The gases of the atmosphere allow most of the solar energy in the visible light
wavelengths to pass through the atmosphere but tend to absorb the longer wave-
length heat energy that is emitted from the earth's surface. Carbon dioxide and
Surr
water vapor in the atmosphere are important in trapping this heat. This phenom-
enon of the transmission of incoming solar light energy and the trapping of out-
going heat energy is known as the greenhouse effect. Without this atmospheric
effect, the earth would be too cold to support life.
The distribution of climatic conditions on the earth is not random but fol-
lows a predictable pattern that is controlled mainly by latitude, elevation, and the
Ni~
distribution of continents and oceans. As we shall see later, if we can predict the
climatic conditions at a certain place, we can also make certain predictions about
the type of organisms we will find there. Latitude is the most important control
on the overall distribution of climatic conditions. Latitude determines the
amount of solar energy that is intercepted over a given area of the earth's surface
(Fig. 2.5). The earth orbits the sun at an average distance of about 150 million km.
The orbit is elliptical, so that in January the earth is about 2 1/z million km closer
to the sun. This is referred to as the perihelion. The aphelion occurs in July when
the earth is an additional 2 1/z million km away from the sun. At these distances,
the electromagnetic energy from the sun can be assumed to be traveling as paral-
lel waves with a constant energy of 1400 watts per square meter. During March
and September, at the equator, these rays strike the earth at a 90° angle at noon,
so that a square meter of incoming solar radiation is intercepted by a square
meter of earth's surface. At higher latitudes, the angle of incidence of the sun's
rays decreases. As a result, a square meter of solar energy is spread over an area
much greater than a square meter of the earth's surface. Due to the decrease in
the angle of insolation, temperatures should be cooler at higher latitudes because
they receive less solar energy. However, that is only part of the story.
The latitudinal distribution of solar energy varies with season as well as lat-
itude. Because the earth's axis is tilted about 23 1/z 0 relative to the plane of the FIG I
orbit around the sun (Fig. 2.5), the amount of solar energy received at different distr
latitudes varies throughout the year. At the December solstice, the northern at th
hemisphere is tilted away from the sun so that the angle of incidence of solar the i
energy is decreased and areas north of 66 1/ 2° (the Arctic Circle) do not receive
any direct sunlight. The point of the earth's surface where the sun's rays strike the dur
earth at a 90° angle lies at 23 1/ 2° south latitude (the Tropic of Capricorn). During lati<
June and the summer months, the northern hemisphere is tilted toward the sun, insc
and the angle of incidence in the north is increased. North of the Arctic Circle the add
sun never sets. The sun's rays strike the earth at a 90° angle at the Tropic of Can- inez
cer (23 1/z0 north latitude). The tropics are defined as the region lying between 23 pol.
1/ ° north and south latitude. During the equinoxes, on March 21 and September ada
2
23, neither hemisphere is tilted towards or away from the sun. The sun's rays poi
strike the earth at a 90° angle at the equator, and there are 12 hours of daylight at ing
all latitudes.
The seasonal distribution of insolation has important implications for tem- the
peratures and life on earth. The long days of insolation during May, June, July, and alle
August at the high latitudes of the northern hemisphere provide large amounts of me
solar energy. This energy results in temperatures warm enough for ice and snow to SIDt
melt and plant life to exist. In the southern hemisphere, the long days of high inso- fen
lation are experienced in the months of November, December, and January. Thus, ave
 PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 25

gths of -5.0 to 20.0 microns. Vernal Equinox North Pole

March 21
u energy in the visible light
d to absorb the longer wave-
surface. Carbon dioxide and Circle of
'ping this heat. This phenom-
:rgy and the trapping of out-
Summer Solstice
June 21 ---------- ""---......'·-........
Day ·-
mum nation

ct. Without this atmospheric

Sun
earth is not random but fol-
lY latitude, elevation, and the
:e later, if we can predict the
_.,. Winter
tke certain predictions about /.· Solstice
; the most important control Earth's orbit ~/ December 22
s. Latitude determines the _\ ___.---
en area of the earth's surface Autumnal Equinox
mce of about 150 million km. September 23
about 2 1h million km closer
600
aphelion occurs in July when
11 the sun. At these distances,
500
.-
1med to be traveling as paral- 'E f~ ~c ..
Q;
-
-0 ~fl ~ ..""\ ~
square meter. During March a. 400
earth at a 90° angle at noon,
en oY I 'II ·~ I\ [\_
'iii
s: 300
-'fj,l I If : ··~\ \ f'..,_
1 is intercepted by a square ..L 0
' \\ I\
c0 --f
gle of incidence of the sun's '\\ \
~ 200 f--1 o I
:nergy is spread over an area 0 !""' -i&J. f ~ 1\..
en
rface. Due to the decrease in E 100 f - - i l ~
~·
lc.'"" \ ~
=r at higher latitudes because f - ~-&
f-' _,'
10

part of the story.
ies with season as well as lat-
a relative to the plane of the
energy received at different
~mber solstice, the northern
~ angle of incidence of solar
Arctic Circle) do not receive
vhere the sun's rays strike the
Tropic of Capricorn). During
here is tilted toward the sun,
'l" orth of the Arctic Circle the
)
0
angle at the Tropic of Can-
s the region lying between 23
on March 21 and September
'rom the sun. The sun's rays
:re are 12 hours of daylight at
portant implications for tem-
Jn during May, June, July, and
tere provide large amounts of
m enough for ice and snow to
:re, the long days of high inso-
December, and January. Thus,
. ~ 1-
0
JFMAMJJASOND
FIGURE 2.5 The seasonal orbital characteristics of the earth and the seasonal
distribution of insolation at different latitudes (after Strahler and Strahler, 1997). Notice how
at the poles (90° latitude) there is less total insolation and greater seasonal variation than at
the equator (0° latitude).
during the 24 hours of sunlight in June, the North Pole actually receives more inso-
lation than the equator (Fig. 2.5). However, over the course of the year, the average
insolation at the equator is much higher than at polar regions. A very important
additional aspect of insolation distribution is the way that seasonal differences
increase with latitude (Fig. 2.5). Organisms existing in the Arctic and Antarctic
polar regions not only have to contend with less total insolation, but they have to
adapt to large seasonal differences between winter and summer. A final interesting
point to note, is that the high and midlatitudes experience one insolation peak dur-
ing the summer, while the equator experiences two peaks in spring and fall.
If incoming insolation was the only control on temperatures at the surface of
the earth, we would see surface temperatures decreasing in a smooth fashion, par-
allel with latitude, from the equator to the poles. However, if we look at maps of
mean January and July temperatures, we see that temperatures do not decrease
smoothly in parallel with latitude (Fig. 2.6). Some of this can be explained by dif-
ferences in elevations. As we discussed earlier, high-elevation sites are cooler on
average than low-elevation sites. Upon inspection of temperature maps, it is clear
26 CHAPTER 2 SOME BASICS

W<
wl
th
lat
fa<
dr
eli.
00
be
int
ab
on

pn
January ph
wh
of
air
pe1
wa
dec
wit
dec
is c
lap
rat'
en<

pre
sur
OC(
July
sur
FIGURE 2.6 Average monthly temperatures {°C) at the surface of the earth in January pn
and July (after Strahler and Strahler, 1997). ovc
sto
wh
fas
on
that summer temperatures are higher over land surfaces than over the sea. Win- inf
ter temperatures are lower on land relative to the sea. This phenomenon is par-
ticularly notioeable in interior Alaska and central Siberia where average January sue
temperatures range from - 25° C to -50° C, while January temperatures in the ah
adjacent North Pacific range from 0° C to - 15° C. In fact, central Siberia is the of
coldest place in the northern hemisphere, with winter temperatures commonly to1
far below those encountered at the North Pole. The coldest recorded temperature re1
in North America occurred in the village of Snag in the Yukon Territory of su
Canada. In July the average daily temperatures in Siberia and Alaska rise to 10° frc
C- 15° C, while the air temperatures over the oceans increase to only soC- 10° C. ap
The reason for these differences in surface temperatures of continents and W<
oceans is related to the differential rates of heating and cooling of land and Tl

urface of the earth in January
faces than over the sea. Win-
;ea. This phenomenon is par-
beria where average January
January temperatures in the
In fact, central Siberia is the
1ter temperatures commonly
:oldest recorded temperature
g in the Yukon Territory of
;iberia and Alaska rise to 10°
; increase to only soC-10° C.
peratures of continents and
ing and cooling of land and
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 27
water. The solid land surface heats and cools more rapidly than the liquid sea
where energy is dispersed in depth. Thus, the land responds more dramatically to
the seasonal increases and decreases in insolation. In addition, the oceans circu-
late heat laterally through currents. Such transfers do not occur on solid land sur-
faces. Sites that are located in the interior of landmasses and are subject to
dramatic seasonal differences in temperature are said to experience continental
climates, whereas islands and coastal sites, that have climates moderated by the
ocean, are said to have marine climates. In the case of Siberia, the difference
between the mean January and July temperatures can be as high as 60° C. In the
interior of Alaska and Canada, this difference in seasonal mean temperatures is
about 4S 0 C. In contrast, the difference between January and July temperatures
on islands and at coastal sites in the tropics is less than 3° C.
Now, let's consider the geographic distribution of precipitation. The term
precipitation is used here to denote moisture that is deposited from the atmos-
phere to the land or water surface as rain, sleet, hail, or snow. Precipitation occurs
when the water vapor in the atmosphere exceeds the moisture-holding capacity
of the air. The amount of water that can be held as vapor increases rapidly with
air temperature. Air with a temperature of ooC can only hold about S g of water
per kilogram of air, whereas air with a temperature of 30° C can hold over 2S g of
water per kilogram of air. When air cools, its ability to hold moisture as vapor
decreases and eventually precipitation forms. As we discussed, air generally cools
with increasing altitude in the troposphere. The average rate of temperature
decrease is about 6.SO C for every 1000 m of altitude, although this varies. Air that
is dry cools at a rate of about 10° C per 1000 m. This is called the dry adiabatic
lapse rate. Air in which condensation is occurring cools at the wet adiabatic lapse
rate, which is approximately so C per 1000 m. So, as air rises it cools, and if it cools
enough, condensation and precipitation occur.
Three processes cause air to rise and generate precipitation. The first
process is convective precipitation, which occurs when the warming of air at the
surface causes the air to rise until it becomes cool enough for precipitation to
occur. This phenomenon causes the thunderstorms and rain showers typical of
summer weather in many parts of the world, particularly the tropics. The second
process is frontal precipitation, which occurs when warm air masses rise up and
over denser masses of cool air. An example of this phenomenon are the winter
storms that occur in the middle latitude regions. Finally, air may be forced to rise
when it encounters physical barriers such as mountains. Rain generated in this
fashion is referred to as orographic p recipitation. The high rainfall encountered
on the western slopes of the Coast Ranges from northern California to Alaska is
influenced by orographic conditions.
The global distribution of precipitation is controlled by surface conditions,
such as the location of mountains and oceans, and by the general circulation of the
atmosphere. The general circulation of the atmosphere is the large-scale patterns
of winds that develop in response to differences in radiation and heat at the equa-
tor and the poles. Precipitation, due to convection, is typical of the equatorial
regions which, due to high insolation, experiences vigorous heating at the earth's
surface. This surface heating causes air to"warm and rise. High rates of evaporation
from the warm sea surface provide abundant moisture to the atmosphere. At
approximately 30° north and south latitude, the tropical air begins to subside and
warm as it descends (Fig. 2.7). As the air warms, it can carry more water as vapor.
Thus, areas that are typified by descending air are dry. The descending air causes a
28 CHAPTER 2 SOME BASICS

band of calm and dry climate in the subtropics at around 30° north and south lati-
tude. These regions are referred to as the doldrums or horse latitudes because sail-
ing ships often were becalmed in these latitudes and the sailors would jettison
horses and other livestock to conserve water. At the surface, air flows from the
region of the horse latitudes to replace the rising air in the tropics (Fig. 2.7). The
flow of the air is deflected to the right due to the Coriolis effect imparted by the
rotation of the earth. As a result, there is a zone of winds from the northeast and
the southeast in the equatorial region that are referred to as the tropical trade
winds. The region where the trade winds from the northern and southern hemi-
sphere converge is called the Inter-Tropical Convergence Zone (ITCZ). The con-
verging winds at the ITCZ replenish moisture for convective precipitation in the
tropical zone. North and South of the horse latitudes, the surface air flows away
from the zone of subsidence in an easterly direction, and this is referred to as the
midlatitude westerlies. These westerlies dominate the temperate areas of the earth
between 30° and 60° north and south latitude. Storms develop along the belt of
westerlies as the atmosphere picks up moisture over the oceans and produces
frontal precipitation. North and south of the midlatitude westerlies, polar air
masses dominate. The polar air masses are generally very cold and very dry.
Elevation also plays a role in the distribution of precipitation. Because of
orographic lifting and cooling of air, high elevations typically have higher
amounts of precipitation than adjacent low-elevation sites. In areas such as
coastal Oregon, Washington, and British Columbia, the frequent occurrence of
midlatitude frontal storms brought ashore by the westerlies, coupled with oro-
graphic precipitation due to high mountain ranges along the coast, produces
large annual totals of precipitation. Air also heats as it descends down the lee side
of mountains. This promotes dry conditions, which are referred to as rainshadows.
In the region of westerlies, rainshadows are often found on the east side of moun-
tain ranges. The dry eastern slopes and the adjacent deserts of the Sierra Nevada
Mountains of California are a result of the rainshadow effect. The warm and dry
winds, experienced as air flows down mountains, are called Chinook Winds in the
northern Rocky Mountains and Fohn Winds in Europe.
The general distribution of annual precipitation on the earth reflects the
processes outlined above (Fig. 2.7). The greatest amounts of precipitation generally
occur in the tropics where over 500 em of rainfall may fall in a given year. Similar
magnitudes of rainfall are experienced on the western slopes of coastal mountains
in western North America, Norway, and western South America. A band of low pre-
cipitation is centered on 30° north and south latitude. This includes some of the
world's great deserts such as the Sahara in northern Africa, the Namibian in south-
ern Africa, the Arabian Desert, the coastal deserts of Chile and Peru, the Australian
Outback, and the deserts of the U.S. Southwest and adjacent Mexico. Average
annual rainfall in some of these regions may be less than 2.5 em per year. In some
areas, such as the Sahara Desert of southern Sudan, 100 years may pass between
occurrences of measurable rainfall. In the midlatitudes, very dry conditions and
deserts may extend as far as 45°-50° north and south latitude in the rainshadows of
mountain ranges. The Great Basin Desert of California, Nevada, and Utah is an
example. The arctic and antarctic regions also experience low annual precipitation..
However, we generally do not think of these regions as deserts. Low precipitation is
not the only factor in creating a desert. In true deserts, the rate of evaporation
greatly exceeds precipitation, and there is a deficit in soil moisture at most times. The
arctic and antarctic regions have low precipitation, but because of low temperatures..
md 30° north and south lati-
horse latitudes because sail-
::1 the sailors would jettison
: surface, air flows from the
in the tropics (Fig. 2.7). The Cii
iolis effect imparted by the cOC
tnds from the northeast and ....
~o

uco
~
r--
·c:!:: C1l
C1l
red to as the tropical trade 0 .9-
-u
'-'aJ ...:-
)rthern and southern hemi- >-~
()c. ~
.t::
nce Zone (ITCZ). The con- ~
Lvective precipitation in the Ul
"0
: the surface air flows away c
md this is referred to as the "'
~
.t::
temperate areas of the earth ~
LS develop along the belt of Ul
r the oceans and produces Qi
4=
ttitude westerlies, polar air ~

ery cold and very dry. E

~
)f precipitation. Because of c
.ons typically have higher .g
~
.on sites. In areas such as ·a.
·c:;
the frequent occurrence of Q)

esterlies, coupled with oro-

c.
-ro:::l
along the coast, produces c
c
. descends down the lee side
"'
Q)
referred to as rainshadows. Cl

1d on the east side of moun- "'w

>
eserts of the Sierra Nevada "'c
"0
w effect. The warm and dry
:ailed Chinook Winds in the
"'
Q)
w
.t::
)e. c.
(/)
0
n on the earth reflects the
1ts of precipitation generally
s
"'
Q)
1 fall in a given year. Similar
slopes of coastal mountains
America. A band of low pre-
-.g
£
0
c

~- This includes some of the ~

:::l
Erica, the Namibian in south- ~
·c:;
bile and Peru, the Australian -ro....
I adjacent Mexico. Average Q)
c
Q)
tan 2.5 em per year. In some <!)

.00 years may pass between .....

es, very dry conditions and c-.i
ttitude in the rainshadows of w
a:
tia, Nevada, and Utah is an ::J
C)
1ce low annual precipitation. ii:
deserts. Low precipitation is
rts, the rate of evaporation
1moisture at most times. The
>ecause of low temperatures,
29
30 CHAPTER 2 SOME BASICS

they also have low evaporation rates. Therefore, these polar regions cannot be con-
sidered to be true deserts in most cases.
In addition to geographic variations due to latitude and orographic fea-
tures, seasonal differences in precipitation are important influences on the distri-
bution of life. The ITCZ does not stay fixed at the equator but changes seasonally. -·
During June and July the ITCZ moves north toward 23 1/z 0 north latitude. During
December and January it moves toward 23 1/ 2° south latitude. As the ITCZ
moves north and south, it brings precipitation to the arid subtropical regions near
30° north and south latitude. In addition, the increased warming of the land sur-
face in the subtropical regions during the summer produces a pattern of
enhanced convective uplift over the land and surface flow of moist air from the
oceans. This creates a rainy season called the Summer Monsoon, which is impor-
tant for bringing moisture to areas such as India, Southeast Asia, and the South-
western United States. Thus, the areas bordering the tropics experience their
highest rainfalls during the summer months and often have very dry winters. In
the midlatitudes, the westerly storm tracks are also displaced. In the summer,
westerly storms do not generally travel as far south as California or Spain, and FIC
this brings dry summers to these regions. In contrast, during December and Janu-
ary, the storm tracks move south and bring winter rains.
Because of geography's role in controlling temperature and precipitation,
the earth can be divided into broad climatic zones based on temperature and
es1
precipitation regimes. One such scheme, often used by biogeographers, is based
tio
on the 1918 work of Wladimir Koppen as later modified by Geiger and Pohl.
co
Koppen was a knowledgeable biogeographer as well as a climatologist, so his sys-
tem was designed to reflect the relationship between world climate and vegeta-
icy
tion zones. The climatic zones he proposed are based strictly on long-term
A]
climatic averages. In determining the boundaries of the climatic zones, Koppen
lru
va
also used vegetation boundaries as guides. He reasoned that large-scale vegeta-
su
tion boundaries were likely related to climate. The complete Koppen system has
de
5 major climate zones and 13 climate types, with a mechanism for further subdi-
ni
visions. A simplified global version of the Koppen system excluding highland
so
areas is presented in Figure 2.8. The five major climate zones are:
we
A. Tropical Rainy Climates found in the equatorial regions. This zone has
monthly average temperatures of 18° Cor higher, with little seasonal varia- N
tion. Rainfall is abundant throughout the year and always exceeds evapora-
liJ
tion.
H
B. Dry Climates found mainly in the subtropical zone. Temperatures are gen- Vf
erally warm, and evaporation exceeds precipitation throughout all or most di
of the year. e1
C. Mild Humid Oimates found in the midlatitudes. The temperature of the fa
coldest month falls between 18° C and - 3° C, with a clear difference in win- S(

ter and summer seasons. Precipitation exceeds evaporation. aJ

D. Snowy Climates found in the mid- to high latitudes. The coldest month has n
an average temperature below -3° C, but the average for the warmest c
month is greater than 10° C. Precipitation exceeds evaporation.
tl
E. Polar Climates found in the polar regions. The average temperature of the
Sl
warmest month is less than 10° C. Precipitation is low, but evaporation is
3
very limited.

:se polar regions cannot be con-
> latitude and orographic fea-
,o rtant influences on the distri-
quator but changes seasonally.
d 23 1/zo north latitude. During
south latitude. As the ITCZ
e arid subtropical regions near
ased warming of the land sur-
lmer produces a pattern of
1ce flow of moist air from the
n.er Monsoon, which is impor-
outheast Asia, and the South-
the tropics experience their
'ten have very dry winters. In
;o displaced. In the summer
h as California or Spain, and
1. during December and Janu-
lins.
mperature and precipitation,
s based on temperature and
by biogeographers, is based
odified by Geiger and Pohl.
as a climatologist, so his sys-
n world climate and vegeta-
based strictly on long-term
the climatic zones, Koppen
•ned that large-scale vegeta-
Jmplete Koppen system has
techanism for further subdi-
system excluding highland
e zones are:
rial regions. This zone has
~r. with little seasonal varia-
ld always exceeds evapora-
me. Temperatures are gen-
ion throughout all or most
:s. The temperature of the
h a clear difference in win-
'aporation.
les. The coldest month has
average for the warmest
s evaporation.
'erage temperature of the
is low, but evaporation is
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 31
Mild humid
Polar
FIGURE 2.8 Simplified Koppen climate classification (after Strahler and Strahler, 1997).
Before we conclude this review of climate, we will consider some inter-
esting properties of local climate related to mountains. Under normal condi-
tions, temperatures decrease with altitude. Thus, high elevations are generally
cooler than low elevations. The impact of this can be seen with particular clar-
ity when we consider how the high elevations of mountains such as the
Appalachians or Alps experience lower temp eratures than the adjacent low-
lands during both the summer and winter. However, in some localized instances,
valley bottoms in the mountains can experience colder temperatures than the
surrounding slopes and peaks. The enhanced cooling of the valleys is caused by
dense cold air which drains downward from higher elevations, particularly at
night and in the early morning. The effect is known as cold air drainage. In
some instances, valley bottoms may be too cold to support trees, which grow
well on adjacent slopes.
Microclimate
In our discussion we have looked at relatively large-scale climatic conditions.
However, significant differences in climate can occur at very small scales. Such
very local conditions are referred to as microclimates. For example, aspect is the
direction that a slope faces. A south-facing slope in North A merica can experi-
ence daytime temperatures that are over 10° C warmer than an adjacent north-
facing slope. Evaporation rates are also higher on the warm south-facing slope, so
soils are drier. Thus, the south-facing slope may support plants and animals that
are typical of hot and dry climates, such as desert species, while the adjacent
north-facing slope may be wooded. South-facing slopes in the Yukon Territory of
Canada sometimes support plants typical of the northern Great Plains, while
adjacent north-facing slopes support cool subarctic forest species. Sites close to
the ground are generally warmer than elevated and exposed sites. In the arctic,
summer daytime temperatures can decrease by over soC from the soil surface to
30 em above the surface. Temperatures decrease rapidly below the soil surface.
32 CHAPTER 2 SOME BASICS

The air temperature in rodent burrows, which are only a few tens of centimeters
below the surface, may be over 10° C cooler than the exposed top soil.
Forests create profound differences in microclimate. The amount of sun-
light received at the floor of a dense forest may be less than 10% of the amount
received at the top of the forest canopy. Temperatures at the forest canopy can be
well over 10° C higher than those found on the forest floor. Decreased evapora-
tion rates and the transpiration of leaves cause forest floors to be much damper
than at the canopy level. Finally, wind speed can decrease by as much as 90%
between the top of the canopy and the forest floor.

World Soils
Soil is the uppermost layer of mineral and organic matter found on the earth's sur-
face. Soil supports and sustains almost all plant life on which terrestrial ecosystems
depend. In addition, many organisms, ranging from bacteria to mammals, live
within the soil. Soil structure, texture, mineralogy, water content, and chemistry
can exert a strong control on the distribution of organisms. Soil is formed by the
physical and biological weathering of rock and the addition of organic matter. The
formation of soils is dependent on many factors and can take thousands of years.
The type of regolith (weathered rock and mineral matter) from which soil is
developed is important. Plants and animals may depend on soils, but they also
modify regolith to create soils. In this manner soil and life on the earth have
evolved together. Climate is also extremely influential in determining the type of
soil that will develop in a given locale. Both temperature and moisture conditions
can have a great influence on soil development. Thus, soils vary in structure, tex-
ture, and chemical properties both locally and globally. A gradient in soil charac-
teristics and types across landscapes is referred to as a catena.
One means of describing soils is by texture. This refers to the amount of
sand, silt, and clay in the soil. Loam soils, with a relatively equal mixture of silt,
sand, and clay, are often best for plant growth because they hold and exchange
water easily and provide mineral nutrients in a form useful to plants. Water runs
quickly through sandy and rocky coarse soils. In addition, plants cannot obtain
nutrients from coarse particles. Water runs slowly through clay-dominated fine
soils and is difficult for plants to acquire. Clay and silt soils that lack sand can FIGU
impede root penetration. maj01
Soils can be divided into stratigraphic units called soil horizons. The bound- and S
aries between the horizons generally run parallel to the soil surface. Each horizon
has distinctive physical and chemical properties. Often, color can be a useful
means of discriminating soil horizons and different soil types in the field. An horu
objective means of describing soil color is provided by the use of a standardized grad
color chart provided by a Munsell Color Book. A vertical section through the
various horizons is called a soil profile, and comparing profiles provides another tract
means of classifying soils. The principal horizons of a soil profile are generally tied
designated as A , E, B, and C (Fig. 2.9). clayl
The C horizon is the unconsolidated rock at the base of the soil profile. It beer
can retain traces of the structures and bedding of the original rock. The C horizon thes'
has undergone little chemical transformation and most closely resembles the bloc
chemical and mineral composition of the original rock. However, silica, carbon- of ir
ates, and salts, weathered from the overlying horizons, can accumulate in the C bloc
 PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 33

ly a few tens of centimeters

exposed top soil.
limate. The amount of sun-
~ss than 10% of the amount
s at the forest canopy can be
;t floor. Decreased evapora-
.t floors to be much damper
!Crease by as much as 90%

\ , I ·
---~~-·- ·----L ___ ,L __ _
;__
.i

:ter found on the earth's sur-

which terrestrial ecosystems
bacteria to mammals, live
ater content, and chemistry
.nisms. Soil is formed by the
1ition of organic matter. The
;an take thousands of years.
matter) from which soil is
Jend on soils, but they also
and life on the earth have
1 in determining the type of
ure and moisture conditions
, soils vary in structure, tex-
y. A gradient in soil charac-
t catena.
hls refers to the amount of
tively equal mixture of silt,
tse they hold and exchange
useful to plants. Water runs
lition, plants cannot obtain
crough clay-dominated fine
silt soils that lack sand can
d soil horizons. The bound-
e soil surface. Each horizon
'ten, color can be a useful
soil types in the field. An
y the use of a standardized
ertical section through the
g profiles provides another
a soil profile are generally
:!base of the soil profile. It
•riginal rock. The C horizon
1ost closely resembles the
k. However, silica, carbon-
s, can accumulate in the C
Soils of the World
[ ] Spodosols
• Boralfs1
• Udalfs Alfisols
• Ustalfs
Xeralfs
Mollisols lay and calcium
.Aridisols compounds
• Tundra soils Parent material
0 Highlands Grassland soil
(Mollisol)
FIGURE 2.9 A profile of a typical grassland soil horizon and the world distribution of
major soil types accordi ng to the Comprehensive Soil Classification System {after Strahler
and Strahler, 1997).
horizon. The base of the C horizon is often difficult to determine exactly as it
grades into the unweathered underlying rock.
The B horizon overlies the C. It is highly weathered and does not retain any
traces of the original bedding or structures of the bedrock. The B horizon is typi-
fied by illuviation (the accumulation of material) and contains substances such as
clays, calcite, aluminum and iron oxides, and humus (organic matter) that have
been weathered and leached downward from the overlying horizons. Some of
these substances bind particles together, imparting structures of granules, plates,
blocks, and columns to the B horizon. Under certain conditions the accumulation
of iron oxides, gypsum, salts, silica, and clay can form impermeable layers that
block the movement of water and nutrients in the soil profile. When such barriers
34 CHAPTER 2 SOME BASICS

consist of uncemented clays, they are called fragipans or claypans. When they
consist of cemented material, they are called duripans or hardpans.
The A and E horizons are formed by eluviation (loss of material) and are
depleted in clays, aluminum, and iron oxides relative to the B horizon. When
leaching is particularly intense, the E horizon can be bleached of color, while
organic matter in the A horizon makes it darker. Because biological activity such (
as rooting and soil-dwelling organisms is important, the A horizon contains more
humus than the E. The A horizon can consist of up to 30% humus. The decay of
humus releases organic acids that promote weathering and leaching in the soil.
The replacement of cations such as calcium, magnesium, potassium, and sodium
in the A and E horizons by hydrogen can reduce the ability of the soil to support
plant growth. In some cases an extremely organic-rich layer of plant matter, the
0 horizon, overlies the A horizon. 1
As is the case with climate, the soils of the world can be classified into broad
categories to produce world soil maps. The pioneers in the area of soil descrip-
tion, classification, and mapping were the Russian scientists V. Dokuchaiv and K.
Glinka and Americans such as E. Hilgard and C. Marbut who worked in the late
nineteenth and early twentieth centuries. Broad classification systems take into
account the structure, texture, chemical properties, and profiles of the soils.
Unlike biological taxonomy, however, there is no international standard system
for the classification of soils. Even in North America different systems are used.
In the United States the Comprehensive Soil Classification System is the accepted
norm. Canada has a different system, the Canadian System of Soil Classification. .·
Many other countries have developed their own systems or use terms from world
soil classification schemes developed by Russian or British pedologists.
The Comprehensive Soil Classification System divides soils into 11 soil
orders. The different orders are discriminated on the basis of criteria such as soil
composition and texture, degree of horizon development, presence or absence of
diagnostic horizons, and degree of weathering of soil minerals. The global distri-
bution of these soil types has been mapped by the U.S. Department of Agricul-
ture (Fig. 2.9). The distribution of some soil types do correspond with Koppen's
climatic regions. The soil orders are as follows:

Entisols are mineral soils that have undergone little to no alteration. They lack
distinct soil horizons and are often found on fresh deposists from rivers, gla-
ciers, and sand dunes. Because of low weathering, it may be difficult for
plants to obtain nutrients from some entisols. Many tundra and desert soils
are entisols.
Inceptisols are young soils that possess enough soil moisture to support plants
for at least three months of the year, have one or more discernible horizons,
are fine to loamy in texture, and contain weathered minerals and associated
plant nutrients. Some tundra soils are inceptisols. Cryaquept inceptisols are
typically found in polar climate regions and high mountains. Aquept incep-
tisols are found in bogs and marshes throughout the world. Inceptisols are
also found on floodplains where significant sedimentation is no longer
occurring.
Tt
Histosols are generally moist and have up to 30% organic matter in the upper
portion of the soil. The organics can come from mosses and other plants La
growing in water, such as in bogs, and from deep forest litter. These soils are the

ms or claypans. When they
s or hardpans.
n (loss of material) and are
;e to the B horizon. When
>e bleached of color, while
wse biological activity such
1e A horizon contains more
) 30% humus. The decay of
ng and leaching in the soil.
urn, potassium, and sodium
:tbility of the soil to support
h layer of plant matter, the
can be classified into broad
in the area of soil descrip-
~ntists V. Dokuchaiv and K.
but who worked in the late
sification systems take into
and profiles of the soils.
emational standard system
different systems are used.
ztion System is the accepted
ystem of Soil Classification.
ms or use terms from world
ritish pedologists.
divides soils into 11 soil
)asis of criteria such as soil
ent, presence or absence of
minerals. The global distri-
.S. Department of Agricul-
correspond with Koppen's
to no alteration. They lack
1deposists from rivers, gla-
ng, it may be difficult for
my tundra and desert soils
moisture to support plants
more discernible horizons,
~d minerals and associated
Cryaquept inceptisols are
mountains. Aquept incep-
the world. Inceptisols are
dimentation is no longer
rganic matter in the upper
1 mosses and other plants
orest litter. These soils are
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 35
commonly acidic and low in nutrients. High water content may produce
reducing conditions and an abundance of iron oxides in the mineral soil
beneath the A horizon. Histosols are common in the coniferous forests of
the snowy climate region. They can also be found in other regions on wet,
vegetated sites.
Oxisols are characterized by the deep weathering of most minerals except quartz,
aluminum, and iron. The soils have water, but the nutrient content is often low.
They generally present little evidence of soil horizons, except that the surface
is darker owing to the presence of organics. Hard concretions (plinthites) and
layers (laterites) of iron concretions form in oxisols. Oxisols are found in the
warm and moist areas of the tropical moist climate zone.
Ultisols have a prominent E horizon and a B horizon in which clay minerals
accumulate and can form plinthite. These soils are low in plant nutrients.
The mean soil temperature for formation of ultisols is 8° C, and they are
associated with climates that produce an abundance of moisture in one sea-
son and dry conditions during another.
Vertisols have a high content of clay and shrink and swell depending on water
content. They have cracks and other evidence of soil movement. Soil hori-
zons may not be apparent. These soils have good nutrient availability but
are difficult to work for agriculture. Vertisols form under grassland and
savanna vegetation in the tropical and subtropical regions.
Alfisols have a gray to reddish horizon close to the surface that lack darkening
by humus and an underlying layer of clay accumulation. Alfisols have good
nutrient compositions for plants. Alfisols form under forests (Boralfs and
Udalfs) and mixed forest-grassland, shrub cover (Xeralfs and Ustallfs) at
sites ranging from the conifer forest in the snowy climate region to season-
ally dry sites in the midlatitudes and sub tropics.
Spodosols often have a bleached and light-colored E horizon that overlies a
dark layer of illuviated humus, aluminum, and iron in the B horizon. Spo-
dosols are generally acidic and low in nutrients. In many cases these soils
are relatively young, despite the clear development of horizons. They are
found under coniferous forests in the snowy climate zone.
Mollisols have a very thick and dark A horizon with a soft structure. Vertical
cracks form in the soils due to cycles of wetting and drying out. These soils
have high amounts of calcium and generally high nutrient content. Mol-
lisols are perhaps the most fertile soils in the world. They form under grass-
lands in the dry climate regions of the midlatitudes.
Aridisols are dry soils that have low humus content but clear horizons. They can
have high amounts of calcium and salt layers. They form in the arid climate
zone where water content in the soil is too low to support much plant growth.
Andisols are soils formed on fresh deposits of volcanic ash. They have a small
geographic distribution and can be very fertile.
The Physical Environment of Lakes
Lakes cover about 2% of the earth's surface and contain only about 0.01 % of
the world's water. However, lakes provide habitats for many types of plants,
36 CHAPTER 2 SOME BASICS

invertebrates, fish, amphibians, birds, and mammals and are an important focus
of biogeographic research. Lake basins can be formed by a number of natural
processes, including glacial activity, volcanic activity, tectonism, landslides, E
changes in river channels, and changes in ocean shorelines. Differences in light,
temperature, acidity, and the chemical composition of water produce different
physical environments in different lakes and within the same lake. These differ-
ences can have a profound impact on the geographic distributions of lake-
dwelling organisms.
Aquatic organisms are sometimes classified by their habitat. Plankton are
organisms that exist by passively floating in the waters of a lake or ocean. These
include many microscopic plants (phytoplankton) and small invertebrate animals
(zooplankton). Organisms such as fish that can propel themselves through the
water are referred to as nekton. Finally, benthic plants and animals are those that
exist on the bottom of lakes and oceans. The water environment is called the
pelagic zone, while the bottom is called the benthos.
Plants that live in lake waters require sunlight for photosynthesis. When
sunlight from the PhAR portion of the electromagnetic spectrum strikes the sur-
face of a lake, some of it is reflected directly back skyward and does not enter the
water. When light strikes the water at a 90° angle, only a small percentage of the
energy is reflected. However, at a 10° angle, about 40% of the light is reflected off
the lake waters. Once the light enters the water, its energy is dispersed by absorp-
tion and scattering. The transmission of PhAR is optimal in clear distilled water.
In such water, light in the blue wavelength is transmitted best. Longer wave-
length light, from the red portion of the spectrum, transmits poorly. About 53%
of the light is absorbed and emitted as heat in the first meter or so of the water.
However, lakes do not contain pure distilled water. Instead, their waters contain
suspended and solid organic and inorganic substances and organisms such as
plankton. The more matter and organisms in the lake water, the less light that can
be transmitted through the water. In small, productive lakes with high amounts of
suspended matter and organisms, the water _is murky, and little light is transmitted
below the first meter of water. In large, low-productivity lakes, such as Lake
Tahoe in California or Crater Lake in Oregon, significant amounts of light can be
transmitted tens of meters through the clear water. However, at greater depths,
most of the light that is transmitted is in the blue portion of the spectrum. The
portion of the lake water column that receives sufficient sunlight for photosyn-
thesis is called the euphotic zone. Depths beneath this that receive enough light
for fish and other organisms to see, but not enough for sustained photosynthesis,
are referred to as the disphotic zone. Depths that are totally dark are called the
aphotic zone. Photosynthetic plants can only live in the photic zone. Sometimes
limnologists classify parts of lakes according to their ability to support photosyn-
thetic plants. The shallow margins of the Jake where plants can root and receive
adequate light for photosynthesis is called the littoral zone, while the deep dark
portion where rooted plants cannot exist is called the profunda! zone (Fig. 2.10).
The impact of solar radiation coupled with warm air temperatures in the
summer often produces a thermal stratification where temperatures are higher in
the upper waters. Mixing of the surface waters by winds and currents generates
isothermal conditions in the upper few meters of the lake waters. Below this,
there is a gradient of decreasing temperature called a thermocline. At depth, the
lake waters are cool and isothermal (Fig. 2.10). The difference in temperature
between the surface and bottom waters can be as much as 20° C in relatively

, and are an important focus
ned by a number of natural
ivity, tectonism, landslides,
'relines. Differences in light,
1 of water produce different
the same lake. These differ-
lphic distributions of lake-
1 their habitat. Plankton are
~rs of a lake or ocean. These
d small invertebrate animals
'pel themselves through the
:sand animals are those that
r environment is called the
tt for photosynthesis. When
~tic spectrum strikes the sur-
ward and does not enter the
.ly a small percentage of the
:Yo of the light is reflected off
1ergy is dispersed by absorp-
imal in clear distilled water.
;mitted best. Longer wave-
ansmits poorly. About 53 %
·st meter or so of the water.
nstead, their waters contain
ces and organisms such as
water, the less light that can
:lakes with high amounts of
md little light is transmitted
ctivity lakes, such as Lake
:ant amounts of light can be
-Iowever, at greater depths,
>rtion of the spectrum. The
ient sunlight for photosyn-
s that receive enough light
r sustained photosynthesis,
totally dark are called the
be photic zone. Sometimes
bility to support photosyn-
>lants can root and receive
zone, while the deep dark
rJrofundal zone (Fig. 2.10).
m air temperatures in the
temperatures are higher in
tds and currents generates
e lake waters. Below this,
thermocline. At depth, the
difference in temperature
uch as 20° C in relatively
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 37
r-~~~~--~--------------------------~1 ------ro
Epilimnion '
Euphotic \
Metalimni;;-- ~... 2
·~ ........~.
\
en
Q) Disphotic \
c
0
N
E Hypolimnion j
Ol
:::; Aphotic ~
8
PC 02
FIGURE 2.10 Life zones, light, temperature, and oxygen distributions in a small lake
during summer.
deep lakes. If you have dived and swam in small northern lakes, you may have
experienced this thermal gradient as a feeling of increasing chill below a certain
depth. These three thermal zones are called the epilimnion, metalimnion, and
hypolimniom, respectively. The metalimnion produces a thermal barrier, and
there is little mixing of waters from the epilimnion with the hypolimnion. Among
other things, this phenomenon affects the distribution of oxygen. There are rela-
tively high amounts of dissolved oxygen in the epilimnion and low amounts in the
hypolimnion (Fig. 2.10). In some cases the bottom waters are anoxic and cannot
support organisms that require oxygen for respiration. This stratification of tem-
perature and oxygen breaks down seasonally in lakes located in cold regions and
allows oxygenation of bottom waters. Some lakes that are deep relative to their
surface area never experience oxygenation of bottom waters.
The degree of thermal stratification in lakes can be related to the relation-
ship between lake area and depth and climatic conditions-particularly air tem-
peratures. This provides a means of lake classification linked to climate:
Amictic Lakes never lose their cover of ice and do not stratify. They are found
only in the antarctic, a few places in the arctic, and on very high mountains.
Cold Monomictic Lakes never exceed 4° C water temperature and do not usu-
ally stratify. These lakes are found in the polar regions and on high moun-
tains.
Dimictic Lakes are the stratified type described in the preceding discussion.
They stratify in summer and mix in spring and fall.
Warm Monomictic Lakes never have water temperatures below 4° C. They
stratify directly in summer. They are found in warm temperate areas and
subtropical mountains.
Oligomictic Lakes have water much warmer than 4° C and irregular circula-
tion events. There may be little temperature difference between the epil-
imnion and hypolimnion. These lakes are found in tropical regions.
38 CHAPTER 2 SOME BASICS

Two categories of lakes can be found in both warm and cold climates. -
Polymictic Lakes are shallow and circulate continuously.
Meromictic Lakes are deep lakes in which the bottom waters remain unoxy-
genated, even during lake overturn.

Lakes can also be classified in terms of chemistry, nutrient status, and produc-
tivity. Many lakes have freshwater; however, some saline lakes contain salt concen-
trations that are as high or higher than that of the ocean. Such saline lakes,
generally develop in arid regions and basins that lack outlets. Water is lost from
these lakes mainly through evaporation, resulting in the accumulation of salts.
Deep, cold lakes, with high amounts of oxygen and low amounts of phosphorus and
other nutrients, typically have low primary productivity. These are referred to as
oligotrophic lakes. In contrast, shallow, warm lakes, with high amounts of nutrients
have high primary productivity. These are called eutrophic lakes. However,
eutrophic lakes generally have low amounts of oxygen due to high rates of respira-
tion and aerobic decomposition. In some cases, the oxygen levels can decrease to
the point where catastrophic die-offs occur for fish and other organisms.

The Physical Environment of Oceans

About 71% of the earth's surface is covered by ocean. This, coupled with the
depth of the ocean, provides an area of habitat for marine life that is 300 times
greater than the area provided by the terrestrial surface of the earth.
As is the case with freshwater, light does not penetrate far down in ocean
waters. In marine waters near the shore, sunlight may only reach the upper few
meters of the water column. In clear midocean sites, light can penetrate many
tens of meters, and 1% of the incident light may reach depths of 150 m. Photo-
synthetic plants such as microscopic phytoplankton, or large plants such as kelp,
can exist only in this upper photic zone. So, despite its great depth and volume,
the marine environment must depend on a thin surface layer for its primary pro-
ductivity.
As is the case with lakes, the ocean environment can be divided into pelagic
and benthic zones (Fig. 2.11). In addition, in the sea the pelagic environment is
subdivided into a nearshore neritic zone that extends to the edges of the conti-
nental shelves and a deepwater oceanic zone. The pelagic zone is also subdivided
by depth into five regions (Fig. 2.11 ). All photosynthesis takes place in the upper-
most epipelagic environment. Because the water is clearer in the midocean areas,
the photic zone is deeper there than it is near the continents. Light sufficient for =
photosynthesis may penetrate for well over 10 m in the mid-ocean. The benthos
of the ocean is also subdivided into a number of subzones extending from the
intertidal supralittoral to the very deep hadal. The substrate of the benthos is
very important in determining the organisms that can live there. Substrate varies
by depth and proximity to land. Shallow areas near the shore may have substrates
ranging from rocks, gravel, sand, and living coral. Areas in the deep sea away
from land and the continental shelves, often have substrates dominated by very
fine silt and clay sediments. E
A major difference between the marine environment and the freshwater :- l

environment of lakes and rivers is salinity. In general, the salinity of the ocean is
about 35 g of inorganic salts per 1 kg of sea water, the major salt being sodium

and cold climates.
nuously.
'ottom waters remain unoxy-
ry, nutrient status, and produc-
'iine lakes contain salt concen-
he ocean. Such saline lakes
ck outlets. Water is lost fro~
in the accumulation of salts.
w amounts of phosphorus and
vity. These are referred to as
rith high amounts of nutrients
I eutrophic lakes. However,
n due to high rates of respira-
)xygen levels can decrease to
td other organisms.
cean. This, coupled with the
marine life that is 300 times
·ace of the earth.
Jenetrate far down in ocean
ay only reach the upper few
~s, light can penetrate many
1ch depths of 150 m. Photo-
or large plants such as kelp,
its great depth and volume,
.ce layer for its primary pro-
t can be divided into pelagic
the pelagic environment is
s to the edges of the conti-
agic zone is also subdivided
sis takes place in the upper-
~arer in the midocean areas,
1tinents. Light sufficient for
he mid-ocean. The benthos
bzones extending from the
mbstrate of the benthos is
live there. Substrate varies
~shore may have substrates
reas in the deep sea away
,strates dominated by very
nment and the freshwater
the salinity of the ocean is
e major salt being sodium
PHYSICAL GEOGRAPHY AND THE FUNCTIONING OF THE EARTH 39
~--~_/'-~~--~~_/~~~~'-~~--~~-/'-~ 0
Littoral Oceanic 200
1000
2000 0
CD
~
3000 ~
Abyssal
2.
4000
Temp.°C
_ 1_ _ _ _
Salini~y
,_
Oxygen
__
-~:
0 20 33 35 0 300 5000
- .- .
Benthic
Hadal 6000
Marine life zones, salinity, temperature, and oxygen distibutions in the
FIGURE 2.11
ocean.
chloride: However, the concentration of salts varies with both latitude and depth.
In general, the salinity of surface waters is greatest in the regions between 20°
and 40° north and south latitude where evaporation rates on the ocean are high
relative to precipitation (Fig. 2.12). This corresponds with the dry climatic zone of
terrestrial deserts discussed previously. In addition, in the low and midlatitudes,
salinity decreases with depth between the surface and about 1000 m. Below 1000-
m depth, the salinity remains fairly constant.
The temperature of the ocean is an important control on planktonic and
shallow nektonic organisms. As we might suspect, sea surface temperatures
(SSTs) are highest in the low-latitude regions and decrease poleward (Fig. 2.12).
Water temperature also varies with depth, and where the surface waters are rela-
tively warm, a thermocline is usually found between the surface and 1000 m
depth. Deep waters are not prone to seasonal changes in temperature. Waters
below 2000 to 3000 m never rise above about 4° C, and in the deepest portions of
the ocean water temperatures can be below 3° C. In the deep ocean waters, there
is little temperature difference between the equator and the poles.
Oxygen in the oceans is highest in the surface waters and declines to mini-
mum values at 500 to 1000 m depth. In some cases, oxygen concentrations may
drop close to zero. The oxygen minimum zone is attributed to high rates of bio-
logical activity such as respiration and decomposition coupled with a lack of oxy-
gen replenishment by photosynthesis because of the absence of light at this
depth. Below the oxygen minimum zone, oxygen then increases with depth (Fig.
2.11). However, in some cases, such as off the coast of Santa Barbara, California,
the sea bottom may be permanently anoxic. In other cases, such as the Gulf of
Mexico, seasonally anoxic conditions may develop on the ocean floor.
The oceans display complex surface current systems that are largely driven
by winds and the impact of the Coriolis effect (Fig. 2.12). Such systems work to
bring warm waters poleward and moderate climates in areas such as northern
Europe and western Alaska. These currents are also an important means for dis-
persing plankton and aiding in the migration and dispersal of nektonic organ-
isms. In addition to surface currents, subsurface currents produce upwelling zones
where cool nutrient-rich waters rise from depth to provide zones of high oceanic
 EFER£N(

• Major upwelling zones

FIGURE 2.12 Ocean surface temperatures, salinity, currents, and major upwelling zones
(from a number of sources including Lalli and Parsons, 1997; Ross, 1992; Thurman, 1990).

40
 REFERENCES AND FURTHER READING 41

productivity (Fig. 2.12). Fisheries in areas such as coastal California owe their
high catches to such upwelling.
A final important physical factor controlling the distribution of life in the
oceans is pressure. Because the oceans are so deep, there are great differences
between the ambient pressure that is exerted on organisms at the surface and
pressure in the hadal zone. Pressure is determined by the weight of the overlying
water. Ocean pressure is measured in atm units (atmospheres). One atm equals
the average pressure of the atmosphere at sea level. The pressure of the ocean
increases by 1 atm for every 10 m of depth. So, in the deepest parts of the ocean,
at 11,000 m, the weight of water is equal to 1100 times the atmospheric pressure
we experience at sea level. These pressures present special challenges for organ -
isms that contain gas-filled sacks such as the swim bladders of fish or the lungs of
marine mammals. In addition, pressure makes the exploration or collection of the
biota of these depths extremely difficult. This is why the depths of the ocean rep-
resent one of the great unexplored frontiers of biogeography.

KEY WORDS AND TERMS

~rm Dicotyledon Photosynthesis
~mblage Domain Phyla
-.osphere Ecosystem Population
-..ecology Family Primary productivity
Aalotrophs Food web Respiration
OOlaSS Genus, genera Soil
Greenhouse effect Species
_ Olpbere G uild Stomata
Pathway Gymnosperm Synecology
Pathway Heterotrophs Systematics
Cllanosyn thesis Hydrosphere Taxon, Taxa; Taxonomy
Kingdom Trophic level
..Oer Lithosphere
..ununity Metapopulations
~acean acid metabolism Monocotyledon
CAM) photosynthesis Order

5iEFERENCES AND FURTHER READING

artman, 0 ., and Berry, J. {1973). High efficiency
-botosynthesis. Scientific American 229, 80--93.
--= _ E. M. (1970). World Soils. Cambridge
Jniversity Press, Cambridge.
lllpenter, S. R. (1988). Complex Interactions in
lAke Communities. Springer-Verlag, Berlin.
-ruttee of the Canadian Soil Survey (1978).
Canadian System of Soil Classification.
Major upwelling zones Department of Agriculture, Ottawa.
..aaft, J. (1983). Species concepts and specia-
ts, and major upwelling zones
- analysis. In Current Ornithology, vol. 1
Ross, 1992; Thurman, 1990).
(ed. R. F. Johnston), pp.159-187. Plenum
Press, New York.
DeAngelis, D. L. (1992). Dynamics of Nutrient
Cycling and Food Webs. Chapman and Hall,
New York.
Elton, C. (1927). Animal Ecology. Sidgwick and
Jackson, London.
Futuyma, D. J. (1979). E volutionary Biology. Sin-
auer Associates, Sunderland, MA .
Gage, J. G., and Tyler, P. A. (1991). Deep-Sea B iol-
ogy: A Natural H istory of Organisms at the
42 CHAPTER 2 SOME BASICS

Deep-Sea Floor. Cambridge University Press,

Cambridge.
Hutchinson, G. E. (1967). A Treatise on Limnol-
Odum, E. P., and Kuenzler, E. J. (1963). Experi-
mental isolation of food chains in an old-field
ecosystem with use of phosphorus-32. In
TER 3
ogy, vol. 2: Introduction to Lake Biology and Radioecology (ed. V. Schultzard and A. W.
the Limnoplankton. Wiley, New York.
Hutchinson, G. E. (1975). A Treatise on Limnol-
Klement), pp.113-120. Van Nostrand, Rein-.
hold, New York.
£ Ph
ogy. Wiley,NewYork. Pimm, S. L. (1982). Food Webs. Chapman and , ~JR(
Ingmanson, D. E., and Wallace, W. J. (1973). Hall, London.
Oceanology: An Introduction. Wadsworth, Polis, G. A., and Winemiller, K. 0. (1996). Food
Belmont, CA.
Kinne, 0. (1971). Marine Ecology: A Comprehen-
Webs: Integration of Patterns and Dynamics.
Chapman and Hall, New York.
STRIJ
sive, Integrated Treatise on Life in Oceans and Raven, P. H., Evert, F., and Eichhorn, S. E. (1999).
Coastal Waters. Wiley Interscience, New York. Biology of Plants, 6th edition. W. H. Freeman,
Krebs, C. J. (1994). Ecology, 4th edition. Harper- New York.
Collins College Publishers, New York. Raven, P. H., and Johnson, G. B. (1999). Biology, -::-::~;: an
Lalli, C., and Parsons, T. R. (1997). Biological 5th edition. McGraw-Hill, Boston. o!'ll""lTODII
Oceanography, 2nd edition. Pergamon Press, Ricklefs, R. E. (1990). Ecology, 3rd edition. W. H. - I
New York. Freeman, New York.
Larsen, C. P. S., and MacDonald, G. M. (1993). Ross, D. A. (1982). Introduction to Oceanogra- "' are
Lake morphometry, sediment mixing and the phy, 3rd edition. Prentice-Hall, Englewood
selection of sites for fine resolution palaeoe- Cliffs, NJ.
cological studies. Quaternary Science Review Service, R. S. (1997). Microbiologists explore life's
12,781-792. rich, hidden kingdoms. Science 275, 174~1742.
Lindemann, R. L. (1942). The trophic-dynamic Staff of the Soil Survey (1975). Soil Taxonomy: A
aspect of ecology. Ecology 23, 399-418. Basic System of Soil Classification for Making
Lipscomb, D. (1996). A survey of microbial diver- and Interpreting Soil Surveys. U.S. Depart- . ~ tern]
sity. Annals of the Missouri Botanical Garden ment of Agriculture Handbook 436. different
83,551-561. Strahler,A., and Strahler, A. S. (1997). Physical assurn
Maitland, P. S. (1990). Biology of Fresh Waters, Geography: Science and Systems of the _etermir
2nd edition. Chapman and Hall, New York. Human Environment. Wiley, New York.
-~ ortai
Mayr, E. (1942). Systematics and the Origin of Thurman, H. V. (1990). Essentials of Oceanogra-
Species. Columbia University Press (Reprint phy, 3rd edition. Merrill Publishing Co., ences in
edition: Dover Publications, New York, 1964), Columbus, OH. - ~ cies. ]
New York. Wetzel, R. G. (1983). Limnology, 2nd edition. -ontrolli
Nybakken, J. W. (1997). Marine Biology: An Eco- Saunders College Publishing, New York. - ·- th~
logical Approach, 4th edition. Addison Wesley Whittaker, R. H. (1975). Communities and Ecosys- ...: -ound
Longman, Menlo Park, CA. tems, 2nd edition. Macmillan, New York. c:ontrolli
Oil
mapped
_ aYera
~a• 1
-C F
:......_ distr
. ;: phys
_,_ the1
:.... :: disti
- -runen1
; - grap
-::ore tl:J
:-elated
meanJu
peratun
enzler, E. J. (1963). Experi-
Df food chains in an old-field
se of phosphorus-32. In
!. V. Schultzard and A. W.
CHAPTER
l- 120. Van Nostrand, Rein-.
THE PHYSICAL
'ood Webs. Chapman and
ENVIRONMENT AND THE
erniller, K. 0. (1996). Food
of Patterns and Dynamics.
tll, New York.
DISTRIBUTION OF LIFE
=<'. ,and Eichhorn, S. E. (1999).
6th edition. W. H. Freeman,
mson, G. B. (1999). Biology,
:aw-Hill, Boston.
). Ecology, 3rd edition. W. H.
Irk.
1troduction to Oceanogra-
>rentice-Hall, Englewood
.1icrobiologists explore life's
Dms. Science 275, 1740--1742.
ey (1975). Soil Taxonomy: A
Jil Classification for Making
oil Surveys. U.S. Depart-
re Handbook 436.
bier, A. S. (1997). Physical
:e and Systems of the
ent. Wiley, New York.
). Essentials of Oceanogra-
lerrill Publishing Co.,
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i). Communities and Ecosys-
,facmillan, New York.
3
There are certain plants and animals that we instinctively associate with specific
environments. For example, we link elephants and palm trees with hot tropical
regions. In contrast, we picture polar bears in cold arctic habitats. These associa-
tions are based on the real geographic distributions of these species. We don't
find palms and elephants living in the arctic or polar bears in the tropical
savanna. Why not? For one reason, plant and animal species require certain phys-
ical conditions related to temperature, moisture, chemistry, and light in order to
survive. Exposure to temperatures that are too high or too low, or to conditions
with too much or too little moisture, can prove lethal. As we have seen in Chap-
ter 2, the physical environment of the earth varies greatly according to location.
The temperature regime experienced by organisms dwelling in the tropics is very
different from the conditions experienced by organisms in the arctic. It is logical
to assume that the differences in the physical environment must have a role in
determining where certain species can be found and where they do not exist. An
important goal for biogeographers is to understand how these geographic differ-
ences in the physical environment influence the geographic distributions of
species. In this chapter we will investigate the role of the physical environment in
controlling the geographic distributions of species. We should, however, keep in
mind that physical conditions are just one factor in determining where a species
is found and where it is absent. Both biological factors and history play a role in
controlling geographic distributions.
One way of embarking on our current investigation is simply to compare the
mapped distributions of species with mapped distributions of physical factors such
as average July temperatures. We might discover that the white spruce (Picea
glauca) is not found in regions where average July temperatures are cooler than
10° C (Fig. 3.1). On this basis, we could speculate that July temperature controls
the distribution of spruce trees. However, unless we have a good understanding of
the physiological impact of temperature on plants and animals, we cannot be cer-
tain there really is a cause and effect relationship between July temperature and
the distribution of spruce. The similarity in the distribution of the physical envi-
ronmental variable and the species could simply be a coincidence. In many cases, a
geographic correlation may exist between the range limits of the species and far
more than one environmental variable. This is particularly true for variables
related to climate. Quite often, there is a high correlation between factors such as
mean July temperature, maximum daily summer temperature, mean annual tem-
perature, potential evaporation rates, and so on. In one example, it was shown that
43
44 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

exarr
physi
situ a
grapl
grapl
descr
and t

UGHT
Ligh
for p
tion
eartt
in ten
unde
their
grow
oblig
cans
ares
into '
the t
faun
FIGURE 3.1 A stunted white spruce (Picea glauca) at the northern treeline in the
Northwest Territories of Canada. Although such plants are often less than 1 m tall, they can shap·
be over 100 years old. A combination of low summer temperatures, short summer growing
read
season, and damage by blowing snow in winter determines the northern range limits of
tosyr
spruce at the northern treeline. Summer temperature and growing season length appear to
be the most important factors, and the northern limit of spruce in Canada occurs wh ere
syntl
average July temperatures lie between 10° and 12.5° C. light
res pi
on p
its s<
the eastern limits of holly (/lex aquifolium) in Europe corresponded with the geo-
graphic distribution of at least 23 different climatic variables. Many of these
aspects of climate may not, in fact, play any role in controlling the geographic dis-
tribution of the species. As we will see later, the geographic distributions of some
species may also be influenced by biological factors such as interactions with other
species. Finally, the present geographic distributions of some species may actually
be in a state of change as the species disperse into new territory. Consequently,
biogeographers cannot be content with merely identifying correlations between
the spatial distributions of species and the spatial distributions of environmental
variables. Biogeographers must base their analysis of these geographic distribu-
R
tions on an understanding of the impact of the physical environment on the phys- phc
iological functioning of plants and animals. R
In this chapter we will examine how physical conditions such as light, tem- phc
perature, moisture, and chemistry impact the physiological functioning of plants
and animals. We will also look at some of the remarkable adaptations that allow FIGl
organisms to cope with stresses produced by the physical environment. Some phot
>F LIFE
LIGHT
rthern treeline in the
n less than 1 m tall, they can
:ures, short summer growing
e northern range limits of
ving season length appear to
in Ca nada occurs where
)ITesponded with the geo-
variables. Many of these
rolling the geographic dis-
phic distributions of some
as interactions with other
;orne species may actually
N territory. Consequently,
1ing correlations between
butions of environmental
hese geographic distribu-
environment on the phys-
jitions such as light, tem-
ical functioning of plants
1le adaptations that allow
sical environment. Some
LIGHT 45
examples of close links between the geographic distributions of species and the
physical environment will be presented. We will then examine some examples of
situations where two or more physical controls act together to determine the geo-
graphic distributions of plants and animals. Finally, we will consider how biogeo-
graphers and ecologists have developed conceptual frameworks for organizing,
describing, and analyzing the relationships between the physical environment
and the distribution and abundance of organisms.
Light in the PhAR portion of the electromagnetic spectrum provides the energy
for photosynthesis. The amount of available light is a key control on the distribu-
tion of plants. Although the amount of sunlight received at the surface of the
earth varies with latitude, most research on the impact of differences in light
intensity on plants has focused on local conditions such as the distribution of light
under forest canopies. Plants may be classified into two categories with respect to
their light requirements. Heliophytes grow best in full sunlight, and sciophytes
grow best in shade. Plants that can only grow on sunny sites are referred to as
obligate heliophytes and are also said to be shade intolerant. Heliophytes that
can survive on shaded sites are referred to as facultative heliophytes. These plants
are shade tolerant but do best in full sunlight. Sciophytes can similarly be divided
into obligative and facultative categories. Most house plants are sciophytes from
the tropics and subtropics. This allows them to live under the low light conditions
found indoors.
The response of photosynthesis to light generally has a roughly hyperbolic
shape (Fig. 3.2). Photosynthesis increases as light intensity increases until it
reaches a maximum.After that, increasing light causes declines in the rate of pho-
tosynthesis. At low illuminations, there is a linear increase in the rate of photo-
synthesis in response to increasing fluxes of radiant energy. The point at which
light intensity and photosynthesis are enough to offset the loss of energy through
respiration is called the comp ensation point. The influence of increased radiation
on photosynthesis declines at higher fluxes of radiant energy as the leaf reaches
its saturation point for light relative to its capacity to absorb and use light for
(/)
'iii
Q)
~
c
~
0
0
~
c.
0
Q)
<a
a:
Respiration <
photosynthesis
Respiration >
photosynthesis
FIGURE 3.2 A generalized model of th e relationship between light intensity and rate of
photosynthesis.
46 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

photosynthesis. For many plants, maximum net photosynthesis occurs when light tha
intensities reach one-third to two-thirds the strength of full sunlight. However, pla
there are large differences in the light intensities at which plants reach saturation. to c
Herbs found on forest floors and shade-dwelling mosses often experience satura- wh
tion at light intensities as low as 5% full sunlight or less. Some shade tolerant ple
trees such as red maple (Acer rubrum) and American beech (Fagus grandifolia) a til
may also reach light saturation at levels as low as 5% to 10% of full sunlight. The
low light saturation level of sciophytes allows them to survive on very shady sites pia
where other plants would not be able to conduct enough photosynthesis to bal- alrr
ance energy lost through respiration. In fact, when most true sciophytes are the
exposed to full sunlight, they cannot conduct photosynthesis at nearly the same aru
rate as heliophytes. High light intensities can actually cause decreased rates of wit
photosynthesis in some shade tolerant plants. This phenomenon, called photoin- flm
hibition, can be due to various factors including shrinking of chloroplasts, loss of can
chlorophyll, and loss of C02 due to photooxidation. Photooxidation occurs only can
in C3 plants and produces a condition called photorespiration, where C02 is mo
released without the production of chemical energy for the plant. The rate of zie,
photorespiration is positively correlated with light intensity and temperature. ligt
Since C4 plants do not suffer from photorespiration, they have higher rates of unl
photosynthesis at high light intensities and temperatures. However, C4 plants are of ~
less efficient at photosynthesis under low light intensities and temperatures than 92
C3 plants. Indeed, C4 plants are generally obligate heliophytes and are restricted
geographically to areas of high insolation and heat. cha
Aquatic vascular plants and algae are adapted to the low light intensities fall
of their watery environment. They only need low levels of light ( -1%- 3% of full the
sunlight) to produce rates of photosynthesis that equal energy losses due to res- en<
piration. Many aquatic plants and planktonic algae reach the saturation point at ver
5% of full sunlight. Despite this relatively low saturation point, there is not ear
enough light to support photosynthetic plants beyond the relatively thin upper Se[
zone of the oceans. In general, maximum photosynthetic activity occurs in the wh
top 20 to 100m of the ocean. The absence of light at depth is one of the strongest
controls on the vertical distribution of photosynthetic organisms in the ocean a f
and lakes. (Pi
In addition to physiological adaptations, certain morphological (physical) cor
adaptations take place in plants that make them uniquely suited to living in of
either high or low light intensities. For example, the leaves of plants adapted to an<
live on sites with high light intensity tend to be small and thick and possess a thick tra
cuticle (outer covering). In addition, the leaves of some high light intensity plants rul
are slightly curled and possess a reflective surface layer. These plants also tend to the
have a high number of stomata, which often are found on both the upper and unc
lower surface of the leaf. The stomata allow for high rates of C02 intake from the tie1
atmosphere while cooling the leaf by transpiring water vapor. Leaves growing
under conditions of very high light conditions, such as the manzanita bushes
(Arctostaphylos patula) of the Southwest are often oriented so that they do not TEMPERA
receive high rates of insolation. In contrast, the leaves of plants living in low light
intensity tend to be large and soft and have high amounts of chlorophyll. The Te1
leaves of low-insolation plants also live longer than those on plants adapted to ha1
high light intensity. Leaf adaptation to light is not restricted to land plants. For tht
example, species of the Caribbean seaweed genus Sargassum that live in deeper de.
waters with low light intensities have wider leaflike blades to capture more light gn
lf LIFE
;ynthesis occurs when light
of full sunlight. However,
ich plants reach saturation.
es often experience satura-
less. Some shade tolerant
beech (Fagus grandifolia)
:o 10% of full sunlight. The
survive on very shady sites
ugh photosynthesis to hal-
most true sciophytes are
·nthesis at nearly the same
if cause decreased rates of
:!nomenon, called photoin-
;:ing of chloroplasts, loss of
'hotooxidation occurs only
respiration, where C02 is
for the plant. The rate of
ntensity and temperature.
they have higher rates of
·es. However, C4 plants are
:ies and temperatures than
ophytes and are restricted
to the low light intensities
s of light ( -1%- 3% of full
1l energy losses due to res-
~ch the saturation point at
ration point, there is not
l the relatively thin upper
1etic activity occurs in the
:pth is one of the strongest
c organisms in the ocean
morphological (physical)
liquely suited to living in
~aves of plants adapted to
d thick and possess a thick
: high light intensity plants
·.These plants also tend to
td on both the upper and
es of C02 intake from the
er vapor. Leaves growing
as the manzanita bushes
ented so that they do not
>f plants living in low light
ounts of chlorophyll. The
Lose on plants adapted to
tricted to land plants. For
~assum that live in deeper
1des to capture more light
47
TEMPERATURE
than species that grow in shallower waters where light intensity is high. Some
plants have heliotropic leaves and flowers that change their angle during the day
to either increase or reduce insolation. The shifting flower heads of sunflowers,
which seem to turn and follow the sun during the course of the day, are one exam-
ple. The changing position of heliotropic leaves or flowers are caused by light cre-
ating differential rates of cell growth on stems.
In addition to morphology, the life history and reproductive behavior of
plants can sometimes be related to light conditions. Shade tolerant plants are
almost always perennials that live for more than one year. The rate of photosyn-
thesis in shady spots is not enough to support the high-energy requirements of
annual plants which must germinate from seed, grow, flower, and produce seed
within a single year. Plants often require high light intensities in order to produce
flowers. In many temperate and northern forest stands, only the trees in the
canopy are able to flower profusely, for the low light intensities beneath the
canopy restrict flowering of understory trees. The production of cones by com-
mon North American coniferous trees such as Douglas fir (Pseudotsuga men-
ziesii) and white spruce (Picea glauca) has been shown to be promoted by high
light intensity. In addition, the seeds of many plant species will not germinate
unless exposed to relatively high light intensities. For example, only about 3-4%
of Virginia pine (Pinus virginiana) seeds will germinate in dark conditions, while
92 to 94% of the seeds germinate when exposed to light.
The duration of daylight often serves as a stimulus for different seasonal
changes in plant growth, such as flowering in spring or shedding of leaves in the
fall. This relationship between phenology (the timing of changes in the growth of
the plant) and light is called photoperiodism. A familiar example of this phenom-
enon in eastern North America is the late persistence in the fall of leaves on sil-
ver maple trees (Acer sacharinum) planted near streetlights compared to the
earlier shedding of leaves from trees planted away from artificial light sources.
Seasonal photoperiodism becomes more important at the mid- and high latitudes
where there are pronounced changes in day length during the course of the year.
A classic example of the impact of light requirements on the distribution of
a plant comes from the forests of southeastern North America. Loblolly pine
(Pinus taeda) is found growing from eastern Texas to southern New Jersey. It is a
common tree and an important timber species. Anyone familiar with the woods
of the Southeast will know that the pine seedlings are numerous in open areas
and at the edge of stands, and occur less frequently within closed forests. In con-
trast, the seedlings of some deciduous trees such as eastern red oak (Quercus
rubra) can be found within the forest. Loblolly pine has lower rates of photosyn-
thesis than oak at all light intensities, but this difference is most pronounced
under the low to moderate intensities typical of forested sites. Low light intensi-
ties under the forest canopy restrict reproduction by loblolly pine to sunny sites.
TEMPERATURE
Temperature exerts a strong physiological impact on both plants and animals. It
has been said that "with regard to life on earth, temperature is-next to light-
the most potent environmental component" (Kinne, 1970). Accordingly, a great
deal of research has been conducted on the impact of temperature on the geo-
graphical distribution of species. Since the physiologies of plants and animals are
48 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

so vastly different, it makes sense to divide our discussion of temperature into

separate sections for plants and animals.

Plants
For plants, temperature can limit geographic distribution by causing metabolic
difficulties and physical damage. On the whole, plants are poikilotherms, which
means that they assume the temperature of their environment. It should not be
assumed, however, that the temperature of a plant is the same as the air temper-
ature. Because of the transpiration of water, leaves can be· as much as 15° C
cooler than air temperature, while portions of plants that intercept high amounts
of sunlight may be 30° C warmer than the air. There are three amazing plants that
have been shown to regulate their temperature biochemically. They are philoden-
dron (Philodendron selloum), skunk cabbage (Symplocarpus foetidus), and
sacred lotus (Nelumbo nucifera). These plants can raise the tissue temperatures
of their flowers up to 34° C above the air temperature for periods ranging from
FIGl
18 hours to two weeks. The biochemical generation of this heat is not yet com- Larc
pletely understood.lt is thought that the heating of the floral parts may aid in the
process of attracting and keeping pollen-carrying insects in contact with fertile
flowers. to lc
Plants depend on photosynthesis for food production. In turn, the rate of c. s
photosynthesis is dependent on temperature. If the temperature is too high or atur
low for effective photosynthesis, the plant essentially starves to death. In addition cole
to starving the plant, low rates of photosynthesis restrict the rate of growth and diet
reproductive functions. There is wide variability among different plant species in but
the optimum temperature for photosynthesis. Some arctic and alpine plants can func
conduct photosynthesis at temperatures slightly above freezing, while tropical susc
plants often require temperatures greater than 15 to 20° C for efficient photosyn-
thesis. In general, tropical plants reach peak rates of photosynthesis at higher
temperatures than plants from cooler regions (Fig. 3.3).
Low temperatures cause slow metabolic activity due to the kinetics of
enzymes such as rubisco and depress the rate of photosynthesis. The circumarc-
tic treeline of the northern hemisphere is a classic example of how low tempera-
tures during the growing season appear to be responsible for determining the
geographic range limits of plant species, particularly coniferous trees. The north-
·-!!::
ern limit of trees such as white and black spruce (Picea glauca and Picea mari-
ana) in North America, and Scots pine (Pinus sylvatica), Norway spruce (Picea
abies), Siberian spruce (Picea obovata), Siberian larch (Larix sibirica), and
Dahurian larch (Larix dahurica) in northern Eurasia corresponds to the 10°
C- 12.SO C July temperature isotherm (Fig. 3.4). It has long been proposed that
north of this region the summer temperatures are too low and the growing sea-
son too short to support the growth of trees. It has been demonstrated that the
rate of photosynthesis for trees such as black spruce declines rapidly at temper-
atures less than 10° C. In addition, the low temperatures during the summer
appear to be insufficient for the development of fertile pollen and seeds. Finally,
the low rates of photosynthesis and growth may make the trees more susceptible
to damage due to desiccation and mechanical damage by blowing snow and ice
during the winter.
Cold temperatures can physically damage plants through chilling stress and FIG I
freezing stress. Plants do not have to actually freeze to suffer severe damage due te rn
~OF LIFE TEMPERATURE 49

;cussion of temperature into Temperate Trees Tropical Trees

E
:I 75
.!!! E
Jution by causing metabolic "'
Ql ·-
X
.<:::: Ol
1ts are poikilotherms, which c>--oE
50
tvironment. It should not be >
0"' Ql
0 Q;
; the same as the air temper- .<::::"'
O...o
:s can be· as much as 15° C 0
~ 25
~
that intercept high amounts
1re three amazing plants that
temically. They are philoden-
0
ymp locarpus foetidus), and -10 0 10 20 30 40 50
aise the tissue temperatures Temperature (°C)
Ire for periods ranging from
FIGURE 3.3 The relationship between temperature and rate of photosynthesis (after
of this heat is not yet com-
Larcher, 1969; Kramer and Kozlowski, 1979).
he floral parts may aid in the
tsects in contact with fertile
to low temperatures. Chilling stress takes place at temperatures that are above oo
duction. In turn, the rate of
C. Some tropical plants have been shown to experience chilling stress at temp~r­
temperature is too high or atures as high as 20° to 15° C. Chilling can cause reduced plant growth, leaf dis-
starves to death. In addition
coloration, curling, crinkling and lobing, surface lesions on fruit, splitting and
;trict the rate of growth and
dieback of stems, and death. In some tropical plants, temperatures that are cool
>ng different plant species in
but not cold enough to damage leaves and other tissue may disrupt reproductive
arctic and alpine plants can
functions, such as the formation of pollen grains. Young plants are often more
ove freezing, while tropical
susceptible to chilling damage. In addition, plants are more prone to chilling
zoo C for efficient photosyn-
of photosynthesis at higher
3).
vity due to the kinetics of
•tosynthesis. The circumarc-
ample of how low tempera-
msible for determining the
coniferous trees. The north-
icea glauca and Picea mari-
'ica), Norway spruce (Picea
larch (Larix sibirica), and
sia corresponds to the 10°
as long been proposed that
o low and the growing sea-
'een demonstrated that the --...10 Mean July 1

declines rapidly at temper- temperature

(oC)
·atures during the summer
le pollen and seeds. Finally, D Distribution
of white spruce
~ the trees more susceptible (Picea glauca)
~e by blowing snow and ice in Canada

; through chilling stress and FIGURE 3.4 The relationship between the northern limits of spruce and July
o suffer severe damage due temperatures in Canada.
50 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

stress when they are not dormant and during the daylight hours when they are rna
most physiologically active. The primary causes of chilling injury are a matter of rna
debate. Cool temperatures can cause damage to plant membranes, decreased alp
rates of nucleic acid and protein synthesis, and decreased photosynthesis. ice
It is not just tropical land plants that are susceptible to damage or death due pe1
to chilling stress. Tropical seaweeds are extremely sensitive to water temperature, tha
and the geographic distributions of certain species can be linked to this. For ha1
example, the tropical seaweeds Botryocladia spinulifera and Haloplegma duer- ph~
reyi have northern limits that coincide with the 20° C sea surface isotherm. It has
been shown that both these species suffer severe damage when exposed to water dm
temperatures of 18° C and die after two weeks in water that has a temperature of SUf

15° C. A clear linkage has been found between their inability to survive cool pre
waters and the relationship between their distribution and the 20° C isotherm. car
Freezing stress occurs at temperatures below ooC. Tropical plants that are cac
susceptible to chill stress usually die when temperatures drop below freezing. In we:
contrast, some arctic plants can withstand temperatures as low as -70° C without ·'st,
damage. Aside from causing death, freezing can result in depressed growth rates, tha
discoloration of leaves, fissuring of woody stems, malformed leaves and flowers, tivt
dieback of roots, stems, leaves, and flowers, and delays in sprouting and flowering. Ca.
Damage to plants from freezing is caused primarily by the intracellular freezing ifc
of water. When ice crystals form inside cells, they cause ruptures to the cell wall, S01
and this is almost always lethal to the cell. The formation of extracellular ice can ten
also occur in plant cells. As this process draws water out of cells, it can cause cel- fou
lular damage due to dehydration and shrinkage of the cell walls. However, extra- shr
cellular ice formation is not always harmful to the cells or plant tissue of many kef
temperate and arctic plants as long as the freezing occurs in the dormant period. it e
There are many interesting adaptations that allow plants to survive in cool fro
environments. One obvious strategy is dormancy during the cold season. Most of to:
the common tree genera in the forests of northeastern North America, western mo
Europe, and eastern Asia, such as the maples, oaks, beech, birches, and ashes, are a pi
deciduous trees that lose their frost-sensitive leaves during the cold winter sea- is I•
son. In most of these genera, the leaves suffer damage at temperatures of freezing tus
or just below. The new leaves arise in the spring from winter buds that can remain cac
viable at colder temperatures. wit
Most of the needle-leaved conifers of the northern and alpine forests, such
as pines, spruces, and firs, do not lose their leaves during the winter. How do such Wll

evergreen plants escape intracellular freezing and tissue destruction when tem- ser
peratures may drop to -40° C or colder? In these plants, the onset of cool tem- reg
peratures causes physiological changes that allow plant tissue to either avoid of
freezing or restrict freezing to extracellular areas. For plants to avoid freezing, (A
they must chemically alter their liquids into a form that is analogous to antifreeze 3.6
in automobiles. The liquids in these plants can be cooled far below ooC and will of
not freeze. This process is called supercooling and is achieved by the metabolic COl

synthesis of sugars and other molecules which, when in solution in the plant's Flc
tissue, lower the temperature for ice formation to far below ooC. Supercooling res
seems to be the prevalent mechanism of frost resistance in herbs. For woody ha·
plants, supercooling is augmented by declines of cellular water content, greater spf
cellular accommodation to deformation, and processes that allow water to accu- be1
mulate and freeze in extracellular spaces. The loss of water from the cells to res
extracellular areas increases the solute content of the remaining cell water, me
j OF LIFE
aylight hours when they are
hilling injury are a matter of
))ant membranes, decreased
ased photosynthesis.
tible to damage or death due
1sitive to water temperature,
; can be linked to this. For
ifera and Haloplegma duer-
~ sea surface isotherm. It has
nage when exposed to water
:er that has a temperature of
eir inability to survive cool
n and the 20° C isotherm.
1° C. Tropical plants that are
ures drop below freezing. In
res as low as -70° C without
lt in depressed growth rates,
ilformed leaves and flowers,
s in sprouting and flowering.
by the intracellular freezing
1se ruptures to the cell wall,
ttion of extracellular ice can
out of cells, it can cause cel-
e cell walls. However, extra-
ells or plant tissue of many
curs in the dormant period.
ow plants to survive in cool
ing the cold season. Most of
:rn North America, western
eech, birches, and ashes, are
during the cold winter sea-
at temperatures of freezing
winter buds that can remain
ern and alpine forests, such
ng the winter. How do such
;sue destruction when tern-
ants, the onset of cool tem-
llant tissue to either avoid
)r plants to avoid freezing,
it is analogous to antifreeze
1led far below 0° C and will
achieved by the metabolic
11 in solution in the plant's
r below oo C. Supercooling
tance in herbs. For woody
ular water content, greater
!S that allow water to accu-
of water from the cells to
the remaining cell water,
TEMPERATURE 51
making it more resistant to freezing. The cell walls can accommodate the defor-
mations caused by water freezing on the exterior of the cell. For northern and
alpine evergreens such as pines and spruces, both supercooling and extracellular
ice formation play a part in allowing the plants to withstand extremely cold tem-
peratures. One interesting facet of these physiological adaptations to freezing is
that most of these plants will still be damaged by cold temperatures if they do not
have a period of cooling in which to adjust to the onset of winter. This process of
physiological preparation for the onset of winter cold is called frost hardening.
Some members of the cactus family ( Cactaceae) appear to resist freezing
during cool nights by radiating heat stored during the day in their thick, moist tis-
sue. The greater the mass of the cactus, the more heat it can store and the less
prone it will be to freezing damage during the night. How is it then that these cacti
can survive cold temperatures when they are young and small? The giant saguaro
cactus (Carnegiea gigantea) is perhaps the most well-known symbol of the south-
western desert (Fig. 3.5). In the popular lore of North America, the distinctive
"stovepipe" shape of the multistemmed saguaro is a universally recognized icon
that is used to represent deserts in movies, television, and comics. Yet, this distinc-
tive and widely recognized plant is actually only found in the Sonoran Desert of
California, Arizona, and adjacent Mexico. The saguaro cactus is damaged or killed
if exposed to prolonged freezing temperatures. Desert climates in the northern
Sonoran Desert are typified by warm days but sometimes experience nighttime
temperatures that are below freezing in the winter. Young saguaro that survive are
found sheltered beneath more frost-tolerant desert shrubs. The cover of these
shrubs acts as a thermal blanket, capturing heat radiated from the ground and
keeping the microclimate of the small saguaro warm at night. As the cactus grows,
it eventually rises above the cover of the protective shrub. The radiation of heat
from the stalk of the large mature cactus prevents freezing. This strategy works up
to a point. The range of the saguaro is restricted to areas that do not experience
more than about 12 to 24 continuous hours of air temperatures below oa C. It
appears that after 24 hours of freezing air temperatures, not enough heat reserve
is left in the saguaro to keep the tissue from freezing. Not all members of the cac-
tus family are limited to regions with relatively mild winters. A small prickly pear
cactus (Opuntia fragilis) grows as far north as central Alberta in Canada and can
withstand temperatures as low as -50° C.
The broad geographic ranges of many plants can in part be explained by
winter temperatures. For instance, most tropical and subtropical plant species are
sensitive to temperatures at or below freezing and are restricted to low-latitude
regions where air temperatures remain warm throughout the year. One example
of this comes from a very familiar plant-the palm. Species of the palm family
(Arecaceae) are found throughout the world in the tropics and subtropics (Fig.
3.6). Despite this wide longitudinal distribution, they rarely grow north or south
of the sub tropics. The cabbage palmetto (Saba! palmetto) is common in the
coastal regions of the southeastern United States from North Carolina to
Florida. A careful analysis of the distribution of palms shows that they are
restricted to areas where temperatures never drop below -10° C to -15° C. Palms
have only one growing point, and if it is damaged, the palm fails to grow. All palm
species that have been tested experience freezing damage at temperatures
between -3° C and -13.5° C. This incapacity to survive freezing temperatures
restricts their geographic range to warm regions. Some organs on the palms are
more prone to freezing damage than others. Leaves and the inner stems of
52 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

FIGUI
parts <
Winte
and st

conif,
rons)
cold
FIGURE 3.5 Saguaro cacti (Carnegiea gigantea) growing in the Organ Pipe Cactus Oreg
National Monument of Arizona. Water stored in stems and CAM photosynthesis help these buds
large succulents to survive the aridity of the Sonoran Desert. The saguaro cannot survive dron
more than 12 to 24 hours of air t emperatures that are below freezing, and this restricts t heir the s1
northern distribution to areas such as southern Arizona and adjacent Mexico. tures
big-If
wintE
mature plants appear to be the most resistant, while reproductive organs are the furth
least resistant (Fig. 3.6). For example, the fronds of some adult palms can survive trees
temperatures of less than - 10° C, while the flowers are killed by temperatures of
- 1o C. Thus, mature palms can be planted and survive in such unlikely places as cally
the mild coastal areas of the British Isles or the Pacific Northwest of North read
America but are unable to reproduce in such regions. plant
Many plants in both the northern and southern hemispheres have geo- tosyn
graphic limits that are controlled by winter temperatures. Some well-known parti<
I OF LIFE
n the Organ Pipe Cactus
<\M photosynthesis help these
The saguaro cannot survive
freezing, and this restricts their
1djacent Mexico.
·eproductive organs are the
me adult palms can survive
e killed by temperatures of
e in such unlikely places as
'acific Northwest of North
rn hemispheres have geo-
ratures. Some well-known
TEMPERATURE 53
Trachycarpus fortunei
Lethal Temp. Lethal Temp.
(DC) (DC)
-1 .0~ -10.5
- 1.0 ~ t5 \< ; ' - - - --10.0
-10.0 _r--'i<;--- - - - 6 .0
-2.0~
v---'1<--.,.--- -11.5
- 1.0_____. 9
• Global distribution of palms
- - Annual monthly minimum temperature (0C)
FIGURE 3.6 The natural global distribution of palms and the susceptibility of different
parts ofthe palm plant Trachycarpus fortuneito cold temperatures (after Larcher and
Winter, 1981 and Sakai and Larcher, 1987). The relative resistance to freezing by the fronds
and stems, and sensitivity to freezing by the flowers are typical of palms.
coniferous trees, including the coastal redwood of California (Sequoia sempervi-
rons), suffer frost damage at temperatures of -15° C to -25° C. This intolerance to
cold appears to keep the coastal redwood from growing north of southernmost
Oregon or at higher elevations in the coastal ranges of California. Similarly, the
buds of most midlatitude deciduous tree species such as the tulip tree (Liroden-
dron tulipifera) of North America, the gingko tree of China (Gingko biloba), or
the southern beech (Nothofagus antarctica) of Chile suffer damage at tempera-
tures below -25° C to -35° C. There are exceptions. For example, the buds of the
big-leafed oak (Quercus macrocarpa) of eastern North America can withstand
winter temperatures of less than -60° C. Not suprisingly, this tree is distributed
further north in eastern North America than many other temperate deciduous
trees such as the tulip tree.
Most plants also suffer from declining rates of photosynthesis once criti-
cally high temperatures are reached. The temperature at which plant species
reach maximum photosynthetic rates often depends on the latitude at which the
plant species is found. For some arctic, alpine, and temperate plants, rates of pho-
tosynthesis decline with temperatures above 10° C. Tropical and desert plants,
particularly those utilizing the C4 photosynthetic pathway, may not attain peak
54 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

rates of photosynthesis until temperatures reach 30° C. The decline in photosyn- suc.IJ
thetic gains at high temperatures is due to many factors. Respiration rates o<i'
increase more rapidly than photosynthesis at high temperatures. Some enzymes,
critical to photosynthesis, are intolerant of high temperatures and break down. prol
Cell structure may be altered, and cell membranes may become thin and leak at and
high temperatures. Finally, high temperatures can cause stomatal closure and havf
inhibit the exchange of C02, water, and oxygen needed for photosynthesis. For peri<
many midlatitude and arctic plant species, heat injury and death occur at temper- .:am
atures of 30-40° C. Tissue temperatures greater than 60-70° C are lethal to even abo1
the most heat-tolerant plant, although cacti can withstand such heat for short deer
periods of time. Tern
It has been suggested that the southern range limits of many arctic plants
might be caused by the depletion of carbohydrate reserves due to high respira- pera
tion rates generated by warm temperatures. In order for plants to survive the low rure
temperatures and short duration of the arctic growing season, they must be able
to grow rapidly in the spring using stored reserves of carbohydrates. When sub- tern]
jected to long and warm growing seasons, however, these plants will deplete all inert
reserves of carbohydrates over the summer and die. For example, the arctic inert
shoreline plant called Scots lovage (Ligusticum scoticum) reaches its southern relat
limits on the Scottish and Irish coastlines where mean July temperatures reach totic
about 14.4° C. Comparative physiological studies show that the respiration rate pera
of Scots lovage is much higher than that of species of shoreline plants found to prod
the south. This difference in respiration rates increases as temperature climbs. dow
Scots lovage can lose 50% of its carbohydrate reserves in just 17 days if tempera- mati
tures remain at 25° C or higher. may
Adaptations to withstand heat include features that decrease leaf tempera- in tal
ture such as the presence of reflective coatings and small hairs on leaf surfaces. dest1
Leaves may also be oriented vertically to reduce insolation or be heliotropic. ternI
Where water is plentiful, leaves may cool down through high rates of transpira- ingi
tion. When seasonal conditions are dry and hot, some plants lose their leaves dur-
ing those times of the year. An interesting adaptation to high temperatures in dry is ge
regions is the development of dimorphic leafing patterns. In hot and dry periods, mals
these plants are typified by small leaves, while in the cool season, they support man:
larger leaves. A number of plants from the deserts of the southwestern United low.
States and adjacent Mexico, such as honey mesquite (Prosopsis glandulosa), have Thill
dimorphic leaves. arou
notl

Animals bod)
Like plants, the so-called cold-blooded animals such as fish, reptiles, amphibians, heat
and insects are poikilotherms. Their body temperature generally reflects the shoVI
ambient temperature of the environment. In contrast, birds and mammals are ex an
homeotherms that maintain a relatively steady body temperature through the dese
metabolic generation of heat. Animals that are able to metabolically produce thro1
heat are also known as endotherms, while cold-blooded animals are referred to bod)
as ectotherms. Both homeotherm and poikilotherm animals can be damaged and unbt
killed by exposure to high and low temperatures. When they are in an active the r
state, insects, fish, reptiles, amphibians, birds, and mammals must keep their body Smai
temperatures within a relatively narrow temperature range. The lowest body area.
temperatures for active animals are found in fish from the polar regions. Species cool
I OF LIFE
C. The decline in photosyn-
'{ factors. Respiration rates
~mperatures. Some enzymes,
1peratures and break down.
rray become thin and leak at
cause stomatal closure and
~ded for photosynthesis. For
rand death occur at temper-
. 60-70° C are lethal to even
thstand such heat for short
limits of many arctic plants
eserves due to high respira-
tor plants to survive the low
tg season, they must be able
,f carbohydrates. When sub-
these plants will deplete all
lie. For example, the arctic
ticum) reaches its southern
an July temperatures reach
ow that the respiration rate
Jf shoreline plants found to
:1ses as temperature climbs.
~sin just 17 days if tempera-
that decrease leaf tempera-
small hairs on leaf surfaces.
1solation or be heliotropic.
ugh high rates of transpira-
plants lose their leaves dur-
to high temperatures in dry
~rns. In hot and dry periods,
e cool season, they support
Jf the southwestern United
Prosopsis glandulosa), have
as fish, reptiles, amphibians,
1ture generally reflects the
st, birds and mammals are
y temperature through the
~ to metabolically produce
jed animals are referred to
nimals can be damaged and
Vhen they are in an active
1mals must keep their body
re range. The lowest body
n the polar regions. Species
TEMPERATURE 55
such as the antarctic icefish (Trematomus borchgrevinki) can remain active at
body temperatures of -1.9° C. In contrast, the desert pupfish (Cyprinodon
nevadensis), found in hot springs near Death Valley, California, can withstand
prolonged temperatures of up to 42° C. Ants of the species Ocymyrmex barbiger
and spiders of the genus Seothyra from the Namib Desert of southern Africa
have been shown to survive body temperatures of 52° C to 55° C for very short
periods. However, these are exceptional species. The vast majority of animals
cannot tolerate prolonged temperatures that are even close to freezing or above
about 40° C. Temperatures that are too high or low for the organism can cause
decreases in metabolic activity, physical movement, growth, and reproduction.
Temperatures that are lethal and the range of temperatures that are optimum
vary widely for different species. Species that can tolerate a wide range of tem-
perature conditions are said to be eurythermic. Species with restricted tempera-
ture ranges are referred to as stenothermic.
In both homeotherms and poikilotherms, many metabolic functions are
temperature dependent and increase exponentially with temperature. The
increased rates of respiration and oxygen consumption generally result from
increasing temperatures. However, at some point on the temperature scale the
relationship between temperature and metabolic functioning becomes asymp-
totic. The physiological functioning of the organism no longer increases with tem-
perature, and eventually further warming causes death. High body temperatures
produce several different effects that can lead to death. Proteins may be broken
down or caused to coagulate. Enzyme activity may be reduced and enzyme for-
mation slowed. The rates of different temperature-controlled metabolic functions
may be driven out of synchronization at high temperatures. Rates of oxygen
intake may not equal respiration needs. Membranes may be distorted or
destroyed. Finally, behavior such as feeding may be negatively altered at high
temperatures. Salmon, for example, exhibit significantly decreased rates of feed-
ing in warm waters.
For homeotherms, lethal core temperatures range from 37° C to 47° C. This
is generally less than 10° C higher than the normal core temperatures of the ani-
mals. The core temperatures and lethal temperatures are higher for birds than for
mammals. In contrast, the lethal temperatures for many cold-water fish are very
low. For example the lethal high temperature for antarctic icefish is only 6° C.
Thus, the geographic range of the icefish is tightly restricted to the cold waters
around Antarctica. The exact causes of such low lethal temperatures in fish are
not understood.
There are many ways in which animals in warm environments regulate their
body temperatures to escape death through overheating. The metabolism of
heat-tolerant fish such as desert pupfish or desert-dwelling rodents has been
shown to differ from related organisms living in less extreme environments. For
example, the overall metabolic rate is lower for desert rodents than for non-
desert species. Large mammals, such as humans and horses, can cool themselves
through the release of sweat. The evaporation of sweat dissipates heat from the
body and cools the animal. The reason why humid nights in the summer seem so
unbearable is that the high amount of water vapor present in the air decreases
the rate of sweat evaporation and precludes us from cooling through sweating.
Smaller mammals do not sweat because of their low ratio of body mass to surface
area. Small animals would have to lose too great a proportion of their moisture to
cool effectively by sweating. Some mammals also pant in order to release heat
56 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

through the evaporation of moisture from the upper respiratory tract and to cool
meet oxygen demands. Marsupials, such as Australian kangaroos, cool themselves them
through the production of saliva, which they distribute on their bodies by licking. "'ilen
Small mammals may seek cool microclimates in the shelter of burrows and when
rock crevasses when temperatures are too high. In this case, small body size can adapt
be an advantage. The antelope ground squirrel (Ammospermophilus leucurus) of Loxt
western North America has been shown to maintain sublethal core temperatures to di~
by running back into its burrows when its body temperature exceeds about 42.4° mete1
C. These squirrels' small bodies rapidly lose heat in the cool microclimate of the
burrows, allowing them to quickly return to above-ground foraging activities. The prove
lack of suitable soils for burrowing can be an effective control on the local geo- and a
graphic distribution of such animals. Birds do not sweat but are known to pant els. 11
and rapidly oscillate the mouth and upper throat in an action called gular flutter- canh
ing in order to cool themselves. Birds such as the ostrich (Struthio came/us) make peratl
their plumage go erect when they are in danger of overheating in order to insu- sunni
late their skin from heat and solar radiation. turef
Reptiles and insects often rely on favorable microclimates in burrows and insect
crevasses in order to avoid overheating. In many cases, the behavior of the organ- The d
isms in selecting and frequenting the burrows is highly developed. If the animal come
stays in the shelter too long, it does not have sufficient time to meet its food minut
needs. In addition, if the shelter is too cool, the low temperature will slow down their
the metabolism of the reptile or insect and hinder it. Reptiles have an amazing ing cc
ability to use body positioning and differences in microhabitat to keep their body raise
temperature within a very narrow optimum range for their metabolism. Reptiles is onl'
can be very selective in choosing sites for thermoregulation. Snakes in California
have been known to select intermediate-sized rocks that are 20 to 40 em thick to Ionge'
hide under. Thinner rocks do not adequately protect the snakes from high tem- find r,
peratures, while thicker rocks provide too cool an environment for the snakes to tolera
maintain proper body temperature. beetl~
Fish are also cold blooded and often must move to different water depths to ature~

maintain optimal body temperature. In eastern North America, the optimal tem- ening
perature for the growth of northern pike (Esox lucius) and walleye (Stizosten- requiJ
odon vitreum vitreum) is about 23° C. These species are found in the upper warm ties w
waters and will preferentially swim to such warm sites. In contrast, lake trout tions
(Salvelinus namaycush) and lake whitefish (Coregonus clupeaformis) have opti- and r
mal growth at approximately 13° C and will actively seek out deep, cool waters. AI ask
The temperature requirements of adult animal species are often different winte
from those of their developing embryos or young, which are often more sensitive ide no
to high temperatures than the adults. For example, the eggs of frogs from the found
genus Rana develop abnormally or die when exposed to temperatures of 25° C north
and above, although the adult frogs can tolerate these temperatures. Pitsii
It is impossible to conclude a discussion of animal adaptations to heat with- tiles <
out mentioning two well-known animals with very distinctive traits that aid in intra-
their thermoregulation-the chameleon and the African elephant. The true mane
chameleons belong to the genus Chamaeleo and are found in semidesert regions becat
of Africa. They are not related to the anole lizards of the genus Anolis, which are ampb
found in the New World and often sold as "chameleons" by pet shops. Like anole out d:
lizards, the true chameleons can change the color of their skin. Although this
might help camouflage the lizard from prey and predators, research suggests that tainin
the color changes also facilitate thermal regulation. When the chameleons are hiber
~OF LIFE
per respiratory tract and to
n kangaroos, cool themselves
tte on their bodies by licking.
n the shelter of burrows and
:his case, small body size can
nospermophilus leucurus) of
sublethal core temperatures
Jerature exceeds about 42.4°
the cool microclimate of the
·ound foraging activities. The
ive control on the local geo-
weat but are known to pant
m action called gular fiutter-
~ich (Struthio camelus) make
>verheating in order to insu-
icroclimates in burrows and
:s, the behavior of the organ-
hly developed. If the animal
cient time to meet its food
temperature will slow down
t. Reptiles have an amazing
rohabitat to keep their body
r their metabolism. Reptiles
Lllation. Snakes in California
that are 20 to 40 em thick to
t the snakes from high tem-
vironment for the snakes to
to different water depths to
1 America, the optimal tern-
ius) and walleye (Stizosten-
,re found in the upper warm
;ites. In contrast, lake trout
us clupeaformis) have opti-
;eek out deep, cool waters.
I species are often different
ich are often more sensitive
the eggs of frogs from the
:d to temperatures of 25° C
~ temperatures.
tal adaptations to heat with-
:iistinctive traits that aid in
\.frican elephant. The true
found in semidesert regions
the genus Anolis, which are
1s" by pet shops. Like anole
,f their skin. Although this
1tors, research suggests that
When the chameleons are
TEMPERATURE 57
cool, they disperse melanin pigment and become darker in color. This increases
the amount of solar energy they absorb and warms them. The process reverses
when they are hot. Similarly, certain tropical tree frogs become almost white
when exposed to high heat and light intensities. Large ears may seem to be an
adaptation for increased hearing. However, in the case of the African elephant
(Loxodonta africana), the large thin ears appear to be an important mechanism
to disperse body heat and cool the animal. The large ears contribute several
meters of surface area for cooling, with very little additional body mass.
Cold temperatures serve to decrease rates of metabolic functioning and can
prove lethal to both homeotherms and poikilotherms. In the morning, reptiles
and amphibians sun themselves to bring their temperatures up to optimum lev-
els. This can be very effective. Peruvian mountain lizards of the genus Liolaemus
can be found living at altitudes of 4000 m in the Andes. Even in the summer, tem-
peratures at such high altitudes are often below freezing for part of the day. By
sunning themselves, these lizards have been shown to raise their body tempera-
ture from 2.5° C to 33° C, even though the air temperature was only 1.5° C. Flying
insects require relatively warm thoraxes in order for flight muscles to function.
The decrease in mosquito activity that occurs when temperatures drop is a wel-
come result of this requirement. In bees and moths, it has been shown that
minute rapid muscular motion, similar to shivering, allows some species to raise
their thorax temperatures. The sphinx moth (Manduca sexta) is often active dur-
ing cool evenings. Studies indicate that by rapid muscular motion the moth can
raise the temperature of its thorax to 35° C even when the night air temperature
is only 10° C.
The only way for most insects, amphibians, and reptiles to survive pro-
longed freezing temperatures is through avoidance and dormancy. Insects may
find refuge underground or in rotting wood. Species in cold climates can often
tolerate very cold temperatures during winter dormancy. For example, Alaskan
beetles of the species Pterosticus brevicornis have been shown to survive temper-
atures of -35° C during winter hibernation. In many cases, a process of frost hard-
ening through exposure to progressively cooler temperatures in the fall is
required for poikilotherms to develop such freezing tolerance. The Alaskan bee-
tles will freeze to death at temperatures of just -6.6° C if exposed to such condi-
tions during the summer. Amphibians, such as the frog species Rana sylvatica,
and reptiles, such as the garter snake, can inhabit cold regions as far north as
Alaska and the Canadian Northwest Territories because they settle down for
winter dormancy. During this time, frogs seek soft mud while the snakes take res-
idence in rock crevasses. In the case of the garter snakes, many thousands can be
found aggregating in rock shelters in the fall and emerging again in spring in cold
northern locations such as Wood Buffalo National Park and the Manitoba Snake
Pits in Canada. Despite sheltered locations, dormant insects, amphibians, and rep-
tiles often experience freezing temperatures in cold climates. The formation of
intra- and extracellular ice would be lethal to many species even during dor-
mancy. A number of species are able to supercool without the formation of ice
because they produce glycerol, which acts as an antifreeze. Dormant reptiles and
amphibians have been shown to supercool to temperatures as low as _go C with-
out damage.
Warm-blooded animals subject to cold temperatures must adapt by main-
taining body temperature while minimizing their expenditure of energy. Going into
hibernation allows the animal to minimize its energy loss by becoming inactive and
58 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

Arctic Smal
400
Frrst, beca
faster rate
g300 in order t<
2 because o
~ .9
-~ -c insulation
0 .§ 200
..0
ctl Cii
implicatio
Q) E should be
:2:
1100 --- --------- ' generally 1

- - Observed - - -Extrapolated
------ -------~ .................................
...............
"~
\. to suppor1
0 mals avoic
-60 -50 -40 -30 -20 -10 0 10 20 30 140 of birds o
Air temperature
Body protected
(oC) temperature
that live ir
related fo
FIGURE 3. 7 A comparison of the relationship between temperature and metabolic rate
body mas:
for tropical and arctic mammals (after Scholander et al., 1950; Schmidt-Nielsen 1997).
Rule. Des
arctic spec
It is
slowing down its metabolism to decrease energy use. However, hibernating mam- graphic di
mals can still freeze to death under extremely cold conditions or starve to death if that are a1
reserves of food energy are insufficient. Two primary adaptations exist to cope with phoebe) i:
cold for birds and mammals that remain active at low temperatures. The first is sponds fai
increased metabolic rates to generate body heat. The metabolic rates for both birds atures (Fil
and mammals are generally higher for arctic than for tropical species (Fig. 3.7). In rates that
addition, the critical air temperature where cold triggers increased metabolic activ- em limits.
ity is much lower for arctic animals than tropical ones. A tropical monkey will expe- boundarie
rience a doubling of its metabolic rate when air temperatures drop from 28° C to
18° C. An arctic ground squirrel would not experience such a doubling of metabolic
activity until air temperature drops from oo C to -4° C. The second strategy
involves increasing insulation to reduce heat losses. Insulation can take the form of
fat, fur, or plumage. Larger birds and mammals can physically support thicker insu- /~
lation; they depend on this strategy to counteract cold more than is the case for
smaller animals.
The polar bear (Thalarctos maritimus) is an example of a large predator
r.
,,.
) \
\
well adapted to a cold environment. Polar bears spend much of their lives in and
out of the water hunting seals on the edges of arctic sea ice. The arctic climate is
severely cold, and polar bears must contend with staying warm both when dry
and when wet. The metabolic rate of the bears is typical of mammals adapted to
arctic environments. The bears are large and present a small surface area for
cooling relative to their mass. However, the bear's fur is coarse and open. It does
not provide as much insulation in the air or in the water as a dense coat of fine fur
\
would. It does, however, dry quickly. This is an advantage given polar bears' ice-
shelf habitat and the severe wind-chill factor of the arctic. In addition to fur, the
bears have a layer of blubber that provides enhanced insulation in air and partic-
ularly in water. It may seem odd that polar bear fur is white. A dark coat would
absorb more solar radiation and aid in heating. However, the white coat provides FIGURE 3 .
important camouflage capability, allowing the bear to approach its prey unde- eastern ph '
tected. In addition, the individual hairs of the fur act as optical fibers that transfer obtain foo<
short-wave solar energy through the coat to heat the skin. their body
 ION OF LIFE
0 10 20 30 t40
Body
temperature
m temperature and metabolic rate
, 1950; Schmidt-Nielsen 1997).
use. However, hibernating mam-
ld conditions or starve to death if
ary adaptations exist to cope with
at low temperatures. The first is
llie metabolic rates for both birds
1 for tropical species (Fig. 3.7). In
riggers increased metabolic activ-
nes. A tropical monkey will expe-
:emperatures drop from 28° C to
~nee such a doubling of metabolic
: to --4° C. The second strategy
~s. Insulation can take the form of
n physically support thicker insu-
~t cold more than is the case for
an example of a large predator
spend much of their lives in and
rctic sea ice. The arctic climate is
th staying warm both when dry
' typical of mammals adapted to
>resent a small surface area for
·'s fur is coarse and open. It does
~ water as a dense coat of fine fur
!dvantage given polar bears' ice-
the arctic. In addition to fur, the
need insulation in air and partie-
. fur is white. A dark coat would
Iowever, the white coat provides
Jear to approach its prey uncle-
act as optical fibers that transfer
t the skin.
TEMPERATURE 59
Small birds and mammals have two disadvantages when dealing with cold.
First, because of their small ratio of body mass to surface area, they lose heat at a
faster rate than larger animals. This places more demands on energy consumption
in order to fuel the high metabolic rates needed to maintain body heat. Second,
because of their small size, they cannot physically support enough fat or fur for
insulation to counteract these demands. It would seem that the biogeographic
implication of the thermal inefficiency of small animals is that arctic animals
should be larger than related temperate and tropical species. However, this is not
generally true. It is possible that available food energy in the arctic is insufficient
to support greatly larger body masses. In addition, many birds and small mam-
mals avoid the rigors of cold winters by migrating to warmer climates in the case
of birds or by finding relatively warm microhabitats under the snow and other
protected locations in the case of small mammals. Generally, however, animals
that live in cold environments have shorter extremities, such as limbs or ears, than
related forms in warm environments. The shorter the extremities are relative to
body mass, the lower the rate of heat loss. This generality is known as Allen's
Rule. Desert rabbit and hare species, for example, typically have longer ears than
arctic species.
It is not surprising that cold climate has been shown to limit the broad geo-
graphic distributions of many animals. Small birds often have winter range limits
that are apparently dictated by low temperatures. The eastern phoebe (Sayormis
phoebe) is one example. Its northern range in eastern North America corre-
sponds fairly well with the --4° C isotherm for average minimum January temper-
atures (Fig. 3.8). Temperatures lower than this likely require increased metabolic
rates that cannot be met by the feeding rates of the eastern phoebe at its north-
ern limits. At least 14 other bird species in North America have northern range
boundaries that coincide with the point at which the metabolic rate of the birds
FIGURE 3.8 The relation between January temperature and the northern limits of the
eastern phoebe (Sayornis phoebe). North of the -4° C January isotherm, the birds can not
obtain food in sufficient quantities to support the metabolic activity required to maintain
their body temperature above lethal levels {after Root, 1993).
60 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

would have to be about 2.5 times higher than their basal metabolic rate in order rele
to maintain body heat during cold winter conditions. the
Cold temperatures, frequently limit the ranges of organisms by affecting no"
reproduction rather than through lethal cooling of adults. Corals, for example, are IC3J]

only found in regions with sea surface temperatures of 20° C or greater. Although ter 4
adult coral can survive in cooler water, it is likely that they cannot breed. The same bet\
is true for other marine organisms. The northern limit of the barnacle species Bal- \<lp
anus amphitrite coincides with the winter sea surface isotherm in summer of 18° C. the
Although the adults can survive temperatures as low as 7.2° C, they require a spa
mean monthly summer temperature of at least 18° C to reproduce. The North ani
American distribution of different species of the frog genus Rana also illustrates COlli
the role that temperature-restricted reproduction plays in limiting the geographic oral
distributions of species. The species Rana sylvatica can be found living and repro- liaD
ducing at sites near the Arctic Circle. This is much farther north than other frog oral
species. It has been shown that Rana sylvatica can tolerate much lower tempera- wbe
tures during breeding and in the embryonic stage than other species of frogs.
As a final point, it is interesting to consider humans in terms of adaptations
to arctic versus tropical conditions. Most humans do not have fur or high amounts
of insulative body fat. In addition, our metabolic rates and lower critical thermal
limits are similar to those of tropical mammals. On the basis of our natural cold IDe\
tolerance abilities, we, like corals and palms, are truly creatures of the tropics and Mao
subtropics. It is only by our ability to manipulate our immediate environment deo
through clothing, artificial heat, and specialized shelter that we can survive in the bee:
colder portions of the planet. turg
denc
furtl
MOISTURE suciJ

As with temperature, the impact of moisture on the physiological functioning and spe<
geographical distribution of animals is of extreme importance and has long been plac
studied by biologists and biogeographers. When examining the role of moisture ~\

in the distribution of life, it is again sensible to divide our discussion into separate Frr.
considerations for plants and animals. ture
5pe{
denl
Plants - 20
Water is essential to plants for a number of reasons. It is, of course, required for
photosynthesis as well as other physiological processes such as cell growth and caJX
protein synthesis. The cells of plants also require sufficient water to maintain tur- ofl
gor (rigidity). The release of water from leaves helps to cool plants. Finally, in vas- extr
cular plants water is essential to the movement of nutrients. Plants are often few
classified by their water demands. Those that can exist in dry environments are posl
called xerophytes. Plants that require moderately moist conditions are referred regi
to as mesophytes, while hydrophytes are those plants found growing in water or Ju~
very wet soils. qui<
In some sense, vascular plants may be thought of as conduits whereby water 5pe(
is taken up through absorption from the soil by fine roots and flows upward leav
through the vascular system. The specialized vascular tissues that conduct water on t
are called xylem, while those that transfer nutrients are called phloem. Eventu-
ally, water that is not retained in the plant or used directly in photosynthesis is of t1
~OF LIFE
Jasal metabolic rate in order
~s of organisms by affecting
iults. Corals, for example, are
of 20° C or greater. Although
they cannot breed. The same
t of the barnacle species Bat-
isotherm in summer of 18° C.
JW as 7.2° C, they require a
C to reproduce. The North
'g genus Rana also illustrates
1ys in limiting the geographic
m be found living and repro-
:arther north than other frog
)lerate much lower tempera-
m other species of frogs.
nans in terms of adaptations
not have fur or high amounts
es and lower critical thermal
the basis of our natural cold
f creatures of the tropics and
our immediate environment
:er that we can survive in the
•hysiological functioning and
1portance and has long been
tmining the role of moisture
our discussion into separate
It is, of course, required for
;ses such as cell growth and
icient water to maintain tur-
to cool plants. Finally, in vas-
: nutrients. Plants are often
'ist in dry environments are
toist conditions are referred
.s found growing in water or
•f as conduits whereby water
ne roots and flows upward
.r tissues that conduct water
are called phloem. Eventu-
directly in photosynthesis is
MOISTURE 61
released to the air through the stomata. Stomata are most commonly found on
the underside of leaves but can also occur on the top of leaves, on stems, and on
flower parts. The number of stomata on a leaf can be very high. The North Amer-
ican basswood tree (Tilia cordata) has about 13,000 stomata per square centime-
ter of leaf area, while the red oak (Quercus rubra) has 68,000. It is the difference
between the low-vapor pressure due to evaporation at the stomata and the high-
vapor pressure at the water-absorbing roots that causes water to move through
the plant. The release of water to the atmosphere by plants is referred to as tran-
spiration. The rate of water loss through transpiration is dependent on water
availability, stomata shape and behavior, air temperature, humidity, and wind
conditions. Hot, dry, and windy conditions increase the rate at which water evap-
orates from the leaf surface. In some regions, the loss of water to the air through
transpiration is much greater than the amount of water lost through direct evap-
oration from the soil surface. Climatologists use the term evapotranspiration
when considering the contribution of both sources of water to the atmosphere.
When the rate of water loss is high relative to absorption through the roots,
water stress occurs. If the water deficit is great enough, the cells and tissue of the
plant lose turgor and exhibit the soft and droopy conditions characteristic of the
wilting point. In cases of severe water loss, the death of cells and tissue is the
inevitable result. However, water stress can have many impacts besides wilting.
Most importantly, when plants are exposed to water stress they experience
decreased rates of photosynthesis. The inhibition of photosynthesis may occur
because the stomata of many plants will close when dehydration causes a lack of cell
turgor. This cuts off the supply of C02 for photosynthesis. There is also some evi-
dence that severe water deficits cause decreases in other physiological functions and
further depress photosynthesis. Support for this theory has been found in species
such as loblolly pine and Engleman spruce (Picea englemannii).
A clear biogeographic relationship exists between the ability of plant
species to conduct photosynthesis at low levels of water availability and the
places where the plant species are found growing. Trees that are typical of rela-
tively moist forest sites in eastern North America, such as alder (A lnus), ash
(Fraxinus), or loblolly pine, experience reduced rates of photosynthesis at mois-
ture deficits of -1 MPa (one MPa = 106 Pa; Pascals are a unit of pressure). Shrub
species of the extreme desert in Death Valley, such as creosote bush (Larrea tri-
dentata), do not exhibit reduced rates of photosynthesis until water deficits of
-2.0 to - 2.9 MPa are experienced.
Water stress can also cause the shedding of leaves and even stems. The bean
caper (Zygophyllum dumosum) of the Negev D esert of Israel may lose up to 90%
of its transpiring leaf and stem surface at the onset of the dry season. In a less
extreme climate, the black sage (Salvia mellifera) of California can lose all but a
few of its terminal leaves during the summer dry season but still maintain a net
positive rate of photosynthesis. Such loss of leaves is not restricted to plants in dry
regions. Well-known trees, such as the buckeye (A esculus glabra) and black walnut
(Juglans nigra) of the eastern North American forests, lose leaves relatively
quickly when subjected .to dry conditions during the growing season. Some oak
species living in the same zones as black walnut and buckeye appear to lose their
leaves less frequently during dry spells. Interestingly, they are found farther west
on the dry fringes of the Great Plains than trees such as walnut and buckeye.
Dry conditions can be particularly lethal to seeds and seedlings. The seeds
of trees such as oaks, willows, bald cypress, and elms are all killed when dried. In
62 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

contrast, seeds of pines and eucalyptus are often quite tolerant of desiccation. ~
The seeds of Eucalyptus siberi can survive several periods of desiccation and wet- oumbe
ting. This ability is not surprising in view of the dry conditions in which most
eucalyptus grow in Australia. ~ -· t
In the southwestern United States and northern Mexico, the desert vegeta- 5ea.SOn.
tion of the lowlands gives way to coniferous forest cover as elevation increases on - ·n
mountains. This lower treeline is a biogeographic boundary that can be traced to ep~
moisture stress. With increasing elevation, temperatures decline. This produces -~ ecies
lower evaporation rates in the upper elevations of the mountains. In addition, the . die
mountains receive higher amounts of rainfall due to orographic precipitation. se;
Thus, the mountains provide sufficient moisture to support tree growth, while the regioru
lower elevations do not. The pinyon pines (Pinus edulis and P. monphylla) occur cuticl~
in the southwestern United States and adjacent Mexico. The mature trees can Anotht
survive extremely dry conditions. The seedlings, however, can generally survive ound
no more than 12 days of wilting water stress, and this helps to exclude the pinyons sess \'e:
from the drier lowland deserts of the region (Fig. 3.9). - sout
A wide variety of adaptations allow plants to cope with dry conditions. = the
Through these strategies plants can be grouped into three broad categories: water amazm
stress escapees, water stress avoiders, and water stress tolerators. Many smaller
plants of seasonally dry environments survive the dry season as dormant seeds ~ arid
and are known as water stress escapees. The seeds of many desert plants can sur- -aobal
vive very long periods of desiccation and remain viable. There are accounts of .. vesrr
~

seeds germinating in the Sahara Desert of Sudan after more than 100 years of .: m ru
dormancy. Annual plants in the Sahara may germinate, grow, flower, and release use ver
seeds in periods as short as eight days following rainfall. These drought-escaping _-out :
annual plants often have very high rates of photosynthesis and conductance of -:l coni
water from leaves during their short lives. Most are not adapted to function well ()go
when water is in short supply. p
cactus
Ameri1
\JOclter ~

~obel
surfaa
contail
enougl

Desert
;) synthe
coveriJ
The cc
face b~
tender
ingant
the CJ
C{}z.l
when I
Pinyon Pine
stomal
(Pinus monophy/la)
- 40 em annual precipitation stomal
very d
FIGURE 3.9 The distribution of the pinyon pine species Pinus monophylla and annual rainfal
precipitation on the desert region of southeastern California. The pines are restricted to CAM
uplands where moisture availability is high compared to that in the dry lowlands. have t
N OF LIFE
Illite tolerant of desiccation.
~riods of desiccation and wet-
ry conditions in which most
rn Mexico, the desert vegeta-
wer as elevation increases on
mndary that can be traced to
ttures decline. This produces
1e mountains. In addition, the
to orographic precipitation.
upport tree growth, while the
lulis and P. monphylla) occur
[exico. The mature trees can
,wever, can generally survive
:helps to exclude the pinyons
)).
:o cope with dry conditions.
three broad categories: water
·ess tolerators. Many smaller
jry season as dormant seeds
f many desert plants can sur-
riable. There are accounts of
tfter more than 100 years of
ate, grow, flower, and release
1fall. These drought-escaping
ynthesis and conductance of
not adapted to function well
'in us monophyl/a and annua l
, The pines are restricted to
t in the dry lowlands.
MOISTURE 63
Many other plants from dry regions are water stress avoiders that employ a
number of strategies that allow individuals to survive during seasonally dry con-
ditions. Certain tropical and subtropical regions experience dry winters, and
many trees and large shrubs lose their leaves and become dormant during the dry
season. The ocotillo plant (Fouquieria splendens) of the southwestern desert of
the United States and Mexico may lose its leaves several times a year in response
to episodes of drought and precipitation. Some smaller plants, such as the sedge
species Carex pachystylis of theN egev Desert, allow their above-ground portions
to die back while their underground portions remain alive and dormant in the
dry season. A similar adaptation is found among grasses of dry subtropical
regions. Plants that retain their leaves in the dry season often have hard and waxy
cuticles that decrease moisture loss. Such leaves are called sclerophyllous leaves.
Another strategy to avoid water stress is to possess roots deep enough to obtain
ground water when the surface soil is dry. Even relatively small shrubs can pos-
sess very deep roots. For example, the chamise shrub (Adenostoma fasciculatum )
of southern California has roots that can extend to 8 m in depth, while the roots
of the mesquite shrubs (Prosopis) of the southwestern desert can extend an ,
amazing 53 m. Another interesting adaptation for water stress avoidance is the
ability of some plants to store water in their tissues. Several types of trees found
in arid regions have enlarged trunks that can store water. These include the
baobab (Adansonia digitata) of Africa and the bottle tree (Brachychiton
rupestris) of eastern Australia. Baobabs may have a circumference of more than
20m and store up to 120,000 liters of water (Fig. 3.10). Some water stress a voiders
use very little water in photosynthesis. The agave (Agave deserti) transpires only
. about 25 g of water for every gram of carbon that is fixed during photosynthesis.
In contrast, many plants from more mesic environments may transpire 250 to
600 g of water for every gram of carbon they fix.
Perhaps the most interesting water stress avoiders are the members of the
cactus family which are found in the deserts and dry regions of North and South
America. These plants possess a number of adaptations to help them survive
water stress and have been extensively studied by the American botanist Park
Nobel and his colleagues. Cacti possess extensive fine root systems near the soil
surface that quickly intercept any rainfall. The stems of the cacti are enlarged and
contain tissue to store water. The barrel cactus (Ferocactus acanthodes) can store
enough water for 50 days of photosynthetic activity. The stems of cacti are photo-
synthetic, and the leaves have been reduced to hard thorns. Some cacti have thick
coverings of light-colored thorns that protect against the sun and drying wind.
The covering of spines on the cholla cactus (Opuntia bigelovii) can cool its sur-
face by as much as 11 o C. In other cases, the thorns serve to defend the moist and
tender stems against herbivores. The exteriors of cactus stems have a waxy coat-
ing and deeply inset stomata that help further reduce water loss. Finally, cacti use
the CAM photosynthetic pathway, whereby the stomata open at night to take in
C02 . The C02 is converted to malic acid, which is decarboxylated during the day
when C3 photosynthesis converts the C02 to sucrose and starch. By keeping the
stomata closed during the heat of the day, evaporative water loss through the
stomata can be reduced. D espite these adaptations, cacti are not found in the
very driest parts of the New World deserts. To survive, cacti must have sufficient
rainfall to replenish their water supplies from shallow ground water. In addition,
CAM plants tend to have slow growth rates and are limited by the fact that they
have the lowest photosynthetic rates of the flowering plants.
64 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

found ale
beyond 51
water tui
water tup

Animal
Here we'
to the me
can be dE
animals, i1
maintain
aboutlO~
In contra5
vive. Watf
water los:
worms or
other seer
The1
animals li
insects an1
skin evap
genus Naz
FIGURE 3.10 A large baobab tree (Adansonia digitata) growing in Kenya. Baobabs may tion and 1
have a circumference of more than 20 m and can store up to 120,000 I of w ater. The stored
the genus
water helps them cope with the dry subtropical winter.
36% thrOl
animals fc
three tim<
All of these adaptations allow plants to survive and conduct photosynthesis conditiom
when water is in short supply in the environment. However, very few plants can greater th
tolerate desiccation of tissue to the point of being near completely dry. True evaporati<
water stress tolerators are able to tolerate severe dehydration down to < -13 bodyweig
MPa. Some tropical C4 grasses and members of the pea family can withstand this of water 1<
degree of desiccation. More amazingly, individuals of some mosses, lichen, algae, Watt
and fungi, such as the so-called resurrection plants Selaginella lepidophylla and areas whe
Craterostigma plantagineum, can actually survive almost complete desiccation Water can
and begin functioning rapidly following rehydration. directly at
Too much water can also be problematic for plants. The diffusion of 0 2 animals n1
through water-saturated soils is about 10,000 times slower than through dry soils. leaves anc
Plants that grow in water-saturated soils may not be able to obtain sufficient oxy- leaves tha
gen through their root systems to survive. Plants that grow in water-logged soils, many anin
however, often develop longitudinal air spaces within their roots. These spaces The
are called aerenchyma and aid in the storage and transfer of oxygen. Some plants adapted tc
have the ability to develop new roots from their stems if they are partially buried tration of;
by wet soils following flooding. These roots are referred to as adventitious roots ica. The k
and are present in herbs such as the common dock weed (Rumex) as well as many water. Wh
trees and shrubs including the coastal redwood. Even plants that are normally through o;
associated with moist soils can differ markedly in their ability to tolerate very wet be 6 ml. In
conditions. Both sycamore (Platanus occidentalis) and water tupelo (Nyssa aquat- small qua1
ica) grow best on moist soils in southeastern North America. These trees are trated am<
N OF LIFE
rowing in Kenya. Baobabs may
) 120,000 I of water. The stored
and conduct photosynthesis
[owever, very few plants can
~ near completely dry. True
dehydration down to < - 13
)ea family can withstand this
f some mosses, lichen, algae,
Selaginella lepidophylla and
lmost complete desiccation
plants. The diffusion of 0 2
ower than through dry soils.
1ble to obtain sufficient oxy-
t grow in water-logged soils,
in their roots. These spaces
tsfer of oxygen. Some plants
s if they are partially buried
Ted to as adventitious roots
ed (Rumex) as well as many
~n plants that are normally
r ability to tolerate very wet
I water tupelo (Nyssa aquat-
1 America. These trees are
MOISTURE 65
found along waterways and in damp areas. However, as soil moisture increases
beyond 50% saturation, the growth rate of sycamore begins to decline while the
water tupelo reaches maximum growth under water-saturated conditions. The
water tupelo is, therefore, found on wetter sites than the sycamore.
Animals
Here we will consider the impact of moisture on terrestrial animals. Water is vital
to the metabolic functioning of all animals. It has been said that "living animals
can be described as an aqueous solution contained within a membrane." Most
animals, including insects, are composed of two-thirds water by weight and must
maintain this high volume in order to survive. Mammals can withstand a loss of
about 10% of their body water, while losses of 15% to 20% generally prove fatal.
In contrast, some frogs can lose as much as 40% of their body water and still sur-
vive. Water can be lost directly through evaporation from the skin. Evaporative
water loss can be particularly high for animals with permeable skin such as
worms or amphibians. Water can also be lost through respiration, excrement, and
other secretions.
There is often a relationship between the environmental conditions that
animals live in and their rate of water loss through evaporation. In general,
insects and reptiles from dry environments have lower rates of water loss through
skin evaporation than through respiration. For example, water snakes of the
genus Natrix lose approximately 88% of their body moisture through evapora-
tion and 12% through respiration. In contrast, desert-dwelling gopher snakes of
the genus Pituophis lose only 64% of their body water through evaporation and
36% through respiration. In addition, total water loss rates tend to be lower in
animals found in arid environments. Water snakes lose moisture at a rate that is
three times faster than that for gopher snakes when subjected to the same dry
conditions. Pond-dwelling turtles lose moisture at a rate that is eight times
greater than the rate of moisture loss for desert tortoises. In desert rattlesnakes,
evaporation and respiration combined account for only a 0.5% loss of their water
body weight per day. At this rate, these reptiles could survive two to three months
of water loss.
Water is generally replenished by drinking, and many animals are limited to
areas where free water exists in lakes, streams, springs, and as dew on leaves.
Water can also be obtained through food and, in the case of some insects, it is
directly absorbed through the skin. In fact, a number of insects and some other
animals never drink water but obtain sufficient moisture from food. Succulent
leaves and fruits may contain up to 90% water. Some insects can subsist on dry
leaves that contain only 5% to 10% water. Even if free water is limited in food,
many animals can obtain water through the oxidation of organic matter.
The kangaroo rat (Dipodomys spectabilis) is a classic example of an animal
adapted to an arid environment. This animal must maintain a body water concen-
tration of about 66% while living in the dry deserts of southwestern North Amer-
ica. The kangaroo rat can live entirely on dry food without ever drinking free
water. When eating 100 g of dried seed, the rat can produce 54 ml of water
through oxidation-even though the amount of free water in the grain may only
be 6 ml. In order to avoid water losses through excrement, the rats produce very
small quantities of very dry feces and urine which contain very highly concen-
trated amounts of urea relative to the water content. The rats do not sweat, and
66 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

they lose only small quantities of water through evaporation. Despite these adap- Of
tations, the kangaroo rat still faces difficulty in compensating for moisture loss. In of worm:
hot and dry atmospheric conditions, typical of desert days, the rats will lose more able area
water through respiration and evaporation than they gain in eating dry food. (concent
These potential losses are mitigated by the fact that the rats are nocturnal and will lose I
spend the daylight hours in cool and relatively moist burrows. In addition, female taking in
rats have increased water demands when they are in their reproductive period urine. Sh
and producing milk. At this time, they must be able to increase their uptake of sure bee~
free water by eating green and moist plants. Despite these limitations, the kanga- Be<
roo rat can survive in much drier environments than many other rodents. For of marin
example, packrats (Neotoma) also occur in the southwestern deserts but are water sp
restricted to moister regions than is the case for kangaroo rats. In contrast to spend pa
kangaroo rats, packrats cannot obtain sufficient moisture through the oxidation as well a~
of dry foods and are unable to produce urine which has the same high concentra- this abili
tions of urea to water. freshwat'
as tides a
cellular <
OTHER PHYSICAL FACTORS ents. Sue
As
Plants and animals are sensitive to many other aspects of the physical environ- than tho:
ment besides light, temperature, and moisture. Plants require chemical nutrients trations 1
in the soil such as nitrate, phosphorus, and potassium in order to grow. Aquatic Few aqw
organisms may be sensitive to the acidity of the water in which they live. Such notsupp
physical factors are often extremely important at a local scale but less apparent in salina) a
controlling broad global distributions. Detailed investigation of these physical salinity I<
factors falls more into the realm of the ecologist and physiologist rather than that Bo1
of the biogeographer. Nonetheless, several of these physical properties of the of free o
earth's environment are important for controlling large-scale geographic distri- anoxic (l
butions of organisms. We will examine a few examples of these physical factors in occur in
this section. surface ,
The most important chemical factor controlling the global geographic dis- organisrr
tribution of aquatic organisms is salinity. Most species of aquatic plants, crus- certain SJ
taceans, fish, and other life can only be found in either fresh or salt water. How do basins ar
differences in salinity control the distributions of aquatic organisms? coast of
Organisms must maintain a relatively stable cellular chemistry. Increases lusks, en
or decreases in the salt content of the cells can cause changes in the reactions of have eve
proteins, enzymes, and other metabolic functions. This disruption can lead to The so-c:
death. If water salinity changes, an osmotic pressure gradient is formed. The (Chiron<
osmotic pressure gradient causes water to flow through the permeable mem- extreme!
brane of the organism. During this flow, pure water moves toward the more cient tra1
saline water. If a salt water worm is placed in freshwater, the purer water will that don
flow across the body wall into the worm. This leads to an imbalance in the salin- Pla
ity of the worm's cells. The opposite flow occurs for a freshwater organism water. H
placed in salt water. Under these circumstances, the freshwater species loses plants. S;
water and suffers desiccation. For this reason, it is impossible for freshwater ate irrig~
amphibians, such as most frogs, to cross salt water barriers. Indeed, most tain soil:
amphibians are restricted to freshwater. There is, however, a crab-eating frog in growth<
Southeast Asia (Rana cancrivora) that spends long periods in the ocean. It main- shown t<
tains an osmotic gradient against a lethal absorption of salt water by having a cover on
high urea content in bodily fluids. ticularly
N OF LIFE
Joration. Despite these adap-
'ensating for moisture loss. In
t days, the rats will lose more
bey gain in eating dry food.
1t the rats are nocturnal and
: burrows. In addition, female
in their reproductive period
e to increase their uptake of
these limitations, the kanga-
tan many other rodents. For
mthwestern deserts but are
:angaroo rats. In contrast to
isture through the oxidation
bas the same high concentra-
ects of the physical environ-
ts require chemical nutrients
m in order to grow. Aquatic
tter in which they live. Such
'cal scale but less apparent in
·estigation of these physical
physiologist rather than that
~ physical properties of the
:trge-scale geographic distri-
~s of these physical factors in
g the global geographic dis-
cies of aquatic plants, crus-
r fresh or salt water. How do
tatic organisms?
ellular chemistry. Increases
~ changes in the reactions of
rhis disruption can lead to
re gradient is formed. The
ough the permeable mem-
~r moves toward the more
water, the purer water will
o an imbalance in the salin-
for a freshwater organism
te freshwater species loses
impossible for freshwater
ter barriers. Indeed, most
.vever, a crab-eating frog in
~riods in the ocean.lt main-
1 of salt water by having a
OTHER PHYSICAL FACTORS 67
Of course, fish and shelled crustaceans do not have the water-permeable skin
of worms. In these organisms, osmotic exchange occurs through exposed perme-
able areas such as gills. Surprisingly, most marine fish have osmotic concentrations
(concentrations of solutes) that are one-third to one-quarter that of sea water. They
will lose body water through areas such as their gills and must avoid desiccation by
taking in large amounts of sea water and evacuating salts through their gills and in
urine. Sharks also have low concentrations of salts but maintain high osmotic pres-
sure because they have high amounts of urea in their body fluids and cells.
Because of their sensitivity to osmotic pressure changes, the vast majority
of marine plants and animals cannot survive in freshwater. Similarly, most fresh-
water species will die if placed in salt water. There are, however, species that
spend part of their lives in salt water and part of their lives in freshwater. Salmon,
as well as certain eels and sharks, found in Lake Nicaragua and in Malaysia, have
this ability. In addition, organisms that live in estuarine environments where
freshwater rivers enter the sea must deal with daily to seasonal swings in salinity
as tides and river flow vary. These species have a special capacity to regulate their
cellular chemistry and avoid the development of lethal osmotic pressure gradi-
ents. Such organisms are said to be euryhaline.
As a final point, there are certain lakes that have salinities much higher
than those of the ocean. For example, Great Salt Lake in Utah has salt concen-
trations that are more than seven times higher than concentrations in the ocean.
Few aquatic organisms can tolerate such high salinities. The Great Salt Lake can-
not support fish. Only two macroscopic invertebrates, a brine shrimp (Artemeria
salina) and a benthic brine fly (Ephydra cincera), are able to tolerate the high
salinity levels.
Both freshwater and marine animals obtain oxygen from the water. A lack
of free oxygen in the water is lethal. In some lakes, the bottom waters become
anoxic (low in oxygen) either seasonally or permanently. This condition may
occur in deep lakes where current activity is not effective in mixing bottom and
surface waters to replenish oxygen. Under such conditions, bottom-dwelling
organisms such as worms and mollusks are excluded. A similar situation occurs in
certain small deep ocean basins. Two well-known examples of such anoxic ocean
basins are the Santa Barbara Basin off California and the Cariaco Basin off the
coast of Venezuela. The bottoms of these basins are devoid of the worms, mol-
lusks, crustaceans, and ground fish typical of the ocean floor. Some organisms
have evolved to accommodate life in waters with relatively low oxygen content.
The so-called freshwater blood worms are actually the larval stage of midge flies
(Chironomidae). The red color of these freshwater organisms is due to the
extremely high hemoglobin content in their blood. This allows for a more effi-
cient transfer of oxygen and gives blood worms the ability to live in lake waters
that do not have enough free oxygen to support most other types of organisms.
Plants are often very sensitive to the chemical conditions of the soil and
water. High concentrations of certain chemical elements may be deleterious to
plants. Saline soils are lethal to most plants, and few terrestrial species can toler-
ate irrigation by sea water. High concentrations of magnesium are found in cer-
tain soils, particularly those derived from dolomite and serpentine rocks. The
growth of many plants, from grasses and herbs to coniferous trees, has been
shown to decrease on soils with high concentrations of magnesium. Vegetation
cover on such soils is often sparse and may be dominated by species that are par-
ticularly adapted to tolerating high concentrations of magnesium.
68 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

Acidic precipitation and fog caused by the burning of fossil fuels can
directly injure leaves and also lead to the mobilization of aluminum and other
trace elements in some soils, and this can prove toxic to plants. It is estimated that
between the years 1960 and 2000 about half of the high-elevation red spruce
(Picea rubens) trees in the Adirondack Mountains of New York died due to acid
precipitation.
In freshwater ecosystems, particularly lakes and ponds, the acidity of the
water exerts a strong control on plankton and larger plants. Lakes that have
become acidified due to air pollution experience great changes in plant species
and animal species. Severely acidified lakes are often almost lifeless. Such lakes
can be found in portions of eastern North America and Europe. It is estimated
that 15% of all lakes in New England are suffering from human-caused acidifica-
tion. In the Adirondack Mountains, approximately 40% of all lakes are suffering
from acidification. About 24% of the lakes in the Adirondacks have become so
acidic that fish can no longer survive in them. Most of the acidic precipitation that
is affecting vegetation and lakes in the northeastern United States is due to sulfur
and nitrogen released into the atmosphere by coal-burning power plants and
industry in the Ohio River Valley. Increased reliance on coal in the future may
well accelerate the problem of acid precipitation.

INTERACTING PHYSICAL CONTROLS

ON GEOGRAPHIC DISTRIBUTIONS

In this chapter, we have explored how individual physical factors such as light,
temperature, or moisture affect the physiological functioning of organisms and AGUR
limit the geographic distributions of species. Of course, frequently more than after~
one physical factor may be responsible for determining whether a plant or ani- subtro
mal can live at a given location. For example, in desert environments, the combi- grasslc
nation of low water availability, high temperatures, and high light intensities may
all be important factors that work in combination to preclude the survival of
most plant species.
The geographic distribution of plant species that employ the c4 photosyn- :-Ian!.!
thetic pathway provides a good example of how several different physical factors latitUt
contribute to the geographic limits of species. The percentage of dicot species expla
that use the C4 pathway ranges from almost 5% in the southwestern United tropi<
States to less than 1% in the Canadian grasslands. The percentage of C4 species in higbe
the North American grass flora ranges from over 80% in the Southwest to 0% in aren
northern Canada and Alaska. The C4 plants can make up an even higher propor- ties a
tion of the flora in tropical and subtropical regions. In northern tropical Australia, c~gr
C4 species account for 80% to 100% of the grass flora in the dry interior. In the hot.(
cooler and moister temperate portions of southern Australia and Tasmania, C4 condi
species make up only 2% to 10% of the grass flora (Fig. 3.11). In North America
there is a very high positive correlation between summer evaporation rates and tions
the abundance of C4 species in the flora. There is also a strong positive relation- ior e:
ship between maximum daily temperature and the abundance of C4 plants in the entg
grass flora. C4 plants appear to be better adapted to warm, dry, and sunny condi- ofsp
tions for several reasons. First, C4 plants are more efficient in C02 bonding at mer1
high light intensities and temperatures than are c 3 plants. Thus, through their thes
stomata, C4 plants can take in more C02 relative to the water they lose than C3 sumr
IN OF LIFE INTERACTING PHYSICAL CONTROLS ON GEOGRAPHIC DISTRIBUTIONS 69

: burning of fossil fuels can

1tion of aluminum and other
c to plants. It is estimated that
b.e high-elevation red spruce
:lf New York died due to acid

md ponds, the acidity of the

rger plants. Lakes that have
reat changes in plant species
en almost lifeless. Such lakes
1 and Europe. It is estimated
'rom human-caused acidifica-
~0 % of all lakes are suffering
\dirondacks have become so
,f the acidic precipitation that
United States is due to sulfur
il-burning power plants and
ce on coal in the future may

U>90% C4
D 50-90%C4
l<:il)<''l 20-50% c.
<20% C4

hysical factors such as light,

mctioning of organisms and
FIGURE 3.11 The relative abundance of C3 and C4 grass species in the flora of Austral ia
mrse, frequently more than
(after Henderson et al., 1993). The C4 species are most abundant in t he wa rm tropical and
aing whether a plant or ani- subtropical zones of the continent. A similar situation is found in the North American
:rt environments, the combi- grasslands.
nd high light intensities may
to preclude the survival of

at employ the c4 photosyn-
ral different physical factors
percentage of dicot species
n the southwestern United
: percentage of c4 species in
b in the Southwest to 0% in
~ up an even higher proper-
northern tropical Australia,
·a in the dry interior. In the
Australia and Tasmania, C4
"ig. 3.11). In North America
tmer evaporation rates and
J a strong positive relation-
undance of c4 plants in the
mrm, dry, and sunny condi-
fficient in C02 bonding at
plants. Thus, through their
he water they lose than c3
plants. Second, C4 plants can benefit from the higher light intensities in the lower
latitudes because they do not lose carbon through photorespiration. This
explains, in part, why C4 plants can survive and dominate in sunny and dry sub-
tropical and tropical grasslands. Why, however, are C 4 plants not abundant in the
higher latitude vegetation of Canada or southern Australia? It appears that they
are restricted from the higher latitudes because they require greater light intensi-
ties and higher temperatures for optimum photosynthesis than are required by
c3 grasses. Thus, there is a tradeoff between the ability of c4 plants to tolerate
hot, dry, sunny conditions and their ability to survive in the cooler and less sunny
conditions found at high latitudes.
When we expand our examination of physical controls on species distribu-
tions to include different geographic areas, southern and northern range limits,
for example, we almost always find that different factors are important at differ-
ent geographic locations. Consider the northern and southern range distributions
of spruce. In the north there appears to be a good correlation between low sum-
mer temperatures and the range limits of the species. However, when we examine
the southern range limits in western Canada, we find little evidence that low
summer temperatures control the distribution of the spruce species. Instead, it
70 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

appears that moisture stress, caused by a combination of high summer tempera- range.
tures and low precipitation, excludes spruce from growing on the prairies. These
as its •
is oftE
ENVIRONMENTAL GRADIENTS land a
AND SPECIES' NICHES
geogr:
As we have seen, many different factors in the physical environment can limit specie
the geographic distributions of plant and animal species. Are there some general densit
models that biogeographers use to try and organize, conceptually at least, the tions
relationships between different species and the physical environmental factors tions i
that affect them? In this section we will look at some ideas from ecology that specie
provide useful conceptual frameworks to organize our observations on how the the d(
physical environment controls the distributions of species. First, let us consider port 1=
how we can describe the geographic distributions of species. Two basic funda-
mental measures for describing the geographic distribution of a species are physi<
range and density. In the preceding sections, we used the term range limits to pies c
describe the geographic distribution of species. The geographic range of a species ment<
is the entire area where the species can be found regardless of whether it is com- moist
mon or rare. For example, the range of the eastern bluebird (Sialia sialis) extends dient
from roughly the U.S.-Canadian border southward to the Gulf Coast and south- tionin
ern Florida. In the west, it extends to the Great Plains (Fig. 3.12). Within its form '
that I

High
Q)

~
u
'a;
.J::
c>.
(/)
0
0
.J::
a..
Low

High

Low

FIGURE 3.12 The range and population density of eastern Bluebird (Sialia sialis) in
North America. Notice how population density is greatest in patches near the ce nter of t he FIGU
geographic range (after Bystrak, 1979 and Brown and Gibson, 1983). speci '
 \1 OF LIFE ENVIRONMENTAL GRADIENTS AND SPECIES' NICHES 71

on of high summer tempera-
owing on the prairies.
ysical environment can limit
:cies. Are there some general
~e, conceptually at least, the
ysical environmental factors
•me ideas from ecology that
)Ur observations on how the
5pecies. First, let us consider
of species. Two basic funda-
istribution of a species are
5ed the term range limits to
:eographic range of a species
;ardless of whether it is com-
uebird (Sialia sialis) extends
o the Gulf Coast and south-
)lains (Fig. 3.12). Within its
range, a species may be found to be common in some areas and rare in others.
These differences in the abundance of the species within its range are referred to
as its density. D ensity is a measure of population abundance per unit of area and
is often reported as the number of individuals per square meter or hectare on
land and the number of individuals per cubic meter in aquatic systems.
The highest densities for species often occur in the central portions of their
geographic ranges. Lowest densities are usually found toward the edges of
species' ranges. We can see that this is generally true for the eastern bluebird. The
density of the species is greatest where the environment provides the best condi-
tions to support individuals of the species. This often occurs in the central por-
tions of the geographic range of the species. In contrast, at the range limits of the
species, the environment is unable to support a large number of individuals and
the density decreases. The number of individuals that the environment can sup-
port per meter or hectare is called its carrying capacity.
Now, let's link these observations on geographic ranges and density to the
physical environmental controls we have discussed. We have seen many exam-
ples of how the physiological functioning of species can be related to environ-
mental conditions along the environmental gradients of light, temperature, and
moisture. The physiological functioning of the organism increases along the gra-
dient to an optimal level and then declines. The increase in physiological func-
tioning along these physical environmental gradients frequently occurs in the
form of a Gaussian curve (Fig. 3.13). In m any cases, the environmental conditions
that promote optimum physiological functioning produce the highest carrying

Zone of Zone of Zone of the Zone of Zone of

High intolerance stress optimum stress intolerance

Q)

~
u
il
.<:::.
c>-
(/)
0
0
.<:::.
a..
Low
ensity of Low High
Jluebird Air temperature
)pulation
High
alia sialis)
~
0 "iii
c::
Q)
1-2 "0
c::
0
3-9
~
:;
10 - 29 a.
~
~ 30
Low
Low High
Air temperature
Bluebird (Sialia sialis) in
atches near the center of the FIGURE 3.13 The photosynthetic rate and population density of a hypothetical plant
1983). species along an environmental gradient of low to high air temperature.
72 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

capacities for the species. For example, a hypothetical plant species might reach
its highest capacity for photosynthesis at about 20° C. At this temperature the asp
high rate of photosynthesis produces rapid and robust growth, allowing plants to bi<>t'
mature quickly and produce more viable seeds, thereby providing better survival fact•
potential for the seedlings. The population size of the species is therefore higher distl
in geographic regions with a temperature of 20° C than in areas where the tem- the
perature is warmer or cooler than 20° C. The geographic differences in species func
density can be related to increasing and decreasing physiological functioning thet
along the environmental gradient of temperature. In general, the density of a enct
species along an environmental gradient also takes the form of a Gaussian curve rv.-o
(Fig. 3.13). The study of how species distributions relate to the environment, WID(

including how species ranges and densities are distributed along environmental rese
gradients, is called ordination analysis. refe
\!he environmental gradients for the physiological functioning and den- men
sity of a species can be divided into several subzones. The zone of the optimum spec
represents the best conditions for the species. In this zone, physiological func- sion
tioning and density are at a maximumjThe zones of stress provide conditions enVl
that allow the species to persist but at low carrying capacities. In the zones of used
intolerance, environmental conditions are too harsh for the species to survive. tom
For our hypothetical plant species, average growing season temperatures cons
above say 30° C or below 10° C are lethal and define the zones of intolerance expl
(Fig. 3.13). The zones of tolerance for species may shift depending on the life dam
stages of the plant or animals. For example, the seeds of a plant might tolerate
temperatures of below freezing or above 40° C with no apparent damage. In acte1
contrast, seedlings frequently have much more restricted tolerances than seeds men·
or adult plants. v.ide
Sucb
ronJJ
brea
v.idc
High geo!
cal I
defu

1:EY WORt

Low
Low High
Air temperature

FIGURE 3.14 The niches of a hypothetical generalist plant species and a specialist plant
species as defined by two gradients, air temperature and soil moisture.
~OF LIFE
:al plant species might reach
' C. At this temperature the
.st growth, allowing plants to
~by providing better survival
e species is therefore higher
han in areas where the tem-
~aphic differences in species
tg physiological functioning
In general, the density of a
he form of a Gaussian curve
relate to the environment,
ibuted along environmental
>gical functioning and den-
os. The zone of the optimum
is zone, physiological func-
>f stress provide conditions
: capacities. In the zones of
1 for the species to survive.
.ving season temperatures
1e the zones of intolerance
shift depending on the life
is of a plant might tolerate
~h no apparent damage. In
icted tolerances than seeds
igh
:pecies and a specialist plant
1oisture.
KEY WORDS AND TERMS 73
In the preceding section, we have seen that the geogr aphic distribution of
a species may be controlled by more than one physical factor. Ecologists and
biogeographers recognize that many environmental gradients, representing
factors such as temperature, precipitation, and light intensity, can control the
distribution of species. Species exist in a multidimensional space defined by all
the different physical environmental factors that affect their physiological
functioning and ultimately their abundance. If we again consider our hypo-
thetical plant species, we might find that its range and abundance are influ-
enced both by growing season temperature and annual precipitation. These
two gradients form the axes of a two-dimensional environmental space in
which the species can be found growing (Fig. 3.14). This two-dimensional rep-
resentation of the environment in which the species can survive can be
referred to as its niche. Of course, there are always more than two environ-
mental variables that control the survival of an organism. So the real niches of
species include many different gradients and are formally defined as n-dimen-
sional hypervolumes that are defined by all (n) of the physical and biological
environmental conditions the species requires to survive. The term niche was
used by J. Grinnell in 1917 and C. Elton in 1927. G. E . Hutchinson, however,
formulated its presently accepted definition in 1957. The niche should not be
considered as synonymous with the habitat of the species. Habitat is the
explicit spatial environment in which species can be found. For example, the
damp surface soil beneath a rock is a habitat.
Different species have different niches. The niche provides a means of char-
acterizing and comparing the general relationship between species and environ-
ment. Species, which can tolerate a wide range of environmental conditions, have
wide ranges of tolerances on environmental gradients and wide niche breadths.
Such species are referred to as generalists. Species, which have very narrow envi-
ronmental tolerances, have more restricted gradient distributions and niche
breadths and are referred to as specialists (Fig. 3.14). Since they can tolerate a
wide variety of physical environmental conditions, generalists often have large
geographic ranges compared to specialists. Although the concept of the ecologi-
cal niche is easy to envision, determining all of the environmental factors that
define the real niches of species is extremely difficult.
JtEY WORDS AND TERMS
-'s rule Habitat Sciophytes
_.:J Heliophytes Sclerophyllous leaves
~g capacity Homeotherm Specialist
a:iduous Hydrophyte Stenothermic
-llmity Mesophyte Transpiration
-.orphic leaves Niche Xerophyte
~e Ordination analysis Xylem
IIIJthermic Perennial
apotranspiration Phenology
il:a:ralist Phloem
graphic range Poikilotherm
74 CHAPTER 3 THE PHYSICAL ENVIRONMENT AND THE DISTRIBUTION OF LIFE

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_ In Nor
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conditions. Tree Physiology 2, 131- 142. gists
Toole, V. K., Toole, E. H., Hendricks, S. B., Borth- actio
wick, H . S., and Snow, A. B., Jr. (1961).
distr
Responses of seeds of Pinus virginiana to
light. Plant Physiology 36, 285- 290. a COl
Walter, H. (1985). Vegetation of the Earth, 3rd sider
edition. Springer-Verlag, New York. ent c
Woodward, F. I. (1987). Climate & Plant Distribu-
tion. Cambridge University Press, Cambridge.
PREDATIO'
Pre1
com
nivc
rate
OF LIFE
>82). Areography: Geographi-
Species. Pergamon Press,
. F., and Eichhorn, S. E. (1999).
s, 6th edition. W. H. Freeman,
:rgy constraints on avian distri-
Idances. Ecology 69, 330-339.
1er, W. (1987). Frost Survival
er-Verlag, Berlin .
. (1997) . Animal Physiology:
Environment, 5th edition.
'ersity Press, Cambridge.
)Ck, R ., Walters, V., Johnson,
(1950). Heat regulation in
tropical mammals and birds.
'in 99, 237-258.
~ipid deposition in intercellu-
203, 164-165.
7). Plants that warm them-
4merican March, 104-109.
:-irst and second season effect
me initiation from a single
nadian Journal of Forest
435 .
eri, J. A. (1978). The geo-
ion of c4 species of the
relation to climate. Ameri-
2, 609-623.
ve, L. G. (1976). Climatic pat-
tribution of C4 grasses in
'Jecologia 23, 1-12.
J. A. , Samuelson, L. J. , and
(1986). Stomatal and non-
Jns to net photosynthesis in
tder different environmental
-:>hysiology 2, 131-142.
~ . H., Hendricks, S. B., Barth-
now, A. B., Jr. (1961) .
ds of Pinus virginiana to
ology 36, 285-290.
?getation of the Earth, 3rd
Verlag, New York.
:7). Climate & Plant Distribu-
University Press, Cambridge.
CHAPTER 4
BIOLOGICAL INTERACTIONS
AND THE DISTRIBUTION
OF LIFE
Sex is not the first thing that comes to most people's minds when they watch a
hummingbird move from flower to flower. Yet that is precisely what the hum-
mingbird is engaged in-on behalf of the flowers, that is. With each sip of nectar
the visiting bird takes away some pollen, which it then transports to the next
flower. Some of the pollen picked up by the hummingbird will be transferred to a
flower of the same species where the male sperm nucleus within the pollen grain
will fertilize the female ovule to form a seed. The nectar produced by the flowers
feeds pollinating birds or insects. Sexual reproduction would be impossible for
many flower species without the intervention of specific pollinators such as the
hummingbird. If the pollinating species is not present, the flower species that
depends on it will also be absent.
We have explored how variations in the physical environment of the earth
affect the physiological functioning of organisms and ultimately influence the
geographic distributions of species. However, plants and animals also interact
with other species, and these biological interactions can control the geographic
ranges and population densities of species. In addition to flowers and their polli-
nators, there are many forms of such interaction. One species of insect may prey
upon another, or two bird species may compete for the same nesting sites. Some
interactions between species can be important factors in controlling geographic
ranges. The study of interspecies interaction has long been a focus for both ecolo-
gists and biogeographers. In this chapter we will explore different types of inter-
actions between species and how these interactions influence geographic
distributions. We will examine how geographic distributions can be controlled by
a combination of biological interactions and physical factors. We will then con-
sider how biological interactions are accommodated within the concepts of gradi-
ent distributions and species' niches.
DATION
Predation occurs when one organism consumes another. In this section we will
consider predation to include both herbivory (animal-plant predation) and car-
nivory (animal-animal predation), although many ecologists treat these as sepa-
rate relationships.
If a predator depends on only one particular species for its prey, the geo-
graphic distribution of the predator is necessarily limited to the geographic
77
78 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

distribution of the prey species. Some predators are indeed highly selective in :t:te va ~
terms of their prey species. Predators that have a very narrow range of prey us ing
species are called stenophagous, or selective predators. In California, the geo- pp ear:
graphic distribution of some races of the checkerspot butterfly (E uphydryas d r ea ~
~eso u r c
editha) are geographically limited to areas where their only food plant, a plantain
species called Plantago hookeriana, is found. Interestingly, this plantain species ,·ore he
itself is restricted to soils formed from serpentine rock. At a continental scale, the aging a
northern limits of the monarch butterfly (Danaus plexippus) corresponds exactly is play
with the northern range limits of its food plant (Fig. 4.1) , milkweed (Asclepias xpectE
spp.). How common is such restricted stenophagy? If we look at all butterfly rela tive
species in the Los Angeles area, we find that about one-third of the 106 recorded io n.lar
species are restricted to one species of food plant or several closely related an d itt
species of the same genus. Within North America, about 80% of all butterfly the se V<
species are relatively stenophagous and restricted to a single family of plants for w
food. Worldwide, it appears that about 90% of all plant-eating insects (phy- racticE
tophagous species) are restricted to food plants from one or two plant families. and the
So, for plant-eating insects, stenophagy is extremely common and must be a re sorr
considered an important factor in controlling geographic distribution. Is this also resource
true for other herbivorous animals and carnivores? There are some famous ontrast
instances of mammals that are highly stenophagous. Two of the best known are ,·ery sm
the Australian koala bear (Phascolartus cinereus), an arboreal (tree-dwelling) i ms live
marsupial that only eats the leaves of the bluegum eucalyptus. The second, the an live
Chinese giant panda bear (Ailuropoda melanoleuca), eats mainly the foliage of Ste
certain bamboo species. In North America, Albert's squirrel (Sciurus alberti) gen- reli ance
erally subsists on seeds from the ponderosa pine (Pinus ponderosa). However, only the
erate a r
'"hv are
mon in r
\\ ill out!
Yide the
This is c
represer
plants tf
fo r usinE
Se<
particula
This allo
vores. In
also mak
caterpill<

,.
\~.,
·~ ·r.o
<\'"-~i!. ·' /
to preda
plants co
more pa
Summer range
Th
of monarch butterfly th ey req
(Dana us plexippus) efficient
.
• Winter areas neural c
a very re
FIGURE 4.1 The correspondence between the northern range limits of the monarch
butterfly (Oanaus plexippus) and the northern range limits of its food plant, milkweed
It
(Asclepias spp.) (after Brower and Malcolm, 1991 ). geograp
:LIFE
: indeed highly selective in
very narrow range of prey
ors. In California, the geo-
pot butterfly (Euphydryas
r only food plant, a plantain
tingly, this plantain species
~.At a continental scale, the
:ippus) corresponds exactly
. 4.1) , milkweed (Asclepias
If we look at all butterfly
e-third of the 106 recorded
or several closely related
::tbout 80% of all butterfly
1 single family of plants for
plant-eating insects (phy-
one or two plant families.
nely common and must be
hie distribution. Is this also
? There are some famous
fwo of the best known are
n arboreal (tree-dwelling)
ucalyptus. The second, the
eats mainly the foliage of
uirrel (Sciurus alberti) gen-
nus ponderosa). However,
~ mon in phytophagous insects? There are many possible explanations, but here we
~ will outline just a few. First, it benefits predators to focus on prey species that pro-
~
~ Second, many plants produce toxins to deter herbivores. Some herbivores,
~
.
,,~~ ".
%
F plants containing toxic and unpalatable chemicals. Euryphagous caterpillars were
r
ge limits of the monarch
ts food plant, milkweed
PREDATION 79
the vast majority of terrestrial birds and mammals are relatively euryphagous
(using many different plant species for food). The degree of food selectivity
appears to be less restrictive for large herbivores than for small ones. The
decrease in selectivity for large herbivores likely reflects their need for a broad
resource base to acquire enough energy to maintain themselves. A large herbi-
vore has a better chance of meeting its energy requirements if it has a large for-
aging area and can eat many different species of plants. In general, carnivores
display a much lower degree of stenophagy than do herbivores. This is to be
expected, since carnivores are generally at the top trophic level and must have a
relatively broad food resource base to acquire enough energy to persist. In addi-
tion, large carnivores will usually transect many different habitats when foraging,
and it benefits them to take advantage of different species of prey that occupy
these various environments.
We can therefore see a biogeographic relationship between the feeding
practices of large herbivores and carnivores, the range of their foraging activities,
and the number of different environments that they use for feeding. Such animals
are sometimes referred to as fine-grained because their foraging is made up of
resources from an array of many fine-scale environments within their range. In
contrast, some insects may spend their entire life on one leaf. Animals that have
very small ranges and restricted diets can be called coarse-grained. These organ-
isms live in a world that is coarsely divided between the limited areas that they
can live in and areas that they cannot.
Stenophagy appears to be a relatively poor strategy for a species. The
reliance on one or two species of prey limits the distribution of the predator to
only those areas where the prey species can exist. The predator may perhaps tol-
erate a much wider range of physical environmental conditions than the prey. So
why are some animals stenophagous, and in particular why is stenophagy so com-
vide the highest ratio of food energy relative to the energy required for foraging.
This is called the optimal foraging theory. The theory assumes that food choices
represent the result of predator evolution. The focus of herbivores on specific
plants that provide high nutrition is the outcome of an evolutionary adaptation
for using only the most nutritious food sources.
particularly insects, evolve resistance to the specific toxins of just one plant species.
This allows the insect to feed on that plant with little competition from other herbi-
vores. In addition to foraging benefits, ingesting high levels of such plant toxins can
also make the insect unpalatable to carnivorous predators. In one experiment, the
caterpillars of 70 different butterfly species from Costa Rica were offered as food
to predatory ants. The ants avoided caterpillars that were stenophagous and ate
more palatable to the ants and were selectively eaten.
Third, small insects may find that all of the microenvironmental conditions
they require during their life cycle are provided by one individual plant, and it is
efficient to restrict foraging to that plant. Finally, insects have relatively simple
neural capabilities and may be behaviorally limited to identifying and consuming
a very restricted number of plant species.
It is easy to see how the presence or absence of prey species can control the
geographic distribution of a predator. Can the presence of a voracious euryphagous
80 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

predator restrict the geographic distribution of prey species? In some instances, the in theBahar
introduction of a generalist predator by humans has led to the extinction or near duced to so
extinction of prey species. In eastern North America, lake trout (Salvelinus namay- lizards had '
cush) are driven towards extinction in lakes that contain lampreys (Petromyzon species as cc
marinus). Lampreys are fearsome predators that spend much of their life at sea but Outsic
come into freshwaters to breed. They use rasping mouths to attach themselves to evidence th
prey fish and literally suck the life out of them. Formerly, sea lampreys could not species. The
enter the Great Lakes beyond Lake Ontario. They were prevented from moving far- may be dep
ther up the Great Lakes system by the presence of Niagara Falls. However, with the sure becom'
building of shipping canals to Lake Erie, lampreys were able to enter the upper turn causes
Great Lakes. The result was the almost complete eradication of lake trout (Salveli- will decline
nus namaycush) in the upper Great Lakes (Fig. 4.2). Populations of lake trout now ator popula1
have to be sustained by lamprey control programs and the breeding of trout in fish increase aga
hatcheries for release into the lakes. An interesting set of long-term experiments has and their p
been conducted on the impact of predation on orb spiders that live on small islands exemplified
populations
can be expn

c;;:----~ In this equa

over time (t,
~zt;ec hunting effie
For predat01

Lake Huron

In this equa1
the efficienc
Finally, q is t
a set of sinu
rises and de,
that explaim
but are less 1
the predator
Lake Michigan DoLo
populations '
prey populal
such populat
bivores. For
moths (Tyria
the abundan
changes in th
nation condi1
Lake Superior sic study of l
the fur tradi1
1935 1940 1945 1950 1955 1960 provide data
Year main prey, sn
FIGURE 4.2 The devastating impact of the accidental introduction of sea lampreys Bay Compan
(Petromyzon marinus) on lake trout (Sa lvelinus namaycush) in Lake Michigan. At present, tury (Fig. 4.3:
lamprey control programs and the breeding of trout in hatcheries are used to maintain lake natural abun
t rout populations in the Great Lakes (after Baldwin, 1964; Krebs, 1994). in the lynx a
IFE
:ies? In some instances, the
i to the extinction or near
:e trout (Salvelinus namay-
lin lampreys (Petromyzon
nuch of their life at sea but
hs to attach themselves to
'ly, sea lampreys could not
•revented from moving far-
,ra Falls. However, with the
·e able to enter the upper
1tion of lake trout (Salveli-
mlations of lake trout now
ile breeding of trout in fish
long-term experiments has
rs that live on small islands
Jction of sea lampreys
Lake Michigan. At present,
es are used to maintain lake
'1994).
PREDATION 81
in the Bahamas. Predatory lizards of the genera Anolis and Leiocephalus were intro-
duced to some of the islands. Subsequent monitoring showed that islands with
lizards had about one-tenth the spider population density and one-half the spider
species as comparably sized nonlizard islands.
Outside of these artificial circumstances, there remains surprisingly little
evidence that predators control the broad geographic range limits of prey
species. There are several possible explanations. First, stenophagous predators
may be dependent on the abundance of a single prey species. If predation pres-
sure becomes intense, the population size of the prey will decrease, and this in
turn causes a decrease in predators. Due to lack of prey, the predator population
will decline because of starvation and decreased fertility. The decline in the pred-
ator population relieves the pressure on the prey, and the prey species is able to
increase again. We would logically expect populations of stenophagous predators
and their prey species to undergo periodic fluctuations in population size as
exemplified in Figure 4.3. This concept of linked oscillations in predator-prey
populations is called the Lotka-Volterra model after its originators. The model
can be expressed mathematically for prey populations as:
dN/dt = rN- a'PN
In this equation, dN/dt refers to the rate of change in the prey population (N)
over time (t), r is the natural rate of growth of the prey population (N), a' is the
hunting efficiency of the predator, and P is the size of the predator population.
For predators the model is expressed as:
dP/dt = fa'PN- qP
In this equation, dP!dt is the rate of growth of the predator population, and f is
the efficiency by which the predator turns captured prey into new offspring.
Finally, q is the death rate for the predator populations (P). The model produces
a set of sinusoidal curves in which the curve for the prey population leads the
rises and declines of the predator population. It is a simple and elegant model
that explains why selective predators can cause changes in prey population size
but are less likely to cause extinction. When the prey population size gets too low,
the predator population size crashes.
Do Lotka-Volterra cycles occur in nature? Laboratory experiments using
populations of mites and beetles have produced cyclic variations in predator and
prey populations. Some short-term observational studies in nature suggest that
such population cycles in prey and predators can exist for both carnivores and her-
bivores. For example, it has been shown that populations of English cinnabar
moths (Tyria jacobaea) climb and fall one year behind increases and decreases in
the abundance of their prey plant, the ragwort (Senecio jacobaea). However, the
changes in the abundance of the ragworts mainly reflected changes in seed germi-
nation conditions, and not pressure by predation from the cinnabar moths. A clas-
sic study of Lotka-Volterra oscillations between predator and prey comes from
the fur trading records of the Hudson Bay Company in Canada. These records
provide data on the number of pelts of Canada lynx (Lynx canadensis) and its
main prey, snowshoe hare (Lepus americanus), that were obtained by the Hudson
Bay Company from the early nineteenth century through the mid-twentieth cen-
tury (Fig. 4.3). It is assumed that the number of pelts can be reliably related to the
natural abundance of the lynx and hare. These records show an 8- to 11-year cycle
in the lynx and hare population size. The lynx populations do indeed follow the
82 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

160 prey species.

are very com
"'~ 90
C1l
removal of a 1
.r:
0
.gj80
c
C1l PETITION
"'::J
_g 40
I- Interspecific <
vi duals of two
1925 the mortality
Year tion, competi1
- - Snowshoe hare on the role of
-----Lynx
It has long be
tionist, Charle
FIGURE 4.3 Historical cyclic variations in the population sizes of the snowshoe tial importanc
(Lepus americanus) and Canada lynx (Lynx canadensis) (after MacLulick, 1937; Began data indicate
et al., 1996).
common occu
determining tl
There m
increases and decreases in the hare population size. Interestingly, these cycles tion in which 1
seem to be synchronous across northern Canada. For many years, this record of or interaction.
lynx and hare populations was used as evidence in support of the classic Lotka- compete for I=
Volterra model. However, recent experimental manipulations of Yukon hare pop- foraging. The
ulations show that both food availability and the presence of predators influence tors actually i
the development of cyclic increases and decreases in hare populations. If preda- might establis
tors are excluded, but food supplies are allowed to vary naturally, periodic cycles rate and physi
in the size of the hare population remain. It has been shown that high populations some competi
of hares quickly deplete the supply of early easy-to-digest twigs of willow and ous to anothe1
birch. The subsequent regrowth shoots contain toxins that make them less palat- some suspectc
able. So the hare-lynx cycle is also an echo of the plant-hare cycle. example, it is
Despite its shortcomings, the Lotka-Voterra model provides a theoretical common frogs
model that helps us understand why selective predators do not seem to be impor- ing natterjack
tant in restricting the geographic ranges of prey species. So, is there any evidence Some bi<
to suggest that predators are important in restricting the geographic distributions competition fo
of species in nature? The answer appears to be yes-under certain circumstances. that are toxic t
This conclusion is based on studies of how the artificial exclusion of predators spatial distribu
changes the species diversity of prey. Experimental studies show that rocky role of allelop<
shorelines along the Pacific Coast of North America and coral reefs off Hawaii ral (shrub land
experience increased numbers of algae species following the artificial exclusion around many
of animals that graze on algae. These species of algae are apparently not able to strated that fa
survive at the sites when grazers are present. However, these and other experi- these areas of
ments, particularly in marine and freshwater ecosystems, have also shown that sage (Salvia le
the long-term impact of excluding predators can cause the decline and extinction nica), and boll
of many prey species. This result is counterintuitive but makes perfect sense if we have phytotox
consider it further. In some cases, predators may be keystone species that signifi- shown to be pi
cantly influence the whole composition of ecosystems by controlling the popula- toxins inhibit s
tion sizes of prey species. When the predators are removed, certain prey species of established 1
are able to outgrow or outcompete other species and eventually eradicate some be a limiting fi
species. In some instances, the presence of generalist predators may allow species the soils aroun•
of prey to grow in areas they would be excluded from by competition with other thought. The c
LIFE
><
91
\t/I\·13~~
0
OM PETITION
6
925
sizes of the snowshoe
er Maclulick, 1937; Began
Interestingly, these cycles
many years, this record of
1pport of the classic Lotka-
!lations of Yukon hare pop-
~nce of predators influence
hare populations. If preda-
ry naturally, periodic cycles
hown that high populations
digest twigs of willow and
that make them less palat-
t-hare cycle.
)del provides a theoretical
~s do not seem to be impor-
~s. So, is there any evidence
he geographic distributions
nder certain circumstances.
cial exclusion of predators
l studies show that rocky
and coral reefs off Hawaii
ving the artificial exclusion
are apparently not able to
er, these and other experi-
ems, have also shown that
~ the decline and extinction
t makes perfect sense if we
~ystone species that signifi-
by controlling the popula-
lloved, certain prey species
eventually eradicate some
redators may allow species
by competition with other
COMPETITION 83
prey species. From these studies we can derive an important lesson. Ecosystems
are very complex. Seemingly simple changes that we might make, such as the
removal of a predator species, can have unexpected and undesirable results.
Interspecific competition can be broadly defined as the interaction between indi-
viduals of two or more species in which the growth and/or fertility is decreased or
the mortality is increased for both species. In the case of intraspecific competi-
tion, competing individuals are within the same species. In this section, we focus
on the role of interspecific competition in determining the range limits of species.
It has long been a topic of intense interest to biogeographers. The great evolu-
tionist, Charles Darwin, placed particular emphasis on competition and its poten-
tial importance in limiting the distributions of species. Extensive reviews of field
data indicate that some degree of interspecific competition between species is a
common occurrence in all ecosystems. How important might competition be in
determining the geographic distributions of species?
There are two general forms of competition. The first is resource exploita-
tion in which both species compete for the same resources without direct contact
or interaction. For example, two species of birds that eat the same insect species
compete for prey, although the birds might never encounter each other during
foraging. The second form, interference competition, occurs when the competi-
tors actually interact either physically or chemically. For example, two plants
might establish as seedlings very close together. One may have a faster growth
rate and physically exclude the other in the rooting zone and in the canopy. In
some competitive interactions, one organism exudes a chemical that is deleteri-
ous to another organism. This is referred to as allelopathy. In animals there are
some suspected instances of interference competition through allelopathy. For
example, it is thought that algae that are found in the feces of the tadpoles of
common frogs (Rana temporaria) can inhibit the growth and survival of compet-
ing natterjack toad (Bufo calamita) tadpoles.
Some biogeographers believe allelopathy is a major feature of interference
competition for plants. It is known that many plants produce chemical substances
that are toxic to other plants. However, the true importance of allelopathy in the
spatial distributions of plants continues to be debated. Classical research into the
role of allelopathy in plant competition comes from studies of California chapar-
ral (shrubland) and grassland communities. Halos of bare ground are found
around many chaparral shrub species. Careful field experiments have demon-
strated that factors such as root competition or canopy shading cannot explain
these areas of bare ground. Common chaparral plants such as purple and black
sage (Salvia leucophylla and S. mellifera), coastal sagebrush (Artemisia califor-
nica), and holly-leafed cherry (Prunus ilicifolia) all produce terpenes that may
have phytotoxic (poisonous to plant) properties. These substances have been
shown to be present in soils around these species. It is thought that these phyto-
toxins inhibit seed germination and thus limit competing plants from the vicinity
of established plants. This is particularly important in chaparral where water can
be a limiting resource for plants. However, some experiments have shown that
the soils around chaparral shrubs may not be as toxic to other species as was first
thought. The causes of the chaparral halos remain debated. Other experiments
84 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

appear to provide evidence for the importance of allelopathy in other vegetation

communities. For example, research conducted in Oklahoma showed that several
competing grass species had low seed germination rates and grew poorly on soil
taken from areas dominated by the grass species Sporobolus pyramidus. It is
known that Sporobolus pyramidus produces p-coumeric and ferulic acid, which
can act as phytotoxins.
{
I
Competition also occurs between individuals of the same species (intraspe-
cific competition). In theory, competition should be most intense between individ-
uals of the same, or closely related, species because these related organisms will
likely have the greatest similarities in the resources they require to survive. The
impact of both interspecific and intraspecific competition on population size has
been demonstrated in many laboratory experiments for over 60 years. One such
experiment was conducted by growing two diatom species in water enriched with
silica. Diatoms are unicellular photosynthetic organisms that build protective
shells out of silica. When the species Asterionella formosa was grown alone, the
population initially grew rapidly. The growth then tapered off as silica abundance
declined. Finally, a stable population size was reached and maintained. A similar
pattern occurred when the competing diatom species Synedra ulna was grown
alone in the silica-enriched water. What happens when we observe the popula-
tions of competing species that are grown together in the laboratory? When the . Oipc
two diatom species are grown together, the rate of population increase for both . o.d,
species is slower. Eventually, Asterionella formosa is driven to extinction by Syne- O.SJ
dra ulna. In this case Synedra ulna is a superior competitor. Such interactions have . o.n,
been observed in many laboratory experiments using competing species. .O.e1
Interspecific competition may be a factor in limiting the geographic distri-
bution of species. Examination of maps showing the range limits of some species FIGURE4
of plants and animals provides evidence in support of competition as an impor- Mexico sh<
tant biogeographic force. For example, the distribution of different species of species in I
kangaroo rats (Dipodomys spp.) in the desert of the southwestern United States Brown and
and adjacent Mexico shows little geographic overlap in the ranges of these
closely related species (Fig. 4.4). A similar pattern of nonoverlapping ranges is D. specta.
apparent in the geographic distributions of subspecies of chuckwalla lizards cold tem1
(Sauromalus obesus) in the same region or subspecies of rat snakes (Elaphe the north
obsoleta) in the southeastern United States. It is possible to suggest, from such The
mapped distributions, that competitive exclusion of one species by another mackinla;
species may be an important factor in determining range limits. However, in tunity to:
many cases closely related species do overlap in range limits. One example is pro- not affect
vided by the South American distribution of species of caiman (Caiman), a rela- occur tog
tive of alligators and crocodiles. In the case of the caiman, only two of the six exclusion
South American species have very sharp range limits with each other, whereas in also been
most cases species overlap significantly in their geographic distributions. It is pos- tions of t
sible that the species with sharp range boundaries with no overlap exclude each range lim
other through competition for similar resources such as prey or nesting sites. The tion has a
overlapping species may avoid competition by making use of different prey or Hur
habitats, thereby lessening competition pressure. that inter
Even when two closely related species do have sharp range limits and no species. P
geographic overlap in distribution, a number of factors other than competition ment occ1
may still influence their range limits. For example, the winter climate in the north- ifornia by
ern regions where the kangaroo rat species Dipodomys deserti and D. ingens are Pric
found is much colder than the climate in Mexico where the species D. nelsoni and Californi<
LIFE
apathy in other vegetation
thoma showed that several
es and grew poorly on soil
1orobolus pyramidus. It is
ric and ferulic acid, which
he same species (intraspe-
st intense between individ-
tese related organisms will
tey require to survive. The
ion on population size has
)r over 60 years. One such
::ies in water enriched with
sms that build protective
"!osa was grown alone, the
red off as silica abundance
and maintained. A similar
; Synedra ulna was grown
n we observe the popula-
the laboratory? When the
pulation increase for both
iven to extinction by Syne-
tor. Such interactions have
ompeting species.
ting the geographic distri-
mge limits of some species
competition as an impor-
on of different species of
mthwestern United States
p in the ranges of these
nonoverlapping ranges is
:ies of chuckwalla lizards
ies of rat snakes (Elaphe
ible to suggest, from such
one species by another
range limits. However, in
imits. One example is pro-
f caiman (Caiman), a rela-
timan, only two of the six
;ith each other, whereas in
)hie distributions. It is pos-
h. no overlap exclude each
s prey or nesting sites. The
g use of different prey or
sharp range limits and no
rs other than competition
;inter climate in the north-
s deserti and D. ingens are
the species D. nelsoni and
COMPETITION 85
• Dipodomys ingens
111 0. deserti
D. spectabilis
_ O. nelsoni
.D.elator
FIGURE 4.4 The distribution of kangaroo rat species in the southwestern United States and
Mexico shows sharp range boundaries that suggest the importance of competition with other
species in limiting the geog raphic ranges of each species (after Bowers and Brown, 1982 and
Brown and Lomolino, 1998).
D. spectabilis live. It is possible that the northern species are better adapted to
cold temperatures. Cold temperatures could be the factor that ultimately controls
the northern range limits of some kangaroo rats.
The distribution of the closely related cuckoo-dove species Macropygia
mackinlayi and M . nigrirostris on small islands off New Guinea provides the oppor-
tunity to study competition in a region where such large differences in climate do
not affect the species. It has been found that the two species of cuckoo-doves never
occur together on the same island. This observation suggests that competitive
exclusion keeps the two species from inhabiting the same islands. However, it has
also been proposed that random chance alone could lead to the observed distribu-
tions of the two bird species. All in all, we can conclude that analysis of species'
range limits from maps cannot be considered unequivocal evidence that competi-
tion has a primary importance in determining geographic distributions.
Human introductions of alien plant and animal species do provide evidence
that interspecific competition can cause changes in the geographic distribution of
species. Perhaps one of the most striking results of such competitive displace-
ment occurred through the introduction of Eurasian annual grass species to Cal-
ifornia by the Spanish and other European colonists.
Prior to European contact, the grassland in the low and mid-elevations of
California was dominated by native annual and perennial grass and herb species.
86 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

Europeans introduced a variety of annual grass species from Eurasia which are
able to occupy the same habitat as the native herbs and grasses. The Eurasian Nevada
annual grasses grow rapidly with the onset of the winter rains, flower and set
seed, and then die during the waterless summer months. The next winter, the
seeds germinate and begin the cycle again. These aggressive competitors are well
adapted to California's seasonally wet and dry climate and the native species
have been displaced throughout most of the state's grasslands. The golden hills
associated with summers in California are largely an artificial landscape domi-
nated by competitively superior alien invaders.
Although the California grasslands are a dramatic example of competitive
/ Ruby Mour

New Mexico
displacement, they represent a situation created by human intervention. Direct
evidence of the competitive interactions controlling natural distributions is more
difficult to find . One classic study that provides such evidence is that of the distri-
bution of chipmunk species in the mountains of the southwestern United States.
There are some 20 species of chipmunks in North America. Like the caiman in
South America, some species do not overlap and have very distinct range bound- Organ Mou1
aries, whereas others have overlapping distributions and appear to utilize different
habitats within those ranges. The isolated mountains in the southwestern United FIGURE 4.5
States are cool and moist compared with the surrounding desert. These mountains in the Great E
support coniferous forests that provide a suitable habitat for chipmunks. The chip- quadrivittatu.
elevation site
munks cannot survive in the dry desert, so the mountains serve as islands of habi-
tat that support isolated populations of chipmunks (Fig. 4.5). The distribution and
behavior of four chipmunk species, Eutamias dorsalis, E. minimus, E. quadrivitta-
tus, and E. umbrinus, have been studied in detail. Many mountains in the region -v MBIOSIS: Ml
have the right environment to support all four species. However, in many cases ::l RASITISM, A
one or more of the species is absent. It is thought that competition may play a role
in excluding some species from the mountains. When the two species, E. dorsalis Symbiosis i:
and E. umbrinus, occur together, E. umbrinus is found only in the dense forests of through cot
the upper elevations and E. dorsalis only in the open woods at low elevations. A the survival
relatively narrow zone separates the ranges of the two species. However, if one of Different Sf
these species is present on a mountain and the other is absent, the species that is both specie1
there can be found at both lower and higher elevations. This indicates that the two (commensal
species can tolerate the range of physical and biological conditions found at both the other. P:
high and low elevations but are excluded from certain elevations when the com- for nutrient:
peting chipmunk is present. The elevational distributions of E. quadrivittatus and hosts. Becat
E. dorsalis also appear to be strongly influenced by competition between these of biologica
species (Fig. 4.5). Finally, when other species of chipmunks are present, E. minimus species invo
is always restricted to the low-elevation shrubby habitat. Observations suggest Some
that interference competition plays a direct role in these elevational distributions. many plant:
E. dorsalis is very aggressive in physically excluding other chipmunks within its flower to ftc
range. This strategy is particularly effective in the open and scattered woods of transferring
lower elevations. At higher elevations, E. dorsalis expends much energy trying to beak within
defend habitat in relatively continuous forest. In contrast, the two species E. that the bin
umbrinus and E. quadrivitattus do not waste as much energy in defending territory distributiom
in the closed forest, and this gives them a survival advantage over E. dorsalis. The west, moths
species E. minimus is less aggressive and behaviorally subordinate to all of the the Yucca dt
other species. As a final point, although competition appears to be clearly impor- tionships ne
tant in the elevational distribution of the chipmunk species, it must be remem- support the
bered that habitat conditions in the form of patchy versus continuous tree cover of different J
ultimately determine which species wins the competition. by the distri
IFE
s from Eurasia which are
md grasses. The Eurasian
nter rains, flower and set
1ths. The next winter, the
ssive competitors are well
te and the native species
asslands. The golden hills
artificial landscape domi-
c example of competitive
1man intervention. Direct
:ural distributions is more
dence is that of the distri-
Jthwestern United States.
erica. Like the caiman in
1ery distinct range bound-
appear to utilize different
the southwestern United
g desert. These mountains
t for chipmunks. The chip-
IS serve as islands of habi-
4.5) . The distribution and
~- minimus, E. quadrivitta-
1 mountains in the region
However, in many cases
mpetition may play a role
1e two species, E. dorsalis
nly in the dense forests of
·oods at low elevations. A
pecies. However, if one of
absent, the species that is
[his indicates that the two
conditions found at both
;;levations when the com-
IS of E. quadrivittatus and
1mpetition between these
cs are present, E. minimus
tat. Observations suggest
• elevational distributions.
her chipmunks within its
1 and scattered woods of
1ds much energy trying to
trast, the two species E.
:rgy in defending territory
1tage over E. dorsalis. The
subordinate to all of the
)ears to be clearly impor-
'ecies, it must be remem-
ms continuous tree cover
l.
SYMBIOSIS: MUTUALISM, COMMENSALISM, PARASITISM, AND MIMICRY 87
Nevada
Eutamias
dorsalis
Ruby Mountains Snake Range Pilot Peak
New Mexico
Eutamias
dorsalis
Organ Mountains Mt. Taylor Magdalena Mountains
FIGURE 4.5 The distribution of three different chipmunk species on isolated mountains
in the Great Basin of the western United States. Where Euatamias umbrinus or E.
quadrivittatus are present, they competitively exclude the species E. dorsalis from high-
elevation sites (after Brown and Lomolino, 1998).
SYMBIOSIS: MUTUALISM, COMMENSALISM,
ARASITISM, AND MIMICRY
Symbiosis is the close association between two species that generally develops
through coevolution. In many cases, the symbiotic association is obligatory for
the survival of one or both species. There are several different types of symbiosis.
Different species may interact closely with each other in a manner that benefits
both species (mutualism) or benefits one species and has no impact on the other
(commensalism). In the case of parasitism, one species benefits at the expense of
the other. Parasites are organisms that are wholly dependent on other organisms
for nutrients and, in many cases, microhabitat. They do not immediately kill their
hosts. Because of the obligatory nature of most symbiotic relationships, this type
of biological control determines the geographic distribution of one or both of the
species involved.
Some examples of mutualistic relationships are very obvious. For example,
many plants require insects, birds, or mammals to transfer their pollen from
flower to flower. Many plants produce nectar that provides food for the pollen-
transferring organism. When, as we have already noted, a hummingbird places its
beak within the flower, some pollen rubs onto it and is carried to the next flower
that the bird visits. Pollinating relationships can tightly control the geographic
distributions of the organisms involved. For example, in the desert of the South-
west, moths of the genus Tegeticula feed only on the desert plant Yucca. In turn,
the Yucca depends on the moth to transfer their pollen. In some mutualistic rela-
tionships neither species can extend its geographic range into areas that cannot
support the other. Most hummingbirds, however, are able to feed from a number
of different plant species and do not have geographic ranges that are constrained
by the distribution of a single flower species. Indeed, the relationship between
 ·----- -----~ - -

88 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

plant species and the animals that pollinate them or disperse seeds is often not
sufficiently obligative to be the primary control on the geographic distribution of The 1
the species. are r<
In the ocean, clownfish of the genus Amphiron benefit from a close and made
often commensalistic association with sea anemones without adversely affecting gene1
the anemone (Fig. 4.6) . Although they look like plants, anemones are animals vide 1
that use their poisonous tentacles to stun or kill fish. The fish are then drawn in in tim
for digestion. The clownfish are impervious to the poison of the anemone and
feed on detrital material left from the anemone 's meals. In addition, the clownfish cific 1
can escape predators by staying within the tentacles of the anemone. Some proce
clownfish appear to be immune to the poison of one species of anemone but are sites ;
killed when coming into contact with other species. However, other species of defen
clownfish can survive in a relationship with a number of different anemone perha
species. In some cases, the clownfish might serve to attract prey to the anemone, sider
in which case the relationship between clownfish and anemone is mutualistic.
Some forms of mutualism are extremely intimate. For example, lichens, tunat(
which form the splotchy black and yellow growths on rocks and old stone struc- and p
tures, are a symbiotic organism composed of a fungus and a photosynthetic part- There
ner such as algae, cyanobacteria, or both. The fungus is the visible part of the that h
lichen and provides a microenvironment in which the photosynthetic organism host r:
can grow, while the algae or cyanobacteria capture energy through photosynthe- (Mon.
sis. The algal partner would never be able to survive on the rocky substrates, nutrie
favored by many lichens, if the fungus did not provide protection against desicca-
tion. The fungus, in turn, absorbs nutrients from the photosynthetic partner. their ~
cases,
the h(
Hum a
ness. I
region
humar
specie
specie
tolerat
I
its life
ing ho
distrib
impor
dance
in the
beeno
Scotla
that ca
the grc
p
FIGURE 4.6 Species of clownfish (Amprion) are not harmed by the poison of sea
impor
anemones . Some species of clownfish are immune to the poison of only one or two species presen
of anemones and have a similar geographic distribution to those anemone species. Other nated
clownfish species can coexist with a number of different anemone species and do not have like bi
a geographic distribution that is tied to just one anemone species. honey
LIFE
disperse seeds is often not
: geographic distribution of
1 benefit from a close and
without adversely affecting
nts, anemones are animals
The fish are then drawn in
oison of the anemone and
s. In addition, the clownfish
=s of the anemone. Some
pecies of anemone but are
However, other species of
ber of different anemone
tract prey to the anemone,
memone is mutualistic.
late. For example, lichens,
rocks and old stone struc-
and a photosynthetic part-
s is the visible part of the
= photosynthetic organism
=rgy through photosynthe-
= on the rocky substrates,
protection against desicca-
lOtosynthetic partner.
I by the poison of sea
on of only one or two species
se anemone species. Other
one species and do not have
ies.
SYMBIOSIS: MUTUALISM, COMMENSALISM, PARASITISM, AND MIMICRY 89
Corals perhaps provide the most striking example of intimate mutualism.
The reef-building organisms in corals are members of the class Anthozoa. They
are related to sea anemones. Many of these species exude tough outer skeletons
made of calcium carbonate that produces the hard reef structure. The Anthozoa
generally harbor symbiotic coralline algae within their tissue. The Anthozoa pro-
vide protection for the algae and benefit from the photosynthate produced. Such
intimate symbionts must, of course, share identical geographic ranges.
The majority of parasites are restricted to one host species. The highly spe-
cific relationship between parasites and hosts is the result of an evolutionary
process whereby hosts evolve chemical or physiological defenses against para-
sites and parasites in turn evolve specialized adaptations that circumvent these
defenses. Upon reflection, one can see how stenophagous herbivorous insects are
perhaps more akin to parasites than predators. Conversely, some ecologists con-
sider parasitism to be a form of predation.
Almost all multicellular organisms, including humans, harbor parasites. For-
tunately, many of these are not lethal. Microparasites include viruses, bacteria,
and protozoa. Macroparasites include parasitic worms, ticks, fleas, lice, and mites.
There are also some interesting plants that are macroparasites. These are plants
that have no photosynthetic tissue and live by securing water and nutrients from
host plants. One example from the forests of North America is the Indian Pipe
(Monotropa uniflora). This small white plant lacks chlorophyll and obtains its
nutrients from the roots of other plants.
Since almost all parasites are very tightly restricted to certain host species,
their geographic ranges are controlled by the distribution of their hosts. In some
cases, the geographic distribution of the parasite may be smaller than the range of
the host. There are several good examples of this among human parasites.
Humans are host to the parasitic protozoans that cause malaria and sleeping sick-
ness. However, the distribution of these parasites is largely confined to tropical
regions. The reason for this distribution is that both parasites are transferred to
humans by a second host. In the case of malaria, the other hosts are certain
species of mosquito. Sleeping sickness is hosted by the tetse fly. The mosquito
species and tetse flies that commonly host malaria or sleeping sickness cannot
tolerate low temperatures and are therefore restricted to warm regions.
Parasites must have hosts that live long enough for the parasite to complete
its life cycle and transfer its progeny to new hosts. Given this dependence on a liv-
ing host, parasites may not be key factors in directly determining the geographic
distributions of host species. However, parasites are probably one of the most
important factors in determining mortality and natality rates, and thus the abun-
dance of natural populations of plants and animals. Interestingly, cyclic variations
in the abundance of parasites and hosts, similar to predator-prey cycles, have
been observed in nature. Red grouse (Lagopus scoticus) in northern England and
Scotland are afflicted with a parasitic nematode worm (Trichostrongylus tenuis)
that causes death or decreased fertility. It has been observed that populations of
the grouse and the nematode rise and fall on a five-year cycle.
Prior to concluding our discussion of parasites, we should consider one
important, and extremely sad, case of geographic range being controlled by the
presence of a parasite. The native bird fauna of the Hawaiian Islands is domi-
nated by a family of birds called honeycreepers (Drepanididae ). They are finch-
like birds, of which some 30 species are known from the islands. The Hawaiian
honeycreepers evolved in isolation and are found nowhere else in the world.
90 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

During the end of the twentieth century, naturalists noticed a steady decline in
the honeycreeper population. On the island of Oahu six species became extinct
by 1900. It was noted that species found on mountains above 600-m elevation
persisted, while the lowland honeycreepers became extinct. Eventually, it was
shown that the honeycreepers were being killed by a form of avian malaria car-
ried by mosquito species, Culex pipens fatigans. The mosquito had likely been
introduced to Hawaii in the 1820s in water carried on European sailing ships. Not
having evolved with this parasite, honeycreepers have no defense against it. For-
tunately, the mosquito is a tropical species and cannot physiologically tolerate the
cool temperatures of the Hawaiian mountains. Some honeycreepers have there-
fore escaped extinction but are limited in their geographic distribution to eleva-
tions above 600 m. Most species of honeycreeper that were restricted to the
lowlands have been lost forever.
One particularly interesting form of biological interaction is mimicry in ays below water
which one species evolves the appearance or behavior of another species. In
many cases, the mimic species may not interact directly but may share common
predators with which they both interact. Muellerian mimicry occurs when a
species that is poisonous or unpalatable to predators possesses the same coloring
or shape as another species that is also poisonous or unpalatable. Both species FIGURE 4 .
benefit since predators avoid either species. A striking example of the role of stellatus in
Muellerian mimicry in geographic distributions is provided by the ranges of dif- factors (de:
ferent races of the butterfly species Heliconius melpomeme and Heliconius erato of the two I
from South America. Both species are unpalatable to predators. Each species has
a number of different races that are discernible by differences in wing coloration.
The different races of the two species have almost completely overlapping ranges
with the similarly colored races of the other species. In this case, both species northwest
benefit since predators, recognizing the coloring of either species, avoid eating uppermos
them. The geographic distributions of the races appear to reflect this. although 1
Batesian mimicry occurs when one species that is not poisonous or unpalat- balanoide
able has the same coloring or shape as a species that is poisonous or unpalatable. can occur
The palatable species benefits from coexisting with the unpalatable species anoides g1
because predators mistake it for the other species and avoid it. This can be con- species oc
sidered a form of commensalism. A good example of the potential influence of ditions of
Batesian mimicry on geographic distributions comes from the biogeography of and drier
poisonous coral snake (Micrurus) species and nonvenomous colubird snake the mean
(Pliocercus) species in Central America. The different colubird species possess algae and
coloring that mimics the markings of the coral snakes that live in the same geo- of Balanu.
graphic region. species rei
tion and b

COMBINED PHYSICAL AND BIOLOGICAL

CONTROLS ON DISTRIBUTION 310LOGICAL II
RADIENTS,'
Here and in Chapter 3, we have focused on cases where individual physical or
biological factors appear to be important in controlling the geographic distribu- Based on
tions of certain species. However, for the vast majority of nonsymbiotic species certain tyj
the actual geographic range limits are probably controlled by a combination of tant in cor
several physical and biological factors. This is illustrated by a classic study of life and comp•
on the rocky intertidal zone of Scotland. Two species of barnacles, Balanus bal- this chaptt
anoides and Chthamalus stellatus, are common on rocky intertidal shorelines in into the er
tF LIFE
; noticed a steady decline in
u six species became extinct
:ains above 600-m elevation
e extinct. Eventually, it was
a form of avian malaria car-
e mosquito had likely been
t European sailing ships. Not
re no defense against it. For-
t physiologically tolerate the
~ honeycreepers have there-
raphic distribution to eleva-
that were restricted to the
:1.1 interaction is mimicry in
1vior of another species. In
ctly but may share common
an mimicry occurs when a
possesses the same coloring
>r unpalatable. Both species
:ing example of the role of
ovided by the ranges of dif-
Jmeme and Heliconius erato
' predators. Each species has
'ferences in wing coloration.
npletely overlapping ranges
s. In this case, both species
either species, avoid eating
1r to reflect this.
is not poisonous or unpalat-
is poisonous or unpalatable.
th the unpalatable species
td avoid it. This can be con-
>f the potential influence of
s from the biogeography of
1Venomous colubird snake
:nt colubird species possess
:s that live in the same geo-
·here individual physical or
ng the geographic distribu-
ity of nonsymbiotic species
rolled by a combination of
ted by a classic study of life
s of barnacles, Balanus bal-
tcky intertidal shorelines in
BIOLOGICAL INTERACTIONS, GRADIENTS, AND NICHES 91
Balanus Chthamalus
Desiccation
• • ··ays below water Predation Intraspecific I Adults larvae IPredatio.n Interspecific
by Thais competition by Tha1s competition
with Balanus
Distribution Limiting factors Distribution Limiting factors
FIGURE 4. 7 Distribution of the barnacle species Balanus balanoides and Chthamalus
stellatus in the intertidal zone ofthe Scottish seacoast. The relative effects of physical
factors (desiccation) and biological factors (competition and predation) on the distributions
of the two barnacle species are indicated (after Connell, 1961 ).
northwestern Europe. The adult Chthamalus stellatus is always found in the
uppermost portion of the intertidal zone when the two species occur together,
although the larvae can also be found at greater depths (Fig. 4.7). When Balanus
balanoides is removed from the rocks, it has been found that Chthamalus stellatus
can occupy both the upper and lower parts of the intertidal zone. Balanus bal-
anoides grows very quickly and smothers out Chthamalus stellatus when the two
species occur together. However, Balanus balanoides cannot tolerate the dry con-
ditions of the upper intertidal zone. Chthamalus stellatus can survive at higher
and drier sites than Balanus balanoides but dies from desiccation at sites above
the mean spring high-water level. In the lower intertidal zone, competition with
algae and predation by the snail Thais lapillus determines the lower range limits
of Balanus balanoides. Thus, the simple spatial distribution of these two barnacle
species reflects a combination of physical control through intolerance of desicca-
tion and biological controls through competition and predation.
OLOGICAL INTERACTIONS,
DIENTS, AND NICHES
Based on the examples we have looked at in this chapter, we can conclude that
certain types of biological interactions, such as symbiosis, are extremely impor-
tant in controlling geographic ranges, while other interactions, such as predation
and competition, are more variable in their impact on distributions. To conclude
this chapter, we might look at how these biological interactions are incorporated
into the environmental gradient and niche concepts.
 92 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

u In the cases of predation and symbiosis, we can consider the abundance of

prey or hosts as being part of the environmental variability that determines the
carrying capacity of a site. Just as we might have a gradient for soil moisture and
alti
of 1
pia:
incorporate this gradient as an axis in defining the niche of a species, so too we \ \ 'O J
might have a gradient and niche axis representing prey density. In some cases, we pet
might also consider incorporating competition in a similar manner. However, it is tha
very difficult to define the actual impact of competition in a numerical manner !S ill
and make an appropriate gradient or niche axis to represent it. The impact of res<
competition has been an area of special interest and accommodation within the- poe
ories of gradient distributions and the niche. gis t
The impact of competition on the distribution of species is often studied by na t
examining the abundance of different species along environmental gradients. If
we graph the population density of several species along an environmental gra- por
dient and find that the distribution of one species is abruptly truncated by tior
another species (Fig. 4.8), we might conclude that competition between the two spe,
species leads to an abrupt replacement of one species by a competitor. Alterna- pet
tively, we might find a high degree of overlap between the distributions of the ace
species. We could then assume that competition between the species is not an call
important factor in controlling their distribution. Analyses of the elevational nicl
distribution of dominant tree species in the Great Smoky Mountains of east- peti
ern North America has been used to study competitive interactions of tree Pllll
species along an environmental gradient. In this case, elevation represents a par.
gradient of increasing moisture and decreasing temperature from low to high hav
the
tior
nicl
High

i?:' tior
"Cii Ho·
c:
Q)
Competing species
"0 sue]
c:
0 con
~
::;
a.
hab
~
due
Low
trib
Low Air Temperature High soil

High
High
i?:'
'Cii
c: Non-competing species
Q)
"0
c:
0
~
::;
a.
~
Low ./

Low High Low

Air temperature
FIGURE 4.8 Hypothetical gradient distribution of four com peting species and four
noncompeting species. FIG I
. LIFE
consider the abundance of
iability that determines the
1dient for soil moisture and
iche of a species, so too we
y density. In some cases, we
nilar manner. However, it is
ion in a numerical manner
represent it. The impact of
1ccommodation within the-
f species is often studied by
~nvironmental gradients. If
ong an environmental gra-
; is abruptly truncated by
mpetition between the two
s by a competitor. Alterna-
:en the distributions of the
ween the species is not an
.nalyses of the elevational
Smoky Mountains of east-
~t itive interactions of tree
1se, elevation represents a
perature from low to high
:ies
High
>pecies
~
High
peting species and four
BIOLOGICAL INTERACTIONS, GRADIENTS, AND NICHES 93
altitudes in the mountains. The data suggest that there is a gradual replacement
of tree species along the elevational gradients and that competition does not
play an important role in these distributions. Some ecologists, particularly those
working with plant communities, feel that there is not much evidence that com-
petition is a strong force in the modern ranges of plant species. They consider
that because competition has a negative impact on both species involved, organ-
isms should evolve in such a way as to avoid or reduce direct competition for
resources. The perceived lack of competition in modern communities has been
poetically referred to as the "ghost of competition past." However, not all ecolo-
gists agree with the conclusion that competition is unimportant in controlling
natural distributions.
The impact of competition on the niches of species is often examined by
portraying competing species within the same niche space (Fig. 4.9). Competi-
tion should be manifest as a reduction in the niche breadth of one or both
species where they overlap. The broad niche of a species in the absence of com-
petition is referred to as its fundamental niche. The more restricted niche that
occurs if competition excludes the species from certain portions of niche space is
called the realized niche. Within the natural environment, the fundamental
niche would define all the environmental conditions and resources used if com-
petition did not exist. In theory, no two species should have completely overlap-
ping niches. This is called the competitive exclusion principle and can be
paraphrased as "complete competitors cannot coexist." That is, if two species
have absolutely identical requirements and relationships to the environment,
the high degree of competition between them will drive one species to extinc-
tion. Ecologists have long sought to understand how much differentiation in
niche is required for species to coexist.
It might be concluded that competition can exert a control on distribu-
tions, particularly among closely related species or members of the same guild.
However, it is also likely that small-scale heterogeneity in the environment,
such as differences in slope, exposure, and soil type might allow for potentially
competing species to coexist geographically by taking advantage of different
habitat types.
An interesting study of competition between pine species has been con-
ducted in the Sierra Nevada Mountains of California. The study looked at the dis-
tributions of closely related pines on sites that had similar slope, exposure, and
soil type. By controlling for these habitat factors, the researcher was able to study
High
~
:::l
"'
'6
E I
'6 Realized'
(/)
niche A '
Low High
Low Air temperature
FIGURE 4.9 The realized and potential niches of three hypothetical plant species.
94 CHAPTER 4 BIOLOGICAL INTERACTIONS AND THE DISTRIBUTION OF LIFE

how the pines were distributed along an elevational gradient, without local habi- c um. A. (1991). A
tat differences influencing the study. In this case, increasing elevation represented r.ary Ecology. Bla
_ ~ London.
a gradient of increasing moisture and decreasing temperature. The study found
- -ell. J. H. (1961). Tl
that when one looked only at sites with similar slope, exposure, and soil type,
::f..:: ompetition an<
there was an abrupt replacement of pine species by other pine species along the --:oution of the barr
elevational gradient. However, if the distributions of pines on different slopes, ..:~-ology 42,710-723
exposures, and soil types are included in the analysis, the actual elevational -e ll . J. H. (1980). D
ranges of the pines are seen to overlap greatly. The elevational ranges of the pines ~ of competitors,
overlap because they are able to coexist by taking advantage of different habitat. past. Oikos 35,
In the context of geographic distributions, we expect that when species face sig- -ell . J. H. (1983). 0
nificant competition they will have more restricted or fragmented distributions e importance of i1
than when they do not face competition. One potential example of this is the dis- _ -: j ence from field •
tribution of the yellow warbler (Dendroica petechia). In North America this -.=.<tralist 122, 661-t
~- o r. E. F. , and Simt
insectivorous bird is found in a wide variety of habitats and geographic locations.
However, the bird also ranges into the tropics of Central and South America. In .,_::s.emblage of speci<
_.;::;:Jpetition. Ecolog
these tropical regions, the yellow warbler is restricted to coastal sites and islands
--~ste r , J.P. , and La1
formed by mangrove trees. It is possible that competition from the large number :- ~ :mla tion model fc
of tropical insectivorous bird species excludes the yellow warbler from other .xxl plant, ragwort.
habitat types within the tropical portion of its range. ~--o logy 48, 143-164
- ond, J. M. (1975) ..
=: ::.nities. In Ecolog}
:.~.n i ries (ed. M. L. C
KEY WORDS AND TERMS ~- 342-444. Belkna1
:::-sity Press, Cambr
Allelopathy Interspecific competition Parasites : n. A. P. , and Hun
Batesian mimicry Intraspecific competition Parasitism :_oa and stability of
Commensalism Keystone species Predation •_.-s1em: Trichostron[5
Competitive exclusion Lotka-Volterra model Realized niche ;:-ouse.II. Populati01
principle Muellerian mimicry Stenophagous -.J/ Ecology 61,487
Euryphagous Mutualism Symbiosis ~ - -..: .:: b . P.R. (1961). In
Fundamental niche Optimal foraging theory :~ in the checkersp(
- - :J 109.
~ -- ..: .::h . P.R. (1965). Th
:..:e butterfly Euphyc
. ..:~e of the Jaspar Ri
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radient, without local habi-
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Wh
that
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ban-
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rare
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5
CHAPTER
DISTURBANCE
In 1988 a huge series of fires burned almost half of Yellowstone National Park.
Thousands of hectares of forest were destroyed, and countless animals were either
killed or driven from their original habitat. Despite the millions of dollars spent to
fight this fire, and the damage it caused, biogeographers recognize that fires are a
natural part of the Yellowstone ecosystem. Today the areas that were burned and
apparently left lifeless in 1988 are alive with hundreds of species of wildflowers,
shrubs, and tree seedlings. The bird and mammal populations of the park are thriv-
ing. Rather than long-term destruction, the effect of the 1988 fire has led to rejuve-
nation. Biogeographers recognize that we cannot understand the functioning and
distribution of organisms without understanding the role of natural disturbance.
Almost all ecosystems are prone to periodic disturbances such as the Yel-
lowstone fires and, in many cases, depend on disturbance for long-term function-
ing. The term disturbance is used to describe short-term physical or biological
events that significantly alter ecosystems. These events include fires, wind storms,
landslides, floods, avalanches, and outbreaks of diseases or pests. Rather than
being thought of as unnatural events that act to destroy ecosystems, disturbances
are generally natural and vital parts of the environment and important for the
long-term functioning of ecosytems. The frequency and impact of disturbances
vary widely between and within ecosystems. Giant sequoia (Sequoia gigantea)
forests of the southern Sierra Nevada Mountains of California experience fires in
the low vegetation of the ground surface as frequently as once every seven to
nine years. These fires kill ground vegetation and seedlings but rarely damage
mature sequoia trees. In contrast, hurricanes may only impact the forests of the
southeastern Great Lakes region less than once every century but can destroy
most of the tree canopy in their paths when they do occur.
A useful formal definition of disturbance and its impact is provided by
White and Pickett (1985): "A disturbance is any relatively discrete event in time
that disrupts ecosystem, community, or population structure and changes
resources, substrate availability, or the physical environment" (p. 7). They suggest
that disturbances can be described in terms of cause, frequency, predictability,
spatial extent, magnitude of impact on ecosystems, synergism with other distur-
bances, and seasonal timing. Some disturbances, such as small fires generated by
lightning, may frequently occur at almost random intervals, while other distur-
bances, such as outbreaks of pathogenic insects, appear to occur at more regular
intervals. Volcanic eruptions and other catastrophic disturbances may be very
rare and unpredictable. Some disturbances are linked together in synergistic rela-
tionships. For example, in many mountainous regions fires destroy vegetation and
produce unstable soils that lead to landslides and further vegetation disturbance.
The adjustments and changes that occur in ecosystems following distur-
bances have long been topics of great interest to biogeographers and ecologists.
Let's start by considering some of the classic terms and concepts applied to the
97
98 CHAPTER 5 DISTURBANCE

recovery of an ecosystem following disturbance. We will then reexamine these water lily
concepts in light of more recent research. Succession is the term used to describe tiona] sed
the changes in physical and biological conditions that follow disturbances. There sediment
are two major types of succession. Primary succession occurs when a previously willows(~
lifeless surface is first colonized by plants and animals. Specifically, it takes place enough to
when new islands or land surfaces are formed by volcanic activity, when sea-level cessions b
changes or shoreline deposition create new land surfaces, or when glaciers melt secondary
and expose new areas of land. Secondary succession occurs when an existing event, anc
ecosystem recovers from a disturbance such as fire or flood. The
In some cases, species that colonize a disturbed site change the environ- succession
ment and allow establishment of yet other species. For example, the establish- Clements
ment of a tree canopy during succession provides a cool, moist, and shady research o
environment that supports forest floor species. The process by which the estab- worked w
lishment of one species changes the environment and allows the subsequent was much
establishment of other species is called facilitation. of the coe
The process of succession is often divided into different stages. A typical Clements'
sequence of these stages for a given vegetation type is called a sere. Seral stages facilitatior
can be observed when agricultural fields in the southern Great Lakes region are caused by
allowed to return to a forested state (Fig. 5.1). The initial successional vegetation communit
consists of annual grasses and herbs such as quackgrass (Agropyron repens) and the absenc
ragweed (Ambrosia artemisiifolia). Within the first 10 years, perennial herbs and and these
grasses such as Kentucky bluegrass (Poa p ratensis) dominate the site. By 40 years, disturbanc
there is a marked increase in dominance by woody shrubs such as roses (Rosa mined laq
spp.) and bramble (Rubus spp.). The seedlings and saplings of white pine (Pinus northeastE
strobus) and red and white oak (Quercus rubra and Q. alba) become more domi- ofmid-ele
nant during this time. If no further disturbance occurs, a mixed oak and pine forest Asr
will occupy the former agricultural field within 100 years. The final vegetation type original te
developed during succession is sometimes referred to as a climax community. climax co1
A special type of succession is recognized when ponds and wetlands fill in models of
with sediment and become terrestrial environments. This process is referred to as
hydroseral or hydrarch succession. As water depths in ponds and lakes decline Toleran
due to natural infilling by mineral sediment and organic debris, plants such as are e
thee
grow
domi
Canopy ment
longt
Inhibiti•
Lower
canopy bane
trees domi
tors 1
Understory OCCUJ
trees
inhib
Tall shrub Randou
understory
simp
Annuals Perennials Shrubs Young forest Mature forest randt
Time-
FIGURE 5.1 An idealized "classical" secondary succession on an abandoned agricultural Man
field in the North American Midwest. cated and

will then reexamine these
> the term used to describe
follow disturbances. There
1 occurs when a previously
. Specifically, it takes place
nic activity, when sea-lev~
tees, or when glaciers melt
1 occurs when an existing
'food.
I site change the environ-
or example, the establish-
a cool, moist, and shady
·ocess by which the estab-
nd allows the subsequent
different stages. A typical
called a sere. Sera! stages
rn Great Lakes region are
ial successional vegetation
;s (Agropyron repens) and
years, perennial herbs and
1inate the site. By 40 years,
1rubs such as roses (Rosa
'lings of white pine (Pinus
alba) become more domi-
mixed oak and pine forest
s. The final vegetation type
s a climax community.
ponds and wetlands fill in
tis process is referred to as
1 ponds and lakes decline
mic debris, plants such as
Canopy
Lower
canopy
trees
Understory
trees
Tall shrub
JO yrs. understory
eforest
m an abandoned agricultural
DISTURBANCE 99
water lily (Nuphur spp.) and cattail (Typha spp.) can root in the mud, trap addi-
tional sediment, and produce increased organic debris. Eventually, the level of
sediment rises to the point where it can be colonized by terrestrial plants such as
willows (Salix spp.) and alders (Alnus spp.). Over time, the site will become dry
enough to support forest cover. Hydroseral successions are not true primary suc-
cessions because there is a preexisting aquatic ecosystem at the site. Nor are they
secondary successions because the process of infilling is a long-term progressive
event, and not the result of a discrete disturbance.
The seral successional patterns outlined above follow the classic model of
succession as proposed by the American ecologists Henry Cowles and Frederic
Clements at the beginning of the twentieth century. Cowles conducted much of his
research on the primary succession that occurs on newly stable beaches. Clements
worked with secondary succession in deciduous forests and grasslands. Clements
was much influenced by the writings of Darwin on evolution and the potential role
of the coevolution of species in the development of modern flora. According to
Clements' model, seres are highly predictable sequences that are controlled by
facilitation. The organisms in the later seres depend on the environmental changes
caused by the establishment of the earlier invading plants and animals. The climax
community is the predictable end product of succession and is self-perpetuating in
the absence of further disturbance. In a sense, the climax community is the norm,
and the sera! stages are anomalies in nature caused by the destructive impact of
disturbances. The exact nature of the climax community was thought to be deter-
mined largely by regional climate. A deciduous forest was the natural climax in
northeastern North America, whereas a coniferous forest was the natural climax
of mid-elevation sites in the Rocky Mountains.
As more data have been collected on disturbance and succession, some
original tenets of the Clementsian model of facilitative succession followed by
climax conditions have been questioned and often refuted. Several alternative
models of succession have been proposed:
Tolerance Model In many instances, species of both early and late seral stages
are established very quickly following disturbance. The plants that dominate
the early successional period simply are those that grow fastest. The slower
growing plants that dominate the later stages eventually displace the early
dominants. All of the species can tolerate the early successional environ-
ments, but as the succession progresses, some of the early dominants can no
longer tolerate the increased competition for light and moisture.
Inhibition Model In some ecosystems, particularly following small distur-
bances, it is very difficult to predict which species will invade and eventually
dominate a site. It appears that proximity to seed sources and chance fac-
tors related to seed dispersal play a major role in determining which species
occupy the site. Once the site is occupied, the presence of the early invaders
inhibits establishment of other species.
Random Model Finally, it has been suggested that succession can be typified
simply as the random arrival and survival of species at a disturbed site. This
random model assumes that there is no facilitation or inhibition.
Many studies have shown successional pathways to be much more compli-
cated and unpredictable than the unidirectional seres leading to a stable climax.
1 00 CHAPTER 5 DISTURBANCE

For example, much work has been conducted on the primary succession that
occurs as glaciers have melted and exposed land surfaces at Glacier Bay, Alaska.
Since the rate of glacial retreat is known, the ages of different land surfaces can
be estimated. By comparing the vegetation on young surfaces, intermediate-aged Tree-ril
surfaces, and old surfaces, the temporal pattern of succession may be recon- Tree-ring
structed. This technique of using different aged patches of the landscape to bance an
reconstruct a general pattern of succession is called the chronosequence analysis. '
approach and is widely used. The chronosequence approach is often referred to Manyco1
as a "space for time" method of inferring succession patterns. climates
Based on chronosequence data, it was initially thought that the sites on annual ri
Glacier Bay represented a classical case of facilitative succession through pre- wood. W
dictable seres reaching stable climax. On sites deglaciated within the last 20 years, light-cole
the vegetation on the rocky surfaces includes rock mosses (Racomitium growth s1
canescens and R. lanuginosum), herbs such as fireweed (Epilobium latifolium) cells and
forests, \\
and mountain avens (Dryas drummondii), and small shrubs such as arctic willow
tiona! xy
(Salix arctica). Slightly older sites are dominated by larger species of willow and
xylem gr
mountain alder (Alnus crispa). The alder may form large, almost pure thickets on late-grov
sites that are 50 years old. On still older sites, alder thickets are replaced by cot- of winter
tonwood trees (Populus trichocarpa). Sitka spruce (Picea siitchensis) eventually when an
comes to dominate the vegetation on sites that have been ice-free for 120 years. and late '
By 200 years, a forest dominated by sitka spruce and western hemlock (Tsuga wood to !
heterophylla) is established. The importance of alders in succession at Glacier found in
Bay has been of particular interest because their roots contain nodules that host of the tn
symbiotic bacteria. The bacteria have the ability to transform nitrogen from the side edgt:
atmosphere into nitrate, which is required by plants. It has been argued that the inside ed
nitrogen-fixing ability of the alders made them important facilitators of further If you
succession by coniferous trees. centric c
wood is
More recent work shows that the unidirectional successional model devel-
outermo
oped from the chronosequence studies at Glacier Bay is oversimplistic. Research
injure tr•
has been conducted using tree-ring analysis to examine the actual ages and year of By coun
establishment of woody plants on the different sites around Glacier Bay. Contrary years car
to earlier conclusions, the tree-ring-based study shows that sitka spruce invaded cal tech1
the older sites very quickly after deglaciation and did not appear to require facili- which dt:
tation by the nitrogen-fixing alders. In fact, alder appears to have been important cross-da1
in the successional vegetation of only some of the sites. The early invasion of sites mate the
by spruce appears to be positively correlated with the proximity of spruce stands. vide am
Sites that were near stands of spruce received high seed falls and were more or other
quickly dominated by spruce following deglaciation. Cottonwood is only an became
important component in the seral vegetation at some of the sites, notably those Some
deglaciated in the past 100 years. Finally, although spruce and hemlock form a cli- fires wit
kill a pa1
max community on some sites, on poorly drained locations Sphagnum moss and
tually m
shore pine (Pinus contorta ssp. contorta) eventually replace this forest. Thus, the
produce
primary succession at Glacier Bay follows multiple pathways and can lead to very gists can
different seral and climax communities. Because stochastic factors such as proxim- aged car
ity of seed sources and environmental factors such as slight differences in drainage pondero
or soil type can be very important in determining successional communities on dozens c
different sites, the results of chronosequence studies must be interpreted with for fire
great caution. For this reason, tree-ring analysis of the long-term history of indi- be recon
vidual sites has become increasingly important for reconstructing vegetation dis-
turbance and succession in forested ecosystems (Technical Box 5.1).

te primary succession that
tees at Glacier Bay, Alaska.
different land surfaces can
;urfaces, intermediate-aged
succession may be recon-
tches of the landscape to
1lled the chronosequence
proach is often referred to
at terns.
thought that the sites on
1e succession through pre-
ted within the last 20 years,
JCk mosses (Racomitium
~ed (Epilobium latifolium)
hrubs such as arctic willow
trger species of willow and
ge, almost pure thickets on
ickets are replaced by cot-
'icea siitchensis) eventually
'een ice-free for 120 years.
j western hemlock (Tsuga
·s in succession at Glacier
; contain nodules that host
msform nitrogen from the
t has been argued that the
·tant facilitators of further
successional model devel-
is oversimplistic. Research
the actual ages and year of
1und Glacier Bay. Contrary
; that sitka spruce invaded
ot appear to require facili-
trs to have been important
The early invasion of sites
xoximity of spruce stands.
seed falls and were more
. Cottonwood is only an
of the sites, notably those
ce and hemlock form a cli-
tions Sphagnum moss and
place this forest. Thus, the
1ways and can lead to very
stic factors such as proxim-
ght differences in drainage
cessional communities on
must be interpreted with
long-term history of indi-
nstructing vegetation dis-
eal Box 5.1).
DISTURBANCE 10 1
TECHNICAL BOX 5.1
Tree-ring Analysis, Stand History and Fire Reconstruction
Tree-ring analysis is one of the most widely used techniques to study forest distur-
bance and succession in nontropical regions. Many biogeographers use tree-ring
analysis. The science of dating past events by tree-rings is called dendrochronology.
Many coniferous and dicotyledonous trees growing in areas with seasonally variable
climates produce annual growth rings. Monocots, such as palm trees, do not produce
annual rings. In most conifers, the rings are composed of couplets of light and dark
wood. Wood consists of the xylem cells (water-conducting tissue) of the tree. The
light-colored wood is produced by rapid xylem growth at the start of each year's
growth season. The light-colored wood is composed of thin-walled and large xylem
cells and is called early wood. The start of the growth season might be the spring in
forests, which experience cold winters. As the tree grows during the summer, addi-
tional xylem cells form and add to the width of the ring. As the autumn approaches,
xylem growth slows, and the cells are smaller and have thicker walls. These dense,
late-growth cells are generally dark in color and are called late wood. With the onset
of winter, the tree becomes dormant, and xylem growth ceases until the next spring
when a new ring couplet begins to grow. Somewhat similar couplets of early wood
and late wood can be detected in dicotyledonous trees, but the transition from early
wood to late wood is often not as clear. The only living cells in the trunk of a tree are
found in this narrow tree-ring-forming zone just beneath the bark. This living zone
of the trunk is called the cambium. The phloem cells that form each year at the out-
side edge of the cambium are crushed outward by the growing xylem formed at the
inside edge and are incorporated into the bark.
If you cut down a tree with annual rings and look at the trunk you will see con-
centric circles of couplets of early wood and late wood. The innermost point of
wood is called the pith. The pith represents the tree's first year of growth. The
outermost ring represents the most recent year of growth. So as to not kill or
injure trees, special small corers are generally used to obtain tree-ring samples.
By counting the rings from the pith to the outermost ring, the age of the tree in
years can be estimated. Dendrochronologists use a number of checks and statisti-
cal techniques to verify that the rings they count are annual. The process by
which dendrochronologists measure, count, and assign an age to a ring is called
cross-dating. By cross-dating the rings from many trees in a stand, one can esti-
mate the age of the patch that the stand comprises. The age of the trees can pro-
vide a minium estimate of the last time a major flood , avalanche, wind storm, fire,
or other disturbance created the gap in which the present trees of the patch
became established.
Some types of tree, such as many species of pines, can preserve evidence of past
fires within their tree-rings. Surface fires often burn away portions of bark and
kill a part of the cambium. The dead cambium does not recover but can be even-
tually overgrown by living cambium at its edges. These areas of dead cambium
produce fire scars that are preserved in the tree-ring record. Dendrochronolo-
gists can cross-date the rings prior to the scar and the rings formed by undam-
aged cambium around the scar to estimate the age of the fire scar. Trees such as
ponderosa pine (Pinus ponderosa) from western North America can preserve
dozens of scars from individual surface fires. By sampling a large number of trees
for fire scars, spatially and temporally detailed histories of past fire activity can
be reconstructed.
1 02 CHAPTER 5 DISTURBANCE

Glacier Bay represents a relatively simple system. An example of greater com- As a

plexity in multi-path successional sequences is provided by primary succession on now shown
sand dunes along the shores of Lake Michigan. The level of the lake relative to selves in tl:
the land has been falling over the past 12,000 years. As a result, chronosequence shading by
studies of ancient dune surfaces and newly created ones can provide evidence of species and
successional and climax communities of many different ages. At a typical site, there often requi
is a sequence of vegetation succession that starts with marram grass (Ammophila senescence
breviligulata) colonization of freshly stabilized dunes. Marram grass is a very inter- Clementsia
esting species that is particularly adapted to primary succession on dunes. It seldom great a sim1
reproduces by seeds, and it spreads mainly through underground rhizomes and Over
pieces of leaf and stem. It aggressively colonizes and stabilizes dune surfaces within stages that
a few years after colonization. If further sand accumulates, marram grass grows rap- cession lea<
idly upward and keeps pace with deposition. Interestingly, the grass seems to die out or greater s
naturally after the dune's surface is stabilized and after deposition of new sand trum of ecc
ceases. The stabilized dunes are eventually occupied by jack, red and white pine of Alaska i~
(Pinus banksiana, P. resinosa, P. strobus) forest within 100 to 200 years. Finally, decid-
uous forest dominated by black and white oaks (Quercus velutina, Q. alba) and/or
sugar and red maple (Acer saccharum, A. rubrum) occupy the site. However,
because of very small differences in initial topography and drainage conditions, a
number of other successional pathways can be identified, and at least six different
"climax" vegetation communities can come to dominate individual sites (Fig. 5.2).

Cottonwood

Years stable 350 600 850 1,1 00

Time----
... ------ ~~

Ql
Initial
E
i= conditions

l processes

Pioneer
vegetation

Conifers
(mostly
temporary)
FIGURE 5.3
coniferous fc
"Climax" different age
dominants imparted by
outline of ea<
FIGURE 5.2 The multiple successional pathways evident for succession on sand dunes vegetation a1
along the shoreline of Lake Michigan (after Olson, 1958; Krebs, 1959, and Krebs, 1994). was burned

/
\n example of greater com-
1by primary succession on
:vel of the lake relative to
s a result, chronosequence
:s can provide evidence of
:J.ges. At a typical site, there
narram grass (Ammophila
arram grass is a very inter-
;ession on dunes. It seldom
nderground rhizomes and
'ilizes dune surfaces within
:s, marram grass grows rap-
/, the grass seems to die out
:r deposition of new sand
1 jack, red and white pine
to 200 years. Finally, decid-
tS velutina, Q. alba) and/or
•ccupy the site. However,
and drainage conditions, a
j , and at least six different
individual sites (Fig. 5.2).
~
3es m tt m
635 3 193 .12
620 189 ~
600 183 ..§!
580 177 w
10,000- 12,000-
succession on sand dunes
1959, and Krebs, 1994).
DISTURBANCE 1 03
As a final point regarding classical succession theory, many studies have
now shown that so-called climax communities cannot in fact propagate them-
selves in the absence of disturbance. This is particularly true in forests, where
shading by the mature canopy has been shown to exclude both the early seral
species and many of the so-called climax trees. Regeneration in these instances
often requires the formation of canopy gaps due to external disturbances or the
senescence and mortality of stand dominant trees. In short, although the
Clementsian model of facilitative succession and climax is elegant, it is far too
great a simplification and too often misleading.
Over the years, many properties have been attributed to the various seral
stages that occur during succession. It has been suggested that the process of suc-
cession leads invariably to higher biomass, higher growth rates, higher diversity,
or greater stability. Such generalizations cannot really be applied to a broad spec-
trum of ecosystems. For example, plant species diversity in the coniferous forests
of Alaska is often highest during the early stages of succession. During this time,
FIGURE 5.3 Different-aged patches that make up a portion of the Rocky Mountain
coniferous forest in Alberta. The map shows how the environment is made up of patches of
different ages and sizes caused by disturbances. In this case, most of the patchiness is
imparted by different forest fires that have occurred over the period 1590 to 1969. The
outline of each patch and the age of the fire that created it are indicated on the map. The
vegetation and microclimate of a recent patch are very different from those of a site that
was burned hundreds of years ago (after Johnson and Larsen, 1991 ).
1 04 CHAPTER 5 DISTURBANCE

annual and perennial grasses, herbs, and shrubs are intermixed with tree seedlings compos
and small saplings. The biomass of small mammal communities is highest in these which r
early successional communities and decreases as shrubby vegetation and conifer- climb f1
ous forest are established. In contrast, the species diversity of small mammals is limbs tl
highest during the period when shrubby vegetation is dominant. such as
Biogeographers are often interested in disturbances because the landscapes genus a
they study consist of a mosaic of patches that represent sites at different stages of Some p
successional recovery from disturbance. When you stand atop a mountain and look other v•
out upon the forest below, you are seeing a patchwork of vegetation made up of than otl
units of different sizes and ages that were created by disturbance-generated gaps C. Jack
(Fig. 5.3). Some gaps, such as those created by large fires, are huge in size, while oth- forests
ers, perhaps created by the wind toppling a single tree, are quite small. The physical associat
and biological environments within gaps of varying ages and sizes can be quite dif- (Betula
ferent. For example, a large area that has been recently burned experiences signifi- nated b
cantly higher light intensities, higher air temperatures, and lower humidity than to struc
adjacent forested sites. Some disturbances and the patch environments they create enhanc•
are detrimental to certain species and thus limit geographic distributions. Fre- drying<
quently, disturbances create patches that allow some species to persist in areas 11
where they might not otherwise be found. A closed forest canopy does not provide determi
enough insolation at ground level for many light-demanding herbs and grasses. A has bee
recently opened gap in the forest provides the proper physical environment for grass bi'
such plants and allows them to persist. Thus, biogeographers who try to understand sive or i
the distribution of species without considering the impact of disturbance and suc- small m
cession do so at great risk. In this chapter we will examine some common forms of branchE
physical and biological disturbance, associated successional patterns, and the influ- one-thi1
ence of disturbance on geographic distributions. of livin~
of subst
tion reg

-
FIRE

Fire is the most pervasive and well-studied form of disturbance in terrestrial envi-
lands, s
within •
souther
ronments. Wildfires have been recorded at places ranging from arctic Alaska and intense
Canada to the depths of the South American tropical rainforests. Despite expen- grasslan
ditures of millions of dollars on fire prevention and suppression, fire remains a 10 to 20
potent form of disturbance even where protection efforts are employed. For
example, Canada is a world leader in the development and application of equip-
400
ment and techniques to fight forest fires. However, each year in Canada approxi-
mately 10,000 fires occur in coniferous forests and burn approximately 2 million 6
~
ha, about 0.6% of the total forested area of the country. At the other end of the ~
::::J
300
continent, a single fire in April1999 burned approximately 60,000 ha in the Ever- "§
Q)
glades region of south Florida, despite the proximity of the blaze to major cities g- 200
and a large fire-fighting infrastructure. In this section we will look at the behavior, 2
natural occurrence, and impact of fire. We will also examine the interesting fire - E
::::J

related adaptations that some plants possess and consider the positive impacts E 100
·x
ell
that fire has on certain environments. :2
The occurrence of fire requires adequate fuel and an ignition source. The 0 0
fuel for wildfires is provided by vegetation, while the ignition source for most nat-
ural fires is lightning. Physical factors such as topography, climate, and the type of
vegetation that serves as fuel, all influence the behavior of fire. Different types of / FIGURE
vegetation have inherently different fuel characteristics. Both plant structure and Texa s gr

:rmixed with tree seedlings
nunities is highest in these
by vegetation and conifer-
~rsity of small mammals is
iominant.
:es because the landscapes
• sites at different stages of
l atop a mountain and look
of vegetation made up of
isturbance-generated gaps
are huge in size, while oth-
re quite small. The physical
; and sizes can be quite dif-
burned experiences signifi-
and lower humidity than
t environments they create
graphic distributions. Pre-
species to persist in areas
>t canopy does not provide
1ding herbs and grasses. A
physical environment for
1ers who try to understand
tct of disturbance and suc-
'le some common forms of
nal patterns, and the influ-
trbance in terrestrial envi-
ng from arctic Alaska and
1inforests. Despite expen-
lppression, fire remains a
fforts are employed. For
and application of equip-
1 year in Canada approxi-
1 approximately 2 million
y. At the other end of the
ely 60,000 ha in the Ever-
f the blaze to major cities
: will look at the behavior,
tmine the interesting fire-
:ider the positive impacts
d an ignition source. The
tition source for most nat-
y, climate, and the type of
of fire. Different types of
. Both plant structure and
FIRE 105
composition determine the fuel characteristics of vegetation. Trees and shrubs,
which maintain dead lower branches, provide fire ladders that allow flames to
climb from the ground to the canopy. Many species of pine possess dead lower
limbs that serve as fire ladders. The lack of dead branches on self-pruning trees
such as Douglas fir (Pseudotsuga spp.) is one reason why the adult trees of this
genus are not killed by the frequent surface fires that burn through their stands.
Some plants, including pines and eucalyptus, contain high amounts of resins or
other volatile oils and ignite much more easily and burn at higher temperatures
than other plants. Pines, for example, typically burn at temperatures of up to 800°
C. Jack and lodgepole pines (Pinus contorta and P banksiana) of the northern
forests of North America have been found to be much more flammable than
associated deciduous trees such as aspen (Populus tremuloides) and paper birch
(Betula papyrifera). As a result, the chance of high-intensity fires in stands domi-
nated by pine is generally higher than in deciduous or mixed stands. In addition
to structure and chemical composition, the flammability of vegetation can be
enhanced by climatic factors. Low precipitation coupled with strong winds causes
drying of living and dead biomass and greatly increases flammability.
The total amount of flammable biomass available for burning is important in
determining fire occurrence and intensity. For example, a very direct relationship
has been shown between maximum temperature of fires in Texas grassland and
grass biomass (Fig. 5.4). Areas with patchy vegetation are unable to support exten-
sive or intense fires. Many deserts do not experience large fires because plants are
small and sparsely distributed. Dead biomass, in the form of leaf litter or dead
branches, is an important fuel for fires in forested regions. Deadwood can make up
one-third of the standing biomass in some California shrub communities. Removal
of living and dead biomass by fires decreases fuel load and reduces the probability
of subsequent burning. The chance of a severe fire increases over time as vegeta-
tion regrowth leads to an accumulation of living and dead fuel. In annual grass-
lands, such as those found in California, high amounts of fuel can accumulate
within one growing season after a fire. The fire-prone chaparral shrublands of
southern California regain pre-fire biomass within 15 to 20 years following an
intense burn. Post-fire accumulation rates of fuel in forests are often slower than in
grasslands and shrublands. However, in Australian eucalyptus forests, it only takes
10 to 20 years for the highly flammable leaf litter to build up to pre-fire levels.
400
• •
~
~
:::l
300
. :. •• ..,••
••••• • ••
~ • •• •
Q)
g- 200 ••
2 •
E
-~X 10ol • •
<1l
:2
0 0 2 4 6 8
Fine fuel (1000 kg/ha)
FIGURE 5.4 The positive relationship between f uel biomass and fire temperature in
Texas g rasslands (after Stinson and W right, 1969).
1 06 CHAPTER 5 DISTURBANCE

The principal ignition source for natural fires is lightning. At any one time flammal:
as many as 1800 lightning storms may be occurring on earth. Lightning is gener- long per
ated by unstable atmospheric conditions associated with convective storms and that is \\
the uplift of warm air along cold frontal systems. In tropical regions, convective dry win<
storms can occur throughout the year, whereas in the arctic they are extremely intensity
rare at any time. In North America lightning storms are most common in the with flar
warm and humid Southeast and least common in the cool arctic and along the quickly<
Pacific Coast. There are roughly 10 times as many thunderstorms per year in the Be
southeastern United States as there are in the Pacific Northwest. Many of the weather
southeastern storms occur in the summer. In coastal California lightning can region a
develop with winter frontal storms, but there are few convective storms during fires to
the summer. Lightning storms can develop in southern California when warm, wildfire~
monsoonal air flows northward, usually in late summer. 100,000
A lightning strike does not always ignite a fire ; fuel conditions must be right than 10,1
for a lightning strike to create a fire. The conditions that favor ignition include each ye:
prolonged dry conditions, high air temperatures, low humidity, and high winds. A and hot
lightning strike will not ignite a fire in a damp forest . Highly resinous pines are enced a
unlikely to ignite if they have moisture contents that are greater than 35% of Canada.
their dry weights. Even in summer, the relatively moist conditions typical of the were bu
eastern United States preclude ignition of large fires by lightning. Therefore, the lion ha'
number of fires does not necessarily correspond to the number of lightning Th
strikes. The southwestern United States receives frontal precipitation in the win- differen1
ter and monsoonal precipitation in the summer months. The late winter and early forests (
spring tend to be extremely dry. Most fires occur in the months of May and June, sonally <
despite the fact that the highest thunderstorm activity occurs in July and August. through
The winter months experience some lightning but almost no fires because of the regions,
cool and damp conditions. north we
Humans are the leading ignition source in inhabited areas. Even in sparsely atures, a
populated areas, such as Alaska, human activity is an extremely important cause ecosyste
of fires. For the period 1940 to 1978, humans caused 6595 recorded fires, while while pc
lightning ignited 4001 recorded fires. Similarly, in Canada approximately 65% of rior exp1
all forest fires are started by humans. Interestingly, the 35% of fires started by coastal~
lightning account for 85% of the total area burned. This is because most of the more de
human-caused fires are started in areas that are easily accessible to fire fighters the low-
and can be controlled quickly. Fn
If ignition does occur, what determines whether a fire will scorch the ground ing the r
over a few hectares or destroy all vegetation over thousands of square kilometers? and sava
The impact of a fire is determined by its rate of spread and intensity. The rate of relief, ar
spread of a fire is simply the distance it advances during a given amount of time. high wir
Fires spread through the progressive advance of the burning front and through the will exp
ignition of spot fires by flying embers. Fire fronts can advance at rates of several South A
kilometers per hour, while embers can generate spot fires kilometers beyond the to five Y
main fire front. Fire intensity is sometimes described in terms of its impact on the has man
vegetation. In forested settings, low-intensity surface fires destroy ground vegeta- cold, mo
tion while leaving trees intact. Fires that are sufficient to cause tree canopies to Southen
ignite produce high-intensity crown or canopy fires. Fire intensity can also be quite sm
measured in terms of kilowatts of energy generated per meter along the fire front. Plc
Surface fires reach intensities of < 260 kw per square meter, whereas crown fires exhibit a
may reach intensities of > 2800 kw per square meter. Several factors govern rates species I
of fire spread and intensity. First, the vegetation has to provide adequately foliage c
 FIRE 107

ghtning. At any one time flammable and closely spaced fuel to carry a fire. Fuel flammability is increased by
=arth. Lightning is gener- long periods of dry, hot weather. Second, fire spread is promoted by topography
th convective storms and that is without natural fire breaks such as lakes, rocky ridges, and rivers. Finally,
lpical regions, convective dry winds provide an important mechanism to increase the rate of spread and
:trctic they are extremely intensity. Winds drive the fire forward and bring the flame front down into contact
ue most common in the with flammable ground fuels. For the same reasons, fires can also spread more
:ool arctic and along the quickly and burn with greater intensity when moving upslope.
jerstorms per year in the Because fire spread is affected by fuel conditions, topography, and
Northwest. Many of the weather, there is wide variation in the average size of fires from region to
California lightning can region and year to year. In some damp tropical forests, it is not unusual for most
:onvective storms during fires to be restricted to individual trees that are hit by lightning. In contrast,
1 California when warm, wildfires in the coniferous forests of western Canada can frequently exceed
100,000 ha in size. Although only 3% of the fires in Canadian forests are greater
l conditions must be right than 10,000 ha in size, these large fires account for 90% of the total area burned
tat favor ignition include each year. Such extensive fires are generally associated with particularly dry
midity, and high winds. A and hot summers. During certain hot dry years, very high fire activity is experi-
-Iighly resinous pines are enced across a broad belt extending from Alaska to the Atlantic coast of
are greater than 35% of Canada. During the dry hot summer of 1990, for example, about 8 million ha
conditions typical of the were burned. In contrast, during the cool moist summer of 1978, less than 1 mil-
· lightning. Therefore, the lion ha were consumed by fire.
the number of lightning The frequency of fire is also influenced by fuel and climate. There are large
precipitation in the win- differences between the frequency of fires in moist tundra and eastern deciduous
The late winter and early forests (where fires may occur less than once every 300 to 1000 years) and sea-
months of May and June, sonally dry areas found in western North America (where surface fires can burn
·ccurs in July and August. through pine stands and grasslands almost annually). Even within subcontinental
st no fires because of the regions, there are significant differences in fire frequency. For example, in the
northwestern United States, a clear relationship exists between fuel type, temper-
~d areas. Even in sparsely atures, and moisture stress, and the frequency of fires in forest and woodland
:tremely important cause ecosystems (Fig. 5.5). Cool-moist coastal forests experience little fire activity,
595 recorded fires, while while ponderosa pine (Pinus ponderosa) and oak-juniper woodlands in the inte-
ia approximately 65% of rior experience fires every 1 to 25 years. Interestingly, when fires do occur in the
: 35% of fires started by coastal and high-elevation forests of the Pacific Northwest, they are often much
is is because most of the more destructive to the tree cover than the frequent surface burns experienced at
accessible to fire fighters the low-elevation and inland sites.
From this discussion we may draw some geographic generalizations regard-
ire will scorch the ground ing the role of fire as an agent of disturbance. Areas such as subtropical grasslands
ads of square kilometers? and savanna (mixtures of grasses and woodly plants), which have low topographic
and intensity. The rate of relief, are subject to prolonged periods of aridity, low humidity, high temperatures,
a given amount of time. high winds, and frequent lightning storms, and possess fast-growing vegetation,
ing front and through the will experience frequent large fires. In portions of Africa, Asia, Australia, and
ivance at rates of several South America, savanna ecosystems experience fires on an average of every one
es kilometers beyond the to five years. In contrast, the arctic tundra (low herb and small shrub vegetation)
erms of its impact on the has many lakes and rivers that serve as topographic fire breaks; the tundra is also
s destroy ground vegeta- cold, moist, has few thunderstorms, and supports sparse, slow-growing vegetation.
o cause tree canopies to Southern tundra areas burn perhaps once every 1000 years, and fires are generally
re intensity can also be quite small. Most of the northern high arctic almost never experiences fires.
neter along the fire front. Plants that exist in regions that are frequently disturbed by fire often
eter, whereas crown fires exhibit adaptations that allow them to cope with the effects of burning. Some tree
veral factors govern rates species possess fire-resistant bark. Mature trees can be killed by direct burning of
to provide adequately foliage or by lethal heating of the cambium layer beneath the bark. A tree with a
1 08 CHAPTER 5 DISTURBANCE

(Quercus
cormic sp
(Increasing Som
temperature)
that OCCUI
,
buds and •
I 3
~, I atures dm
Coastal Ponderosa grow. Am
1 1 , pine
t7mperate adjacent l
Mixed _--I
conifer 2 from lign<
I

I tremuloid,
o,I 6 5 I 4
Most sava
I An
\
scarificati1
\ parent pic:
advantage
' following
Subalpine germinate
1
and boreal seeds can
dormant s
' pines, pos
- - - - - - - - - - - - - - - (Increasing moisture stress)
Moisture stress index
heated by
0 Little fire influence 4 Short return interval crown/ tain viablt
1 Infrequent light surface fires severe surface fire (25-1 00 Yr.) at temper.
(Above 25 Yr.) 5 Long return interval crown/ The seeds
2 Frequent light surface fires severe surface fire (1 00-300 Yr.)
(1-25 Yr.) 6 Very long return interval crown/
short peri'
3 Infrequent severe surface fire sever surface fire (300+ Yr.) and lack c
(Above 25 Yr.) lodgepole
FIGURE 5.5 An idealized representation of the relationship between temperature,
ing severe
moisture stress, fuel type, and fire frequency in the Pacific Northwest of North America Alth
(after Agee, 1981). environmt
to release
clear awa'
bark thickness of 0.6 em can survive external heating of 500° C for about 1 nate and
minute, whereas a tree with a bark thickness of 2.6 em can survive the same tem- canopy by
peratures for 20 minutes. Douglas fir (Pseudotsuga menziesii and P macrocarpa) , soils enric
western larch (Larix occidentlis), and ponderosa pine (Pinus ponderosa) stands Pattt
in western North America experience frequent fires. All of these trees have bark forests. Fe
that can grow to over 10 em in thickness. These species survive significantly forests of.
longer periods of heating than trees with thinner bark. Fires can also kill root sys- Within tw
tems, and all four of these species have relatively deep root systems that are insu- seeded plz
lated by soil cover against high temperatures. Temperatures during fires decline by heat a1
quickly with soil depth. A surface fire burning at a temperature of 600° C may from surv
produce a temperature of only 80° Cat the soil surface and 33° Cat 10 em depth. such as g'
A number of tree and shrub species display epicormic sprouting that allows roses such
the regrowth of foliage from trunks and branches after fire. Epicormic sprouting ana. How<
occurs from latent buds that lie behind the bark. In some cases, only buds pro- erous tre
tected by the thick bark of the trunk may survive, and the new foliage will appear neverthelt
as a fire column growing from the standing trunk. Coniferous trees, such as pitch During th1
pine (Pinus resinosa) in eastern North America and coastal redwood (Sequoia and the s:
sempervirons) in western North America, and angiosperms, such as live oak paper birc

·~-j"o;p"
~~odlands
1moisture stress)
I crown/
:2s-100Yr.)
crown/
100-300 Yr.)
lrval crown/
:00+ Yr.)
between temperature,
ihwest of North America
ng of 500° C for about 1
can survive the same tem-
niesii and P macrocarpa) .
(Pinus ponderosa) stands
Jl of these trees have bark
·ecies survive significantly
Fires can also kill root sys-
root systems that are insu-
atures during fires decline
:mperature of 600° C may
• and 33° Cat 10 em depth.
1rmic sprouting that allows
r fire. Epicormic sprouting
orne cases, only buds pro-
he new foliage will appear
iferous trees, such as pitch
coastal redwood (Sequoia
>sperms, such as live oak
FIRE 109
(Quercus agrifolia) in California and Eucalyptus spp. in Australia, display epi-
cormic sprouting following fires.
Some plants can produce new growth from lignotubers, which are swellings
that occur at the interface between the roots and shoots. The lignotubers contain
buds and reserves of food. Soil cover protects the lignotubers from lethal temper-
atures during fires. Moderate heating by the fire may even stimulate the buds to
grow. A number of shrubs found in the shrub lands, or chaparral, of California and
adjacent Mexico, such as chamise (Adenstoma fasiculatum), sprout vigorously
from lignotubers following fires. Other plants, such as trembling aspen (Populus
tremuloides) and paper birch (Betula papyrifera), sprout suckers from root buds.
Most savanna and prairie grasses resprout from root systems following fires.
A number of plants produce seeds that remain dormant in the soil until
scarification by heating causes them to germinate following fires. Although the
parent plants may be destroyed by the fire, the newly emerged seedling can take
advantage of the high soil nutrients and competition-free environment that exists
following the burn. For example, the seeds of deer brush shrubs ( Ceanothus spp.)
germinate best after experiencing temperatures of 75 to 100° C. The deerbrush
seeds can remain dormant and viable in the soil for several decades. The store of
dormant seeds in the soil is referred to as a soil seed bank. Some plants, especially
pines, possess serotinous cones that remain closed and hold their seeds until
heated by a fire. Jack and lodgepole pine cones can be held on the trees and con-
tain viable seeds for 25 to 75 years. The resin keeping the cone scales closed melts
at temperatures of about 60° C, and this releases seeds during and after the fire.
The seeds can remain viable when exposed to temperatures as high as 370° C for
short periods. The pine seedlings take advantage of the favorable soil conditions
and lack of competition that results from fires. It is not unusual to find jack and
lodgepole pine seedling densities of over 1 million individuals per hectare follow-
ing severe fires (Fig. 5.6).
Although we may think of fire as a destructive and negative aspect of the
environment, it is clear that fire plays a beneficial role in many ways. Fires serve
to release nutrients from dead and living vegetation back into the soil. Fires also
clear away leaf litter and decadent vegetation cover. Most plants cannot germi-
nate and grow under the shade of a dense forest canopy. Destruction of the
canopy by fire allows shade intolerant plants to regenerate and take advantage of
soils enriched by the minerals in the ash from the fire.
Patterns of post-fire succession have been intensively studied in many
forests. For example, stands dominated by white spruce (Picea glauca) in the
forests of Alaska and northwestern Canada burn every 80 to 400 years (Fig. 5.7).
Within two to five years following a fire, the site is usually dominated by light-
seeded plant species, plants with seeds that lie dormant in the soil until stimulated
by heat and chemical changes associated with fire, and plants that can sprout
from surviving roots and corms. These early dominants include grasses, herbs
such as geranium (Geranium bicknellii), fireweed (Epilobium angustifolium),
roses such as Rosa acicularis, and willows such as Salix bebbiana and S. scouleri-
ana. However, even at this very early stage, the seedlings of deciduous and conif-
erous trees are often present in the vegetation. The tree seedlings are
nevertheless very small and grow too slowly to dominate the early vegetation.
During the next 25 years, large shrubs such as willows and birches (Betula nana)
and the saplings of deciduous trees such as aspen (Populus tremuloides) and
paper birch (Betula papyrifera) come to dominate the vegetation. Over the next
11 0 CHAPTER 5 DISTURBANCE

50 years, dt:
herbs, and "
been presen
years it don
the ground,
forest.
The ht
fires are imr
shoe hare. I1
lynx. Fires ;
burned stan
fallen and d·
itats are use•
beetle founc
trees. There
have special
and larvae s
use standing
As bio
FIGURE 5.6 A dense stand of young lodgepole pines (Pinus contorta) established
geographic <
following a fire in Glacier National Park, Montana. Fire in this forested ecosystem is a northern di5
natural event. The destruction of the forest canopy allows light-demanding species such as and oaks in •
lodgepole pine to become established and grow. North Ame1
forests, whic
larch. In con
ject to frequ
Q)
Q)
0)
Q)
0) Q)
0)
(1) Q)
appear to bt:
(1) El
en
0)
(1) Cii (.)
2 Q) range of ash
Q) Cii Cii ""0
;: Q)
E 0)
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>< ~ 0
are lower th
Q) Q) en .1:: 0
E en ""0 (1) (1)
:s: 0 ~ g.~ are able tot
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c
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>-
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.1:: <t!
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cen u~ co ~I
Europt
Q)
z
Q)
I
.c Q) · -
o@.
~
(j)
activity in a
provides fun
butions of p
since 1860 (1
tlers has bet:
reduction in
uted to fire s
expansion of
ures, includir
southern Ar
eight years. I
allowed the
\ recent incre<
Canadian an
Age (yr)
the United S
FIGURE 5.7 Idealized post-fire succession in a boreal white spruce (Picea glauca) forest the northern
(after Van Cleve and Viereck, 1981 ). while in the
 FIRE 111

50 years, deciduous trees dominate the site, and the shade intolerant grasses,
herbs, and willows are replaced by more shade tolerant species. Spruce, which has
been present all along, becomes increasingly important over time, until after 100
years it dominates the vegetation. After 200 years, a dense moss cover forms on
the ground, and most of the deciduous trees have died and disappeared from the
forest.
The herbs, shrubs, and deciduous trees that regenerate following northern
fires are important sources of food for animals such as moose, beaver, and snow-
shoe hare. In turn, these herbivores serve as food for predators such as wolf and
lynx. Fires are therefore important in supporting the mammalian fauna. The
burned stands also provide a number of different microhabitats encompassing
fallen and dead trees, older surviving trees, and new vegetation. These microhab-
itats are used by many different animal species. As one example, the Melanophila
beetle found in North America deposits its eggs under the bark of freshly burned
trees. There the larvae feed on the dead cambium of the trees. The adult beetles
have specialized organs to sense heat and find freshly burned stands. The beetles
and larvae serve as food for a number of birds such as woodpeckers, which also
use standing dead trees as nesting sites.
As biogeographers, we are interested in the role of fire in controlling the
contorta) established
geographic distributions of species. One example of this role is provided by the
Jrested ecosystem is a northern distribution of black ash (Fraxinus nigra) and some species of maple
-demanding species such as and oaks in Canada. The trees are typically found in eastern deciduous forests of
North America. The trees are almost never found in the more northern boreal
forests, which are dominated by coniferous trees such as spruce, fir, pine, and
larch. In contrast to the deciduous forest, the northern coniferous forests are sub-
ject to frequent intense fires. Ash trees, for example, are thin barked and do not
appear to be particularly well adapted to frequent fires. However, the northern
Q) Q) range of ash extends far into the boreal forest in moist areas and along river val-
'"'
r.lcu Ol
S:Q)(j) leys. Among other things, these areas provide habitat in which fire frequencies
""0(..)(1)
Q) :::J (/)
X ~ 0
are lower than in the surrounding boreal forest. It is possible that the ash trees
~ 5}:2; are able to tolerate the colder northern climates of the southern boreal zone as
long as they are protected from frequent fires.
European land-use practices in North America have led to decreased fire
activity in a number of areas. The response of vegetation to this decrease in fire
provides further evidence of the role of burning in controlling geographic distri-
butions of plants. The area of grassland in southern Texas has declined sharply
since 1860 (Fig. 5.8). Much of the grassland first encountered by American set-
tlers has been replaced by mesquite (Prosopis) shrubland and woodland. The
reduction in grassland and expansion of shrubland and woodland can be attrib-
uted to fire suppression following American settlement. Cattle ranchers view the
expansion of mesquite as a menace to rangelands and undertake various meas-
ures, including setting fires to battle it. Recent work in the semiarid grasslands of
southern Arizona shows that the natural fire frequency there is about four to
eight years. Decreased frequency of fire due to grazing and fire suppression has
allowed the advance of pine and oak forest at the expense of grassland. Similar
recent increases in woodland at the expense of grassland are evident on the
Canadian and American plains, and in dry portions of the Pacific Northwest of
the United States. For example, poplar and spruce have invaded the periphery of
;pruce (Picea glauca) forest the northern prairies in Canada, pines have invaded grasslands in Colorado,
while in the Pacific Northwest juniper (Juniperus spp.) woods have invaded
112 CHAPTER 5 DISTURBANCE

After fire, c:
trial ecosys
regimes in 1
agent of dis
Wind is als
forests. Exp
rounding lo
that betwee
Mexico Mexico events that
least eight s1
Forest~
1860 1960
Winds in for
5.10). The UJ
Grasslands
soils. Winds 1
FIGURE 5.8 The expansion of mesquite (Prosopsis) shrubland at the expense of ple them. T<
grassland between 1860 and 1960. Decreases in burning due to human land-use change responsible J
and fire control are thought to be a major contributor to the decline in grassland cover (after speeds as lm
Johnston, 1963; Brown and Lomolino, 1998).
tain conditio
by hurricane
upland grasslands. The exclusion of shrubs and trees by fire is crucial in maintain- hour, cause
ing grassland and savanna in many other settings throughout the world. Suppres- affect very s
sion of fires can also change the species dominance in wooded sites. Studies in and adjacent
northern Florida indicate that the Ocala sand pine (Pinus clausa) regenerates canes causes
poorly beneath its own canopy. Frequent fires serve to open the canopy and allow August 1992
new pines to establish. Fire control measures over the past century have allowed impacted ov<
the pine stands to be invaded by shrubs and evergreen oaks which were formerly in Everglade
excluded by fires. The shade cast by these plants makes it even more difficult for 5 million tre
the sand pines to regenerate.
Although large fires are still capable of burning beyond our control, fire
suppression methods have been relatively successful in the United States and
Canada over the past 100 years. One consequence has been the buildup of a dan-
gerously high level of fuel in some forest ecosystems. As a result of this artifically
high fuel loading, some recent fires in coniferous forests and chaparral-domi-
nated portions of the western United States have been particularly intense and
large. To mitigate this situation, government authorities in the United States and
Canada have instituted programs of prescribed burns that are set under carefully
controlled conditions in order to imitate natural disturbance regimes and
decrease fuel loads. In many cases, such efforts are useful. However, on some
occasions these prescribed fires have gotten out of control, with disastrous
results. For example, a prescribed fire that went out of control in the spring of
2000 caused severe damage to the town of Los Alamos, New Mexico, despite
intensive efforts to control it. Given the artificially high fuel loads that remain in
many western North American forests, similar near catastrophes and catastro-
phes will likely continue into the foreseeable future. It will take many years of
prescribed burns and other fuel management techniques to remedy the situation
in North America. A study in the Boundary Waters Canoe area of Minnesota
concluded that it would take 50 to 75 years of prescribed burning to return the FIGURE 5.9
forest structure of that area to its natural state. (after Runkle, 1~
 WIND 113

IND

After fire, catastrophic wind is the most frequent and widespread agent of terres-
trial ecosystem disturbance (Fig. 5.9). Although fires dominate disturbance
regimes in grassland, savanna, and coniferous forests, wind is a more important
agent of disturbance in many temperate deciduous forests and tropical forests.
o;:~
Wind is also a major cause of disturbance in many high-altitude coniferous
forests. Exposed alpine sites generally experience much higher winds than sur-
rounding lowlands. Research near alpine treeline in western Oregon suggests
that between 20% and 70% of the fir stands were established after windthrow
.~Jf.l
"~-1.;
1'1
.:ii·"'·Y events that occurred in the past 60 years. Documentary evidence shows that at
;,it: ~~

~
0
land at the expense of
to human land-use change
lecline in grassland cover (after
y fire is crucial in maintain-
ughout the world. Suppres-
in wooded sites. Studies in
(Pinus clausa) regenerates
open the canopy and allow
: past century have allowed
1 oaks which were formerly
es it even more difficult for
least eight severe wind storms occurred during that period.
Forests are more sensitive to wind damage than other vegetation types.
Winds in forests strip foliage, break branches, shear trunks, and uproot trees (Fig.
5.10). The uprooting of trees causes soil disturbance and the exposure of mineral
soils. Winds of over 100 to 130 km per hour typically strip trees of foliage and top-
ple them. Tornado-induced winds of 160 km per hour have been shown to be
responsible for reducing tree cover by 68% in affected stands in Minnesota. Wind
speeds as low as 60 km per hour can still produce significant destruction under cer-
tain conditions. Catastrophic winds, with gusts over 200 km per hour, are generated
by hurricanes and tornadoes. Wind micro blasts, with velocities reaching 400 km per
hour, cause small linear patches of intense tree mortality. Tornadoes generally
affect very small areas. However, they can be frequent occurrences in the plains
and adjacent forests of interior North America. In contrast, individual large hurri-
canes cause severe damage to forests over thousands of kilometers. For example, in
August 1992 Hurricane Andrew created a 20-km-wide swath of wind damage that
impacted over 1600 sq km of southern Florida and disturbed 80% of the vegetation
in Everglades National Park (Fig. 5.11). It is estimated that Hurricane David killed
5 million trees on the island of Dominica. Hurricanes and associated tropical

tg beyond our control, fire

l in the United States and
been the buildup of a dan-
'\s a result of this artificall y
>rests and chaparral-domi-
en particularly intense and
es in the United States and
that are set under carefully
disturbance regimes and
useful. However, on some
•f control, with disastrous
of control in the spring of
mos, New Mexico, despite
F =Fires
~h fuel loads that remain in G =Gap creation by
catastrophes and catastro- individual events of
It will take many years of tree mortality
1es to remedy the situation
Canoe area of Minnesota
ibed burning to return the FIGURE 5 .9 The major disturbance regimes of eastern North American forest types
(after Runkle, 1990).
114 CHAPTER 5 DISTURBANCE

or cool oc1
the tropic~
Puerto Ri,
Sri Lanka
ricanes (c<
ricanes di
Nonethele
experience
tury, hurri'
eastern Cc:
bance as f~
Alth
areas of la
of trees ha
age gap si<
vidual tret
reported f<
of North A
trees. Mos1
trees actua
FIGURE 5.10 A wind microblast destroyed the coniferous tree canopy and allowed The!
light-demanding grasses, herbs, shrubs and trees saplings to become established at this is highly v~
site in Idaho. swaths. Ht
Exposed s
most susce
storms are categorized by their wind speeds. The class 4 and 5 storms, like Hurri- the most p
cane Andrew, with sustained winds of over 200 km per hour cause the most damage size and ar
to forest cover. Hurricanes develop in late summer and early fall over warm tropi- impact of I
cal seas. The convective energy that drives hurricanes dissipates quickly over land 70% mort~
canopy tre
forests sho
large canol
Not a
small gaps
winds. Sud
forests and
one study i
472 trees P'
the death o
mortality, h
cause of trc
As is
tioning of e
the lower'
growth of s
dense cane
action of ur
FIGURE 5.11 The path of major destruction due to Hurricane Andrew (1992) in southern
Florida and the paths of other historical hurricanes in Florida. The inset maps show some
plants requ
typical hurricane paths apparent from compilations of historical records (after Roman et al. gaps, while
1994; Strahler and Strahler, 1997). growth ofp

tree canopy and allowed
) become established at this
; 4 and 5 storms, like Hurri-
hour cause the most damage
j early fall over warm tropi-
dissipates quickly over land
me Andrew (1992) in southern
The inset maps show some
~al records (after Roman et al.
WIND 115
or cool oceans. The frequency of ecosystem disturbance by hurricanes is greatest in
the tropics and subtropics. It has been estimated that 83% of the forested area of
Puerto Rico has been disturbed by hurricanes over the past 100 years. A study in
Sri Lanka reports that 73% of the forest there has been similarly disturbed by hur-
ricanes (called cyclones in that region) over the past century. The frequency of hur-
ricanes diminishes relatively quickly as one moves away from the tropics.
Nonetheless, the Gulf of Mexico and Atlantic coastlines of the United States also
experience hurricanes relatively frequently. Remarkably, during the twentieth cen-
tury, hurricanes have traveled as far inland and north as the province of Ontario in
eastern Canada. Thus, hurricanes may be an important force of vegetation distur-
bance as far north as the forests of New England and adjacent Canada.
Although catastrophic wind events such as hurricanes can impact very large
areas of land, the actual size of the gaps (contiguous areas where large numbers
of trees have been toppled) created by wind disturbance is highly variable. Aver-
age gap size created by wind disturbances generally ranges from the size of indi-
vidual trees to around 30 ha. In some cases, gaps of over 3000 ha have been
reported for hurricanes. In contrast, in the coastal forests of the Pacific Northwest
of North America, 46% of the windthrow gaps are created by the felling of single
trees. Most wind events create a number of different gap sizes. The percentage of
trees actually toppled in such gaps can range from 1% to over 90%.
The geographic pattern of tree destruction during catastrophic wind events
is highly variable. Tornadoes tend to destroy relatively narrow and discontinuous
swaths. Hurricanes and large cyclonic storms cause damage over wider areas.
Exposed stands on high ground, at the edges of forests, or on shallow soils are
most susceptible to windthrow. Within continuous forests, tall canopy trees are
the most prone to destruction. They have bigger areas of foliage relative to trunk
size and are more exposed to the shear stresses caused by wind. Data from the
impact of Hurricane Andrew on Florida showed that canopy trees suffered up to
70% mortality, while smaller trees suffered less than 20% mortality. When large
canopy trees fall, they may crush smaller trees and shrubs. One study in palm
forests showed that 70% of the tree mortality during wind storms was due to
large canopy trees crushing smaller subcanopy individuals.
Not all wind-generated gaps are associated with catastrophic events. Many
small gaps are caused when old and senescent trees are felled by relatively mild
winds. Such events are part of the background mortality that occurs naturally in
forests and causes losses of perhaps 5% of the canopy each year. Interestingly,
one study in Puerto Rico showed that hurricanes could be expected to kill about
472 trees per hectare per century. In contrast, background mortality likely causes
the death of 1973 to 2650 trees per hectare per century. Thus, in terms of general
mortality, hurricanes cause devastating short-term effects but are not the primary
cause of tree mortality in this tropical forest regime.
As is the case with fire, wind disturbance plays a crucial role in the func-
tioning of ecosystems. The clearing of canopy trees increases the light received by
the lower vegetation layers. In many cases, this facilitates the germination and
growth of seedlings that would have been unable to grow under the shade of a
dense canopy. The dead vegetation supplements soil nutrients. The churning
action of uprooted trees can produce areas of exposed mineral soil which certain
plants require for seedling establishment. Soil temperature generally increases in
gaps, while humidity decreases. These changes aid in the establishment and
growth of plants that cannot survive under dense forest canopies.
116 CHAPTER 5 DISTURBANCE

There are key differences between the impact of fires and wind disturbance. be accommod.
Surface fires may destroy low vegetation but leave canopy trees intact. Wind rivers in semi
always impact the canopy more than the subcanopy layer. Although falling tree along the ocea
can damage the subcanopy vegetation, the damage to the lower vegetation layer with high tide~
during wind events is much less than during fires. Finally, while fires can clear the The seas
litter layer and return nutrients to the soil as ash, winds often add undecomposed areas along m.
organic matter to the litter layer. land surface a
Tropical and subtropical areas, which are prone to hurricane disturbance. When such re~
appear to show remarkable rates of succession and recovery. Closed forest replaced by a t
canopies at sites in Central America damaged by Hurricane Joan in 1988 devel- are available, i
oped within five years. The height of the canopy, however, was still much lower magnitude flo<
than its pre-hurricane condition. Hurricanes also kill wildlife and cause severe intervals. The 1
damage to nesting sites. Of particular concern is damage to nesting sites for birds. 1997 flooding ,
However, studies in the Everglades following Hurricane Andrew showed that Dakota in mos
populations of many wading birds remained stable or increased slightly following ond or less. Th<
the storm. Bald eagles have been shown to have recovered quickly despite the mated that su<
loss of about 44% of their nesting sites in areas of South Carolina impacted by planning and 1
Hurricane Hugo. called the "100
Wind is certainly a factor in controlling the geographic distribution of trees The 100-year fl
in alpine environments. High winds cause limb breakage and the destruction of tury. For the F
foliage and buds by blowing snow and ice crystals during the winter. Cold and dry water per secoJ
winter winds can also cause the death of needles and buds by desiccation. There Flooding
are many examples where the elevation of treeline on exposed and windy slopes can drown ani
is considerably lower than on protected sites. The gaps caused by wind are Decreased roo
undoubtedly important in allowing shade intolerant plant species to exist in production. In
regions of continuous forest cover. Tree species such as Sitka spruce (Picea the ocean carr
sitchensis) require the relatively continuous creation of gaps of at least 800 to can also produ·
1000 sq m in order to regenerate and remain codominant with coastal hemlock conditions in "'
(Tsuga heterophylla) in the Pacific Northwest. Hemlock is more successful than rous and mang
Sitka spruce on shady sites and in very small gaps. In the absence of large wind loads that are d
gaps, the spruce would be excluded in many areas. ment to the sc
Floods also kill
~tation by push
FLOODING Ecosyster
periods of inun
Flooding is an important form of disturbance impacting low-lying areas near terns in arid an
rivers and shoreline ecosystems along oceans and lakes. A flood is formally short periods o
defined as a high flow of water that overtops normal confinements and covers ment are the m
land that is normally dry. In areas of flat topography, river flooding can expand lated that a cm
over thousands of square kilometers. Despite many control measures, widespread the Great Plai1
river flooding still occurs throughout the world. One recent example was the most floodplain
destructive flooding of communities and large areas of farmland along the Red floods are activ·
River in North Dakota and adjacent Manitoba during the spring thaw of 1997. In additio
Three types of floods are recognized. First, river and associated lake flood- ecosystems. Hig
ing takes place when high seasonal precipitation and/or spring snowmelt forests to grow
exceeds soil moisture-holding capabilities and produces high amounts of runoff. are referred to
River flooding can also be caused by channel blockages owing to factors such as riparian trees fo
river ice. The area of low-lying land that is typically inundated when a river deltoides) stand:
floods is referred to as the floodplain. Second, flash floods are short-duration lensis forests in
events that occur when individual storms produce more runoff water than can savanna of Ken'

'fires and wind disturbance.
canopy trees intact. Winds
ayer. Although falling trees
the lower vegetation layers
tlly, while fires can clear the
Is often add u~decomposed
e to hurricane disturbance,
1d recovery. Closed forest
rricane Joan in 1988 devel-
vever, was still much lower
I wildlife and cause severe
ge to nesting sites for birds.
cane Andrew showed that
increased slightly following
overed quickly despite the
outh Carolina impacted by
;raphic distribution of trees
~age and the destruction of
1ng the winter. Cold and dry
buds by desiccation. There
1 exposed and windy slopes
gaps caused by wind are
t plant species to exist in
tch as Sitka spruce (Picea
t of gaps of at least 800 to
inant with coastal hemlock
xk is more successful than
1 the absence of large wind
tcting low-lying areas near
lakes. A flood is formally
1l confinements and covers
, river flooding can expand
mtrol measures, widespread
te recent example was the
of farmland along the Red
the spring thaw of 1997.
' and associated lake flood-
t and/or spring snowmelt
ces high amounts of runoff.
;es owing to factors such as
ly inundated when a river
1 floods are short-duration
10re runoff water than can
FLOODING 117
be accommodated by stream and river channels. In general, steep mountainous
rivers in semiarid regions experience flash floods. Third coastal floods occur
along the ocean margin and are often associated with storm surges that coincide
with high tides and hurricanes.
The season, frequency, and severity of flooding vary greatly. Low-lying
areas along many rivers are flooded annually. In Brazil some 300,000 sq km of
land surface are prone to annual flooding by the Amazon and its tributaries.
When such regular cycles of dry and wet conditions exist, the term flood may be
replaced by a term such as seasonal inundation. Where long records of river flow
are available, it is clear that many flood-prone rivers are subject to frequent low-
magnitude floods, with very high-magnitude flooding recurring at widely spaced
intervals. The flood frequency of the Red River, which was responsible for the
1997 flooding of North Dakota and Manitoba, shows this pattern. Fargo, North
Dakota in most years experiences peak flows of about 1000 cum of water per sec-
ond or less. The flow during the 1997 flood was three times this amount. It is esti-
mated that such high flows and flooding occur only once every 200 years. For
planning and conservation purposes, many resource managers use a measure
called the "100-year flood" as a worst case scenario for anticipating flood impacts.
The 100-year flood is the peak discharge that is expected to occur once each cen-
tury. For the Red River at Fargo, the 100-year flood discharge is 2300 cu m of
water per second.
Flooding impacts terrestrial ecosystems in a number of ways. High water
can drown animals and kill plants by restricting oxygen uptake through roots.
Decreased root activity can also kill plants by causing cessation of carbohydrate
production. In effect, the flooded vegetation starves to death. Flood waters from
the ocean carry salts that are lethal to many plants. Evaporation of freshwater
can also produce salt crusts that inhibit plant establishment and growth. Anoxic
conditions in waterlogged soils also lead to the production of toxins such as fer-
rous and manganese ions and sulfides. Flood waters carry suspended sediment
loads that are deposited on the floodplain. The addition of large amounts of sedi-
ment to the soil surface can kill plants by depriving root systems of oxygen.
Floods also kill plants by eroding soils. In cold climates, flooding can damage veg-
etation by pushing large slabs of ice onto the shoreline.
Ecosystems in humid, low-lying regions often must be able to endure long
periods of inundation by low-energy flood waters. In contrast, floodplain ecosys-
tems in arid and hilly regions often experience high-energy floods that produce
short periods of inundation. In these systems, the erosion and deposition of sedi-
ment are the most important factors in ecosystem disturbance. It has been calcu-
lated that a complete reworking of sediment along the floodplains of rivers on
the Great Plains near the Rocky Mountains occurs every 200 to 300 years. In
most floodplains, both inundation by water and erosion and deposition caused by
floods are active forms of disturbance.
In addition to their destructive potential, floods play a positive role in many
ecosystems. High water associated with flooding increases soil moisture and allows
forests to grow in regions that are normally too dry to support trees. Such woods
are referred to as riparian forests, gallery forests, or riverine forests. Examples of
riparian trees found on very frequently flooded sites include plains poplar (Populus
deltoides) stands found on the North American Great Plains, Eucalyptus camaldu-
lensis forests in arid portions of Australia, and Populus ilicifolia stands in the
savanna of Kenya. Floodplain trees such as those mentioned above are usually fast
118 CHAPTER 5 DISTURBANCE

growing, reproduce at an early age, and can withstand soil inundation. In addition Forest types
to providing moisture, floods can replenish soil nutrients in areas where sediment is
deposited. The irrigation and fertilization of drylands by floods have played aL 1a Active leve
important role in human history in many regions. Perhaps most notable is th 1b Inactive le•
2 Acacia
development of early Egyptian civilization along the floodplain of the lower Nil 3 Clay everg
River. Floods also serve as an important mechanism for seed dispersal. The seeds oi 4a Point-barf
many common floodplain species, such as poplar and willow, can survive transpon 4b Point-bar t
in water. A number of Amazon trees use fish to disperse seeds. In fact, some Ama- 5 Oxbow
6 Dry bush
zon plants require that seeds be chewed by fish in order to release them from their 7 Scrub
hard outer coats. Most relatives of the flesh-eating pirhana fish (Serrasalmus 2/3 Acacia/cia~
rhombeus) of South America are fruit eaters that rely on flooded forests for habit a 2/6 Acacia/dry
and food. Similarly, in Africa, the butter barbel fish (Schilbe intermeddins) is impor- Culitvated/
New sedirr
tant in the dispersal of seeds from the sycamore fig (Ficus sycomorus).
Aside from seeds that are adapted for transport by water, floodplain plants 6
possess other adaptations that allow them to cope with floods. The hollow an ' ~
7 2 2/6
tubular stems of bullrush (Schoenoplectus lacustris) serve to oxygenate water- 5 5

~~
covered roots. Other plants have long cavities in their wood and roots that alloK
downward oxygen flow. The coastal variety of lodgepole pine (Pinus contorta ssp.
contorta) , which grows on wet sites in the Pacific Northwest and Alaska, produces

~
2,
such cavities. Many floodplain plants have metabolic and physical adaptation_ 2 '
that allow them to persist for long periods of time with their roots in anoxic soils.. 6
Floodplain plants generally have high tolerances for ferrous and manganese ions
or root systems that are inefficient at extracting these potentially toxic ions from
the soil. The swamp cypress (Taxodium distichum) of southeastern North Amer-
ica produces elongated vertical structures from their roots. These structure
called pneumatophores, extend above flood level and allow the root systems to
obtain oxygen. Many floodplain plants, such as cattails (Typha latifolia), possess
rhizomes (horizontal stems and associated root systems) that contain large
reserves of carbohydrates and can tolerate submergence and oxygen deprivation
for months at a time. When detached from the parent plants by floods, these rhi- FIGURE 5.12 T
zomes can grow and produce a complete new plant. Following floods, new growth the village of Pum
can sprout from the toppled stems of riparian willows and poplars (willows an · along the river is i,
poplars that grow along rivers or streams). The California coastal redwood
(Sequoia sempervirons) produces new roots from its trunk when buried by flood
deposits. These adventitious roots develop close to the new soil surface wher above the flood;
oxygen is more plentiful. plain support a 1
Flooding controls the geographic distribution of plant species at a number tion on these sitt
of different spatial scales. At the larger scale, the additional soil moisture pro- as long periods <
vided by flooding allows some tree species to extend their range into arid region less than 28 day
At a smaller scale, flooding is important in determining local plant distributions.. sites that do not
Slight differences in elevation produce large changes in floodplain vegetation. allows trees to e
Examples of such local vegetation zonation are apparent from most river sys- development to
tems. As river channels shift, they leave behind new terrain for colonization by Patchy floc
terrestrial plants. Succession on these abandoned river channels is also an impor- throughout the '
tant factor in floodplain vegetation zonation. deposits are cole
A good example of the local geographic vegetation zonation that develop sisting of willow
in floodplains comes from the Tana River of Kenya. Here the floodplain support enough above tt
gallery forests of Acacia, poplar, and a number of evergreen tropical trees (Fig. sam poplar (PoJ
5.12). The channel of the Tana shifts often, and the sediments of the floodplain (Populus tremuh
are completely reworked by erosion and deposition on a 150-year cycle. The area highest land surf
 1 19

:1 soil inundation. In addition Forest types Relative ha

tts in areas where sediment is proportion (%)
:Is by floods have played an 1a Active levee evergreen 1.5 28 • Active river
Jerhaps most notable is the 1b
2
Inactive levee evergreen
Acacia
5.0
44
94
842
D New sediment
floodplain of the lower Nile 3 Clay evergreen 5.1 98 • Sand quarry
1r seed dispersal. The seeds of 4a Point-bar front 3.0 58
willow, can survive transport 4b Point-bar back 2.5 48 • Village sites
5 Oxbow 7.5 139
rse seeds. In fact, some Ama-
6 Dry bush 0.1 1.5 D Cultivated/cleared
er to release them from their
?; pirhana fish (Serrasalmus
7
2/3
Scrub
Acacia/clay evergreen
2.74
5.5
52
103
D Forest (see key for
forest types)
on flooded forests for habita t 2/6 Acacia/dry bush 3.5 69
Culitvated/cleared 16.5 316
hilbe intermeddins) is impor- New sediment 2.75 52
icus sycomorus).
t by water, floodplain plants
with floods. The hollow an d
1 serve to oxygenate water-

r wood and roots that allow

ole pine (Pinus contorta ssp.
:hwest and Alaska, produces
ic and physical adaptations
th their roots in anoxic soils.
ferrous and manganese ions
~ potentially toxic ions from
f southeastern North Amer-
eir roots. These structures.
1d allow the root systems to
ils (Typha latifolia), possess
ystems) that contain large
nee and oxygen deprivation
t plants by floods, these rhi·
'allowing floods, new growth
vs and poplars (willows and
::=alifornia coastal redwoo
trunk when buried by flood
the new soil surface wher
>f plant species at a number
jditional soil moisture pro-
heir range into arid region
ing local plant distributions.
es in floodplain vegetation.
)arent from most river sys-
terrain for colonization b\·
~r channels is also an impor-
tion zonation that develops
Iere the floodplain supports
rergreen tropical trees (Fig
sediments of the floodpla ir:
m a 150-year cycle. The arec:
Poplar (4b)
+ Terminalia brevipes
+ Spirostachys venenifera
Point-bar (4a) + Cordia sinensis
including
Pluchea dioscoridis
young Populus ilicifolia on ridges
~"0
o~....~,...-
..c"'mE
C'd-g.::::-
.E;gi~
OlCJlom
"ij) < 1 ) - -
I
Tana River
FIGURE 5.1 2 The patchy soil conditions and vegetation of the Tana River floodplain near
the village of Pumwani in Kenya. The vegetation typical of different successional stages
along the river is illustrated (after Hughes, 1990).
above the floodplain is dominated by arid shrubland. Lower levels of the flood-
plain support a mosaic of bare soil, small shrubs, and saplings. The sparse vegeta-
tion on these sites results from frequent erosional and depositional events, as well
as long periods of inundation. Forest is restricted to higher areas that experience
less than 28 days of continuous inundation. The most diverse forest is found on
sites that do not experience flood erosion. In the case of the Tana River, flooding
allows trees to extend their range into a relatively arid region but restricts forest
development to very specific sites along the floodplain.
Patchy floodplain zonation and succession are typical of forested regions
throughout the world. Along the Tanana River in Alaska, recently stabilized river
deposits are colonized within a few years by a shrub-dominated vegetation con-
sisting of willow (Salix alaxensis) and alder (Alnus tenuifloia). If the site is high
enough above the current floodplain, the shrubby vegetation is followed by bal-
sam poplar (Populus balsamifera) forest, and then mixed balsam poplar, aspen
(Populus tremuloides), and white spruce (Picea glauca) forest. On the oldest and
highest land surfaces the forest is dominated by white spruce.
120 CHAPTER 5 DISTURBANCE

One of the most interesting examples of how flooding controls the very
large-scale geographic distribution of a species is provided by bald cypress (Tax -
odium distichum) in the southeastern United States (Fig. 5.13). Bald cypresses
are large, striking trees well adapted to flood-prone areas and swamp. They have
trunks that spread out at the base in buttress fashion to provide support in water-
logged soils (Fig. 5.14). They are also noted for their large and abundant pneu-
matophores. The seeds of the bald cypress survive immersion in water and ar
typically transported by floods. The trees are found in the floodplains of slow·
moving streams with gentle gradients, in swamps that experience inundation by
water for a large portion of the year, and in small shallow lakes called oxbows.
Oxbow lakes are formed when the meandering movement of a river or stream
channel experiences a rapid shift that leaves a portion of the channel cut off fro ffi
regular flow. These cutoff channels form small lakes that eventually fill with sedi-
ment and disappear. The geographic distribution of bald cypress in the southeast-
ern United States has a relatively sharp border that coincides with the
physiographic boundary between the flat Coastal Plain of the Southeast and the
steeper terrain of the surrounding uplands. Bald cypress seeds germinate after
being transported by floods and deposited on moist surfaces. They will not germi-
nate on dry soils. The seedlings, however, are killed if they are covered by water
for more than a few days following germination. The floodplains of streams o
FIGURE 5.1•
the uplands are narrow and do not provide much habitat for bald cypress seeds to
United States
settle and germinate. Most importantly, incidences of flooding are more frequen
and erosive in the narrow upland floodplains than on the broad floodplains of th
Coastal Plain. The Coastal Plain, which experiences low-energy floods followec
by long dry intervals, is ideal for bald cypress reproduction. Therefore, althougl::
flooding is essential for the germination of bald cypress, its seedlings cannot tol- reservoir sy
erate frequent large-magnitude floods and the geographic distribution of the ing. Over ti1
species is largely limited to the Coastal Plain. dry out, mi1
Human activity can alter flood regimes by changing the frequency or geo- and organic
graphic extent of flooding. Reservoirs built along river systems can serve as Onee
modulators of river flow by capturing and storing water during periods of natu- is provided
ral peak flow and then releasing it during periods of natural low flow. If such of the Glen
tion of the
addition, th,
the Colorad
ing the com
tation bega1
plants were
floods caust
bars no Ion!
Within the 1
and sedime1
tists, recogn
of the Gran
1996 a total
week perioc
less than hal
FIGURE 5.13 The correspondence between the northern limits of bald cypress (Taxodiu m creation of 1
distichum) and the boundary of the coastal plain. Seasonal flooding on the coastal plain did not have
appears to be crucial for bald cypress regeneration (after Shankman and Kortright, 1994). the lower 0
 FLOODING 121

flooding controls the very

vided by bald cypress (Tax-
(Fig. 5.13). Bald cypresses
reas and swamp. They have
o provide support in water-
large and abundant pneu-
nmersion in water and are
in the floodplains of slow-
t experience inundation by
1allow lakes called oxbows.
ement of a river or stream
of the channel cut off from
1at eventually fill with sedi-
1ld cypress in the southeast-
that coincides with the
[n of the Southeast and the
1ress seeds germinate after
rfaces. They will not germi-
they are covered by water
~ floodplains of streams on
FIGURE 5.14 Bald cypress (Taxodium distichum) trees in parts of the southeastern
tat for bald cypress seeds to United States require regular, low-velocity flood events in order to successfully reproduce.
flooding are more frequen t
he broad floodplains of the
ow-energy floods followed
uction. Therefore, although
~ss , its seedlings cannot tol- reservoir systems are extensive, they can virtually remove any chance of flood-
~raphic distribution of the ing. Over time, this can cause severe problems for floodplain ecosystems as soils
dry out, mineral soil cover ceases to be replenished or eroded, and vegetation
ging the frequency or geo- and organic matter build up on the land surface and in pools (Table 5.1).
·iver systems can serve as One example of the damaging impact of such flood control on vegetation
ter during periods of natu- is provided by the lower Colorado River in the Grand Canyon. The completion
,f natural low flow. If such of the Glen Canyon Dam above the Grand Canyon in 1963 caused the cessa-
tion of the large annual floods that were a feature of the river's hydrology. In
addition, the fine sediment that normally was carried into the Grand Canyon by
the Colorado River became trapped in the still waters behind the dam. Follow-
ing the construction of the dam and the cessation of large floods, woody vege-
tation began to thicken on the lower banks of the river. Many of the successful
plants were nonnative to the canyon. On higher areas, the loss of water from
floods caused the native mesquite vegetation to die back. In addition, sandy
bars no longer received fresh inputs of sediment, and they began to disappear.
Within the river, trout, which had not been able to live in the previously warm
and sediment rich waters, began to flourish. In 1996 the government and scien-
tists, recognizing the changes that the dam was creating in the natural ecology
of the Grand Canyon, began to experiment with controlled flooding. In March
1996 a total of 900 million m3 was released from Glen Canyon Dam over a one-
week period to simulate a natural flood. The rate of discharge was, however,
less than half of a natural flood on the Colorado. The experiment resulted in the
nits of bald cypress (Taxodiu m creation of new sandy bars and other changes in the river channel. However, it
oding on the coastal plain did not have significant impact on the woody vegetation that now lines much of
1kman and Kortright, 1994). the lower Colorado.
122 CHAPTER 5 DISTURBANCE

TABLE 5.1 Impact of Dams on the Physical and Biological force. Ava
Conditions of Downstream Rivers often occu
occurreno
Physical Changes
lanche tra'
Water temperature forested n
Water sediment content erant plan
Water chemistry tant fe edir
Magnitude of flow The
Timing of high and low flow ure. Land
Variability of flow organic m
Changes in river channel morphology from rock
Changes in bank morphology wet or dr)
Changes in floodplain surface and soil water vegetation

Biological Changes

Fragmentation of aquatic populations

Death of individuals at spillways and intakes
Disruption of aquatic migration routes
Increased colonization of banks
Decreased abundance or extinction of flood-dependent species
Increased abundance of invasion by flood intolerant species
Alterations of food webs
Changes in biodiversity

Modification of river drainage basins can also serve to increase river flows
and the frequency and magnitude of flooding. Removal of vegetation and soil
cover causes increased rates of water runoff into rivers. Some scientists argue that
increased agricultural land clearance in the mountainous regions above
Bangladesh has led to more frequent and intense flooding along the lower
Ganges River. The building of levees and dikes along rivers can constrict their
channels and keep high flows from flooding adjacent low-lying areas. Much of the
lower Mississippi River is constrained by artificial levees and controlled by water
diversion channels. Decreased incidents of flooding by the Mississippi is causing a
loss of bald cypress swamps along the lower portions of the river.
Not all human modifications of riparian vegetation make stream banks
more unstable. When compared to natural forest vegetation, the establishment of
pasture grasses has been shown to cause decreased channel erosion rates in some
small stream systems. The dead branches and trunks of trees that fall into
forested streams are important in causing ripples and pools for aquatic life and in
directing erosive river flow. In some instances, clearing forest cover and replacing
it with grass may artificially stabilize floodplains and be destructive to aquatic
ecosystems and riparian vegetation.

FIGURE 5.1
OTHER PHYSICAL DISTURBANCES mature fore!
Smaller aval
Avalanches and landslides are important agents of disturbance in mountainous upper part o
regions. Avalanches are flows of snow and debris that travel with great speed and grasses, shn
 OTHER PHYSICAL DISTRUBANCES 123

I Biological force. Avalanches are capable of destroying entire stands of trees. Avalanches
often occur along the same relatively narrow tracks year after year (Fig. 5.15). The
occurrence of frequent avalanches precludes the growth of mature trees, and ava-
lanche tracks are typically vegetated by grasses, herbs, shrubs, and saplings. Within
forested regions, avalanche tracks provide important open habitat for shade intol-
erant plants. The herbaceous and shrubby vegetation of avalanche tracks is impor-
tant feeding ground for deer, moose, and bear within forested mountain areas.
The term landslide encompasses a number of different types of slope fail-
ure. Landslides can be defined as sudden mass movements of rock, soil, and
organic material downslope along sheer surfaces. These movements can range
from rockfalls off of steep cliffs, to slides and slumping of large coherent blocks of
wet or dry earth, to debris flows of water-saturated material. Landslides destroy
vegetation by removing soil, shearing stems and trunks, and burying plants. The

lS

~rve to increase river flows

>val of vegetation and soil
. Some scientists argue that
untainous regions above
flooding along the lower
g rivers can constrict their
)W-lying areas. Much of the
:es and controlled by water
the Mississippi is causing a
Jf the river.
ation make stream banks
:ation, the establishment of
mnel erosion rates in some
tks of trees that fall into
Jools for aquatic life and in
forest cover and replacing
I be destructive to aquatic

isturbance in mountainous
:ravel with great speed and
FIGURE 5.15 A large avalanche near Taos, New Mexico, has cleared a swath through
mature forest that had become established on the lower portion of the avalanche track.
Smaller avalanches that occur more regularly have kept forest from establishing on the
upper part off the avalanche track. Avalanche tracks can provide habitat for light-demanding
grasses, shrubs, and small trees. They can also serve as natural fire breaks in the forest.
124 CHAPTER 5 DISTURBANCE

frequency and magnitude of landslides are controlled by many factors, including Volcan
slope angles, bedrock type, soil properties, rainfall, and vegetation cover. Human noes can cr'
activities, such as deforestation, grading of slopes, or increasing ground water cover existir
through upslope irrigation, can produce increased landslide activity. A study of During vole<
landslides in northeast Puerto Rico found that 53% of the landslides recorded and depositc
between 1964 and 1989 were associated with road building. for primary ~
Landslides are a particularly important form of disturbance in steep tropi- areas. Hot g<
cal landscapes where deeply weathered rock and high rainfall promote slope volcanic eru1
instability. Hurricanes are important in promoting tropical landslides as they strip prone to ian<
away protective vegetation cover and produce extremely intense rainfall. It is a good exam
estimated that 0.8 to 7.5% of tropical forests are affected by landslides in the Saint Helens
Caribbean region each century. Although huge tropical landslides do occur, most of the cone. I
are small in area. Studies in Jamaica indicate that 80% of the recent landslides tion called tl
were less than 0.08 ha in area, while a similar study in Puerto Rico showed that peak. The bla
77% of recent landslides were less than 0.4 ha in size. stumps of mi
Succession patterns following tropical landslides appear to be highly south side of
dependent on the proximity of seed sources and the degree to which the original river valleys.
soil is disturbed by the landslide. The initial vegetation on tropical landslides i were deposit
usually dominated by shade intolerant plants such as tree ferns, climbing ferns. unable to est
grasses, large herbs, shrubs, and vines. Completely denuded and unstable surface nus lepidus),
may be colonized by mosses and other nonvascular plants. Establishment of fern reserves for t
and shrub thickets leads to decreased soil evaporation, increased moisture avail- tion. These fe
ability, and more soil stability. Increasing moisture and soil stability eventually the bare sites.
facilitate tree establishment. As the trees grow, they shade-out the pioneer vege- (Aster ledopf.
tation. In Puerto Rico, the vegetation on landslides can revert to forest within 50 relied on the
years. However, on sites where no residual soil remains, it may take several hun-
dred years before a mature forest cover is established.
In cold mountain regions, the paths of debris flows often coincide with ava-
lanche tracks. The frequency of both forms of disturbance can keep such areas in
permanent states of disruption when compared to surrounding areas. However.
not all landslides in cold regions are restricted to avalanche paths. Deep unstable
soils, slope angles greater than 30°, and a lack of deeply rooted vegetation cover
contribute to landslides. Debris flows are promoted by the combination of moist
soils due to melting snow and intense rainfall. Analysis of satellite images from a
small section of Glacier National Park in Montana produced evidence of 73
recent debris flows associated with avalanche paths, 48 associated with slope fail-
ures beneath large snow patches, and 28 flows down ravines that were not ava-
lanche paths. The widespread occurrence of landslide activity in the northern
Rocky Mountains has been shown to depress the elevation of treeline by exclud-
ing the growth of forest trees in protected ravines and other sites that would oth-
erwise provide favorable habitat for trees.
Fires can promote landslides by stripping away vegetation cover and desta-
bilizing slopes. In the semiarid chaparral vegetation of California, large fires in
the late summer are often followed by severe landslide activity in the following
winter. Aside from removing the vegetation cover, the fires also decrease the
moisture-repelling properties of the soil surface, leading to greater absorption of
water and increased soil instability. Interestingly, research in the Canadian Rocky
Mountains indicates that the shrubby vegetation associated with avalanche FIGURE 5.16
tracks and debris flow paths form important fire breaks that restrict the spread of angustifolium) i~
fires in the surrounding conifer forests. trees ki lied by th
 OTHER PHYSICAL DISTRUBANCES 125

I by many factors, including
d vegetation cover. Human
'r increasing ground water
ndslide activity. A study of
of the landslides recorded
I ding.
disturbance in steep tropi-
igh rainfall promote slope
Jicallandslides as they strip
!mely intense rainfall. It is
fected by landslides in the
allandslides do occur, most
J% of the recent landslides
n Puerto Rico showed tha t
ides appear to be highly
Iegree to which the original
on on tropical landslides is
s tree ferns, climbing ferns.
!Uded and unstable surfaces
!ants. Establishment of fern
n, increased moisture avail -
md soil stability eventually
hade-out the pioneer vege-
m revert to forest within 50
ns, it may take several hun-
Volcanic activity is one of the most spectacular forms of disturbance. Volca-
noes can create disturbances through several different processes. Lava flows
cover existing ecosystems and provide new landscapes for primary succession.
During volcanic eruptions, large amounts of fine ash, called tephra, are created
and deposited. Tephra can bury existing ecosystems and provide a new surface
for primary succession. Explosive eruptions can denude the landscape over large
areas. Hot gases, lava, and ejected material can cause fires in the vicinity of the
volcanic eruption. Finally, unstable land surfaces left by volcanic eruptions are
prone to landslides. The eruption of Mount Saint Helens in Washington provides
a good example of volcanism as an agent of disturbance. On May, 18, 1980 Mount
Saint Helens was rocked by an explosive eruption that removed 400 m off the top
of the cone. Due to the explosive nature of the eruption, a larger area of devasta-
tion called the blast zone extended some 18 km north of the former mountain
peak. The blast zone was typified by bare soil surfaces and the broken and burned
stumps of millions of dead trees. Debris flows of wet sediment came down the
south side of the mountain and extended considerable distances along adjacent
river valleys. In the years after the eruption, the wind borne seeds of many plants
were deposited on the bare ash and lava surfaces. Most of these seeds were
unable to establish plants on the dry and nitrogen-deficient soils. Lupines (Lupi-
nus lepidus), however, were able to establish. They have seeds that contain starch
reserves for the seedlings and roots that possess nodules that allow nitrogen fixa-
tion. These features allowed lupines to dominate the early vegetation on many of
the bare sites. The patches of lupines were in turn invaded by plants such as asters
(Aster ledophyllus) and fireweed (Epiliobium angustifolium), which apparently
relied on the more favorable soils created by the lupines (Fig. 5.16). In this case,

ws often coincide with ava-

mce can keep such areas in
trrounding areas. However.
anche paths. Deep unstable
)ly rooted vegetation cover
'Y the combination of mois:
is of satellite images from a
t produced evidence of 73
8 associated with slope fail-
ravines that were not ava-
je activity in the northern
•ation of treeline by exclud-
1other sites that would oth-

vegetation cover and desta-

of California, large fires in
de activity in the followin g
the fires also decrease the
ing to greater absorption o~
uch in the Canadian Rock ~
associated with avalanche FIGURE 5.16 Four years after the 1980 eruption, a dense growth of fireweed (Epilobium
cs that restrict the spread o: angustifolium) is established on part of the Mount Saint Helens blast zone. The remains of
trees killed by the blast cover the slopes.
126 CHAPTER 5 DISTURBANCE

the lupines are referred to as nurse plants because their presence facilitates the
increased by 5'
establishment and growth of the other species. In most cases, the lupines were
nutrient-rich w
replaced by other plant species after about five years. The process of succession i
led to declines
still underway on Mount Saint Helens, and it will likely take decades before
fish population
coniferous forest cover is reestablished.
Islands are der
ing El Nino, thE
events recur e"
PATHOGENS turbances. In tJ:
decreases and 1
The outbreak of pathogens, especially plant-destroying insects, is an important
appears to res1
and widespread form of disturbance. For example, tree diseases and destructive
Islands througt
insects destroy some 60 million cum of Canadian timber each year. This is second
Although
only to fire which destroys an average of 80 million cum of timber each year. Per-
range reductior
haps the most destructive insect in the forests of northern North America is the
the corals havE
spruce budworm (Choriostoneura fumiferana). During a peak outbreak in 1978.
processes such
over 72 million ha of forest were affected by this insect. The larvae eat the foliage
ing mollusks, he
and young cones of spruces and firs, but the principal host is balsam fir (Abies bal-
of Millepora co
samea). After three to four years of infestation, even mature trees are killed.
An exam
Severe outbreaks of spruce budworm occur approximately every 29 years and
biological distu
may take 7 to 15 years to subside. During outbreaks, individual trees and whole
pathogen killec
stands may be killed, opening many different-sized gaps in the forest. The dry
loss of the sea l
dead foliage produced during budworm outbreaks promotes the ignition and
grew corals at r
spread of high-intensity fires.
urchins were a
An interesting example of how disturbance by a pathogen can be transmit-
the loss of over
ted through different levels of an ecosystem is provided by outbreaks of anthrax
As in ma1
among grazing and browsing mammals in Africa. Anthrax is a bacterium that
controlled by t
causes death in mammals. Following outbreaks, anthrax can remain dormant in
algae establish '
soils for years. In Tanzania epidemics of anthrax among impalas (Aepyceros
urchins and fish
melampus) caused widespread mortality in 1961, 1977, and 1984. The declines in
dominance. ThE
the impala population led to decreased browsing on shrubs and the invasion of
only destroy m<
some grasslands by shrubby species and umbrella thorn trees (Acacia tortilis). In
coral that event
this manner, the disturbance provided by outbreaks of anthrax among mammals
largely depend<
contributes to maintaining the mosaic of grassland, shrubs, and trees that makes
surfaces and no
up the savanna landscape.
occupancy of t
worms, crustace
sitivity of coral
MARINE DISTURBANCES fine sediments
American coral
Marine ecosystems are also subject to physical and biological disturbances. Phys-
also has a burna
ical disturbances include abrupt changes in water temperature and salinity, sub-
species that fee<
marine landslides, and episodes of rapid erosion and deposition. The impact of
after the sea un
the 1982-1983 El Nino event demonstrates how different marine communities
are impacted by rapid changes in sea surface temperatures. During El Nino
events, there is a rapid warming of surface waters in the eastern equatorial region
of the Pacific Ocean. The 1982-1983 event produced unusually strong warming of
3° C to 4° C off the coast of Central and South America. The higher temperatures EYWORDSAND
rendered reef-building corals more sensitive to ultraviolet radiation and caused
them to expel their symbiotic algae. Within two to four weeks the corals died. The -.:\·entitious roots
~~~· ground mortality
mortality extended to corals growing as deep as 20 m below the sea surface. In
-mopy fire
some reefs, up to 97% of the coral died. Off the Galapagos Islands temperatures ~..:m a x

~ir presence facilitates the
>st cases, the lupines were
he process of succession is
kely take decades before
1g insects, is an important
e diseases and destructive
~r each year. This is second
n of timber each year. Per-
lern North America is the
~a peak outbreak in 1978.
. The larvae eat the foliage
Jst is balsam fir (Abies bat-
a mature trees are killed.
nately every 29 years and
ndividual trees and whole
aps in the forest. The dry
•romotes the ignition and
pathogen can be transmit-
d by outbreaks of anthrax
tthrax is a bacterium that
ax can remain dormant in
nong impalas (Aepycero s
, and 1984. The declines in
.hrubs and the invasion of
n trees (Acacia tortilis). In
· anthrax among mammals
rubs, and trees that makes
logical disturbances. Phys-
perature and salinity, sub-
deposition. The impact of
:rent marine communities
eratures. During El Nino
: eastern equatorial region
msually strong warming of
1. The higher temperatures
iolet radiation and caused
weeks the corals died. The
below the sea surface. In
agos Islands temperatures
KEY WORDS AND TERMS 127
increased by soC. This warming caused a decline in the upwelling rate of deep,
nutrient-rich waters. The warmer water temperatures and decreased upwelling
led to declines in marine plant productivity and a southward movement of many
fish populations. Marine iguanas (Amblyrhynchus cristatus) of the Galapagos
Islands are dependent on seaweed for food. Due to decreased productivity dur-
ing El Nino, the iguanas suffered population declines of 60 to 70%. Since El Nino
events recur every 5 to 11 years, the iguana population has adapted to such dis-
turbances. In the year following an El Nino, the age for first breeding by females
decreases and the average egg clutch size increases. Thus, the iguana population
appears to respond to this periodic disturbance and persists on the Galapagos
Islands through increased fertility following El Ninos.
Although such climatic extremes may be rare, their occurrence can lead to
range reduction and extinction. In contrast to the recovery of the marine iguana,
the corals have suffered much more long-lasting damage. Erosion by physical
processes such as storms and bioerosion, caused by coral-eating fish and burrow-
ing mollusks, has led to complete destruction of some reef systems. Several species
of Millepora coral may have been driven to extinction by the 1982-1983 El Nino.
An example from coral ecosystems in the Caribbean demonstrates how
biological disturbances can impact marine environments. In 1983 an unknown
pathogen killed more than 95 % of long-spined sea urchins (Diadema spp.) . The
loss of the sea urchins decreased grazing pressure on seaweeds, which then over-
grew corals at many sites. The corals that were covered by seaweed died. The sea
urchins were a keystone species in the coral ecosystem, and their decline caused
the loss of over 90% of corals surrounding Jamaica.
As in many terrestrial ecosystems, succession in coral reef systems is often
controlled by the species that reach the site first. In general, large filamentous
algae establish quickly upon bare surfaces and dead coral. Grazing pressure from
urchins and fish generally limits algal growth and allows corals to grow and regain
dominance. The long-term decrease in sea urchins that graze on algae may not
only destroy mature coral, but also inhibit its restablishment. The exact species of
coral that eventually dominates the patch in the latter stages of succession is often
largely dependent on the species of coral larvae that first settle there. The hard
surfaces and nooks and crannies produced by mature coral subsequently facilitate
occupancy of the reef by a wonderful assortment of sea anemone, mollusks,
worms, crustaceans, and fish . Unfortunately, human activity has increased the sen-
sitivity of corals to physical and biological disturbance. Increased deposition of
fine sediments due to land clearance has choked and weakened many Central
American coral communities. The recent destruction of corals in the Caribbean
also has a human component. Extreme overfishing off Jamaica removed many fish
species that feed on seaweed. This made the explosive growth of seaweed possible
after the sea urchins were killed by the unknown pathogen.
WORDS AND TERMS
-•.:s entitious roots Crown fire Pneumatophores
::.::.::kground mortality Disturbance Primary succession
- - opy fire Facilitation Random model
ax Inhibition model Rhizome
128 CHAPTER 5 DISTURBANCE
Riparian forests Sere
Scarification Serotinous cones
Secondary succession Succession
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 6
I.

CHAPTER
Dice
0 Arctic tundr:

COMMUNITIES, FORMATIONS, []Taiga

1111 Pacific coas
AND BIOMES • Palouse pra
.lntermountc
shrub-stepp
woodlands,
0 Califorian fo
and alpine\
Imagine you are taking a road trip across North America. You start driving south- California 91
west from the border between New Brunswick and Maine. At first, you are sur- chaparral ar
rounded by dark forests of coniferous trees such as pine, spruce, and fir (Fig. 6.1). Mojave and
deserts
For hours you drive through stands dominated by these same trees. Long before []Rocky Mour
you cross into Pennsylvania, the forest has become quite different. In place of and alpine\
pines and spruces, the dominant trees are broadleaved deciduous genera such as 0 Central prai
maple, oak, ash, and hickory. Crossing central Kansas you find that the deciduous and plains
forest has been replaced by open grasslands. West of Denver you climb into the • Mixed decic
Rocky Mountains and discover that the vegetation is again dominated by conif- • Chihuahuar
and woodla
erous forests of pine, spruce, and fir. For a short time, near the crest of the Rock- i] Appalachiar
ies, you see treeless meadows of low herbs and grasses. After you have descended
from the mountains and drive across southern Nevada, you pass through a vast [ill] Piedmont o•
expanse of desert with small shrubs and bare ground. Approaching the end of iJl Coastal plai
swamps, an
your trip at the Pacific Coast of central California, you find dry grassy meadows [J Tropical forE
dotted with scattered oaks. As an astute biogeographer, you have noticed how
certain combinations of species and vegetation types seem to dominate the land-
FIGURE 6 . 1
scape over large areas and then disappear altogether. You might wonder what Barbour and Bil
controls such associations of species and causes large changes in vegetation? Bio-
geographers have marveled at these geographic changes in vegetation and con-
sidered these same questions for hundreds of years. Finally, you have noticed that
much of the natural vegetation along your route is restricted to areas such as biomes. Since
parks and reserves. In many cases, urban and agricultural land use has led to the different bion
removal of natural forests or grasslands. The causes and extent of such human elude with a t<
impacts have also long been a topic of concern to biogeographers.
In the preceding chapters, we have seen how physical and biological factors
influence the geographic distributions of individual species. It is clear that certain ..... OMMUNITIES
groups of species are almost always found living together. Species may occur
together because they are dependent on each other or because they have very sim- fbe modern c
ilar tolerances and requirements from the environment. Some biogeographers British ecolo:
concentrate their studies on identifying these recurring groupings of species and species of ani1
understanding why such associations occur. At the smallest spatial scale, such stud- to those foun
ies focus on single communities described from one location. At the largest spatial define the co1
scale, the terrestrial biosphere is often subdivided into a number of supercontinen- place or habil
tal community units called biomes. The biomes are grossly generalized subdivi- munity. In pn
sions of the biosphere based on similarities in vegetation structure and climate. In defined by thE
this chapter we will look at some basic principles of community ecology that all called an eco
biogeographers should be familiar with. We will then consider how vegetation species that is
structure can be systematically described and used to subdivide the biosphere into study all of t

132

IONS,
\ woodlands, and forests
ca. You start driving south-
faine. At first, you are sur-
te, spruce, and fir (Fig. 6.1).
;e same trees. Long before
tuite different. In place of
l deciduous genera such as
•ou find that the deciduous
Denver you climb into the
again dominated by conif-
lear the crest of the Rock -
After you have descended
a, you pass through a vast
!. Approaching the end of
1 find dry grassy meadows
er, you have noticed how
;;em to dominate the land-
. You might wonder what
1anges in vegetation? Bio-
~es in vegetation and con-
ally, you have noticed that
estricted to areas such as
ralland use has led to the
md extent of such human
~ographers .
;ical and biological factors
cies. It is clear that certain
.ether. Species may occur
ecause they have very sim-
:nt. Some biogeographers
groupings of species and
est spatial scale, such stud-
ttion. At the largest spatial
number of supercontinen-
ossly generalized subdivi-
n structure and climate. In
>mmunity ecology that all
consider how vegetation
bdivide the biosphere into
COMMUNITIES 133
I,
Dice
0 Arctic tundra .
0Taiga
B Pacific coastal
• Palouse prairies
.Intermountain deserts,
shrub-steppes,
0 Califorian forests
and alpine vegetation
D California grasslands,
chaparral and woodlands
~ Mojave and Sonoran
deserts
0 Rocky Mountain forests
and alpine vegetation
0 Central prairies
and plains
Mixed deciduous forests .
• Chihuahuan deserts
and woodlands
0 Appalachian forests
miJ Piedmont oak-pine forests
[1] Coastal plain forests, bogs,
swamps, and strand
[]Tropical forests
FIGURE 6.1 The major vegetation communities (formations) of North America (after
Barbour and Billings, 2000).
biomes. Since an understanding of the biome concept and an appreciation of the
different biomes are fundamental knowledge for every biogeographer, we'll con-
clude with a tour of the world 's biomes.
OMMUNITIES
The modern concept of biological communities can be traced to the work of the
British ecologist E . Forbes in the mid-nineteenth century. He described the
species of animals found in British coastal waters and contrasted this assemblage
to those found in the Mediterranean Sea. Today ecologists and biogeographers
define the community as an assemblage of all organisms living in a prescribed
place or habitat. We could also consider the entire biosphere as one large com-
munity. In practice, the area encompassed by an individual community is usually
defined by the researcher. The spatial boundary between adjacent communities is
called an ecotone. The term community type is used to describe a group of
species that is typically found in a specific type of habitat. For example, we could
study all of the organisms living under many different damp stones, note which
134 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

species are generally found in such habitat, and describe this collection of recur- The c
ring coexisting species as the "damp stone community type." ogist H.A.
In many cases, researchers are only interested in the plants or the animal! ceive as bi'
within the community. Plant ecologists employ the term association when refer- coexist bee
ring to groups of plant species commonly found in similar habitats. The fores: resource d
dominated by sugar maple (Acer saccharum), red maple (Acer rubrum), white own indivi,
oak (Quercus alba), red oak (Quercus rubra), ash (Fraxinus), and hickory Since the~
(Carya), found on mesic sites in northeastern North America, is an example of c. thought of
plant association. Forest plant ecologists generally refer to the individual sites as munity cor
stands. To simplify descriptions, plant biogeographers and foresters otter. If thE
describe associations in terms of the dominant plant species and call such associ- cations. Le
ations dominance types. Central America has forested areas that contain anum - in the field
ber of different plant species, but Caribbean pine (Pinus caribaea) is one of tb nance, stn
region's more common canopy-forming tree. This assemblage can be referred to developed
as the Caribbean pine dominance type. removal o
A number of properties can be measured or described for a community. Th impact. If 1
properties can relate to either static features of community structure or dynami tively diffi,
properties of community function. Some of the most important properties o · communit;
community structure include (1) the species that occur, (2) overall species diver- tones, com
sity, (3) the species that is most abundant, (4) the species that is most dominant in The overal
terms of biomass or structure, (5) species evenness (i.e., the degree to which th A co
number of individual organisms are evenly divided between the different species the graphi'
of the community) , (6) the general growth form or structure of the organisms or dient, soil J
vegetation that makes up the community, (7) the number of trophic levels, an species an
(8) primary productivity. Properties of ecosystem function include features such ronmental
as rates of energy cycling, rates of nutrient cycling, or exchange rates of gase_ might be c
between the community and the atmosphere. tight grOUJ
Although the concept of subdividing the geographic distribution of life into figuration
community types seems straightforward and useful, it has been one of the most communit:
contentious problems in biogeography and ecology. The nature of communities species mi.
has been a particularly keen subject of debate among plant ecologists. The prob- between c
lem can be illustrated by examining the two classical and diametrically opposed species ou
views of natural communities. One view was initially outlined by S.A. Forbes in alistically •
1887 in the United States and forcefully developed by the American ecologist tion due tc
and biogeographer F.E. Clements in 1916. From the last chapter we recall that individuali
Clements was also a founder of the classical succession theory and the concept of other spec
the climax community. Clements argued that because of interactions between Inm'
species during the climax phase of succession and because of facilitation during \ ents appe<
sera! stages, plants and animals are bound together into tight communities that in a numl::
behave like superorganisms. In this view, all of the species in a community are where mm
like the different organs of a larger organism. All the species depend on the func- A classic o
tioning of the other species in the community and cannot persist outside of this Oregon a
association. Clements was much influenced by the evolutionary theory pro- vegetation
pounded by Charles Darwin, and implicit in Clements' theory is the assumption menziesii)
that the species in these communities coexisted and evolved (see Chapter 9) dominate(
together over very long periods of time. The Clementsian view of community was (Chamaec
embraced by many famous ecologists of the mid-twentieth century such as A. G. ever, Pacif
Tansley in England and J. Braun-Blanquet in France. One attraction of the super- Douglas f
organism model is that it allows biogeographers to assume that the community species is
types that are described and mapped are stable features of the biosphere. aries betw

:ribe this collection of recur-
y type."
in the plants or the animals
erm association when refer-
similar habitats. The fores t
naple (Acer rubrum), white
;h (Fraxinus), and hickory
America, is an example of a
:fer to the individual sites as
\
phers and foresters often
species and call such associ-
:d areas that contain anum-
'inus caribaea) is one of th e
;emblage can be referred to
cribed for a community. The
nunity structure or dynami c
)St important' properties of
ur, (2) overall species diver-
:ies that is most dominant in
[.e. , the degree to which th e
:!tween the different species
ructure of the organisms or
mber of trophic levels, and
1ction include features such
or exchange rates of gases
phic distribution of life into
t has been one of the most
The nature of communitie
plant ecologists. The prob-
and diametrically opposed
outlined by S.A. Forbes in
by the American ecologist
last chapter we recall tha t
n theory and the concept of
se of interactions between
cause of facilitation during
nto tight communities that
pecies in a community are
pecies depend on the func-
nnot persist outside of this
evolutionary theory pro-
s' theory is the assumption
evolved (see Chapter 9)
an view of community was
1tieth century such as A. G.
ne attraction of the super-
sume that the communitY
es of the biosphere.
COMMUNITIES 135
The opposing view of the community was developed by the American ecol-
ogist H.A. Gleason in the early twentieth century. He argued that what we per-
ceive as biological communities are simply areas of similar habitat where species
coexist because they have somewhat comparable environmental tolerances and
resource demands. Gleason went on to argue that each of these species has its
own individual sets of tolerances and demands and do not always occur together.
Since the species do not always have to occur together, communities cannot be
thought of as superorganisms. Gleason's model is called the individualistic com-
munity concept.
If the superorganism community concept is correct, it has important impli-
cations. Let's make an extreme case. First, communities should be easy to define
in the field. They should be homogeneous in terms of species composition, domi-
nance, structure, and so on, and quite distinctive from adjacent communities
developed in a different habitat. Second, from the view of conservation, the
removal of one or more species from a community should have a significant
impact. If the individualistic concept is correct, then communities should be rela-
tively difficult to define on the basis of species composition. Species from one
community will likely be found in several other habitats. Instead of sharp eco-
tones, communities will blend into each other and be more difficult to discern.
The overall community should be less sensitive to the removal of one species.
A convenient way of summarizing these two opposing views is to consider
the graphical representation of several plant species along an environmental gra-
dient, soil moisture, for example (Fig. 6.2). The examination of the distribution of
species and communities relative to environmental conditions and along envi-
ronmental gradients is known as ordination analysis. On such ordinations, species
might be distributed in three different ways. First, they might be distributed in
tight groupings with sharp boundaries between different communities. This con-
figuration will occur if there is strong dependency between species within the
community and competition with species from outside of it (Fig. 6.2a). Second,
species might be segregated into tight community groupings, but the boundaries
between communities may be less distinct because of low competition with
species outside of the community (Fig. 6.2b ). All of the species may be individu-
alistically distributed along the gradient but have sharp boundaries in distribu-
tion due to competition with other species (Fig. 6.2c). Finally, the species may be
individualistically distributed with no apparent competition or interaction with
other species (Fig. 6.2d) .
In most cases, the natural distribution of species along environmental gradi-
ents appears to fit an individualistic model. Some species are widely distributed
in a number of communities and others are more restricted, but distributions
where many species have similar limits and only occur together are uncommon.
A classic ordination study along a moisture gradient in the Siskiyou Mountains of
Oregon and California illustrates this idea. Based on dominance types, the
vegetation of the mountains can be divided into a Pacific Madrone (Arbutus
menziesii) shrub association on dry sites, a Douglas fir (Pseudotsuga menziesii)-
dominated forest association on intermediate sites, and a Port Orford cedar
(Chamaecyparis lawsoniana)-Douglas fir forest association on moist sites. How-
ever, Pacific Madrone is relatively common on all but the very wettest sites, and
Douglas fir is common on all sites (Fig. 6.2). When the distributions of these
species is graphed along a moisture gradient, it is difficult to detect sharp bound-
aries between the communities. The boundaries between supposed communities
136 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

only the past fev

~~~~~~
argued, the rela1
that they have n•
It is true tl
gradients or on
ern North Amer

~ ~~
vegetation struc
sus a loblolly pi
southern CalifoJ
one ascends froJ
- Pseudotsuga big cone Doug!<
menziesii found at higher
300
different sites. I
*
t5
250
- Chamaecyparis
lawsoniana dominance-type

t
2! 200 scales and provi
- Arbutus menziesii
d 150 . . bution of life.
,s; 100
(/)

~ 50
•
~ ><'
Given the
phers has gone i
en spatial boundar
releve approach
(moist) the researcher
observed in the
FIGURE 6.2 The ordination of plant species along a hypothetical environmental gradient.
(a) Species with strong dependency between species within the community and
Sociability Clas:
competition with species from outside of it. (b) Species with strong dependency but low species; (2) the 1
competition with species outside of the community. (c) Individualistically distributed and (3) the Abu
species exhibiting competition with other species. (d) Individualistically distributed species tation. The Bra1
with no apparent dependency or competition . (e) The actual distribution of dominant tively rapid to u·
shrubs and trees along an altitudinal gradient in the Siskiyou Mountains of Oregon and Another a
California (after Whittaker, 1960) species present
larity of stands.
stands, we can c
any single spec
would become even less clear if nondominant plants such as small shrubs and
square meter 01
herbs were included on the gradient diagram.
contain the spec
In contrast to gradient studies, the geographic mapping of the local distribu-
dominance (whi
tions of plant communities often detects clear boundaries between communities.
sity, frequency,
In some cases, this may be because the landscape does not really provide a contin-
occupied by the
uum of different environmental conditions, but rather is composed of a patchwork
wiqely used set
of sites that offers highly contrasting habitat. For example, a landscape consisting
species (x), thes1
of small rolling hills provides either very dry sites on hills or very moist sites in the
hollows. We observe two very different plant associations for these two sites, but
R
the differences are exaggerated because there are no intermediate semi-moist
sites available to provide a true continuum of environmental conditions.
Perhaps the most damaging evidence against the superorganism concept R
comes from studies of past vegetation change. Research on the past distributions of
plant species based on fossil evidence indicates that most modern plant associations
have only been in existence for the past 10,000 years. Many communities of plants R
have only existed in their current state for relatively short periods. For example,
hemlock trees (Tsuga canadensis) are important species in the northern forests of I
the midwestern United States. Hemlock trees have been a part of these forests for

~ that they have not evolved together as superorganisms over long periods of time.
~
\ one ascends from the dry chaparral shrubs at low elevation to the stands of huge
"
hetical environmental gradient.
he community and
;trong dependency but low
dualistically distributed
Jalistically distributed species
Jistribution of dominant
Mountains of Oregon and
; such as small shrubs and
1pping of the local distribu-
ries between communities.
not really provide a contin-
s composed of a patchwork
1ple, a landscape consisting
Us or very moist sites in the
ons for these two sites, bur
o intermediate semi-moist
rrental conditions.
he superorganism concept
on the past distributions of
t modern plant associations
lany communities of plants
hart periods. For example.
s in the northern forests of
n a part of these forests for
COMMUNITIES 137
only the past few thousand years. As the noted American ecologist, M.B. Davis, has
argued, the relative youthfulness of many of our present plant associations means
It is true that there is much individualism in the distributions of species along
gradients or on the landscape. However, anyone hiking in the forest of southeast-
ern North America can readily see that there is a difference in the plant species and
vegetation structure found in a wet bald cypress (Taxodium distichum) swamp ver-
sus a loblolly pine (Pinus taeda) stand on adjacent dryland. Similarly, a hiker in
southern California cannot help but be struck by the difference in vegetation as
big cone Douglas fir (Pseudotsuga macrocarpa) and jeffrey pine (Pinus jefferyi)
found at higher elevations. Clearly, different associations of species do exist on
different sites. Even if their spatial or environmental boundaries may be fuzzy,
dominance-type communities, can often be delineated at many different spatial
scales and provide a generalized way to describe and study the geographic distri-
bution of life.
Given the often indistinct nature of communities, much work by biogeogra-
phers has gone into identifying communities and associations and mapping their
spatial boundaries. In Europe, many biogeographers use the Braun-Blanquet
releve approach to describe and classify plant communities. Using this approach,
the researcher makes a number of qualitative observations on the species
observed in the community. The observations fall into three major areas: (1) The
Sociability Classes relate to the spatial grouping of the plants belonging to the
species; (2) the Vitality Classes represent the reproductive ecology of the species;
and (3) the Abundance Classes measure how common the species is in the vege-
tation. The Braun-Blanquet technique and similar releve approaches are rela-
tively rapid to use in the field but suffer from their subjective nature.
Another approach that is widely used is to quantitatively survey the plant
species present and then to mathematically compare the similarity and dissimi-
larity of stands. When a quantitative sampling is conducted for a number of
stands, we can calculate a number of measures to describe the community. For
any single species these measures include density (the number of plants per
square meter or hectare), frequency (the percentage of samples or stands that
contain the species) , cover (the amount of ground surface occupied by the plant),
dominance (which can be calculated in many ways including the summing of den-
sity, frequency, and cover, for example), and basal area (the area per hectare
occupied by the cross-section trunks of tree species). For forest trees, there is a
widely used set of quantitative measures of community structure. For a given
species (x), these include:
. . individuals of species x
Relative Density= . d .. d f . x 100
tota 1m IVI ua1s o a11 species
. Fr frequency of species x
R e IatJve equency = X 100
sum of frequency of all species
. D . basal area of species x
R e Iat1ve ommance = X 100
total basal area of all species
Importance Value = Relative Density + Relative Frequency +
Relative Dominance of Species x
138 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES p

In conducting surveys of stands, it is important to examine a large enou ~ ·­ types share son
area to provide a valid sampling of the true species composition. Samples o: and structure. I
species composition are often made by laying out square subsampling areas types to be sub:
called quadrats. If the quadrat is too small, it will not capture all of the trut Alternati-
species diversity of the stand. If the quadrat it is too large, it takes too long tc a reticulate cla
sample and yields no additional information on species diversity. When samplin~ sidered a sepai
large areas, or examining dominant tree species only, line or belt transects ar- might be the fc
often used. A line through the stand is surveyed and all trees touching it, or grO\Y- These associati
ing within a certain distance of the line, are identified and recorded. For ver: erally distinct l
large stands, where only the dominant trees are being studied, point-quarter tran- number of mal
sects are used. For this technique, very long line transects are established an · mations of rec
divided into widely separated points. The trees growing nearest to each point, ir. can help deten
each of the four cardinal directions, are identified and recorded. In addition tC' sidered hieran
the size of the quadrat or transect, the placement of the sampling points is critica: The imp~
in community analysis. In many cases, researchers use random number tables to communities.!-
generate geographic sampling coordinates for quadrats, transects, and nodes. implicitly assu
Once quantitative information is obtained from stands, a number of mathe- dance, and doi
matical techniques are used to compare the species compositions of the samples from that of t
and determine whether there are differences between different communities an marked chang
whether two communities representing the same supposed habitat are really sim- pare commun:
ilar. The degree of difference in the species composition of two different habitats successional st
or community types is often referred to as beta diversity. The degree of differenc
between stands of the same community type is called alpha diversity. One of th
simplest mathematical measures of beta or alpha diversity is the index of similar- -:.ANT PHVSIOG
ity. The index is based on the presence or absence of species in two different STR UCTURE, AN
stands and has the form:
In addition tc
2 x the no. of species that occur in both stands basis of plant 1
Index of Similarity
total species in stand 1 + total species in stand 2 .such schemes,
are used to d(
If there is no similarity between the stands (i.e. , no shared species), the index will vegetation st1
equal 0. If the species present in the two stands are exactly the same, the index America too1
will equal 1. There are many other mathematical forms of this general approach nized on the
using species counts from two stands to calculate similarity. from treeless
The problem with the index of similarity is that it does not consider the dif- southern Me1
ferences between the two stands in terms of the abundance of the different units can be 1
species. One stand may be dominated by a single tree species and contain a few \J.E.B. Warmil
shrubs, while the other might be dominated by shrubs and contain one or two have been inf
trees. The species present in the two stands could be identical, but the structure of since earliest
the stands and habitat would be completely different. In view of this considera- ways to den<
tion, some researchers use more sophisticated mathematical measures that structural cla
include the abundance of individuals of each species. tural classific
The classification schemes that are used to group sites into associations or these classifi<
community types have two general forms: hierarchical and reticulate. For the and, the phy
hierarchical form, we assume that there is one major community type, deciduous insight into tl
forest, for example. This large community can be subdivided into smaller units the plants, m
that are generally similar to each other but show consistent differences in their and structun
species composition. One example might be the grassland association of the mid- ties on differ
western United States. The extensive grasslands can be broken down into a tall Just a~
grass prairie in the East and a short grass prairie in the West. The two grassland species pres

t to examine a large enough
es composition. Samples of
t square subsampling area
not capture all of the true
•O large, it takes too long to
ies diversity. When sampling
tly, line or belt transects are
ill trees touching it, or grow-
tied and recorded. For ven
?; studied, point-quarter tran-
ansects are eStablished and
ing nearest to each point, in
nd recorded. In addition to
he sampling points is critical
;e random number tables to
its, transects, and nodes.
1 stands, a number of mathe-
:ompositions of the samples
1 different communities and
Josed habitat are really sim-
tOn of two different habitats
ity. The degree of difference
l alpha diversity. One of th
~rsity is the index of similar-
of species in two different
:cur in both stands
I species in stand 2
ared species), the index will
exactly the same, the index
ns of this general approach
larity.
it does not consider the dif-
bundance of the different
~ species and contain a few
bs and contain one or two
!entical, but the structure of
. In view of this considera-
tthematical measures that
tp sites into associations or
cal and reticulate. For the
ommunity type, deciduou
Jdivided into smaller units
sistent differences in their
and association of the mid-
Je broken down into a tall
1e West. The two grassland
PLANT PHYSIOGNOMY, VEGETATION STRUCTURE, AND FORMATIONS 13 9
types share some species but also have clear differences in species composition
and structure. In an hierarchical classification,we would consider the two prairie
types to be subsets of the midwestern grassland association.
Alternatively, we might instead consider that the communities are joined in
a reticulate classification network in which each identifiable community is con-
sidered a separate entity and not a subset of a larger association. An example
might be the forest , tundra, and peatland patches found at the northern treeline.
These associations occur in close proximity, may share some species, but are gen-
erally distinct from each other in terms of species composition and structure. A
number of mathematical techniques are used to classify stands and identify for-
mations of recurring species associations. In addition, some of these techniques
can help determine whether the relationship between formations should be con-
sidered hierarchical or reticulate.
The impact of disturbance adds to the difficulty in sampling and describing
communities. Most of the community types and associations that are described are
implicitly assumed to represent "climax" states. The species composition, abun-
dance, and dominance of a recently disturbed site may be significantly different
from that of the pre-disturbance conditions. Over time, succession will lead to
marked changes in the species composition and structure of the stand. If we com-
pare communities along environmental gradients and include stands at different
successional stages, the results can obscure evidence of community structure.
?LANT PHYSIOGNOMY, VEGETATION
STRUCTURE, AND FORMATIONS
In addition to species composition, communities are sometimes defined on the
basis of plant physiognomy (growth form and function) and vegetation structure. In
such schemes, the physiognomy of the plants and the structure of the vegetation
are used to describe the community. Units that are defined on the basis of similar
vegetation structure are called formations. Our imaginary drive across North
America took us through a number of the various vegetation formations recog-
nized on the continent (Fig. 6.1). The major North American formations range
from treeless tundra in the far north of Alaska and Canada to tropical forest in
southern Mexico. The formal use of the term formation to characterize vegetation
units can be traced back to the work of the German ecologists A. Grisebach and
J.E.B. Warming in the nineteenth and early twentieth centuries. Of course, people
have been informally characterizing landscape on the basis of vegetation structure
since earliest times. Informal terms such as meadow or woodland are common
ways to denote different vegetation structures. The value of physiognomic and
structural classifications of vegetation is threefold. First, physiognomic and struc-
tural classifications do not require a specialized knowledge of plant species. Thus,
these classifications can be applied quickly and with little technical expertise. Sec-
ond, the physiognomy of the plants and structure of the vegetation can provide
insight into the climatic and soil requirements of the plants, the habitat provided by
the plants, and the primary productivity of the vegetation. Finally, physiognomic
and structural classifications allow us to compare and group vegetation communi-
ties on different continents where we would not find many shared species.
Just as scientific names help to ensure a standardized way to describe
species present in communities, there are standardized methods for describing
140 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

the physiognomy of plants and the structure of vegetation. The Raunkiaer Syster: Megathenr
developed in the early twentieth century by the Danish botanist C. Raunkiaer is Xerophiles:
a widely used method of systematically describing plants on the basis of physiog-
Mesothern
nomy and life history. The Raunkiaer system divides plants into six main cate-
availabil
gories called life forms:
Microthern
Phanerophytes: Perennial land plants that are trees and very large shrubs wi t~ Hekistothe
persistent above-ground stems and buds. These plants do not require pro-
tected microsites in order to grow or shield buds during cold winters o:- In recent worl
other dormant periods. The phanerophytes are broken down into five sub- on the basis o
categories that include megaphaneorphytes, which are over 30 m tall an · climatic varia
nano-phanerophytes which are less than 2 m tall such as vines (called dim requirements.
ing phanerophytes), of"BIOME"1
Chamaephytes: Perennial land plants that grow as small shrubs. They hay· Vegetati,
persistent above-ground stems and buds. These plants do depend on pro- dividing veget
tected microsites to protect buds in winter or during dormant periods. mations with 1
cover and tho
Hemicryptophytes: Perennial land plants that are small shrubs and herbs. The y
subdivided on
have persistent stems and buds located at the soil surface.
might include
Cryptophytes: Perennial land plants that are small shrubs and herbs. They trees and sapli
depend on subsurface organs such as bulbs and corms to survive winter or divided into
dormant periods. Lomolino (19'
Therophytes: Annual land plants that survive winter or other periods, such as raphers:
summer drought, as seeds only.
Hydrophytes: Water plants. Forest: Site:
Woodland:
By using sampling techniques similar to the strategies used for studying plan dominat'
species composition, we can typify stands by the relative dominance of different Shrub land:
Raunkiaer life forms. For example, tropical forest stands typically have over 70%
Grassland: :
phanerophytes and only 1% therophytes. In contrast, herb-dominated tundra in
the high arctic typically has over 50% hemicryptophytes and no phanerophytes. Scrub: Sites
Although the life form classes developed by Raunkiaer have proven a use- Desert: Sit'
ful method for describing plant physiognomy and characterizing different vege- ground.
tation formations, they have two drawbacks. First, they require some knowledge
of the life history of the plant. Second, they do not truly capture the great diver- In practice, sci
sity of plant forms that can be seen in nature. An alternative is to classify stands tion
I
on the ph
and define formations on the basis of the plant's growth forms. The Canadian number of veg
ecologist, P. Dansereau, developed one system of growth form description that is
widely used. Dansereau's method divides plants into six main classes: Trees.
Shrubs, Herbs, Bryophytes (nonvascular plants), Epiphytes (plants that grow on :COLOGICAL EOt
other plants but do not tap the host plant's vascular system), and Lianas (woody FE ZONES, ANC
vines). These main classes are then subdivided on the basis of their leaf shape,
size, texture, and function. The system includes symbols for each physiognomic Biogeographe
type and can portray plant growth form and vegetation structure graphically. The that embrace t
symbols make the Dansereau method very easy to use in the field. and describing
A final way in which plants may be classified and communities described is biosphere intc
in terms of their functional requirements from the environment. The Swiss structure can t
botanist A. P. de Candolle working in the early nineteenth century subdivided and early nine
plants into different groups based on heat and moisture requirements. The main that similar re
plant functional types that he defined were as follows: and vegetatior

ttion. The Raunkiaer System
ish botanist C. Raunkiaer is
mts on the basis of physiog-
s plants into six main cate-
s and very large shrubs with
plants do not require pro-
uds during cold winters or
broken down into five sub-
tich are over 30 m tall and
such as vines (called climb-
as small shrubs. They have
: plants do depend on pro-
Iring dormant periods.
mall shrubs and herbs. They
>il surface.
all shrubs and herbs. They
corms to survive winter or
er or other periods, such a
~gies used for studying plant
tive dominance of differen t
tds typically have over 70 %
, herb-dominated tundra in
~s and no phanerophytes.
unkiaer have proven a use-
aracterizing different vege-
:y require some knowledge
Lily capture the great diver-
:rnative is to classify stands
owth forms. The Canadian
vth form description that is
to six main classes: Trees.
>hytes (plants that grow on
ystem), and Lianas (woody
e basis of their leaf shape.
,oJs for each physiognomic
n structure graphically. The
in the field.
1 communities described is
; environment. The Swiss
teenth century subdivided
re requirements. The main
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 141
Megatherms: Plants that require high temperatures and abundant moisture.
Xerophiles: Plants that require high temperatures and tolerate drought.
Mesotherms: Plants that require moderate temperatures and moisture
availability.
Microtherms: Plant that have relatively low heat and moisture requirements.
Hekistotherms: Plants that tolerate extremely cold conditions.
In recent work, biogeographers have attempted to define plant functional types
on the basis of more refined quantitative relationships between plant form and
climatic variables such as tolerances to low temperatures and soil moisture
requirements. Perhaps one of the most sophisticated of these studies is the series
of "BlOME" models by I. C. Prentice and his colleagues.
Vegetation structure can be described in many ways. Most schemes start by
dividing vegetation into formations with trees and formations without trees. For-
mations with trees are then subdivided into those that have continuous canopy
cover and those with discontinuous canopy cover. Tree formations may also be
subdivided on the basis of the number of strata or layers in the canopy. These
might include a canopy layer of dominant trees, a subcanopy layer of smaller
trees and saplings, a shrub layer, and a ground layer. Sites without trees might be
divided into shrub-dominated and grass-dominated formations. Brown and
Lomolino (1998) list six basic vegetation structural types recognized by biogeog-
raphers:
Forest: Sites dominated by trees and a generally continuous canopy.
Woodland: Sites typified by widely spaced trees allowing for substantial areas
dominated by shrubs, grasses, or herbs.
Shrubland: Sites dominated by a relatively continuous cover of shrubs.
Grassland: Sites dominated by grasses and herbs.
Scrub: Sites dominated by widely spaced shrubs.
Desert: Sites dominated by sparse xerophytic plant cover with mostly bare
ground.
In practice, scientific descriptions of vegetation structure often include informa-
tion on the physiognomy of the dominant plants, degree of canopy closure, and
number of vegetation strata.
OLOGICAL EQUIVALENTS,
E ZONES, AND THE BIOMES
Biogeographers have long been interested in vegetation classification systems
that embrace the entire earth. To this end, much work has been done on defining
and describing the world 's biomes. The modern biome concept of subdividing the
biosphere into intercontinental formations of similar climate and vegetation
structure can be traced back to phytogeographers working in the late eighteenth
and early nineteenth centuries. At the heart of the biome concept is the premise
that similar regional climates produce similar plant physiognomic adaptations
and vegetation structure. The early phytogeographers noticed that similar plant
142 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

in the Andes.
growth forms and vegetation formations could be found in many parts of th:e
encountered, ~
world, even in cases where the similar formations were far distant from eac'-
particularly in
other and did not share any of the same species. For example, the life forllli.
Upon his retUJ
growth forms, and the vegetation structure of the Amazon rainforest is very sim-
altitudinal veg
ilar to tropical forested sites in Africa and Southeast Asia. However, the speci :
dence and dec
of plants, and indeed in many cases the genera and families found in the Amazo
the climate an
Africa, and Southeast Asia are very different. Such widely separated, but phys-
encountered v
iognomically and structurally similar, species and vegetation formations are-
recall from Cb
called ecological equivalents.
for every.1000
The scientific study of ecologically equivalent vegetation and the causes o:
equatorial site
this phenomenon date back over 200 years. The English scientist, J.R. Foreste ~
4000-m-high rr
sailed around the world with Captain Cook in order to study the natural histor:
his travels in t
of the globe. In 1778 he published his observations on how the earth could b:e
Altai Mountar
divided into latitudinal belts of similar vegetation structure and similar climate
five-volume W<
The German geographer Alexander von Humboldt is often described as th-:
lated concepts
father of biogeography and the originator of the modern ecological equivalen -
In North
concept. Interestingly, he is also considered to be a founder of the science of gecr
tribution of de
physics. Humboldt was a young nobleman, 30 years of age, when he sailed fro c::
the San Franci:
Europe in 1799 and spent the next five years on an expedition to the Americas
occur as one g•
(Fig. 6.3). In his travels, he explored the Amazon, Orinoca, and Magdalena rivers
dian arctic. He
of South America. He also climbed to the 5800-m summit of Mount Chimborazc
altitudes were
latitudes. He c•
the mountains
1200 miles nor
tional and latit
estimated that
equivalent to a
By themi
ural vegetation
with similar pl<
different conti1
from meteorol
possible to ana
mations and eli
SHelford coine<
associated with
iognomy and V
the 1936 Koppt
vegetation ecot
of world clima
earth based on
English botani~
the relationshir
mean annual p
6.5). Modified
for use by clim
FIGURE 6.3 The German geographer Alexander von Humboldt (1769-1859) who result from fut
conducted some of the earliest scientific work linking the geographic distribution of been produced
vegetation formations with climatic gradients . He is considered a founding figure in common them
biogeography and geophysics.

'o und in many parts of the
were far distant from each
'o r example, the life form
tazon rainforest is very sim-
Asia. However, the specie _
nilies found in the Amazon.
widely separated, but phys-
vegetation formations ar
egetation and the causes of
ilish scientist, J.R. Forester.
to study the natural history
on how the earth could be
·ucture and similar climate.
t is often described as the
iern ecological equivalence
under of the science of geo-
)f age, when he sailed fro m
~xpedition to the Americas
rroca, and Magdalena rivers
1mit of Mount Chimborazc
Joldt (1769-1859) who
graphic distribution of
~d a founding figure in
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 143
in the Andes. During his explorations he took measurements of the climates he
encountered, studied ocean currents, and noted the plant and wildlife. He was
particularly interested in the relationship between altitude and temperature.
Upon his return to Europe in 1804, von Humboldt combined his observations of
altitudinal vegetation zonation in South America with Forester's latitudinal evi-
dence and deduced that as one progresses from high elevation to low elevation
the climate and vegetation structure change in a manner similar to the changes
encountered when one travels from the polar regions to the equator. We may
recall from Chapter 2 that air cools in the troposphere at a rate of about 6.5° C
for every 1000-m increase in elevation. Thus, if the sea-level air temperature at an
equatorial site is 31 ° C, it is likely to be a chilly 5° C at the top of an adjacent
4000-m-high mountain. Humboldt's scientific explorations did not conclude with
his travels in the Americas. In 1829, he set off on an expedition to the Ural and
Altai Mountans of Russia. His scientific work culminated in the publication of a
five-volume work called the Kosmos. The Kosmos presents Humboldt's accumu-
lated concepts on geography and geophysics.
In North America, during the 19th century, C.H. Merriam compared the dis-
tribution of desert, coniferous forest, and tundra found at different elevations in
the San Francisco Mountains of Arizona, with similar changes in vegetation that
occur as one goes northward from the southwestern United States to the Cana-
dian arctic. He deduced that the climate and vegetation at progressively higher
altitudes were equivalent to the climate and vegetation at progressively higher
latitudes. He correlated the conifer forest vegetation found at mid-elevations in
the mountains with conifer-dominated forests found growing at low elevations
1200 miles north in Canada. He used the term life zones to refer to these eleva-
tiona! and latitudinal bands of similar climate and vegetation (Fig. 6.4). Merriam
estimated that a one-mile increase in elevations in the Southwest was roughly
equivalent to an 800-mile-northward displacement in latitude.
By the mid-twentieth century, much information had been gathered on nat-
ural vegetation formations from around the world. It was clear that formations
with similar plant physiognomy and vegetation structure could be observed on
different continents. In addition, detailed data on climate was being gathered
from meteorological stations all over the world. From this information, it was
possible to analytically compare the relationship between large vegetation for-
mations and climatic conditions on a global scale. In 1936, F.E. Clements and V.E.
SHelford coined the term biome to describe the dominant vegetation formation
associated with specific climatic conditions. Based on the premise that plant phys-
iognomy and vegetation structure are directly controlled by climatic conditions,
the 1936 Koppen climatic subdivision of the earth (discussed in Chapter 2) used
vegetation ecotones to help infer the location of climatic boundaries for his map
of world climatic regions. Other scientists have devised similar divisions of the
earth based on large climatic zones and associated vegetation types. In 1947 the
English botanist L.R. Holdridge published a subdivision of the earth based on
the relationship between vegetation formations and mean annual temperature,
mean annual precipitation, and mean annual potential evapotranspiration (Fig.
6.5). Modified versions of the Holdridge system are particularly popular today
for use by climate modelers interested in the changes in vegetation that may
result from future climate change. Many formation and biome schemes have
been produced for individual continents or for the entire globe. They all have in
common the use of some measure of heat (such as temperature or potential
144 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

dinal regions
Polar

iill Lower austral - - -----------

.Tropical Subpolar

Boreal
-- -- --------8
Cool
lem perate
.,_....
- - - warrii- -16.oo
Ponderosa pine :emperate [
• . Subtropical
-- - - - - -32.oo- - -
Tropical Oeser
62.5 1;
32.00
Semiparched S
FIGURE 6.4 Merriam's life zones for the classification of North American vegetation based
FIGURE 6.5
on similarities in structure and climate (based upon Merriam, 1890, 1894; Bailey, 1996; Brown
basis of clim.
and Lomolino, 1998).

2000 g per
evaporation) and some measure of moisture (such as annual precipitation or soi. has charact
moisture deficit). Each biome is typified by similar climate and vegetation stru - etation con
ture. Although terminology and details on geographic boundaries and number f era! basis f<
formations may differ, the general patterns of vegetation and vegetation-climar · the steamin
relationships are similar from system to system. dry arctic t
In 1975 the American ecologist R.H. Whittaker published a classification o: mate of eac
global vegetation formations and their relationship to mean annual precipitatior. etation and
and mean annual temperature (Fig. 6.6). One shortcoming of Whittaker"_ also consid<
approach is that it does not incorporate any measure of seasonality. The magni-
tude of seasonal differences in temperature and precipitation can be extreme ! ~
Tropical
important in determining dominant plant physiognomy and vegetation structure.
However, this simple scheme remains widely used as a basic introductory model Tropical rai
for the description of the world's biomes. Whittaker's system divides the bio - south latitu
phere into a number of distinctive biomes, ranging from tropical rainforest to arc- Central an<
tic and alpine tundra. The geographic distribution of the biomes reflects general and the ren
global climatic zones and more regional influences on vegetation such as the mated at 17
presence of cool conditions in high-altitude mountain ranges or the presence of average an
rainshadows. Estimates of the total area, net primary productivity, and mean bio- 200-250 err
mass for the biomes are also available from work by Whittaker (Table 6.1). Th exceeds 40C
biomes exhibit large differences in net primary production, which range from perature ( <
 ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 145

, ~
~
qJ
-s::
0
~
-~

,§'
-~(;:o
iit

0.25
~
~
0.125 62.5 ~

125
'0
'Q),..
\
~/
,2i ~
,# 0.50 ' 250 (.Q%.
' \I
~(f; v ~
I \
~~
3oreal ~ '?
Latitudinal regions q_(f ~
Altitudinal belts
rransition
Polar '2, Nival
Jpper austral
.ower austral
·ropical Alpine
2ooo - - - - - - - - - - - - 3°C

fir
1derosa pine :;
<

62.5
32.00 0.125
Semiparched Arid Semiarid Subhumid Humid Perhumid Superhumid
rth American vegetation based
FIGURE 6.5 The Holdridge System for classification of the world's vegetation on the
1890, 1894; Bailey, 1996; Brown
basis of climate or geographic location (after Holdridge, 1947; Holdridge et al., 1971 ).

; annual precipitation or soi:
limate and vegetation struc-
c boundaries and number o:"
tion and vegetation-climate
published a classification o:"
) mean annual precipitatio;:
hortcoming of Whittaker"'
e of seasonality. The magni-
cipitation can be extreme!:
ny and vegetation structure
a basic introductory mod_,
r's system divides the bi o~ ­
•m tropical rainforest to arc-
the biomes reflects generC.:.
on vegetation such as th.:
n ranges or the presence o:"
productivity, and mean bio-
, Whittaker (Table 6.1) . Th.:
•duction, which range fr oc
2000 g per square meter for tropical rainforests to 71 g for deserts. Each biome
has characteristic soil conditions that form due to the particular climatic and veg-
etation conditions associated with the biome. Using Whittaker's system as a gen-
eral basis for our exploration, let's take a tour of the world's biomes, starting at
the steaming tropical rainforests of the equator and moving poleward to the cold,
dry arctic tundra (Fig. 6.6). We will look at the geographic distribution and cli-
mate of each biome. We will investigate the unique features of each biome's veg-
etation and make a few observations on the fauna. During our journey we will
also consider the impact of human land use on the natural biomes.
Tropical Rainforest
Tropical rainforest occurs in the equatorial regions between about 25° north and
south latitude. About 50% of the total area of tropical rainforest is found in
Central and South America, 30% in southeastern Asia and eastern Australia,
and the remainder in central Africa. The total area of the biome has been esti-
mated at 17 to 19 million sq km. Tropical rainforest is found in areas that have an
average annual temperature of greater than 20° C and receive more than
200-250 em of precipitation per year. In some areas average annual rainfall
exceeds 400 em. This tropical biome has little daily or seasonal variation in tem-
perature ( < so C) and never experiences freezing conditions. Precipitation
146 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

Primary I

·a inforest
:eci duous

Natural vegetation regions

of the world - =~ • a fr om Whittaker an•
D Tundra
-10 D Boreal- coniferous forest
D Temperate forest
-5 ll!!!l Grasslands volcanic depc
G' [!ll!l Shrubland, scrublands, woodland weather and
~
0 (Mediterranean)
~
:::>
The ph)
• Desert
'§Q) 5 D Savanna, tropical deciduous forests, etation refiec
0.
E Temperate scrubland, shrubland, woodland tions to perioo
.l!l 10 rain forest • Tropical rain forest to seasonal d
growth can cc
environment::
to heat and w
rainforests tyj
that is receive
of rainforest
0 50 100 150 200 250 300 350 400 450 Although wat
Mean annual precipitation (em) orative stress
through evap<
FIGURE 6.6 The relationship between the biomes and climate according to Whittaker's 4o C cooler aJ
system and a general map of the world's biomes (after Whittaker, 1975 and a variety, high humiditi
of sources). and decrease
short supply.
the tropical ra
As a res1
occurs throughout the year. Although dry seasons may occur, the rainfall during petition for li~
the driest month in the tropical rainforest biome averages greater than 12 em. an understory
On an annual basis, moisture stress is not a problem as potential evapotranspi- 70% of the p
ration does not exceed actual evapotranspiration. almost exclus
The warm, moist conditions of the tropical climate lead to deep weather- slides. The str
ing and rapid leaching of soluble nutrients from soils. The result is a dominance biomes. In rna
by oxisols. These soils often have a deep red color due to the accumulation of sists of scatter
iron oxides. Oxisols are typically very low in plant nutrients. In tropical areas of canopy cover.
Central America and Southeast Asia, the dominant soils are developed on fresh heights of ove
 ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 14 7

-ABLE 6.1 Primary Productivity of the Major Biomes

Area Net Primary Production Total Biomass

~ - ~ me (million sq kml (grams per meter per year) (kilograms per meter)

- =oical rainforest 17 2000 44

- --: :Jical deciduous
=-: ·est 7.5 1500 36
~.::. an n a 15 700 4
: -:se rt 18 71 0.67
=-~ it er ranean and other 8 600 6.8
=c1rublands
- :-- oe rate grassland 9 500 1.6
- =--:J erate deciduous
::.:·3St 7 1200 30
' - =-De ra te rainforest 5 1300 36
::.: =a I fo rest 12 800 20
m regions <;;l
) _-c ~a
8 144 0.67

_-:e: Data from Whittaker and Likens, 1975.

rous forest
3St
volcanic deposits and have greater nutrient content. Over time, these soils will
rublands, woodland weather and become similar to the more mature oxisols.
n) The physiognomy of tropical plants and the structure of tropical forest veg-
ical deciduous forests, etation reflect several key factors. Temperatures are relatively warm, so adapta-
·ubland , woodland tions to periods of freezing are not required. Soil water is plentiful, so adaptations
Jrest to seasonal dry soil conditions are not needed. Thus, photosynthesis and plant
growth can continue throughout the entire year. However, plants do face several
environmental challenges that are reflected in the tropical vegetation. In addition
to heat and water, the photosynthetic process requires light. The floor of tropical
rainforests typically receives only 1% of the photosynthetically active radiation
that is received at the top of the canopy. Much of the physiognomy and structure
of rainforest vegetation can be understood in terms of competition for light.
Although water is plentiful in the rooting zone, the high temperatures and evap-
orative stress on top of the canopy cause high rates of water loss from foliage
through evapotranspiration. In contrast to the canopy, the forest floor may be 2 to
ate according to Wh ittaker's 4° C cooler and maintain a relative humidity of close to 100% at all times. Such
<er, 1975 and a variety, high humidities can hinder the evaporation of excess water from leaf surfaces
and decrease rates of photosynthesis. In oxisolic regions, soil nutrients are in
short supply. Finally, the diversity of phytophagous insects is extremely high in
the tropical rainforest, and plants must be adapted to herbivory by these insects.
As a result of the year-round growth, relatively plentiful moisture, and com-
y occur, the rainfall during petition for light, the tropical rainforest is dominated by tall evergreen trees with
rages greater than 12 em. an understory of perennial plants. It has been estimated that trees can make up
as potential evapotranspi· 70% of the plant species found in a rainforest stand. Annual plant species are
almost exclusively restricted to forest gaps caused by windthrow, fires, or land-
ate lead to deep weather· slides. The stratification of the rainforest vegetation is the most complex of all
The result is a dominance biomes. In many cases there are five distinct strata (Fig. 6.7). The A stratum con-
ue to the accumulation o sists of scattered supercanopy trees that have trunks rising above the continuous
rients. In tropical areas o: canopy cover. The A stratum trees typically grow to 30 to 50 m but can reach
1ils are developed on fres l:. heights of over 70 m. An 84-m-tall Koompassia excelsa tree growing on Sarawak
148 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

in the soil and

~ trunks and bra
1ii
>.
c.
0
not have to in
Meters c
<1l order to reach
35 ()
of 165m. This i
30 A Epiphyte~

25
ally never have
nutrients from
20 B cular tissue of t
15 c annual rainfall
phosphorus, 18
by relatively sn
5 D and bromeliad:
E phytes. Epiphy
host's foli age f<
Meters 30 20 10 can benefit tre<
FIGURE 6. 7 Tropical rainforest structure and stratification (after Richards, 1996).

Island in Indonesia is reputed to be the tallest tree in the tropics. This is rough2_
the height of a 15- to 20-story building! The B stratum is the continuous canol-_
of dominant trees that typically reach heights of 18 to 27 m. The C stratum ~
made up of subdominant trees and immature trees that are 8 to 14m tall. The =
stratum consists mainly of smaller trees, such as palms and tree ferns, and immc.-
ture canopy trees. On the forest floor, the E stratum contains small herbaceo ~
plants and low shrubs but is often dominated by tree seedlings. There is a con:-
mon perception that tropical rainforests are a tangled mass of dense vegetatio::..
This is only true around openings and gaps where light is plentiful. The low Jig1-·
levels beneath the canopy of an undisturbed rainforest do not support much ve~­
etation, and the understory is often quite open.
The tropical rainforest contains the greatest amount of standing biomass o:
any biome. The complexity of the tropical forest vegetation and the many diffe:-
ent microenvironments in the forest is reflected in the high species diversity typi-
cal of this biome. Although the exact number is unknown, there are well oYe:
100,000 different plant species in the tropical rainforest biome. That means thc.:
40% of all plant species on earth are found in this one biome. Over 170 differe :
tree species have been recorded on single hectares of rainforest in both Indone-
sia and South America. African rainforests have lower diversities, but even there
some 60 to 130 different tree species can be found on a hectare of rainforest lan ·
In parts of the Amazon Basin over 300 species of trees have been found on 1 ha.
Tropical plants have developed some interesting physiognomic adaptatioiE
in response to the low light conditions and different microclimates found withir.
the rainforest. The high growth rates and continuous growing season mean tha:
tree species must reach and dominate the canopy quickly if they are to surviv
Canopy and supercanopy trees commonly have very slender trunks supporte ·
by spreading lower trunks called plank buttresses. Plants in the intermediate
canopy sometimes are supported by stilt roots. The buttresses and stilts provide FIGURE 6.8 A I<
stability to the tall slender trunks. A number of plant species reach canopy anc The fig begins as a
subcanopy heights by using the physical support of other plants. Woody stemmed tendrils that establ
lianas and soft vines are diagnostic plants of the tropical rainforest. They sprout epiphytic plants ha
 ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 149

.l!l in the soil and use tendrils, adhesive surfaces, thorns, and twining to climb the
~
Cii trunks and branches of trees. The advantage for vines and Iianas is that they do
>- not have to invest energy and nutrients into the formation of large trunks in
c.
0
:ers c:
C1l order to reach light. The climbing palms of the genus Calamus can reach lengths
35j (.) of 165m. This is the plant from which rattan furniture and baskets are made.
30 A Epiphytes sprout and grow on the trunks and branches of trees. They usu-
ally never have contact with the soil zone. Epiphytes receive all their water and
25
nutrients from rainfall. Unlike parasitic plants, epiphytes do not tap into the vas-
cular tissue of their hosts. In the tropical rainforest, precipitation is plentiful, and

'T
15 c annual rainfall may contribute as much as 21 kg per hectare of nitrogen, 16 kg of
phosphorus, 18 kg of potassium, and 16 kg of calcium. Epiphytes are dominated
10
by relatively small perennials such as orchids, bromeliads, and ferns. Many ferns
5 D and bromeliads grown as houseplants throughout the world are tropical epi-
E phytes. Epiphytes put mechanical strain on their hosts and compete with the
host's foliage for light. However, nutrients in the humus produced by epiphytes
can benefit tree hosts by adding to the soil nutrient balance. The strangler fig
(after Richards, 1996).

1 the tropics. This is roughly

m is the continuous canop'
to 27 m. The C stratum ~
1at are 8 to 14m tall. The D
ts and tree ferns, and imma-
L contains small herbaceous

e seedlings. There is a com-

d mass of dense vegetatior.
ht is plentiful. The low ligb:
st do not support much veg-

aunt of standing biomass o:

:tation and the many diffe r-
~ high species diversity typi-
known, there are well ove:-
·est biome. That means the:
e biome. Over 170 differer.:
f rainforest in both Indone-
r diversities, but even there
a hectare of rainforest Ian ·
:s have been found on 1 b
5 physiognomic adaptati o ~
microclimates found witb.i::.
; growing season mean the·
tickly if they are to survi' e
y slender trunks supporte.:
Plants in the intermedi at ~
mttresses and stilts pro vi c~ FIGURE 6.8 A large strangler fig (Ficus virens) in the rainforest of Queensland, Australia.
tt species reach canopy ar..: The fig begins as an epiphyte living in the upper branches of a host tree. It sends down long
her plants. Woody stemme.: tendri ls that establish roots. Eventually, the fig kills the host tree . A number of other
>ical rainforest. They sproe: epiphytic plants have established on the upper branches of the strangler fig.
150 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

trees of the genus Ficus combine aspects of epiphyte and liana growth with c. Although 1
deadly result for the host tree (Fig. 6.8). The seeds of the strangler figs ar~ are extremely hi
deposited on the branches of host trees in bird droppings. The young fig begin~ for support of th
life as an epiphyte. However, it also develops woody roots that grow downwar- root systems th~
along the trunk of the host tree. These roots eventually reach the soil. Constri - and released fro1
tion by the fig roots and shading by the expansive fig canopy lead to the death o: nal fungi associa
the host tree. In most cases, the fig tree remains standing, supported by its wood:> sive absorbent s1
root structure. Plants such as the strangler fig, which combine Iiana and epiphy1.: A unique
characteristics, are classified as hemiepiphytes. The bark of most tropical trees ~ These forests an
very smooth, and this may be an adaptation that decreases the ability of epiphytE coastlines. The rr
seeds to adhere to and germinate on them. or Avicennia. Li
With its trees, lianas, vines, epiphytes, and hemiepiphytes, the uppe:- and sclerophyllc
branches and foliage of the continuous B stratum canopy provide a huge habit<:: less in height. M
that can stretch for thousands of kilometers. This surface is broken only by tree- allow them to ob
fall gaps, disturbed areas, and the courses of larger streams and rivers. Many o:: ing facet of man
the plants and animals of the rainforest exist on this leafy surface and may new :- water in mangro
have contact with the ground. The hot sunny world of the canopy is very differe : ities, mangroves
from the cooler and wetter depths of the forest floor. Rainforest plants displa: ter out a certai
many adaptations that result from the different microclimatic conditions found i= undersides of tht
the upper canopy and in the lower strata. Leaves in the upper strata are ofte::. also stored in lea
sclerophyllous in form , being small and having stiff, thick cuticles with waxy coa:- of the genus Rhi;
ings and stomata that are sunk in pits. These features protect the leaves agains. are called vivipa
short-wave radiation and help decrease water loss. Upper stratum leaves ar= tree, they receiv(
often held at high angles relative to incoming sunlight, which decreases radiatim:. from the parent •
and heating of the leaf surface. Some tree species have different leaf types o::. to grow.
their upper and lower branches. Newly formed leaves in the upper stratum rn a~ Tropical ra
be red to pink in color. The color is provided by anthocyanins which appear rc some mammals, 1
protect the leaf against short-wave radiation. Leaves on lower strata plants ofte::. 30 to 50 million c
have sharp elongated tips called drip tips. The presence of drip tips significant!: mates ranging fr
increases water drainage from the surface of the leaf, and this allows for bett :- bus murinus). r
photosynthesis. The drainage of water from the leaf surface also inhibits growr" Rainforest planti
of fungi, algae, and lichens. such as Calumu~
Low windspeeds within the canopy makes wind dispersal of pollen an larger herbivores
seeds inefficient in the rainforest. Instead, more than 90% of the plant species ir: example, manioc
some tropical forests depend on insects and animals for pollination and seed dis- cyanide when ing
persal. The shape, color, and smell of many tropical flowers attract specific polli- At the pres
nators. Beetles typically pollinate bow-shaped flowers, which are greenish o:- and destroyed at
off-white and emit strong odors. Hummingbirds are associated with tubular flO\\-- aged 170,000 sq k
ers that are brightly colored but scentless. Some species of tropical bats feed onl_ forest of the wofl,
on nectar and pollen and are associated with flowers that bloom at night. Th - may average 80,1
Indonesian plant, corpse flower (Rafflesia arnoldii), produces a meter-tall flowe:- some of this clea
that is colored brown-red and emits an odor like decaying meat in order tc lowing clearance.
attract flies as pollinators. Studies in the African rainforest indicate that througt rainforest cover r'
selective eating of soft fruit tissue, droppings, and regurgitation, monkeys dis- parts of the biom
persed about 96% of seeds they encountered while feeding and birds disperse · along its eastern ,
88%. As discussed in Chapter 4, fruit -eating fish are important in dispersing seeds Some estimates c
in seasonally flooded portions of the tropical rainforest. The low seasonal vari- year 2020 unless
ability in climate and the dependence of rainforest plants on different animal an · countries with tht
insect pollinators and seed dispersers result in flowers of different species bloom- lations and limit
ing at different times throughout the year. exploitation of th

te and Iiana growth with a
Is of the strangler figs are
·pings. The young fig begins
roots that grow downward
tlly reach the soil. Constric-
canopy lead to the death of
ing, supported by its woody
::ombine liana and epiphyte
ark of most tropical trees is
eases the ability of epiphyte
hemiepiphytes, the upper
10py provide a huge habitat
face is broken only by tree-
.treams and rivers. Many of
eafy surface and may never
the canopy is very different
r. Rainforest plants displ ay
climatic conditions found in
. the upper strata are often
tick cuticles with waxy coat-
s protect the leaves against
Upper stratum leaves are
t, which decreases radiation
tave different leaf types on
:s in the upper stratum rn a ~·
:hocyanins which appear t
on lower strata plants ofteu
1ce of drip tips significantly
f, and this allows for better
surface also inhibits growth
nd dispersal of pollen anc
90% of the plant species in
:or pollination and seed dis-
lowers attract specific polli-
ers, which are greenish or
ssociated with tubular f!O \\·-
es of tropical bats feed onl\
rs that bloom at night. The
)roduces a meter-tall flowe ~
decaying meat in order tc
forest indicate that througt
·egurgitation, monkeys dis-
feeding and birds dispersec
nportant in dispersing seeds
rest. The low seasonal vari-
mts on different animal anc
>of different species bloom-
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 151
Although soils may be poor in nutrients, decay rates of soft tissue and wood
are extremely high in the rainforest. This allows for quick cycling of nutrients and
for support of the forest vegetation. Most rainforest plants have extensive shallow
root systems that are more efficient at taking up nutrients brought in by rainfall
and released from decaying litter. In addition, many plants have internal and exter-
nal fungi associated with their root systems. The hypha of the fungi produce exten-
sive absorbent surfaces that increase rates of nutrient uptake by the host plants.
A unique type of forest found in the tropical regions are the mangroves.
These forests are found growing in calm ocean waters and estuaries along tropical
coastlines. The mangroves are often dominated by tress of the genera Rhizophora
or Avicennia. Like many tropical forest canopy trees, mangroves have stilt roots
and sclerophyllous leaves. However, mangrove trees are generally only 10 m or
less in height. Many mangrove trees have pneumatophores (see Chapter 3) that
allow them to obtain sufficient oxygen while growing in flooded soils. The interest-
ing facet of mangroves is their ability to grow in saline water. The salinity of soil
water in mangroves stands equals that of sea water. To cope with these high salin-
ities, mangroves have a number of interesting adaptations. Mangrove roots can fil-
ter out a certain amount of salt. Avicennia trees have special glands on the
undersides of their leaves that exude salt to remove it from the tree. Excess salt is
also stored in leaf tissue and eliminated when old leaves drop from the trees. Trees
of the genus Rhizophora produce seed that germinate while still on the tree. These
are called viviparous seedlings. When the seedlings are attached to the mother
tree, they receive nonsaline water from the parent. Eventually, the seedlings fall
from the parent and root. At this point they appear capable of using saline water
to grow.
Tropical rainforests contain the highest diversities of insects, birds, and
some mammals, such as primates and bats, in the world. There may be as many as
30 to 50 million different species of rainforest insects and even 180 species of pri-
mates ranging from gorillas (Gorilla gorilla) to the tiny mouse lemur (Microce-
bus murinus). Many of the insects and other animals are phytophagous.
Rainforest plants have a number of strategies that combat herbivory. Some vines
such as Calumus have thorny stems that assist in climbing and protect against
larger herbivores. Many plants contain chemicals that are toxic to herbivores. For
example, manioc (Manihot ultissima) contains cyanogenic glycosides that release
cyanide when ingested.
At the present time, the tropical rainforests of the world are being cleared
and destroyed at a rapid pace. By the 1990s, the rate of rainforest clearance aver-
aged 170,000 sq km per year. That represents a loss of almost 1% of the total rain-
forest of the world each year. In Brazil alone, the annual rate of Amazon clearance
may average 80,000 sq km. However, recent remote sensing data suggest that
some of this cleared rainforest is being allowed to regenerate by succession fol-
lowing clearance. In areas such as Central America, less than half of the original
rainforest cover remains (Fig. 6.9). Tropical rainforest clearance has occurred in all
parts of the biome. Even Australia, which contains only a tiny area of rainforest
along its eastern edge, has transformed a significant area to pasture and farming.
Some estimates conclude that almost all natural rainforest could be lost by the
year 2020 unless conservation measures are taken now. However, many of the
countries with the highest rates of rainforest loss also have rapidly growing popu-
lations and limited economic resources. For many people in such countries,
exploitation of the rainforest is seen as one of the only means of survival.
152 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

Tropical Sea
1940 (67%) 1950 (56%) 1961 (45%)
Tropical seasonc
the south and n<
the tropics but c;
biome is around
The mean;
20 to 30° C with
sonal variability
forest from the
1977 (32%) 1983 (17%) rainfall pattern,
months. The win1
variations in ten
been reported ir
example. Potenti
The soils i
leached oxisols.
extreme as in tht
the rainforest.
As a result
FIGURE 6.9 The loss of primary rainforest in Costa Rica from 1940 to 1983 (after Sader
and Joyce, 1988, Whitmore, 1990; Richards, 1996).
est differs in rna
most, most of th'
winter dry seaso
Until this century, human impact on many tropical rainforests was relative!~ maintain their le:
limited. Small bands of hunter-gatherers lived in some areas, while a shifting pa - all go dormant ir
tern of slash and burn agriculture was practiced in other regions. Such agricul- and less leathery
ture, also known as swidden farming, consists of clearing small plots of forest an must develop qui
growing crops on them for one to five years. After this time, the soil fertility i5 ture stress becon
generally reduced, and the plot is abandoned to forest succession. This type o:' bloom just befm
farming is sustainable in the rainforest but can only support small populations o:' blooms before c<
people. It has been estimated that the population density of the Yanomami peo- during the wet s
ple of the Amazon, who practice hunting, gathering, and shifting agriculture, i5 dry season.
only one person per every 777 ha. Increasing population pressure and access tc The season
the rainforest in areas such as the Amazon Basin have led to attempts to establisl:. est, with only tb
permanent farm plots on cleared rainforest land. In many cases, the soil fertility i5 cover. The heigh1
quickly reduced, and the farmland is colonized by tropical grasses such as Imper- erally averaging
ata cylindrica and other light-demanding herbs and ferns. Introduction of live - forest, but they a
stock to such areas maintains poor-quality grasslands at the expense of rainfores: are rarer and les
succession. Rainforest around the world has also been cleared or altered by larg · nutrients to persi
plantations growing commercial crops such as coffee ( Coffea spp.), bananas (MusG cal rainforest bee
spp.), and rubber trees (Hevea brasiliensis). The standir
Logging has also had a dramatic impact on rainforests. Even selective log- second only to t
ging can cause widespread damage due to road construction, log felling, and l o~ tropical seasonal
transport. Removal of teak (Tectona grandis) and other valuable hardwoods. 11,000-ha tropica
which make up about 10% of Southeast Asian rainforests, has been shown to lea · species of insects.
to damage to about 65% of the forest. In countries such as Indonesia and Malaysia reptiles and amp
hardwood logging almost quadrupled between 1965 and the 1990s. Once loggin ~ less than 50% of
roads are established, they are often used for access by farmers, and this leads to ber of tree specie
permanently cleared land. Other human disturbances include road building, min- the rainforest. A
ing, and dam construction. 11,000-ha Costa I

1961 (45%)
om 1940 to 1983 (after Sader
:al rainforests was relative]\
e areas, while a shifting par-
)ther regions. Such agricul-
ing small plots of forest anc
his time, the soil fertility ~
est succession. This type o:
upport small populations o:
1sity of the Yanomami peo-
, and shifting agriculture, ~
tion pressure and access tc
: led to attempts to establi s ~
.any cases, the soil fertility ~
pical grasses such as Imp er-
ferns. Introduction of live-
at the expense of rainfore5:
n cleared or altered by large
Coffea spp.), bananas (Mus.:.
nforests. Even selective loE-
truction, log felling, and lo;
other valuable hardwoods
~ ests, has been shown to lea c.
h as Indonesia and Malaysic:..
and the 1990s. Once loggin;
by farmers, and this leads tc
s include road building, mir.-
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 153
Tropical Seasonal Forest
Tropical seasonal forest , also called tropical dry forest, occurs as broad bands to
the south and north of the tropical rainforest biome. It is generally restricted to
the tropics but can be found as far as 30° N latitude in India. The total area of the
biome is around 7.5 million sq km.
The mean annual temperature of tropical seasonal forest region ranges from
20 to 30° C with rainfall averaging 150 to 250 em per year. However, it is the sea-
sonal variability in rainfall that most strongly differentiates the tropical seasonal
forest from the rainforest. The tropical seasonal forests experience a monsoon
rainfall pattern, with the vast majority of precipitation falling in the summer
months. The winter is typified by dry conditions for four to seven months. Seasonal
variations in temperature are not large. Frost generally does not occur but has
been reported in portions of the tropical seasonal forest in southern Africa, for
example. Potential evaporation exceeds precipitation in the dry winter months.
The soils in the seasonal tropical forest tend to be deeply weathered,
leached oxisols. However, owing to the dry season, leaching is often not as
extreme as in the tropical rainforest, and nutrient retention can be higher than in
the rainforest.
As a result of the seasonal differences in precipitation, the dry tropical for-
est differs in many important ways from the adjacent rainforest. First and fore-
most, most of the canopy trees are deciduous. They become dormant during the
winter dry season and lose their leaves (Fig. 6.10). Some understory shrubs may
maintain their leaves during the winter, but in many cases shrubs, vines, and herbs
all go dormant in the dry season. The leaves of the canopy trees tend to be larger
and less leathery than those found in the rainforest canopy. The deciduous leaves
must develop quickly at the start of the wet season, and then they die when mois-
ture stress becomes acute at the start of the dry season. The canopy trees often
bloom just before the onset of the wet season. This allows pollinators to find
blooms before canopy leaves obscure them. Fruit and seeds can then be formed
during the wet summer months and are often dispersed during the subsequent
dry season.
The seasonal tropical forest has a much simpler structure than the rainfor-
est, with only three dominant strata-canopy, shrub and sapling, and ground
cover. The height of the canopy tends to be less than the tropical rainforest, gen-
erally averaging 20 to 30 m. Vines and lianas are found in the tropical seasonal
forest, but they are not as common or as diverse as in the rainforest. Epiphytes
are rarer and less diverse because they often cannot obtain sufficient water or
nutrients to persist. The ground layer vegetation may be denser than in the tropi-
cal rainforest because of greater light penetration.
The standing biomass and species diversity in tropical seasonal forests are
second only to tropical rainforests. In terms of insect and animal fauna , some
tropical seasonal forests have diversities close to those of adjacent rainforests. An
11,000-ha tropical seasonal forest in Costa Rica is estimated to host some 130,000
species of insects, 175 species of birds, 115 species of mammals, and 75 species of
reptiles and amphibians. The floral diversity of tropical seasonal forests is often
less than 50% of the diversity found in nearby rainforests. In particular, the num-
ber of tree species and epiphytes found in the dry forest is much less than that of
the rainforest. A total of 700 angiosperm species is estimated to inhabit the
11,000-ha Costa Rican dry tropical forest.
154 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

C. Most tropic
season can pe1
Reddish
biome are do1
soils are highc
very dry areas
Because of tt
sesquioxides a
horizons or la
mulate to forn
by water and
flow. The orgm
to the low org;
by termites. Th
to be 17.5 mil
organism biorr
tion of dead gr
FIGURE 6.10 Dry tropical deciduous trees in northern Queensland, Australia. These regions.
trees cope with the rainless winter season by releasing their leaves and going dormant. Tropical
When rains return in the summer the canopy will become leafy and green.
shrubs. The rel
grassland can
Venezuela, the
Tropical seasonal forests are even more threatened by human activity thar: of southern Al
tropical rainforests. The biome is generally found in areas of the world wher the savanna ha
increasing human population size and persistent poverty make conservation dif- forest are faun
ficult. The forests are easy to traverse in the dry season and easy to clear usin ~ surrounded by
fires. Some forest is lost to intentional fires, while large areas are also destroyec and the woody
by accidental burning. Once the trees and shrubs are cleared, the long dry seasor: The trees
makes it easy to use fire to preclude forest reestablishment. In many cases, the with sclerophy
soils are better for crops or grazing land than those found in the rainforest. It has mon in Africa.
been estimated that only 2% of the original tropical seasonal forest in Centra: world. Decidu<
America remains intact. Only a paltry 480 sq km of the original 550,000 sq km o: moisture stress
seasonal tropical forest in Central America has official conservation statu branches, causi
Despite its high productivity, biomass, and species diversity, the seasonal tropic<L this water loss.
forest has not received the same public attention and conservation efforts as th loss is found ir
rainforest. forests and sav:
(Adensonia di&
Tropical Savanna America have •
can reach a cin
Tropical savannas and associated thorny woodlands occur in tropical and sub- The ever!
tropical regions, centered on about 20° N and S latitude. The tropical savanna trate through l;
borders tropical forest areas, and often there is a diffuse ecotone between americana tree
savanna and tropical seasonal forest. Over half of all tropical savanna is found in it develops tar
Africa, where this biome comprises 65% of the total land cover. Tropical savan· Attalea exigua i
nas and thorny woodlands have many different regional names; in Africa th the soil surface
savanna may be called bushveld, in Australia it can be called brigalow, in Sout dies back. The 1
America it is known as campos or llanos. The total area of the biome is about 15 trees and fares
million sq km. The tropical savanna is typified by average annual temperatures of Vines are not
between 20 and 30° C and annual precipitation of between about 50 and 150 em. savanna tend tc
Seasonal differences between winter and summer temperatures average 10 to 15c die back during

Jensland, Australia. These
eaves and going dormant.
fy and green.
ned by human activity thaL
1 areas of the world where
~rt y make conservation dif-
;on and easy to clear usin_§:
ge areas are also destroye' ·
:!eared , the long dry seasoc
shment. In many cases, the
,und in the rainforest. It has
! seasonal forest in Centra:
te original 550,000 sq km o:
fficial conservation statu~­
·ersity, the seasonal tropica:
conservation efforts as t b ~
occur in tropical and sub-
tude. The tropical savannc.
1 diffuse ecotone betwee;:
tropical savanna is found i;:
land cover. Tropical savan-
.ional names; in Africa the
Je called brigalow, in SoutZ.
ea of the biome is about 15
age annual temperatures o:
ween about 50 and 150 CIL
tperatures average 10 to 15 =
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 155
C. Most tropical savanna sites experience prolonged winter dry seasons. The dry
season can persist as long as seven months. Frosts are uncommon.
Reddish oxisols are found on moist savanna sites. Drier portions of the
biome are dominated by alfisols with brownish to red surface horizons. These
soils are higher in nutrients than oxisols. Shallow and sandy entisols develop in
very dry areas. The entisols are low in nutrients and may also be relatively saline.
Because of the contrasting wet and dry conditions, bases, clay minerals, and
sesquioxides accumulate in the B horizon of savanna soils. In some cases, saline
horizons or layers of calcium carbonate develop. Iron and aluminum can accu-
mulate to form hardpan layers known as laterite. These layers are impenetrable
by water and block drainage, upward movement of ground water, and nutrient
flow. The organic content of all savanna soils is very low ( <3% ). One contributor
to the low organic content of the soils is the rapid decomposition of plant matter
by termites. The termite population of a savanna site in Zaire has been estimated
to be 17.5 million individuals per hectare. They can make up 80% of the soil
organism biomass. In Africa, termites are responsible for 90% of the decomposi-
tion of dead grass. Termites and their mounds are common in all tropical savanna
regions.
Tropical savanna consists of grasslands with scattered trees and woody
shrubs. The relative abundance and the spatial distribution of woody plants and
grassland can differ markedly. In some areas, such as the Curatella llanos of
Venezuela, the Eucalyptus savanna of eastern Australia, or the Mopane savanna
of southern Africa, scattered individual trees dot the grasslands. In other cases,
the savanna has a parkland configuration in which areas of open or closed canopy
forest are found on well-drained uplands and along rivers. These forest stands are
surrounded by extensive areas of treeless grassland. In some cases, trees are rare,
and the woody vegetation is dominated by often thorny shrubs.
The trees and woody shrubs of the savanna can be either evergreen, often
with sclerophyllous leaves, or deciduous. Deciduous tree species are most com-
mon in Africa, and evergreen species are more common in other parts of the
world. Deciduous leaves and dormancy do not allow trees to completely escape
moisture stress during the dry winter months. Water can be lost through bark and
branches, causing lethal desiccation. The thick bark of some savanna trees reduces
this water loss. Perhaps the most interesting adaptation to dry season moisture
loss is found in species of the Bombaceae family that grow in seasonal tropical
forests and savanna of Africa and South America. These trees such as the baobab
(Adensonia digitata) of Africa and the silk cotton tree (Ceiba pentadra) of South
America have enlarged trunks where they store water in spongy tissue. Baobabs
can reach a circumference of more than 20 m and store up to 120,000 1 of water.
The evergreen trees of the savanna have deep tap root systems that pene-
trate through laterite layers and provide access to the water table. The Curatella
americana tree of the South American savanna reaches a height of only 6 m, but
it develops tap roots as long as 20 m. Some shrubs, such as the palm species
Attalea exigua in Brazil, have large woody organs called xylopdia that lie beneath
the soil surface. During the dry season, the above-ground portion of the plant
dies back. The plants resprout from xyplodia when the rains return. The height of
trees and forest canopies is generally less than 20 m and often less than 10 m.
Vines are not common, and epiphytes are extremely rare. The grasses of the
savanna tend to be perennial species. The above-ground portions of the grasses
die back during the dry season, and the plants resprout from underground organs,
156 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

such as rhizomes and stolons, in the wet season. Most grasses are of the C4 variety
(see Chapter 2 for a description of C4 photosynthesis) and can conduct photo-
synthesis at high temperatures and illumination. For example, many temperate
C3 grass species have maximum rates of photosynthesis at around 24° C. In con-
trast, tropical C4 grass species such as Paspalum dilatum reach peak net photo-
synthesis at 35° C. In general, the percentage of woody species and the height o:
trees and shrubs decrease as one moves from moister to drier portions of th
savanna. In the very driest areas of the savanna biome, annual grasses becom
increasingly important.
What controls the spatial distribution of woody plants and grasslands ir.
tropical savannas has long been a topic of great interest. Why do neither the
woody plants nor the grasses come to dominate the entire landscape? It is
thought that this may be due to a fine balance between the two types of plants.
The grasses have shallow and extensive root systems (Fig. 6.11). They depend or.
efficient interception of rainwater close to the surface. Tree seedlings often can-
not compete with the grass roots for moisture. In addition, the grasses gro\Y
quickly during the rainy season and can attain heights of greater than 2 m. Ir.
these circumstances, the slower growing seedlings of tree and shrub species are Dryfo1
shaded out. Grazing and browsing by animals also likely favors grasses. Th Savanna
perennial grasses can endure relatively close grazing and still resprout and gro\Y
quickly. Recovery of trees and shrubs after intense browsing is slower. Close
grazing will often kill tree and shrub seedlings. Elephants uproot and destroy
small trees when browsing and often destroy several trees in the course of a day.
Thorns and tough leaves are common in woody plants of the savanna. These offer
some protection against browsing. Finally, lightning strikes are common, and the
dry savanna grasses burn easily. Fires can kill tree and shrub seedlings. Deci<

When trees are established, they produce deep intensive roots to obtair.
ground water. Trees tend to limit grassland cover by shading out the grass under-
story, which decreases competition for water and the amount of ground fuel fo
fires. In addition, many trees and shrubs have adaptations to fire, such as thick
bark or the ability to resprout from underground organs. In some cases the loca-
tion of trees and parkland groves appears dependent on small differences in
topography and soils (Fig. 6.11) . Tree roots may not be able to penetrate to the
water table in areas with laterite, and this restricts the survival of woody specie
Low-lying sites, which are underlain by laterites, often flood during the rainy sea-
son, precluding the establishment and growth of trees. Termite mounds some- Grassland
times form small raised areas that are occupied by woody vegetation. In addition
to providing better drainage, the mounds sometimes have higher concentration>:
of potassium, calcium, and magnesium than adjacent soils. The increased nutrient
availability helps to support the woody plants. Areas with coarse, rapidly drained Wooded grov
soils may not hold enough water at the surface to support shallow rooted grasse_
but can support deeply rooted trees and shrubs.
The standing biomass of the tropical savanna biome is only a fraction of
that found in the tropical rainforest. The species diversity for plants, particularly
woody plants and vines, is also much lower. The tree flora over large areas is dom-
inated by a handful of species. Savannas do support a considerable diversity of
birds. Some 708 bird species are known from the African savanna, 521 from Cen-
tral and South America, and 227 from Australia. Large flightless birds including FIGURE 6.11
the ostrich (Struthio camelus) from Africa, the emu (Dromaius novaehollandia) hardpans helps cc
from Australia, and the rhea (Rhea americana) from South America are all foun d savanna. The gra~
intensive systems
 ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 157

~rasses are of the c4 variety

s) and can conduct photo-
example, many temperate
sis at around 24° C. In con-
tum reach peak net photo-
ly species and the height of
er to drier portions of the
ne, annual grasses become

ly plants and grasslands in

terest. Why do neither the
he entire landscape? It is
~en the two types of plants..
(Fig. 6.11). They depend on
e. Tree seedlings often can-
addition, the grasses gro\,.
b.ts of greater than 2 m. In
tree and shrub species are Dry forest
likely favors grasses. The Savanna Savanna Savanna
and still resprout and grow
browsing is slower. Close
phants uproot and destroy
trees in the course of a daY.
of the savanna. These offer
trikes are common, and the
Deciduous fo rest
l shrub seedlings.
J intensive roots to obtain
hading out the grass under-
amount of ground fuel for
ations to fire, such as thick
ans. In some cases the loca-
~ nt on small differences ir.
be able to penetrate to th
~ survival of woody species..
Savanna Wooded grove
L flood during the rainy sea-

~ es. Termite mounds some- Grassland Grassland

'ody vegetation. In additio
have higher concentrations
;oils. The increased nutrie n~
.vith coarse, rapidly drainec Wooded grove
port shallow rooted grasses

biome is only a fraction o:

rsity for plants, particular!\
ora over large areas is dom-
a considerable diversity o:'
::an savanna, 521 from Cen-
0 Hard pan soil layers

ge flightless birds including FIGURE 6.1 1 Small difference in topography and differences in drainage caused by
'Dromaius novaehollandia hardpans helps control the distribution of trees, shrubs, and grasses on the tropical
;outh America are all founc savanna. The grasses have shallow, extensive root systems, while trees have deep,
intensive systems (after Walter, 1985).
158 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

in the savanna. The mammal fauna of the African savanna is distinctive for i= Desert
great number of large grazers, browsers, and their predators. Over 90 differec Arid regions of
species of ungulates (hoofed mammals) alone are found there. Some of the mo r~ earth's land surf
well-known African savanna mammals include elephant, giraffe, rhinoceros, hir- 20 to 30° N and :
popotamus, zebra, gazelle, lion, cheetah, and hyena. Such diversity is not found o:. duces warm and
the savanna of any other continent (Fig. 6.12). The mammal population sizes, pa=-- latitude in area:
ticularly for grazers, can be quite high on the African savanna. It has been es ·- America where
mated that the Serengeti-Mara National Parks of Tanzania and Kenya suppc:: largest single de
some 3 million ungulates in an area of 25,000 sq km. some 9 million s
Human impact on the savanna ecosystem includes the introduction c: Some tropi<
domesticated grazing animals such as sheep and cattle, decimation of populatiocs of 30° C with litt
of native mammals such as elephant, and changes in fire regime. These chang - tral Asia, and So
have occurred in almost all savanna regions that are not protected as parks an.: 15° C, with diffen
preserves. Large savanna preserves are particularly well developed in Africa. I:: Many deserts fou
some areas, control of fire and declines in native grazing populations have led 1~ Diurnal variation
the invasion of grassland by woody species. In other areas, increased rates c- tic of desert clim
hurning to spur grass production have led to declines in woodland cover. Th:: biome is less tha
African elephant (Loxodonta africana) has faced the threat of extinction in i annual precipita1
native savanna at the hands of ivory poachers. In recent years, rigorous prote - while portions of
tion from poachers and an international boycott of new ivory have led to a resu:-- for over 100 year
gence of elephant populations in some parks. The population boom of this no mer when the Int
protected mammals has now led to some destruction of woodland. soon precipitatio
Mojave deserts o

II Great Basin
•Mojave
Sonoran
• Chihuahuan
FIGURE 6.12 The grasslands and short woodlands, and wide diversity of large grazing
mammals on the tropical savanna of Kenya are well illustrated in this picture. The various
mammal species on the African savanna take advantage of different types and layers of
vegetation for feeding and also migrate seasonally to take advantage of different feeding FIGURE 6.13 Th
grounds. MacMahon, 2000) .

tvanna is distinctive for its
redators. Over 90 differen
nd there. Some of the more
mt, giraffe, rhinoceros, hip-
ch diversity is not found or.
mmal population sizes, par-
1 savanna. It has been esti-
nzania and Kenya support
:ludes the introduction o::
, decimation of populatiom
fire regime. These changes
not protected as parks an ·
'ell developed in Africa. lr;
ing populations have led tc
r areas, increased rates o~
es in woodland cover. Th e
! threat of extinction in irs
:ent years, rigorous protec·
w ivory have led to a resur ·
pulation boom of this nO\\
)f woodland.
1ide diversity of large grazin g
~d in this picture. The variou s
lifferent types and layers of
lvantage of different feeding
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 159
Desert
Arid regions of desert over 18 million sq km and comprise about 30% of the
earth's land surface. Most of the world's great deserts are centered on latitudes
20 to 30° N and 20 to 30° S, where descending air from the tropical regions pro-
duces warm and dry conditions. The desert biome extends beyond 40° N and S
latitude in areas such as central Asia , the western United States, and South
America where large mountain ranges produce intense rainshadows. The
largest single desert in the world is the Sahara of North Africa, which covers
some 9 million sq km.
Some tropical portions of the desert experience average annual temperatures
of 30° C with little seasonal variation. Cold desert regions in North America, cen-
tral Asia, and South America can have average annual temperatures of less than
15° C, with differences of as much as 35° C between July and January temperatures.
Many deserts found in higher latitudes experience freezing temperatures in winter.
Diurnal variations in temperature may also be quite high. The defining characteris-
tic of desert climate is severe moisture deficit. Annual precipitation in the desert
biome is less than 50 em per year and is often less than 10 em per year. Average
annual precipitation in the Atacama Desert of Chile averages 0.4 em per year,
while portions of the eastern Sahara Desert may experience no significant rainfall
for over 100 years. Low-latitude deserts generally receive their rainfall in the sum-
mer when the Inter-Tropical Convergence Zone shifts to higher latitudes and mon-
soon precipitation occurs. Some far northern deserts, such as the Great Basin and
Mojave deserts of the United States (Fig. 6.13), experience greatest precipitation in
Ill Great Basin
•Mojave
Sonoran
• chihuahuan
FIGURE 6.13 The desert regions of North America (after Brown and Lomolino, 1998;
MacMahon, 2000).
160 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

winter when they are crossed by midlatitude storm tracks. In the Great Basin a .:
Mojave deserts of California and Nevada, precipitation can come as rain or sno
In some cases, such as the Son or an Desert of southern Arizona and northern Mex-
ico, rainfall occurs during the summer due to monsoon activity and in the wint :
due to midlatitude storms. The Chihuahuan Desert of New Mexico and Mexico :..:
dominated by summer rain. Freezing temperatures or snow is extremely rare :.::.
the Sonoran Desert but occur in other North American deserts. The Great Basi:.
Desert and portions of the Mojave can experience particularly cold temperatures.
In all cases, annual, and generally monthly, potential evapotranspiration rates si=-
nificantly exceed precipitation. In tropical and subtropical deserts, that ratio c:
annual potential evapotranspiration to annual precipitation generally excee ·-
10:1.
Aridisols dominate desert regions. Due to dry conditions, these soils are li:- FIGURE 6.14
tle weathered. They are generally shallow, coarse, and contain negligible orga ;: Desert is well rep
species of Ia rge c
matter. They are often alkaline and can contain bands of calcium carbonate o:
(Carnegiea gigan
salts. Entisols are found in areas of active sand dune activity. These soils are ver:
ground is actuall•
poorly developed and contain very little organic matter of accessible plant nutr:- colder Mojave De
ents. Substrate and soil type are very important for local differentiation of ari.: green plants sho•
land vegetation, and deserts are sometimes classified on the basis of edaphi: Death Valley Nati
(soil) conditions. Hamadas are rocky deserts where fine soil particles have bee::.
washed or blown away. Regs are deserts with a ground cover predominant:.
made of gravel. Sandy deserts, with their characteristic shifting sands and dune>.. rainfall events.
are called ergs. Dry valleys in desert regions are sometimes referred to as wadis.. cases, flowerin
Playas or salt pans are the beds of ancient lakes, or ephemeral desert lakes. The_ tion events. Th
consist of fine sediment, such as clays and silts, and are generally very rich in sal - desert can blo<
and other evaporites. They may have salt crusts at the surface. Oases are areas
where ground water rises to the surface, or at least close enough for access by the
roots of plants. Owing to the availability of water, plants found at oases are ofte::.
not true desert species.
The structure of desert vegetation is characterized by scattered shru s. 30.5
herbs, or grasses. There is usually more bare ground than vegetation. Because o:
the range in precipitation and temperature regimes experienced by differen: 27.9
deserts, there is a wide variety of plant physiognomic features and vegetation den- 25.4
sities (Fig. 6.14). In general, the relative amount of vegetation cover is positive!~
linked to precipitation (Fig. 6.15), but soil conditions are also extremely impor- 22.9
tant. The heavy and saline soils of playas frequently do not support vascula: 20.3
plants. Similarly, few plants can establish or persist on actively moving sand. lr: "E
~ 17.8
deserts with extensive vegetation cover, the dominant plant species often appea;-
~
to be more or less regularly spaced with little growth between them. Some wor · c:
"(ii 15.2
has shown that this is due to root competition for moisture and nutrients. In th a:
12.7
most arid deserts, plants are extremely rare and may only be found in very shel-
tered sites along valleys and in rock crevasses. Trees are generally absent except a 10.2
oases and along river courses. Some desert plants, however, such as the joshua tre
7.6
(Yucca brevifolia) found in the Mojave Desert, the saguaro cactus (Carnegiea
gigantea) of the Sonoran Desert, and the kokerboom (Aloe dichotoma) of Sout 5.1
Africa can grow to heights of over 4 m.
2.5
Desert vegetation generally contains both perennial and annual plants. In
hyper-arid regions, however, with very rare rainfall events such as the interior
Sahara, annual plants may dominate. In these cases, the plants persist for decades FIGURE 6 . 15
as dormant seeds that germinate, mature, and set seed within days or weeks after bushes in the M'
 ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 161

:ks. In the Great Basin and

1 can come as rain or snow.
\.rizona and northern Mex-
1 activity and in the winter
'Jew Mexico and Mexico is
· snow is extremely rare in
.n deserts. The Great Basin
icularly cold temperatures.
1apotranspiration rates sig-
lpical deserts, that ratio of
pitation generally exceeds

nditions, these soils are li t-
contain negligible organi
ls of calcium carbonate or
ctivity. These soils are very
;r of accessible plant nutri-
JCal differentiation of ari
J on the basis of edaphi
ne soil particles have been
lUnd cover predominantly
c shifting sands and dun e
times referred to as wadis.
hemeral desert lakes. The\·
generally very rich in sal ts
e surface. Oases are areas
;e enough for access by th
tts found at oases are ofte
FIGURE 6.14 The relatively diverse succulent and nonsucculent vegetation of the Sonoran
Desert is well represented at Organ Pipe Cactus National Monument in Arizona. The two
species of large cacti are organ pipe (Stenocereus thurberi) in the foreground, with saguaro
(Carnegiea gigantea) in the background. Despite the seemingly dense vegetation, much of the
ground is actually bare of any plant cover (a). In contrast to the Sonoran Desert, the drier and
colder Mojave Desert and Great Basin Desert have much lower plant species diversity. The
green plants shown here are the ubiquitous creosote bush (Larrea tridentata). Areas such as
Death Valley National Park in California have very low amounts of vegetation cover (b).
rainfall events. Both evergreen and deciduous plants are found in deserts. In most
cases, flowering occurs during the wet season or following individual precipita-
tion events. The beautiful coulter hibiscus (Coulter hibiscus) of the southwestern
desert can bloom several times a year whenever a sufficiently heavy rainfall event

rized by scattered shrub

1an vegetation. Because o:
experienced by differen: •
:atures and vegetation den-
getation cover is positive!\·
are also extremely impor-
r do not support vascula;-
n actively moving sand. Ir:: E' • •
-2-
plant species often appear
:§
between them. Some wor'- c
·o;
isture and nutrients. In th
mly be found in very shel-
a: • •
: generally absent except a:
:ver, such as the joshua tre - • • ••
7.6
saguaro cactus ( Carnegiea

••
(Aloe dichotoma) of Soutt 5.1

25
mial and annual plants. Ic ' 2 3 4 5
:vents such as the interi o~ Density (93m 2 )
; plants persist for decades FIGURE 6 . 15 The correlation between annual precipitation and the density of creosote
within days or weeks aft e~ bushes in the Mojave Desert of California (after Woodell et al., 1969).
162 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

occurs. Seedling establishment can be difficult in deserts, and many of the peren- waxes, resins,
nial plants can resprout from below-ground organs if their leaves and stems ar leaves, the ligh
destroyed. The ubiquitous creosote bush (Larrea tridentata) of the Southwes: keep the leaf s
does this readily, and it has been estimated that some patches of this plant may stomata in pitE
represent individuals that have persisted for 11,000 years or more. A number of Some plants, s1
plants can regenerate from portions of detached leaf or stem. The jumping cholla ering of small!
cactus (Opuntia fulgida) of the Sonoran Desert has spiny stem joints that are orative water
very loosely attached to the parent plant. The joints break easily, produce root to keep tempE
and grow into full-sized plants. This species appears to rely more on this mode o- nensis) found
regeneration than on seedlings. same plant us(
The physiognomies and life histories of desert plants generally reflect adap- were long usee
tations to moisture stress. There are many different ways in which such deser Interesti
plants survive in their arid environments. strategy for e:
Drought-escaping annuals germinate, mature, and set seed when soil mois- enough to cot
ture is sufficient to support them. The growing plants have no physiological their leaves dt
mechanism that copes with severe drought. Seeds can remain dormant in the phyllous xero1
upper soil layers, or in some cases, in dried fruiting bodies on the dead paren many plants d
plant. Plants have appeared from the soil seed bank in areas that have no and remain ac
received rain for almost 100 years. Not all drought-escaping seeds are stored i southwestern
the soil. The seeds of the southwestern desert plant Chorizanthe rigida are cidium spp.) c
released from the parent plant over periods of years as the flower and fruiting ratio of surfac
structure that holds them disintegrates. After germination, the growth and see desert broom
set by drought-escapees happen very quickly, generally within six to eight week and rely soh
The annual plant Boerhaavia repens of the Sahara can germinate and set seed mesquite (Pn
within eight days. Many desert annuals can self-pollinate so that viable seeds are terns in order
produced during their short life spans even if they do not come into reproductive Some pc
contact with other members of their species. cope with dr
Some perennial species evade the impact of aridity by remaining dormant leaves are the
until there is sufficient moisture to trigger renewed activity. As in the case o tral, and Sout
drought-escaping annuals, these drought-avoiding plants have no physiological are well-knov
adaptations that cope with arid conditions during their growth periods. Some of tus wislizenii;
these plants persist as bulbs and rhizomes after the above-ground portion of the 70% water. C
plant dies off. They become active very quickly once the soil is moist. The sedge and many ha
species Carex pachystylis of Israel's Negev Desert produces new rootlets and evaporative v
begins spreading within 12 hours after precipitation. A number of woody specie night (see Ch
lose their leaves and become dormant during dry periods. The wolfberry (Lycium limit evapotr:
berlandieri) of North American deserts can lose its leaves several times each year Althou:
and survive for many months until the next rainfall. The white thorn acacia (Aca- as dominant
cia constricta) of the American Southwest loses its leaves in response to drought low root sys
and frost. As a result, the plant can spend much of the year in a leafless state. relatively fre
Perennial species that continue functioning even during the driest portions piration of w
of the year can be divided into three broad classes based on leaf structure. First. day. In add
stenohydric xerophytes, such as members of the genus Euphorbia found in deserts extremely in
of Africa and Asia, shut their stomata to stop water loss during drought and per- Southwest tr
sist for long periods by relying on stored carbohydrate reserves. However. is found in t
because photosynthesis ceases when the stomata are closed, the plant can even- throughout t
tually starve from lack of C02. This strategy is therefore not very useful in summer wet
regions where very long periods occur between precipitation. The sta
Second, sclerophyllous xerophytes, such as agritos (Berberis trifoliolata), Productivity
found in southwestern American deserts, have stiff sclerophyllous leaves with species diver

ts, and many of the peren-
their leaves and stems are
ientata) of the Southwest
patches of this plant may
ars or more. A number of
r stem. The jumping cholla
spiny stem joints that are
reak easily, produce roots.
rely more on this mode of
mts generally reflect adap-
vays in which such desert
d set seed when soil mois-
lts have no physiological
n remain dormant in the
·odies on the dead parent
k in areas that have not
~aping seeds are stored in
Lt Chorizanthe rigida are
as the flower and fruitin g
tion, the growth and seed
' within six to eight week&.
n germinate and set seed
te so that viable seeds are
ot come into reproductive
ity by remaining dormant
1ctivity. As in the case of
nts have no physiological
r growth periods. Some of
we-ground portion of the
1e soil is moist. The sedge
·oduces new rootlets and
number of woody species
js. The wolfberry (Lycium
es several times each year
~white thorn acacia (Aca-
•es in response to drough t
1ear in a leafless state.
during the driest portions
ed on leaf structure. First.
"uphorbia found in deserts
s during drought and per-
jrate reserves. However.
lased, the plant can even-
·efore not very useful in
tation.
os (Berberis trifoliolata).
~lerophyllous leaves with
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 163
waxes, resins, and other coatings that decrease evaporative loses. On agritos
leaves, the light gray color of the waxy coating also helps to reflect sunlight and
keep the leaf surface cool. Other features typical of sclerophyllous leaves, such as
stomata in pits and inrolled leaf margins, are also found on many desert plants.
Some plants, such as wild buckwheat (Eriogonum fasciculatum) , also have a cov-
ering of small hairs on the leaf surface to help reflect sunlight and decrease evap-
orative water loss. Leaves may also be held with their thin edges toward the sun
to keep temperatures and water losses low. The jojoba plant (Simmondsia chi-
nensis) found in Arizona and California has such high-angle leaves. This is the
same plant used today in shampoos, soaps, and waxes. The ground jojoba seeds
were long used by Native Americans for shampoo.
Interestingly, sclerophyllous xerophyty does not appear to be a suitable
strategy for extremely arid deserts. In such areas, the leaf adaptations are not
enough to counter water losses. In more arid settings, plants lose all or most of
their leaves during drought. Such plants belong to the third broad class, malako-
phyllous xerophytes. The term malakophyllous means "soft leafed." However,
many plants do not go dormant. They have green photosynthetic tissue in stems
and remain active. There are many examples of such plants in the deserts of the
southwestern United States and Mexico, including the paloverde plants ( Cer-
cidium spp.) common in the Sonoran Desert. Photosynthetic stems have a low
ratio of surface area and stomata relative to tissue. Some desert plants, such as
desert broom (Baccharis sarothroides) have dispensed with leaves altogether
and rely soley on photosynthetic stems. Finally, some perennials, such as
mesquite (Prosopis spp.), typical of Texas deserts, have extremely deep root sys-
tems in order to reach water during dry periods.
Some perennials store water and have other physiological adaptations that
cope with dry periods. Succulent plants with thick water-storing stems and
leaves are the most common example of this strategy. The cacti of North, Cen-
tral, and South America and the euphorbs typical of African and Asian deserts
are well-known examples of succulents. The saguaro and barrel cactus (Ferocac-
tus wislizenii) of the Sonoran Desert have fleshy stems that are composed of
70% water. Cacti and euphorbs also have waxy coatings on leaves and stems,
and many have spines. Both these features decrease tissue temperatures and
evaporative water loss. The cacti also use the CAM pathway and obtain C02 at
night (see Chapter 2), so they can keep their stomata closed during the day and
limit evapotranspiration.
Although highly adapted to relatively dry conditions, the cacti are not
as dominant in American deserts as one would at first suspect. Cacti have shal-
low root systems, and despite their water storage capabilities must receive
relatively frequent infusions of moisture. The cacti cannot rely on the trans-
piration of water from their stomata to cool leaf and stem surfaces during the
day. In addition, their high water content makes many species of cacti
extremely intolerant of freezing temperatures. It is not surprising that in the
Southwest the greatest diversity of cacti species and their greatest dominance
is found in the Sonoran Desert, with its combination of warm temperatures
throughout the year, and the addition of soil moisture during both winter and
summer wet seasons.
The standing biomass of the world 's deserts is only about 13 billion tons.
Productivity and standing biomass are positively related to precipitation. Plant
species diversity in this harsh biome is also low. Large areas may support only
164 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

one or two species of plant. The entire desert biome in Africa supports approX::- The typic
mately 3000 flowering plant species, but individual desert subregions within the mosaic of valle
continent typically have fewer than 400 plant species. The Tomboctou Desert o:": lands in both tb
Mali supports only 134 plant species. The Australian desert regions have some nated by oaks
1000 different plant species. In North America the deserts support well over 300C Southern beed
different plant species. The most diverse is the Sonoran Desert, which h ~ woodlands. Cor
approximately 2700 species of plants. In contrast, Death Valley supports onl? firs (Abies) , an1
about 279 plant species. Animal diversities are similarly low in desert regions. cone Douglas fi
Human impact on deserts varies greatly. In some areas, such as the centri: tion of the bior
Sahara, the desert's inhospitable conditions preclude human occupation excep: woodland, shru
at oases. It has been suggested that overgrazing in areas such as the souther:: regional rainfal
fringe of the Sahara Desert in Africa may be leading to an increase in desert a: where annual p
the expense of adjacent shrubland and savanna. This conclusion is not universall. colation of wa1
accepted. It has been estimated that the Sahara expanded southward by some annual plants, a
130 km between 1980 and 1990. This represents an increase in the Sahara of some mers and can r
7%. However, during this time there were also northward shifts by as much a;: some of the ext·
110 km owing to climatic variations. Increased salinization of soils due tc overgrazing by
improper irrigation can also cause expansion of desert regions. When large shrublands of tl
amounts of irrigation water are allowed to evaporate from fields, they leay- in California (F
behind salts that can make the soil infertile. In the United States much develo the Mediterran
ment of desert regions has taken place, in areas such as Las Vegas, Nevada, Tu - which is evergn
son, Arizona, and Lancaster, California. This urban growth depends largely oc. consists of shru
imported water. The building of large cities directly destroys local deserts, whil Three em
road building, recreational activity, and other land uses at the fringes of these life history of 1
metropolitan areas cause significant deterioration of desert environments. The low soil nutrie1
smallest North American desert, the Mojave, is particularly threatened by suet then survive th·
activity. One difficulty that faces efforts to preserve desert environments is tha the plant speci
despite their beauty and the many fascinating adaptations of their biota, deserts nate on very dr
are generally perceived as wastelands. mon because t

The Mediterranean Biome

Next we will explore a biome that is not well accommodated within Whittaker's
simple biome model. This is the Mediterranean or winter rain biome. The Mediter-
ranean is the smallest of all the biomes we will consider. It only occupies about 1.
million sq km and is found as narrow strips of land along the Mediterranean Sea.
the southeastern coasts of Africa, Australia, and South America, and the coast of
California. More than half of the total area of the biome is found in the Mediter-
ranean region of southern Europe, western Asia, and northern Africa. In terms of
temperature and annual precipitation, the biome would fall into Whittaker·
Woodland-Grassland-Shrubland category. The biome experiences average annual
temperatures of 15 to 20° C and total annual precipitation of 30 to 100 em. Sum-
mers are warm and winters mild. Frosts are rare. What makes the biome distinc-
tive is the fact that precipitation is limited to the fall , winter, and spring months
only. For four months or so during the summer there usually is no precipitation.
The characteristic soils of Mediterranean regions are formed from the deep
weathering of bedrock and are rich in clay and poor in nutrients. The organic con-
tent of the Mediterranean soils is generally low. In areas of calcareous rocks, a FIG URE 6 . 16 C
deep red color is imparted to the soil by oxidized iron. These distinctive soils are hillside in norther
known as terra rosa. invasive annual h1

n Africa supports appro xi-
:sert subregions within the
The Tomboctou Desert of
desert regions have some
erts support well over 300
moran Desert, which has
'eath Valley supports on! ~
ly low in desert regions.
e areas, such as the centra
human occupation excep
treas such as the southern
to an increase in desert at
)nclusion is not universal!\·
mded southward by som
ease in the Sahara of som
ward shifts by as much a~
linization of soils due to
~sert regions. When large
te from fields, they leav
tited States much develop-
iS Las Vegas, Nevada, Tuc-
:rowth depends largely o
;'!Strays local deserts, while
ses at the fringes of these
desert environments. Th
:ularly threatened by sue
~sert environments is that.
ions of their biota, deserts
odated within Whittaker's
:r rain biome. The Mediter-
. It only occupies about 1. ~
rrg the Mediterranean Sea.
America, and the coast o:
1e is found in the Mediter-
orthern Africa. In terms o:
)Uld fall into Whittaker·s
xperiences average annu ~
tion of 30 to 100 em. Sum-
makes the biome distinc-
winter, and spring months
ually is no precipitation.
are formed from the deep
mtrients. The organic con-
eas of calcareous rocks, 2
These distinctive soils ar
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 165
The typical vegetation structure of the Mediterranean biome includes a
mosaic of valley forests, open woodlands, shrublands, and grassland. The wood-
lands in both the Mediterranean Sea region and in California are typically domi-
nated by oaks (Quercus spp.). In Australia the dominant tree is eucalyptus.
Southern beech (Nothofagus obliqua) and acacia (Acacia) are found in Chilean
woodlands. Conifers also occur in some woodlands. These include pines (Pinus) ,
firs (Abies), and cedars (Cedrus) in the Mediterranean Basin, and pine and big-
cone Douglas fir (Pseudotsuga macrocarpa) in California. The South African por-
tion of the biome is dominated by shrubs and some grasses. The distribution of
woodland, shrubland, and grassland can reflect a number of factors, including
regional rainfall differences, slope aspect, substrate, and disturbance. In regions
where annual precipitation is less than 40 em there is often insufficient deep per-
colation of water to support deep-rooted shrubs. In these areas small shrubs,
annual plants, and grasses predominate. Fire is common during the long dry sum-
mers and can restrict shrub and tree dominance. In addition, it is thought that
some of the extensive shrublands around the Mediterranean Sea are the result of
overgrazing by goats and other livestock, followed by erosion of the topsoil. The
shrub lands of the biome are known by many different regional names: chaparral
in California (Fig 6.16), matorral in Chile, and fynbos in South Africa. In Europe
the Mediterranean shrublands are divided into two main categories: maquis,
which is evergreen shrubs and small trees over 2 m in height, and garrique, which
consists of shrubs between 2 and 0.6 m in height.
Three environmental factors are most important in the physiognomy and
life history of Mediterranean vegetation: the long summer drought, fires, and
low soil nutrients. Annual plants grow during the moist and mild winters and
then survive the dry summers as seeds. Annuals often account for 40 to 50% of
the plant species found in Mediterranean regions. In general, annuals domi-
nate on very dry sites and recently burned areas. Perennial herbs are less com-
mon because they must face severe summer drought on dry sites and must
FIGURE 6.16 Coastal Chaparral shrubs of the California Mediterranean biome cover a
hillside in northern California . The flowering plants in the foreground are dominated by
invasive annual herbs from Europe.
166 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

compete with the roots and canopies of small trees and shrubs that domina:O" bigcone Douglas f
moister sites. Cork used in wim
Interestingly, many trees and shrubs in the Mediterranean biome are eve:-- Many plants can r
green rather than summer deciduous. It has been postulated that, despite tb:: ing following fires.
summer drought conditions, evergreen leaves are favored because they allow th:: ties for sprouting.
plant to take advantage of any soil moisture that is available during the lar:: open only after fir
spring and early fall. This may be an advantage over deciduous species, whi '- soil surface. A nur
have no capacity to benefit from late- or early-growing season moisture. In : temperatures prio
compromise strategy, the black sage (Salvia mellifera) of California loses all b - until activated by
its outermost leaves in the summer. Other species of small shrubs, such as (Teu- Some ecologists 1:
crium polium), possess large leaves during the winter and replace them wi ._ largely a fire-dep<
much smaller leaves during the summer. Nutrient demands are also higher fo:- found in many sh1
species that must produce new leaves each year; this is important in the poor so· - the species diversi
that typify the biome. Interestingly, some trees such as California valley o decrease. A numt
(Quercus lobata) actually lose their leaves in the winter and must rely on dee;- open gaps are ere<
root systems to provide water during their summer growing season. The species
Because of the dry conditions of the summer, many Mediterranean shrub-5 small spatial exten
and trees possess sclerophyllous leaves. Some well-known examples include tb ~ other parts of the
olive (Olea sativa) and the cork oak (Quercus suber) from Europe, manzanita (A rc- Cape Province of :
tostaphylos spp.) from California, and eucalyptus from Australia. The small size an · Human imp;
stiff cuticles of these leaves cut water loss by reducing the evaporative surface arec particularly high. r
and diffusion rates. Stomata also tend to be small and recessed in pits. Species sue agricultural and m
as the California lilacs ( Ceanothus spp.) have stomata that respond quickly to mois- biome but have nc
ture stress by closing. Some plants, such as California chamise (Adenstoma fascicu- duction of exotic
latum), have leaves reduced to needlelike spines, while Spanish broom (Spartiun: Eurasian annual g
junceum) has sclerophyllous photosynthetic stems instead of leaves. lyptus trees introd
Many evergreen shrubs produce aromatic organic substances that ar Shrubs and annua
deposited on the soil around the plant and decrease rates of water infiltration. culus) from Euror
This characteristic may seem odd in a water-stressed environment. However. Control of fires ha
these substances also decrease rates of water evaporation from soils. In addition. shrubs such as co
the morphology of stems on shrubs causes water to flow toward the base of th California. In addi
plant stem. Studies have shown that water infiltration rates are higher at the base fires in California
of chaparral shrubs and that this provides water directly to the root system. natural fires frequ
The root systems of shrubs and trees in the Mediterranean biome are also generally small an
adapted to the summer dry conditions. Plants such as black sage, which are almost small areas of rna
dormant in the summer, typically have shallow and extensive root systems to cap- areas of more rece
ture winter moisture. Evergreen sclerophyllous plants, such as chamise, have very low fuel loads acte
deep root systems that can extend 6 to 8 m and supply moisture during the sum- By trying to excluc
mer. However, even deep-rooted plants tend to have an extensive network of fin e of old chaparral w
roots near the soil surface to maximize water absorption during winter rains. ral fires that burn 1
Adaptations to the low level of nutrients in Mediterranean soils are com-
mon. A number of species have root nodules that contain nitrogen-fixing bacte-
Temperate Gr
ria. Some plants are parasitic and tap into the root systems of adjacent plants with
subsurface organs called haustoria. The haustoria of the Australian plant Nytusia Extensive temperc
fioribunda can reach over 100m in length. Many shrubs in the California chapar- North America, s
ral store nitrogen in old leaves during the winter and then translocate it to new Africa. Temperate
growth during the spring and summer. eastern Europe tt
Fire is common in the Mediterranean biome. Under natural conditions, New Zealand and
many sites will burn every 10 to 30 years. Plant species display strong adaptations North America, pc
to this frequent disturbance by burning. Some trees such as the cork oak and in Eurasia. The tot

and shrubs that dominat e
iterranean biome are ever-
ostulated that, despite the
•red because they allow the
; available during the late
r deciduous species, which
ving season moisture. In a
) of California loses all bu t
small shrubs, such as (Teu-
er and replace them with
!mands are also higher for
important in the poor soils
1 as California valley oak
ter and must rely on deep
:lwing season.
tany Mediterranean shrubs
town examples include the
m Europe, manzanita (Arc-
\ustralia. The small size and
he evaporative surface area
~cessed in pits. Species such
tat respond quickly to mois-
tamise (Adenstomafascicu-
! Spanish broom (Spartium
ad of leaves.
sanic substances that are
rates of water infiltration.
d environment. However.
ion from soils. In addition.
ow toward the base of th e
.·ates are higher at the base
ly to the root system.
literranean biome are also
lack sage, which are almost
ensive root systems to cap-
such as chamise, have very
' moisture during the sum-
1 extensive network of fin e
on during winter rains.
:iiterranean soils are com-
tain nitrogen-fixing bacte-
:ms of adjacent plants with
e Australian plant Nytusia
sin the California chapar-
then translocate it to new
Jnder natural conditions.
display strong adaptations
such as the cork oak and
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 167
bigcone Douglas fir have thick bark that shields the cambium from heat damage.
Cork used in wine bottles is made from the bark of this Mediterranean species.
Many plants can resprout from underground organs or display epicormic sprout-
ing following fires. A number of species of oak and eucalyptus share both capaci-
ties for sprouting. The aleppo pine (Pinus halepensis) has serotinous cones that
open only after fire and distribute the seeds of the species onto the newly burned
soil surface. A number of other species of plants produce seeds that require high
temperatures prior to germination. These seeds may remain dormant in the soil
until activated by fire. Examples include acacias in Australia and California lilac.
Some ecologists believe that the shrublands of the Mediterranean biome are
largely a fire-dependent system. The vegetation structure and the volatile oils
found in many shrubs promote burning. After long periods of time without fire
the species diversity, vegetation productivity, and amount of live vegetation cover
decrease. A number of species seem unable to reproduce without fire, even if
open gaps are created by the removal of other plants.
The species diversity of the Mediterranean biome is high considering its
small spatial extent. The Mediterranean Basin has over 7000 different species. In
other parts of the world, the number of plant species ranges from 6000 in the
Cape Province of South Africa to 2000 in Chile and 900 in California.
Human impact on areas of the biome in Europe and California has been
particularly high. Much of the area in Europe has been impacted by grazing and
agricultural and urban development. Similar pressures exist in other parts of the
biome but have not resulted in the same degree of vegetation change. The intro-
duction of exotic species is particularly problematic in California. Introduced
Eurasian annual grasses have virtually replaced native perennial grasses. Euca-
lyptus trees introduced from Australia have invaded woodlands and grasslands.
Shrubs and annuals, such as Spanish broom and cardoon thistle (Cynara cardun-
culus) from Europe, invade shrublands and grasslands, displacing native plants.
Control of fires has also impacted the biome. Decreases in burning have allowed
shrubs such as coyote bush (Baccharis pilularis) to invade some grasslands in
California. In addition, it is thought that the aerial extent of individual chaparral
fires in California has increased greatly due to fire control measures. Although
natural fires frequently occurred before fire control was introduced, they were
generally small and averaged < 400 ha in area. What resulted was a mosaic of
small areas of mature chaparral with high amounts of fuels, interspersed with
areas of more recently burned chaparral with lower fuel loadings. The areas with
low fuel loads acted as natural fire breaks that served to control the spread of fire.
By trying to exclude fires, we have instead produced a landscape with large areas
of old chaparral with high amounts of fuel. It is not surprising that large chapar-
ral fires that burn over 100,000 ha are becoming more commonplace.
Temperate Grassland
Extensive temperate grasslands are found in the southern and central interior of
North America, small portions of east-central South America, and southern
Africa. Temperate grasslands also occur as a long band extending from south-
eastern Europe through central Asia. Limited areas of grassland are found in
New Zealand and portions of Australia. Grasslands are referred to as prairie in
North America, pampas in South America, grassveld in South Africa, and steppe
in Eurasia. The total area of the grassland biome is around 9 million sq km.
168 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

The mean annual temperature in the grassland biome ranges from over 1Y The phys
C in areas such as South Africa or northern Mexico to 2-3° C in Canada anc pressures: aridi
Siberia. Freezing temperatures are rare in the grassland areas of South Africz enced frequent
and South America. In contrast, average January temperatures in grassland areas have burned al
of western Canada and Siberia are generally less than -10° C. Annual precipita- lands include b
tion averages between 30 to 100 em. The season of highest precipitation varies in Eurasia, and
regionally. Potential evapotranspiration exceeds precipitation during the growing Most tern
season. Grasslands in North America and central Asia also suffer from periodi tion is in Nortl
severe droughts. species of Eurc
Mollisols are the common soil type in the grassland biome. These soils haYe The perennial
a thick dark brown to black surface horizon, high nutrient content, and a loose main stems frm
granular structure. Mollisols are excellent for farming, being particularly suitable thetic stems an<
for cereal crops such as wheat or barley. such as wheat
The structure of grassland vegetation is relatively simple. Grasses anc
herbs dominate. Grasses make up about 90% of the biomass at most grasslan
sites. In terms of species diversity, only about 20% of the plant species found or.
grasslands are grasses; the other 80% are mainly herbs. What at first glance
looks like a monotonous cover of grass may be found to contain a wonderfu~
assortment of wild flowers (Fig. 6.17) . Some trees and shrubs are found in pro-
tected valley sites and as gallery forests along rivers. In some cases, trees such as
limber pine (Pinus flexilis) and ponderosa pine (Pinus ponderosa) can be foun c
growing on rocky outcrops in the northern prairie of central North America.
Scotts pine (Pinus sylvestris) can be found on similar sites in portions of th
0
Eurasian steppes.
50

E 100
~
.c
15.
Q)
0 150

200

250

FIGURE 6.17 A remnant of tall grass prairie on the Nature Conservancy Preserve in FIGURE 6.18 il
Oklahoma . Although the above-ground biomass is dominated by grasses, there is a good 1979) and the tiller
diversity of flowering herbs in the vegetation. 1968 and Archibol

.orne ranges from over 1S:
to 2-3° C in Canada an
md areas of South Africa
;ratures in grassland areas
-10° C. Annual precipita-
ighest precipitation varies
itation during the growinE
t also suffer from periodi-
ad biome. These soils ha,··
rient content, and a loos-
being particularly suitab]-
vely simple. Grasses an ·
1iomass at most grasslanc
he plant species found or:
~rbs. What at first glance
d to contain a wonderfu~
l shrubs are found in pro-
1 some cases, trees such as
·ponderosa) can be fou nc
f central North America.
tr sites in portions of th
onservancy Preserve in
JY grasses, there is a good
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 169
The physiognomy of the plant species reflects the influence of three main
pressures: aridity, fire, and grazing. Prior to human intervention, grasslands experi-
enced frequent fires. Some grasslands in the southwestern United States appear to
have burned almost every year. Native grazing animals found in temperate grass-
lands include buffaloes (Bison bison) in North America, horses (Equus caballus)
in Eurasia, and many smaller mammals and insects such as grasshoppers.
Most temperate grasslands are dominated by perennial grasses. One excep-
tion is in North America in the Central Valley of California, where introduced
species of European annual grasses have displaced the native perennial grasses.
The perennial bunch grasses, such as needle grass (Stipa spp.), have very short
main stems from which arise photosynthetic stems and leaf blades. The photosyn-
thetic stems and blades of grass are called tillers (Fig. 6.18). In sod-forming grasses,
such as wheat grass (Agropyron spp.), tillers arise from lateral underground
Tall Mixed Short
D E F G H J K
0
, , J RI !
0
so
150
I ~~
' '
. .
.~ L
..,
I J
!''Pr '~
i!
I
! ;~_t '(I ! '', ~
\
II i If
11 A ndropogon gerardi
B Panicum virgatum
C Spartina pectinata
D Stipa Spartea
c Koeleria cristata
Andropogon scoparius
200 -1 ~f) ')fiK I G Sporobolus heterolepis
Buchloe dactyloides
I Aristida purpurea
250 ..J I
Agropyron smithii
Bouteloua curtipendula
• Tall-grass prairie
Mixed prairie
Short-grass prairie
IIIII Desert grassland
• Palouse prairie
• California prairie
• Aspen parkland
FIGURE 6.18 Major grassland formations of North America (after Sims and Coupland,
1979) and the tillers, stolons, and root systems of some common grasses (after Weaver,
1968 and Archibald, 1995).
170 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

rhizomes. Some grasses, such as buffalo grass (Buchloe dacyloides), produ ~ teenth century,
above-ground stolons that also grow laterally and produce new tillers. Rhizom : brome (Bromu
and stolons allow the sod grasses to spread over considerable areas and produce c. come to domin
dense mat of vegetation. Such grasses can spread over 20 sq m in as little as t,,-c pied by more b
years. Following heavy browsing or fires, most perennial grasses can resprout fr oc: duced annual g
their roots or rhizomes. Rhizomes and stolons also store carbohydrates and he!; Grass lane
grasses survive during periods of drought. Grassland herb species such as golde:: mary producti'
rod (Solidago spp.) also possess rhizomes. Annual produc
In some cases, the root biomass of grasslands can be three times larger th a= cipitation is 40 1
the living biomass found above ground. The root systems of grassland plant5 em. The diversi
reflect two main strategies for coping with arid conditions. Short grass speci : structurally cor
(with tillers usually< 50 em in height) found on arid sites typically have intensi ,-~ the central pla
networks of dense fine roots within the top 150 em of the soil surface (Fig. 6.1 herbs can be fo
Examples include buffalo grass and wheat grass. Some short bunch grasses ha,·- Human ir
as much as 90% of their root biomass concentrated in the top 15 em of soil. Suet are useful for !
species depend on efficient capture of rainfall at the surface, leaving little wat e~ grassland veget
to percolate down to lower levels in the soil. This strategy is advantageous in d0 North America
areas. Tall grass species (tillers up to 1m in height), such as switch grass (Panciwr. 370 million ha t
vigatum), grow on moist sites and have long roots that can extend over 2m belo' tall grass and m
the surface (Fig. 6.18). These species take advantage of deeper ground water an · and corn have <
are particularly suited to areas where rainfall is sufficient for water to percolat - haps only 1% c
to depth in the soil. Many grassland herb species have similar deep roots that car: the United Stat,
extend more than 3 m in depth. Some species, such as blue-stem grass (Andro- changed the na
pogon gerardi), have a combination of dense and fine extensive root systems in tation cover an1
the top 30 to 50 em of the soil and coarse intensive roots that penetrate more th ac portions of Tex<
2m (Fig. 6.18). over 90% of th
The root systems of grassland species may also be important in determining and herbs are f
the position of the ecotone between grassland and adjacent forest. The concen- grasslands are c
tration of fine roots near the surface allows the grasses to capture precipitation as the southern
before it can percolate downward and provide moisture to the deeper root sy - allowed shrubs
terns of trees and shrubs. The dense root mat also makes it difficult for tree and
shrub seedling to establish. Finally, sprouting from roots and rhizomes allm,-s
Temperate
perennial grasses to survive fires that kill tree and shrub seedlings.
The wide range in temperatures, and to a lesser extent precipitation Temperate fore
regimes, experienced within the grassland biome produce regional differences in ern Asia, southt
vegetation composition and structure. Within North America, the grassland biome occupie~
biome is often divided into several different formations (Fig. 6.18). The relatively deciduous fore~
moist eastern portions of the central plains support tall grass prairie dominated is between 50 :
by species such as eastern blue stem (Andropogon gerardi) and switch gras occurrence of f
(Panicum virgatum). Some of these grasses can grow to heights of 2m. The cen- America and p<
tral portion of the plains supports mixed-grass prairie with some tall grasses. temperatures o
medium-sized species such as little bluestem (Andropogon scoparius), and short States and Ne"
grass species such as buffalo grass. The height of vegetation in the mixed-gras that are above f
prairie generally does not exceed 1.5 m for the taller grasses, with an understory from region to
of short grasses that grow to 30 em in height. The western edge of the plains i with most preci
composed of short grass prairie that includes species such as buffalo grass and has generally hi
blue grama (Bouteloua gracilis). The height of the vegetation is generally 30 to 50 Soils in th
em. The California prairie originally supported more perennial bunch grasses alfisols, commo
from genera such as three awn grass (Aristida spp.), bunch grass (Poa spp.), and of Asia, have a
needle grass (Stipa spp.). Starting with settlement by the Spanish in the late eigh- quite fertile. Ul

chloe dacyloides), produce
)duce new tillers. Rhizomes
ierable areas and produce a
:r 20 sq m in as little as two
al grasses can resprout from
ore carbohydrates and help
herb species such as golden
n be three times larger than
ystems of grassland plants
ditions. Short grass species
ites typically have intensive
·the soil surface (Fig. 6.18).
e short bunch grasses have
1 the top 15 em of soil. Such
surface, leaving little water
tegy is advantageous in dn
::h as switch grass ( Pancium
can extend over 2 m belO\\
,f deeper ground water and
:ient for water to percolat
similar deep roots that ca
1s blue-stem grass (Andro-
:: extensive root systems in
ts that penetrate more tha
e important in determining
ijacen t forest. The concen-
es to capture precipitatio
ure to the deeper root sys-
kes it difficult for tree and
roots and rhizomes allows
1b seedlings.
esser extent precipitation
.uce regional differences in
h A merica, the grasslan,
1s (Fig. 6.18). The relatively
1ll grass prairie dominated
gerardi) and switch grass
to heights of 2 m. The cen-
"ie with some tall grasses.
ogon scoparius), and shon
etation in the mixed-gras"
~rasses, with an understory
:stern edge of the plains is
such as buffalo grass and
~tation is generally 30 to 50
:: perennial bunch grasses
unch grass (Poa spp.) , and
1e Spanish in the late eigh-
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 171
teenth century, European annual grasses of the genera wild oats (Avena spp.),
brome (Bromus spp.) , and fescue (Festuca spp.) have been introduced and have
come to dominate. The Palouse prairie of the Pacific Northwest was also occu-
pied by more bunch grasses but has come to be increasingly dominated by intro-
duced annual grasses.
Grasslands contribute some 14 billion tons of standing biomass. Net pri-
mary productivity is positively correlated with rainfall in the grassland biome.
Annual productivity may range from 200 g per square meter where annual pre-
cipitation is 40 em to over 400 g per square meter where annual precipitation is 80
em. The diversity of plant species found in grasslands is not as high as in more
structurally complex systems, such as tropical and temperate forests. However, in
the central plains of North America, over 100 different species of grasses and
herbs can be found in areas of 1 to 2 ha.
Human impact on the grassland biome has been considerable. Grasslands
are useful for growing crops and grazing livestock. Most of the area of natural
grassland vegetation in North America and Eurasia is now used for agriculture. In
North America the area covered by grasslands in the Great Plains decreased from
370 million ha to 125 million ha between 1830 and 2000. In some cases, such as the
tall grass and mixed-grass regions of the Great Plains, fields of crops such as wheat
and corn have almost completely replaced native prairie. It is estimated that per-
haps only 1% of these grasslands remain intact in certain states and provinces of
the United States and Canada. Within the short grass prairie, extensive grazing has
changed the nature of the vegetation. Heavily grazed fields have less dense vege-
tation cover and a much higher proportion of invasive nonnative plant species. In
portions of Texas, Oklahoma, and Nebraska, nonnative plant species can make up
over 90% of the grass cover on overgrazed fields. Introduced species of grasses
and herbs are found in all North American grassland formations. The California
grasslands are completely dominated by such invasive species. In some areas, such
as the southern and northern edges of the central plains, the exclusion of fire has
allowed shrubs and trees to invade former grasslands.
Temperate Forests
Temperate forests are found in eastern North America, portions of Europe, east-
ern Asia, southern South America, southeastern Australia, and New Zealand. The
biome occupies some 7 million sq km. The mean annual temperature of the
deciduous forest region ranges from 20° C to SO C. Average annual precipitation
is between 50 and 250 em. A distinguishing characteristic of the biome is the
occurrence of frosts in the winter. Some regions, including northeastern North
America and portions of Asia, experience long cold winters with average January
temperatures of less than, - 10° C. Areas of the biome in the southeastern United
States and New Zealand have milder winters, with January mean temperatures
that are above freezing. The annual distribution of precipitation is highly variable
from region to region. Areas of eastern Asia experience a monsoonal regime,
with most precipitation occurring as summer rainfall. In contrast, western Europe
has generally high precipitation throughout the year.
Soils in the temperate forest biome include alfisols, ultisols, and entisols. The
alfisols, common in cooler areas of eastern North America, Europe, and portions
of Asia, have a brown surface horizon that is rich in humus. They are generally
quite fertile. Ultisols are associated with warmer portions of the biome in south-
172 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

rates of evapotr:
onset of dorman
ments such as ar
of the leaves di~
require a period
stimulate bud o
rubrum) from n<
temperatures fol
leafing out. The <
the passing of wi
strategy to be eff
for the trees to p1
duce sufficient n
and spring when
new leaves to rna
The timing
species to specie
stand will lose th
the spring will al:
ering of the cane
spring before full
FIGURE 6 . 19 Autumn foliage in an area of mixed deciduous and coniferous forest in to be transportee
New Hampshire. The loss of leaves and dormancy allows the broadleaved tree species to tors have a bette1
survive the long and cold winter climate. The structu
quite similar. In ;
dominate the for'
deciduous trees s
eastern North America, Asia, Australia, and New Zealand. The ultisols are gener- spp.), hickory (C
ally red to yellow in color. Most nutrients are found in decaying organic matter a: (Quercus spp.). T
the top of the soil and are quickly recycled by the forest vegetation. m. Beneath the c
Within the northern hemisphere, the key characteristic of the temperat saplings of the c:
forest biome is the dominance by winter deciduous trees (Fig. 6.19). Winter tem- large shrubs such
peratures in the portion of the biome that lies within the southern hemispher there is a ground
are milder. Most of the temperate forest regions there are dominated by and ferns. Vines o
broadleaf evergreen trees such as mountain southern beech (Nothofagus solan- or as diverse as ir
dri) in New Zealand and eucalyptus species in Australia. Evergreen conifer evergreen conifer
species including matai (Podocarpus spicatus) and the beautiful kauri tre spp.) and a few ev
(Agathis australis) in New Zealand also occur. The only southern hemisphere for- Deciduous f
est that is dominantly deciduous is found in the drier and colder portions o' America, extendir
Patagonia. Here the forest is dominated by deciduous species of southern beec expanse, differenc
such as lengue (Nothofagus pumilo). ences in the fares
The deciduous temperate forests of the northern hemisphere are distinctive subdivided into a
for the winter loss of leaves from broadleaved trees and shrubs. During the winter. include a southerr
temperatures are too cold for photosynthesis, and freezing conditions damag southern magnoli
exposed soft tissues. As a response to cold conditions, deciduous plants lose their (Pinus taeda) and
leaves and go dormant. It is thought that the deciduous trait may have developed number of species
in plants in response to winter drought in the tropical seasonal forest region in the merly by chestnut
geologic past. The environmental cue that leads to leaf mortality and release trees, including sp
varies for different tree species. In some cases, tree species respond to decreased (Lira dendron tuli
daylight in the autumn. In other cases, increased moisture stress due to decreased diversity of broad

)US and coniferous forest in
3 broad leaved tree species to
tland. The ultisols are gener-
1 decaying organic matter a:
est vegetation.
racteristic of the temperate
rees (Fig. 6.19). Winter ten:-
in the southern hemisphere
s there are dominated b·
n beech (Nothofagus solw:-
ustralia. Evergreen conife~
d the beautiful kauri tre.:
ly southern hemisphere far-
rier and colder portions c:
ts species of southern bee(:
n hemisphere are distincti\·;:-
td shrubs. During the wint e~
freezing conditions damag:-
., deciduous plants lose thee
us trait may have develope "
seasonal forest region in th:-
leaf mortality and releas:-
Jecies respond to decrea s e~
.ture stress due to decrea s e~
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 173
rates of evapotranspiration and water conductance leads to leaf mortality. At the
onset of dormancy, the leaves lose their chlorophyll and turn red or yellow as pig-
ments such as anthocyanins are exposed. The walls of abscission cells at the base
of the leaves disintegrate and release the leaf from the tree. Many tree species
require a period of chilling followed by sustained warm temperatures in order to
stimulate bud opening and the development of new leaves. Red maple (Acer
rubrum) from northeastern North America requires at least a month of freezing
temperatures followed by warm temperatures of 15° Cor more in order to initiate
leafing out. The chilling requirement keeps trees from breaking dormancy before
the passing of winter and the onset of warm spring conditions. For the deciduous
strategy to be effective, the summer growth period must be long and warm enough
for the trees to produce a new crop of leaves, form buds for the next year, and pro-
duce sufficient reserves of carbohyhdrates to sustain the plant during the winter
and spring when new leaves begin to develop. It can take several weeks for the
new leaves to mature and reach peak photosynthetic activity.
The timing for leaf abscission and subsequent leafing out varies from
species to species and from year to year. In general, most tree species within a
stand will lose their leaves during the same 14- to 30-day period. Leafing out in
the spring will also occur at about the same time for all species in a stand. Flow-
ering of the canopy trees and many of the understory plants occurs early in the
spring before full leaf development. As a result, it is easier for windborne pollen
to be transported through the canopy, and insects and other animal pollen vec-
tors have a better opportunity to find flowers.
The structure of deciduous forests in North America, Europe, and Asia is
quite similar. In addition, the same genera of broadleaved and coniferous trees
dominate the forests on the three continents. There is a canopy formed by large
deciduous trees such as beech (Fagus spp.) , maple (Acer spp.), basswood (Tilia
spp.), hickory (Castanea spp.), elm (Ulmus spp.), ash (Fraxinus spp.), and oak
(Quercus spp.). The height of the mature canopy is generally between 20 and 30
m. Beneath the canopy, there is a layer of subdominant trees that may include
saplings of the canopy dominant trees, shade tolerant subdominant trees, and
large shrubs such as dogwood (Comus spp.) and hazel ( Corylus spp.). Finally,
there is a ground layer formed by tree and shrub seedlings, small shrubs, herbs,
and ferns. Vines occur in the temperate deciduous forests but are not as common
or as diverse as in the rainforest. Most temperate deciduous forests also contain
evergreen conifers from genera such as pine (Pinus spp.) and hemlock (Tsuga
spp.) and a few evergreen shrubs such as holly (!lex spp.).
Deciduous forest grows across a wide latitudinal range in eastern North
America, extending from northern Florida to southern Canada. Within this broad
expanse, differences in seasonal temperatures and precipitation produce differ-
ences in the forest composition. The North American deciduous forest is often
subdivided into a number of associations (Fig. 6.20). From south to north these
include a southern mixture of deciduous trees, evergreen broadleaf trees such as
southern magnolia (Magnolia grandiflora), and conifers such as loblolly pine
(Pinus taeda) and shortleaf pine (Pinus echinata); an association dominated by a
number of species of oaks and pines; an association dominated by oaks and for-
merly by chestnut (Castanea dentata); an area of mixed mesophytic deciduous
trees, including species of maples, beeches, elms, basswoods, and the tulip tree
(Lirodendron tulipifera) (this mixed mesophytic association has the highest
diversity of broadleaf trees in North America); an association in the southern
174 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

The stanc
210 billion tom
relatively high.
ranges from 18,
region. Even ir
ferent species c
est, some 2000
herbaceous spe
also has a dive
coniferous gen
Temperate maple alone. T
Forest Regions North America
Mixed Mesophytic
Because c
perate forests t
l!iiJ Western Mesophytic ging has long l
0 Oak-Hickory forests for agri<
0 Oak-Chestnut northwestward
of the arable la
.Oak-Pine ous forests in CJ
0 Southeastern Evergreen ulations in eas
0 Beech-Maple
swidden agricul
burning. Howe·
• Maple-Basswood
American fores
Ill Hemlock-White Pine- forest remained
Northern Hardwoods human activity ;
icantly altered ·
FIGURE 6.20 Eastern North American temperate forest formations (after Archibold, 1995). growth forest. 1
found in the we
forest was clear
Great Lakes region typified by maples and beech; a forest bordering the Great primary forest t
Plains dominated by oaks and hickory; and a mixed wood in the north typified by The accid
deciduous species such as maple and oak in association with conifers such as pogenic pressur
white pine (Pinus strobus) and eastern hemlock (Tsuga canadensis). of beech, chestr
During the winter, about 70% of the light received at the top of the leafles pathogens. For ,
canopy is transmitted to the ground surface. However, during the summer only phonectria para
about 10% of the incident light is transmitted to the forest floor. This light on the sia. Old World r
forest floor is more than that in tropical rainforests; as a result, the understory fungus. By 195C
vegetation in the deciduous forest can be relatively lush. Some trees, such as oaks. and the species
have different leaves on canopy branches as compared to lower limbs. The sun and are killed b
leaves found at the canopy level are smaller in area and thicker, and have more tially removed f
chlorophyll and more stomata than the lower shade leaves. Certain tree genera In recent
including birch (Betula spp.) , cherry (Prunus spp.), and poplar (Populus spp.) that forests. First, pa
occur in the deciduous forest do not grow as high as canopy dominants and are to protect remai
intolerant of shade. They require gaps in the forest in order to establish and grow. origins several t
Due to natural mortality, windthrow, and other disturbances, it is estimated that hunting by Eur
approximately 10% of the deciduous forest consists of canopy gaps at any given northern Great
time. Shade intolerant subcanopy trees such as cherry or poplar can persist 30 to ically viable to p
80 years in such gaps before being shaded out by canopy-dominant trees such as allowed to reve:
maple or beech. When undisturbed, the canopy-dominant trees generally live 300 forest. In genen
to 400 years. sity of large prin

...
):,D
iemperate
=orest Regions
Ill Mixed Mesophytic
~Western Mesophytic
~Oak-Hickory
~Oak-Chestnut
.Oak-Pine
~Southeastern Evergreen
::J Beech-Maple
• Maple-Basswood
l1 Hemlock-White Pine-
Northern Hardwoods
lations (after Archibald, 1995 _
orest bordering the Grea:
,od in the north typified b\-
:ion with conifers such a-'
1 canadensis).
:d at the top of the leafless
·,during the summer only
rest floor. This light on the
iS a result, the understory
1. Some trees, such as oak
d to lower limbs. The sun
1d thicker, and have more
~aves. Certain tree genera
poplar (Populus spp.) tha
:anopy dominants and are
rder to establish and grow_
)ances, it is estimated that
canopy gaps at any give
or poplar can persist 30 to
py-dominant trees such as
mt trees generally live 30
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 175
The standing biomass contributed by temperate deciduous forests is about
210 billion tons. The plant species diversity in the temperate deciduous forests is
relatively high. The tree species diversity in North American deciduous forests
ranges from 180 species in the Southeast to 60 species in the western Great Lakes
region. Even in the Great Lakes region, it is not unusual to find 10 or more dif-
ferent species of trees on a 1-ha study site. Within the southeastern deciduous for-
est, some 2000 species of vascular plants are known. As many as 14 different
herbaceous species can be found within a single square meter of forest floor. Asia
also has a diverse deciduous flora. Over 50 genera of broadleaved trees and 12
coniferous genera are known from China, with over 50 individual species of
maple alone. The diversity of tree species is not nearly as great in Europe as in
North America or Asia, but it is still considerable.
Because of their temperate climate and fertile soils, much of the world's tem-
perate forests have been cleared for agriculture and settlement. In addition, log-
ging has long been practiced in these forests. Significant clearing of European
forests for agriculture commenced some 5000 to 8000 years ago and progressed
northwestward from Turkey and Greece. Between 2000 and 4000 years ago, most
of the arable lands throughout Britain were cleared of forest. Clearing of decidu-
ous forests in China for agriculture began some 5000 years ago. Native Indian pop-
ulations in eastern North America were clearing small sections of forest for
swidden agriculture 2000 years ago and perhaps modified some larger areas by
burning. However, the arrival of Europeans caused massive clearing of North
American forests. By the early twentieth century, only a tiny fraction of original
forest remained in the eastern United States. Stands that have not been cleared by
human activity are called primary forest. If the primary forest has not been signif-
icantly altered by recent natural disturbance, it is sometimes referred to as old
growth forest. The most recently cleared deciduous forest in the United States is
found in the western Great Lakes region. During 1850-1995, almost 99% of that
forest was cleared or altered. In many other areas of the temperate forest biome,
primary forest has been reduced to only 1 to 2% of its natural extent.
The accidental introduction of pathogens has been an additional anthro-
pogenic pressure on the deciduous forest biome in North America. Populations
of beech, chestnut, and elm have all declined due to introduced blights and other
pathogens. For example, in the early twentieth century, the chestnut fungus Cry-
phonectria parasitica was introduced to the eastern deciduous forest from Eura-
sia. Old World relatives of the North American chestnut have a resistance to the
fungus. By 1950 all North American chestnut trees were infected by the blight,
and the species only persists as small suckers that emerge from the root system
and are killed before reaching tree form. Thus, a dominant tree has been essen-
tially removed from a large portion of the forest .
In recent times, two factors have led to some recuperation of deciduous
forests. First, parks and conservation areas have been created around the world
to protect remaining forests. The preservation of deciduous forest patches has its
origins several hundred years ago with the establishment of game parks used for
hunting by European aristocracy. Second, in areas such as New England, the
northern Great Lakes region, and portions of northern Europe, it is not econom-
ically viable to practice farming on some formerly forested sites. Farms have been
allowed to revert back to forest. Such reforested sites are called second growth
forest. In general, they do not have the same species diversity or structural diver-
sity of large primary forest stands that include closed canopy and gap sites. Many
176 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

parts of northern Europe and northeastern North America have far more deciC.-
uous forest today than they did 100 years ago. In fact , it is estimated that owing tc
the increase in deciduous forest, the total rate of reforestation exceeds deforesta-
tion in the United States. Europe has experienced a similar increase.

Temperate Rainforest
The temperate rainforest is a small and scattered biome. The largest conterminoc.s
area of temperate rainforest is found on the northwest coast of North Ameri
extending from northern California to the panhandle of Alaska. Smaller strips o:
temperate rainforest are found in southeastern Australia, Tasmania, New Zealan '
and the southwestern coast of South America. Annual rainfall exceeds 200 err:..
and precipitation occurs throughout the year. In some regions total rainfa:...
approaches 400 em. All areas of temperate rainforest are found close to the oceans
and have mild temperature regimes. Average annual temperatures range from jus:
less than 20° C to about 5° C. Winters may experience freezing temperatures be:
are generally mild. Summers are cool.
Along the California coast, the temperate rainforest vegetation extends
quite far south into the dry summer climatic region of the Mediterranean biom
This extension of the rainforest is due to the prevalence of summer fogs in this FIGURE 6.21 T
National Park in W
region. Condensation along the branches and needles of the redwood tre :
little light to penet
(Sequoia sempervirens) that dominate the forest in coastal California falls <E
relatively sparse al
large droplets at the base of the canopy and provides an important source o:
moisture during the dry summer months. In addition, large amounts of winte:
orographic rainfall are generated by air rising over the coastal mountains, an..:
this produces high soil moisture that helps trees persist over the summer. soils quite acidi<
Soils in the temperate rainforest biome are generally spodosols, which ar= consisting of a c<
acidic and low in nutrients. They often contain a B horizon composed of reddisi:: of early successi
mineral matter overlain by a light gray E horizon that has lost iron and aluminu c: immature conif<
through intensive eluviation to the B horizon. In poorly drained areas, histosols. spp.), together w
made up of poorly decomposed organic matter, are common. These soils ar"'" tered. The canop
extremely acidic and contain almost no nutrients. of the dominant
The temperate rainforest biome is dominated by evergreen trees. In the tressed trunk sys
southern hemisphere, the dominant trees include karri (Eucalyptus diversicolor high as in the te
and other eucalyptus species in Australia, southern myrtle beech (Nothofagus conditions and
cunninghammi), southern mountain ash (Eucalyptus regnans) and the huon pin- mature stands, is
(Dacrydium franklinii) in Tasmania, and southern beeches (Nothofagus spp . . The large t
Patagonian cypress (Fitzroya cupressiodes), and monkey puzzle tree (Araucarin Iarly attractive t
araucana) in South America. The North American portion of the biome is domi- rugged and inac<
nated by evergreen conifers, including the giant coastal redwood in California. ging activities. H
The redwood can reach heights of over 110 m. Douglas firs (Pseudotsuga men- practices such as
ziesii), which are commonly over 50 m tall, grow from California to Alaska. Othe: In addition, log1
massive, dominant trees in the northwestern rainforests include red cedar (Thuja approach in whi<
plicata), sitka spruce (Picea sitchensis), amabalis fir (Abies amabalis), and yellO\\- Approximately ·
cypress (Chamaecyparis nootkatensis). In addition to great size, the dominan between 1940 ar
trees of the temperate rainforests often have very long life spans. Douglas fir remains unlogge
yellow cypress, red cedar, and huon pine all can typically reach ages of 1000 year_ whereas Alaska I
or greater. California redwoods may reach ages of over 2000 years. 55% of the temr:
Light penetration in the mature canopy of the temperate rainforest is very parks or reserve
limited (Fig. 6.21). In addition, the litter from the dominant conifers makes the endangerment oJ

nerica have far more decid-
it is estimated that owing to
restation exceeds deforesta-
:imilar increase.
1e. The largest conterminous
~st coast of North America.
of Alaska. Smaller strips o:
Jia, Tasmania, New Zealand.
1al rainfall exceeds 200 em.
some regions total rainfaE
tre found close to the oceans
~mperatures range from jus
e freezing temperatures bu
nforest vegetation exten d~
f the Mediterranean biom~.
~nee of summer fogs in tllli
dies of the redwood tree ~
coastal California falls ~
:les an important source o:
•n, large amounts of wint e~
the coastal mountains, anc
st over the summer.
terally spodosols, which are
Jrizon composed of reddist
has lost iron and aluminuiL
1rly drained areas, histosol
~ common. These soils are
by evergreen trees. In th
ri (Eucalyptus diversicolo r
myrtle beech (Nothofagus
regnans) and the huon pine
Jeeches (Nothofagus spp. l.
key puzzle tree (Araucaria
1rtion of the biome is domi-
stal redwood in California.
~las firs (Pseudotsuga men-
California to Alaska. Othe
:ts include red cedar (Thuja
1bies amabalis), and yello""
o great size, the dominan:
mg life spans. Douglas fir·
lly reach ages of 1000 yeaE
~r 2000 years.
emperate rainforest is verY
minant conifers makes th -
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 177
FIGURE 6.21 The cool and moist floor of the coastal temperate rainforest of Olympic
National Park in Washington. The dense closed canopy formed by large conifers allows
little light to penetrate to the ground. Vascular plant vegetation on the forest floor is
relatively sparse although there is a dense covering of mosses.
soils quite acidic. As a result, the structure of the forest is very simple, usually
consisting of a continuous conifer canopy and sparse subdominant layer made up
of early successional trees such as California bay (Umbellularia californica) and
immature conifers. A lower layer of shrubs includes huckleberry (Vaccinium
spp.), together with a few herbs and ferns. Thick carpets of moss are also encoun-
tered. The canopy of the forest generally ranges from 40 to 60 m in height. Trunks
of the dominant trees are often massive, reaching 20 m in circumference. But-
tressed trunk systems occur in a number of species. Tree species diversity is not as
high as in the temperate deciduous forest. In addition, because of the low light
conditions and acidic soils, the shrub and herb diversity, particularly within
mature stands, is also very low.
The large trees of the temperate rainforest have made this region particu-
larly attractive to logging operations and forest clearance. For many years, the
rugged and inaccessible nature of much of the Pacific Northwest restricted log-
ging activities. However, the increasing value of quality wood and the advent of
practices such as helicopter slinging to remove timber has changed this situation.
In addition, logging in the coastal rainforest has tended to use the clear-cut
approach in which essentially all trees are removed from large swaths of forests.
Approximately 76% of the Olympic National Forest of Washington was cut
between 1940 and 1990. In Oregon, perhaps only 4% of the coastal rainforest
remains unlogged. British Columbia has seen over 50% of its rainforest logged,
whereas Alaska has had about 11% logged. Worldwide, it is estimated that about
55% of the temperate rainforest has been logged, and only 17% is protected in
parks or reserves. Destruction of large portions of the forest has led to the
endangerment of bird and mammal species adapted to the old growth stands. In
178 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

North America, both the spotted owl (Strix occidentalis) and marbled murrele peratures of 1
(Brachyramphus marmoratus) have been threatened with extinction due to loss atures reachir
of old growth habitat. Efforts to protect the owl led to large areas of old growth precipitation i
forest being closed to logging in the United States. The remaining old growth precipitation i
stands are too small to support the industry for more than a few years in any case. tion in the wir
Ongoing battles between conservationists and logging interests continue ir. Why are
places such as the Carmanah Valley and the Clayoquot Sound of British Colum- northern hem
bia over how much of the remaining portions of this biome should be preserved. mate does no1
nental land in
have conifer-d
Coniferous Boreal (Taiga) and Montane Forests
at high elevati
The coniferous boreal and montane forests are found only in the northern hemi- beech forests <
sphere. The boreal forest, known as taiga in Eurasia, extends as a broad band cen- ical Africa, wo,
tered on 60° N latitude (Fig. 6.22). The boreal forest extends across northern a relative of t
Eurasia from Norway to the Pacific Coast of Russia and the northern islands o: Lobelia kenien
Japan. In North America the boreal forest extends from coastal Alaska to the erous forests d
island of Newfoundland. It is the largest conterminous forest in the western Spodoso
hemisphere. The largest montane portion of the coniferous biome extends down boreal zone. Ir
the Rocky Mountain region of North America into the highlands of Mexico. biome and in :
Small portions are also found at higher elevations in the Appalachian Mountains and on poorly
in the East. In Eurasia areas of montane coniferous forest are found at higher have poorly d<
elevations in alpine regions such as the Alps or the Himalayas. The total area o~ High-latitude ~
the biome is around 12-15 million sq km. It has been estimated that the boreaJ southern bordi
forest constitutes one-quarter to one-third of all forest cover on the planet. forest zone in :
The coniferous boreal and montane forests are found in regions with a ous to sporadic
mean annual temperature of between 5° C and -5° C. Mean annual precipitation of the boreal b
varies between about 20 em and 200 em. In general, precipitation is lowest in the The conif
boreal portions of the biome. Summers are generally mild, with average July tern- by evergreen sr
(Abies spp.). Ir
conifer species,
These trees, tho
during the win
Siberia because
tinental region.
age and desicca
The struc
dominated by c
m in height, an
dron gigantewr
100m. The coni
pines (Pinus ar,
4000 years. Ott
can reach ages
many other con
Recently
deciduous broa
forest canopy i~
FIGURE 6.22 Winter in the boreal forest of the Pelly Mountains in the Yukon Territory of Similarly, mid-e
Canada. The diversity of tree species is very low in this biome. All of the coniferous trees in canopies. Towm
the picture are either white spruce (Picea glauca) or black spruce (Picea mariana). in the mountain

ttalis) and marbled murrele:
d with extinction due to loss
to large areas of old gro\\rtt
:. The remaining old groMt
than a few years in any case
gging interests continue ·
Liot Sound of British Colun::-
biome should be preserve
' Forests
d only in the northern herri-
:::xtends as a broad band ce rr-
est extends across norther::
. and the northern islands 0~
from coastal Alaska to tt::
inous forest in the wester::
iferous biome extends dow::.
o the highlands of Mexicc
the Appalachian Moun taiL.;;
s forest are found at high e~
Himalayas. The total area c:
~ n estimated that the borec__
:st cover on the planet.
ue found in regions with =.
:. Mean annual precipitatio::.
precipitation is lowest in tl:::
mild , with average July terr.-
1ntains in the Yukon Territory c"
1e. All of the coniferous trees ' -
•ruce (Picea mariana).
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 179
peratures of 10 to 20° C. Winters are generally cold with average January temper-
atures reaching as low as - 50° C in central Siberia. The seasonal distribution of
precipitation is regionally variable. Central Alaska receives its greatest amount of
precipitation in the summer months, while Scandinavia has its highest precipita-
tion in the winter.
Why are the boreal and coniferous montane forest biomes restricted to the
northern hemisphere? A similar broad band of cool summer and cold winter cli-
mate does not occur in the southern hemisphere owing to the absence of conti-
nental land in the vicinity of 60° S latitude. The southern hemisphere does not
have conifer-dominated montane forest. For example, eucalyptus trees are found
at high elevations of the Snowy Mountains in southern Australia, and southern
beech forests are found in the mountains of New Zealand and Patagonia. In trop-
ical Africa, woodlands are formed on the subalpine zone of Mount Kilimanjaro by
a relative of the ragwort, Senecio keniodendron, and a relative of the lobelia,
Lobelia keniensis. These remarkable landscapes bear no resemblance to the conif-
erous forests developed on similar cool alpine sites in the northern hemisphere.
Spodosols are often found in coniferous forest regions, particularly in the
boreal zone. In addition, alfisols may be found in southern portions of the boreal
biome and in some montane regions. Histosols are common in the boreal zone
and on poorly drained sites in other portions of the biome. Alpine regions often
have poorly developed rocky soils, which are sometimes referred to as regosols.
High-latitude sites and high elevations sites may have permafrost. In general, the
southern border of continuous permafrost lies at the northern edge of the boreal
forest zone in North America. The boreal forest here is underlain by discontinu-
ous to sporadic permafrost. In Siberia continuous permafrost underlies almost all
of the boreal biome.
The coniferous forests in North America and Europe are generally dominated
by evergreen species from the genera pine (Pinus spp.), spruce (Picea spp.), and fir
(Abies spp.). In Siberia, much of the boreal zone is dominated by the deciduous
conifer species, Siberian larch (Larix sibirica), and dahurian larch (Larix dahurica).
These trees, though related to the evergreen conifer genera, lose all of their needles
during the winter. It is possible that this adaptation allows them to dominate in
Siberia because of the extremely cold winter temperatures experienced in this con-
tinental region. Exposed needles of evergreen species are susceptible to frost dam-
age and desiccation in the extreme cold of the northern Siberian winter.
The structure of the conifer forests is relatively simple. There is a tree layer
dominated by conifers, and the conifer canopy is generally no more than 30 to 40
m in height, and often much less. In the case of the giant sequoia (Sequoiaden-
dron giganteum) of California, the dominant trees can reach heights of close to
100m. The conifers from this biome are the oldest trees in the world. Bristlecone
pines (Pinus aristata) from the White Mountains of California reach ages of over
4000 years. Other species such as giant sequoia and limber pine (Pinus flexilis)
can reach ages of 1000 to 2000 years. Ages of 500 years are not uncommon for
many other coniferous tree species.
Recently burned sites and areas along rivers are sometimes dominated by
deciduous broadleaf genera such as birch (Betula spp.) or poplar (Populus) . The
forest canopy is closed in the southern and middle portions of the boreal biome.
Similarly, mid-elevation sites in coniferous montane forests generally have closed
canopies. Toward the northern limits of the boreal forest and the upper treeline
in the mountains, the forest canopy begins to open. The trees also become smaller
180 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

in stature. At their extreme limits, trees are often restricted to protected site~ rare at the nor
where they grow as low shrubs called krummholtz. The dominance of the conife ~ low rates of e'
biome by a few families of trees with the same physiognomy and stand structure years between
produces very similar landscapes in widely separated regions such as Siberia, Fin- breaks such as
land, and Canada. Some refer to the conifer forests, particularly the boreal por- areas with san
tion, as monotonous. intervals as sh•
The underlying vegetation in the coniferous forest often includes evergree with their var
shrubs such as many members of the heath family (Ericaceae) and junipe~ which patches
(Juniperus spp.). Deciduous shrubs such as willow (Salix spp.), alder (Alnus spp. scape. For exm
and shrub birch are also common. In addition, stunted saplings of the dominan: old, while 75o/c
conifers may be present. In many cases, however, there is little regeneration o: boreal region i
dominant trees beneath their own canopies. Due to dense shade and acidic needle- Many pla
litter, the ground cover can be relatively sparse and consists of a few herb speci e~ lodgepole pine
In some regions, the ground surface is covered by feather moss (Pleurozium schre- cones that ope
beri), peat moss (Sphagnum spp.), and other mosses. Black spruce (Picea mariana burned soils an•
and larch (Larix laricina) stands in the North American boreal forest are ofte;:, grow. Aspen (P.
found on thick organic soils composed of peat mosses. Similarly, the larch forests : species, resprol
Siberia often grow on layers of moss that can be up to 10 m thick. Peat is "mined- suga menziesii)
in many parts of the world as a fuel for fire and a soil additive. The bags of pee:: protects the vas
moss that are purchased in local nurseries generally come from the boreal forest. I.;: The impa
portions of eastern North America, thick growths of Cladonia lichens form a d~ to region. In Ia
yellow crust on the ground. In some dry conifer forests, such as those at lower ele- Canada, there :
vations in the Southwest, frequent burning and lack of soil moisture preclu e mal. In contras
understory growth. Here the ground can be relatively bare, or in some cases it ca:. and pasture Ian
support sparse grass cover if the canopy is open enough to allow sufficient light. are carefully m
Fire is the most common agent of vegetation disturbance in coniferous forest for timbc
forests, although wind, insects, and permafrost activity can also be importar:: boreal forest tb
causes of local to regional vegetation disturbance. It has been estimated that u: ing population
to 2% of the upland boreal forest burns each year. Even in areas where fire su P,. North America
pression has been widely practiced, fires in the boreal zone are numerous an.: for recreational
burn very large areas. The most frequent fires in coniferous forests are very sma.L. to high rates c
consuming a few tens of hectares. However, it is the infrequent large fires that au United States a
often responsible for most of the area burned. In the Canadian boreal forest , fir : low-intensity fir
of >200 ha accounted for only 3% of the total number of fires but 97% of the Such areas rna)
total area. In fact, fires of >10,000 ha are responsible for 90% of the total are=. control. In addi
burned annually in Canada. shrub species 1
Two important factors influence the number and size of conifer fires: higi: regions. For ex::
rates of ignition events and high rates of spread. Fire ignition may be caused b:· ern United Stat
human activity and lightning. Despite the frequency of human-ignited fires, ligh - established spn
ning-caused fires account for 90% of the area burned in the boreal forest. As th- ground fires tha
occurrence of summer frontal activity and lightning storms decreases towar and restricted <
treeline, so too does the number of ignition events. Conifer-dominated stands fuel-thinning ef
provide high levels of fuel in the form of dry litter, fine needles, twigs, bark flake problems. How•
and resinous products. In addition, the conical shape of conifers promotes cro\\ forests to returr
fires and rapid spread. Understory shrubs and lichens are highly flammable an ·
carry fire between trees. Tundra
The average frequency of fires in the biome varies according to climatic set-
ting and site type. Low-elevation ponderosa pine (Pinus ponderosa) stands in the The final biome
southwestern United States experience very dry summers and frequent lighting nated vegetatio
storms. They have ground fires every few years. In contrast, lightning storms are Tundra covers ~

~estrictedto protected sites
he dominance of the conifer
ognomy and stand structure
regions such as Siberia, Fin-
particularly the boreal por-
est often includes evergreen
y (Ericaceae) and junipers
zlix spp. ), alder (Alnus spp. ).
ed saplings of the dominan:
tere is little regeneration o:·
~nse shade and acidic needle
)nsists of a few herb species.
her moss (Pleurozium schre-
llack spruce (Picea marian a
ican boreal forest are ofte;::
Similarly, the larch forests o:
, 10m thick. Peat is "mined-
il additive. The bags of pea:
me from the boreal forest. I.::
Cladonia lichens form a dr.
s, such as those at lower ele-
k of soil moisture preclude
bare, or in some cases it ca.;:_
~h to allow sufficient light.
t disturbance in coniferoe-
vity can also be importar.:
has been estimated that u:
ven in areas where fire su r-
eal zone are numerous an.:.
erous forests are very smaL
1frequent large fires that ar~
Canadian boreal forest, fir es
ber of fires but 97% of t h~
e for 90% of the total are::.
nd size of conifer fires: hi!!]::
e ignition may be caused b:·
)f human-ignited fires, ligh:-
1in the boreal forest. As tt .:
1g storms decreases towar:::
:. Conifer-dominated stan --'-
1e needles, twigs, bark flak es.
of conifers promotes crow::.
ts are highly flammable an-
[es according to climatic se:-
ms ponderosa) stands in tl:::e
1mers and frequent light in~
mtrast, lightning storms ar:e
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 181
rare at the northern treeline of the boreal forest, and cool temperatures produce
low rates of evaporation and keep the landscape moist. It may be thousands of
years between fires at treeline. In the central boreal forest, areas with natural fire
breaks such as lakes and sharp topography may not burn for 100 to 400 years. Dry
areas with sandy soils and little topographic relief can have average fire-return
intervals as short as 28 years. In many regions, the high number of fires coupled
with their variability in size imparts a mosaic pattern to coniferous forest in
which patches of various ages, some large and some quite small, typify the land-
scape. For example, about 10% of the boreal forest stands are less than 20 years
old, while 75% are between 20 and 75 years old. The old growth forest in the
boreal region is very small.
Many plants of the coniferous forest are adapted to disturbance by fire. The
lodgepole pine (Pinus contorta) of western North America produces serotinous
cones that open following heating. The seedlings of this species thrive on freshly
burned soils and require the high insolation provided by an open canopy in order to
grow. Aspen (Populus tremuloides) and birch trees, as well as many other deciduous
species, resprout vigorously following fires. Conifers such as Douglas fir (Pseudot-
suga menziesii), ponderosa pine, and many larches have extremely thick bark that
protects the vascular tissue of the mature tree from heat during ground fires.
The impact of human activity on the coniferous forests varies from region
to region. In large areas of the boreal forest, particularly in Russia, Alaska, and
Canada, there are few settlements or roads, and so human impact has been mini-
mal. In contrast, humans have used the forests of the Alps for timber, firewood,
and pasture land for thousands of years. Many of the conifer stands in this region
are carefully managed resources. In Canada and Russia, harvesting of the boreal
forest for timber and paper is not insignificant. The average area of the Canadian
boreal forest that is harvested each year is roughly 0.7 to 0.8 million ha. Increas-
ing population size has caused increased pressure on some forested regions. In
North America, montane coniferous forests are often seen as prime real estate
for recreational development. In Nepal, increased demands for fuel wood has led
to high rates of deforestation. Fire management practices, particularly in the
United States and Canada over the past century, have led to a decrease in small,
low-intensity fires in many areas. This has allowed fuels to build up to high levels.
Such areas may experience large fires that will be difficult, if not impossible, to
control. In addition, controls on natural fires have allowed fire-sensitive tree and
shrub species to increase at the expense of fire-resistant species in certain
regions. For example, many old growth ponderosa pine stands in the southwest-
ern United States are becoming closed canopied or are being invaded by newly
established spruce and fir. Prior to fire control measures, frequent low-intensity
ground fires that killed seedlings and small trees kept the canopies of pines open
and restricted other species from occupying these sites. Controlled burns and
fuel-thinning efforts have been implemented in some areas to cope with these
problems. However, even with such efforts it can take many decades for conifer
forests to return to their natural state.
Tundra
The final biome we will consider is the tundra. This sparse herb- and shrub-domi-
nated vegetation is found beyond treeline in the arctic and on high mountains.
Tundra covers some 8 million sq km of the earth's surface. The largest areas of
182 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

conterminous tundra are found in arctic portions of Eurasia and North America_
Large areas of continuous alpine tundra are found in the Rocky Mountains, th
Andes, and the Himalayas. Antarctica is largely covered by ice and supports little
vegetation. The vascular plant flora of that continent is dominated by only twc
species: a grass called Deschampsia antarctica and a small cushion-form plan:
called Columbanthus quitensis.
The key climatic characteristic of the tundra biome is low temperature. Mear.
annual temperatures in the biome are less than -15.0° C. Average temperatures o::
the warmest month of the year are generally less than 10° C, whereas average tem-
peratures during the coldest month of the year are typically less than -30° C. Th
growing season is quite short, averaging under three months. Diurnal variations ir.
temperature during the growing season in arctic regions are relatively low owing tc
the long hours of daylight in the summer. In contrast, during the summer, alpin -
regions can experience relatively warm days and below-freezing temperatures a:
night. Radiation is also higher during the day in high-elevation alpine regions..
Alpine regions near the equator have little seasonal variation in temperature. Pre-
cipitation in the arctic is generally quite low, ranging from about 40 em to less thar.
FIGURE 6.23
10 em per year. In contrast, some alpine tundra regions, such as the coastal moun- Canada . The low
tains of Washington and British Columbia, can receive well over 200 em of precipi- and protection b
tation each year. foreground is pu
The very low temperatures of the tundra biome restrict soil development. lL
many areas, the substrate consists of regolith with low organic content and ]O\\
amounts of available nutrients. In the very high arctic, vegetation cover ofteL 10 em across a
increases in the vicinity of the bones of dead animals due to the increased avail- and the cushic
ability of nutrients. Hisotosols are found in some regions and can be several meter5 close to the soi
in depth. Permafrost is continuous in the arctic and in very high alpine regions. insulated frorr
Areas such as the Rocky Mountains may have sporadic permafrost. Frost heavin§: plants have he1
and other cryogenic processes associated with permafrost can cause the develop- head always f
ment of sorted stone polygons in regolithic areas and small hummocks and depres- pollen, ovule,
sions in histosol-dominated sites. The patterned ground surface in such regions ~ flower is benet
often very active and important in determining local vegetation cover. Almost a
The structure of tundra vegetation is extremely simple (Fig. 6.23). In south- annual plants t
ern portions of the arctic and at lower elevations in the mountains, small shru : Most upright s
such as willow (Salix spp.), shrub birch (Betula nana), alder (Alnus spp.), an · plants such as <
heaths form a canopy that is up to 2 m tall. The ground cover in such areas and dryas (D
includes sedges, grasses, diverse herbs, and mosses. This vegetation is sometime: demand less ti
referred to as low arctic tundra. Further north and at higher elevations, the shrut- resources duri
cover becomes sparser, and the shrubs become more prostrate. Areas of poorh must produce
drained soils are often dominated by the sedge species called cotton grass (Erio- greens are pro
phorum vaginatum) . Patches of unvegetated ground become increasingly com- The stan•
mon away from the treeline zone. In the far north and at very high elevations. some 5 billion ·
vegetation cover is extremely sparse. In general, the climate is very cold, precipi- entire arctic tl
tation is low, and soil nutrients are very limited. In the arctic, such areas ar- species of flow
referred to as polar desert or high arctic tundra. In some cases, vascular plants ca L tinents and in 1
only survive in moist microsites in valleys, along wetlands, and adjacent to sum- of flowering r
mer snow fields. Only a few very small prostrate plants, lichens, and mosses exis: known. In con1
in the polar desert. The vascular plants often include low creeping species such as Canada has les
arctic willow (Salix arctica), which is related to willow trees but grows as a lO\\ arctic and in al
woody plant that rises only a few centimeters above the ground. Other plant the plants faun
such as the purple saxifrage (Saxifraga oppositifolia), grow as compact cushions related to arcti

=<:urasia and North America.
1 the Rocky Mountains, the
·ed by ice and supports little
t is dominated by only twc
a small cushion-form plan:
ne is low temperature. Mea:.
C. Average temperatures o:"
10° C, whereas average tem-
pically less than -30° C. The
1onths. Diurnal variations i=.
ts are relatively low owing tc
, during the summer, alpine
)w-freezing temperatures a:
gh-elevation alpine regions.
uiation in temperature. Pre-
·om about 40 em to less tha.:.
ts, such as the coastal mour;-
well over 200 em of precip:-
restrict soil development. ·-
>w organic content and lo
~tic, vegetation cover ofte;:
: due to the increased ava.L-
ns and can be several mete"
in very high alpine regions.
ic permafrost. Frost heavin~
frost can cause the develor-
mall hummocks and depr e ~­
ld surface in such regions ~
egetation cover.
simple (Fig. 6.23). In sout t-
:he mountains, small shru ~
a), alder (Alnus spp.), an.:.
:round cover in such ar e~
his vegetation is sometimQ
higher elevations, the shru::-
: prostrate. Areas of poor!:
~s called cotton grass (£rio-
become increasingly corr:.-
md at very high elevations.
:limate is very cold, precip:-
L the arctic, such areas ar::
ne cases, vascular plants ca.:.
lands, and adjacent to sum-
ts, lichens, and mosses e xi ~:
ow creeping species such ~
'w trees but grows as a lov
~ the ground. Other plan E.
, grow as compact cushio ~
ECOLOGICAL EQUIVALENTS, LIFE ZONES, AND THE BIOMES 183
FIGURE 6.23 The short vegetation cover of the arctic tundra near Churchill, Manitoba, in
Canada. The low stature of the plants provides them with optimum warmth in the summer
and protection beneath snow cover in the winter. The flowering cushion plant in the
foreground is purple saxifrage (Saxifraga oppositifolia).
10 em across and a few centimeters high. The low stature of the prostrate plants
and the cushion form allows plants to grow in a relatively warm microclimate
close to the soil surface (Fig. 6.24). In addition, low stature allows the plants to be
insulated from cold winter temperatures by a covering of snow. Some tundra
plants have heliotropic flowers that rotate as the stems elongate so that the flower
head always faces the sun. Heliotropism keeps the flower warm and aids in
pollen, ovule, and seed production. In addition, the warm microclimate of the
flower is beneficial for insect pollinators.
Almost all tundra plants are perennials. There is simply insufficient time for
annual plants to germinate, mature, and set seed during the short growing season.
Most upright shrubs in the tundra are deciduous, but many of the low prostrate
plants such as crowberry (Empetrum nigrum), arctic heather (Cassiope tetrgona),
and dryas (Dryas integrifolia) are evergreen. Plants with persistent leaves
demand less time to reach maximum rates of photosynthesis and require fewer
resources during the short summer than is the case for deciduous plants, which
must produce new leaves at the start of each summer. The leaves of these ever-
greens are protected beneath snow cover during the winter.
The standing biomass of the world's tundra is extremely low, consisting of
some 5 billion tons. The species diversity of tundra flora is also extremely low. The
entire arctic tundra region of North America and Eurasia has only about 900
species of flowering plants. Many of the same species are found on all three con-
tinents and in the floras of adjacent alpine regions. The highest regional diversity
of flowering plants is found in Alaska where approximately 600 species are
known. In contrast, the flowering plant flora of the northernmost arctic islands of
Canada has less than 100 species. Some plants, such as crowberry, are found in the
arctic and in alpine tundras of Eurasia and South America. In general, however,
the plants found on alpine tundra sites in the southern hemisphere are not closely
related to arctic tundra species. For example, the prostrate shrub Dracophyllum
184 CHAPTER 6 COMMUNITIES, FORMATIONS, AND BIOMES

YWORDSAND

unity
- unity type
- · ance type
..:::... .Dgical equivalent

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FIGURE 6.24 The microclimatic conditions of the arctic tundra in northern Sweden
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environmental pressure from human activity.

mdra in northern Sweden
75). The thermal advantage to
e common at alpine tundr.::.
in the northern hemispher·
,f the tundra biome becaus.:
eindeer herding is practice.::
and trampling have cause.::
as the Yamal Peninsula c:
the alpine meadows of tt.:
North and South Americc_
in many tundra regions. Fe:
n the world lie in the Yamc.. .
oil have been discovered i=
1ds have recently been d ~­
xtract metals such as go! -
number of arctic and alpir.::
1ns from smoke stacks ha\·;;-
sia. Buildings and road co;:-
st soils. A rut created in pe:--
be apparent for decades. :.:::.
the permafrost and the s u e~
called thermokarst erosior..
tundra is facing increas ir.~
KEY WORDS AND CONCEPTS
_-\.Jpha diversity Ecotone
_-\ssociation Formation
3eta diversity Growth form
3iome Individualistic community
~ o mmunity concept
~o mmunity type Life form
Jominance type Life zone
~co logical equivalent Old growth forest
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Change in North America. Association of
-~e rican Geographers, Washington, DC.
:=.::e r, H . (1985). Vegetation of the Earth, 3rd
-=dition. Springer-Verlag, New York.
:-2\·er, J. E. (1968). Prairie Plants and Their
~nv ironment:A Fifity Year Study in the Mid-
, est. University of Nebraska, Lincoln.
Whitmore, T. C. (1990). An Introduction to
Tropical Rainforests. Clarendon Press,
Oxford.
Whittaker, R. H . (1960) . Vegetation of the
Siskiyou Mountains, Oregon and California.
Ecological Monographs 30, 279-338.
Whittaker, R. H. (1975). Communities and Ecosys-
tems, 2nd edition. Macmillan, New York.
Whittaker, R. H . and Likens, G. E. (1975). The
biosphere and Man. In Primary Productivity
of the Biosphere (ed. H . Lieth and R. H . Whit-
taker), pp. 305-328. Springer, New York.
Wilson, E. 0. (1988). Biodiversity, p. 521.
National Academy Press, Washington, DC.
Wolf, J. J., and Mast, J. N. (1998). Fire history of
mixed conifer forests of the North Rim,
Grand Canyon National Park, Arizona.
Physical Geography 19, 1-14.
Woodell, S. R. J., Mooney, H. A., and Hill, A. J.
(1969). The behaviour of Larrea divaricata
(creosote bush) in reponse to rainfall in Cali-
fornia. Journal of Ecology 57, 37-44.
Woodward, F. I. (1987). Climate & Plant
Distribution. Cambridge University Press,
Cambridge.

0) . Ecology, W. H. Freeman .

1. Abiotic controls on prima::

d nutrient cycles in North
.lands. In Concepts of
1ogy ( ed. L. R. Pomeroy anc

'· 115-129. Springer-Verlag.

'·Postglacial Vegetation of
idge University Press,

r-Ellis, J. G. K., Johansson, L

~yden , B. E ., and Sonesson. :'\ ~
n (Abisko) Sweden. Ecologi-
~dish National Science
il20, 265-294.
). Gap dynamics in an Ohio
st and speculations on the
>turbance. Canadian Journ al
ch 20, 632-641.
yce, A. T. (1988). Deforesta-
ends in Costa Rica. Biotrop-

). Canada's Vegetation: A
ve. MeGill-Queen 's Universi:-
and Kingston.
'3d/7 a NV '3WI.l
II ~H~d
CHAPTER ]

CHANGING CONTINENTS
AND CLIMATES

The Latin term terra firma means solid earth, and indeed that is how we generally
think about the continents-solid and unmoving. However, geological and pale-
ontological evidence tells us that the continents and oceans have shifted and
changed their geographic positions over time. For example, if we rearranged the
continents to look as they did some 200 million years ago, the cities of Boston,
Massachusetts, and Lisbon, Portugal, would have almost been neighboring com-
munities with no Atlantic Ocean between them. Although changes in the geogra-
phy of the continents and oceans have occurred at almost imperceptibly slow
rates over millions of years, these changes explain many aspects of the modern
distributions of species. Thus, it is important for biogeographers to understand
and consider the past changes in the geography of the earth when trying to com-
prehend present-day plant and animal distributions.
In addition to the slow movement of continents and oceans, there have
been large-scale changes in the earth's climate. Many of these climatic changes
have occurred on much shorter time scales. In the last 2 million years, climatic
changes have generated repeated episodes of growth and retreat by glaciers at
high latitudes and high elevations. Standing in the sweltering summer heat and
humidity of Chicago, it seems impossible that the region was covered by thick
glacial ice only 20,000 years ago. The climatic and geographic changes that
accompanied these glacial cycles have had a profound impact on the modern dis-
tributions of life. Over the past 10,000 years there have been changes in climate
that, though smaller in magnitude than the glacial cycles, have also affected the
modern distributions of species. Finally, we are now standing on the edge of a
unique event in the climatic and biogeographic history of the earth. For the first
time, human activity has the potential of changing the climate of the planet.
Biogeographers recognize that the modern distribution of life reflects both
present-day environmental conditions and the past history of the planet. Future
climatic changes will also affect the distribution of life, perhaps within our life
time. In this chapter we will review the processes and patterns of past changes in
the earth's geography and climate. We will then look at the potential role of
human activity in altering the future climate of the planet. To begin, however, we
should review a little bit of basic geology.

FE AND THE GEOLOGIC TIME SCALE

For most people, events that occurred even a few decades ago seem like ancient
history. Events that go back thousands of years appear extremely remote, and the
concept of dealing with changes and processes that occur over millions of years
191
192 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

seems overwhelming. Biogeographers, however, know that the modern distrih_- higher strata. Th
tions of species are strongly influenced by events that occurred thousands to · - assumed that tht
lions of years ago. Geologists, paleontologists, and biogeographers must routine:_ past were the sar
consider events of the distant past and deal with huge units of time. The geolo · of erosion and
time scale is an internationally accepted system that serves as the standard m e~ observed today.
of subdividing the many millions of years of earth's history. The geologic tin:::: observations of 1
scale has its origins in the nineteenth century when the sciences of geology ar...: ica, the geologic
paleontology were rapidly developing. twentieth centur
The exposures that geologists study consist of layers of rock with differe : Initially, ge
compositions and textures. These layers are referred to as strata, and their anah - ferent strata rep
sis and classification is known as stratigraphy. Rocks can be classified into thr~= teenth-century ~
broad categories: igneous rocks are formed by the cooling and solidification c: realized that the
molten material called magma; intrusive igneous rocks are formed when tt:: ferent strata to b
cooling occurs deep underground; and extrusive igneous rocks are formed whe:::. of the nineteent
cooling occurs at the surface. Examples of intrusive and extrusive igneous roc -- eluded that the e
include granite and basalt. Over time, physical, chemical, and biological process .: tieth century, so1
cause the igneous rocks to break down into fine particles and soluble chemicals. of unstable isot<
This process is referred to as weathering. Weathered material is transported awa:- age of the earth.
by water, wind, and gravity. The transport of these weathered materials is ca11e;:: to decay to their
erosion. Sedimentary rocks are formed when the weathered material accum - years for half o1
lates. The accumulating substances might include sand grains that are deposite · decay to calciun
into lakes by rivers and form sandstone, or calcium carbonate ooze precipitatec radioactive elem
in the ocean that forms limestone. The accumulated material is buried and ove::- By examining th
long periods of time becomes compacted and cemented into solid rock. The phys- the age of the cr
ical and chemical process by which sediments are turned into rock is called lithi- cons, which are L
fication. Sandstone and limestone are just two examples of the wide range o: years older than
sedimentary rock. Sometimes the nature of the sedimentary rock provides clues we can affix age
to the climate at the time of its formation. For example, glaciers develop unde::- scale is based or
very cold conditions and by eroding underlying rock and soil produce sedimen- and fossils, with
tary deposits called tillites (also called glacial tills), which contain a poorly sorte - encapsulates, in :
mixture of clay, silts, sand, and rock fragments. Metamorphic rocks are formec The model
when igneous or sedimentary rocks are altered by high temperatures and pre - subdivided into <
sures. Marble is a metamorphic rock that is formed when limestone is exposed to 7.1). The names
high heat and pressure. Geologic strata consist of bedded layers of differem Some, such as tht
igneous, sedimentary, and metamorphic rock. diagnostic depm
Some sedimentary strata contain the remains of plants and animals. In drawn from othE
many cases, these remains were altered by chemical processes so that the\ the Silurian Peri
became lithified. Such lithified remains are called fossils. Some researchers co~­ an ancient tribe
sider all preserved organic remains to be fossils, whereas others contend that the We live in t
remains must be lithified. The presence of fossil remains of extinct animals in sed- era of the Phant
imentary rocks has been known for a long time. In ancient China it was though t complex forms c
that some of these remains were mythical beasts such as dragons. In Europe these have occurred d1
fossils were thought to be the remains of animals that had not been able to get to rapid proliferati<
Noah's Ark and were destroyed during the great flood described in the Judea- species is called
Christian Bible. ago. The ancesto
During the nineteenth century, geologists and paleontologists began to clas- the Cambrian Pt
sify strata on the basis of their physical characteristics and their fossil content. The Ordovician
The geologists made two assumptions. First, they assumed that strata that were the development
lower in a stratigraphic exposure were older than overlying strata. Thus, the fos- land plants, lan<
sils in lower strata were assumed to represent older forms of life than those in Silurian and De\

w that the modern distribu-
occurred thousands to mil-
,geographers must routine],-
~ units of time. The geologi
erves as the standard means
; history. The geologic time
the sciences of geology anc
ayers of rock with differen:
to as strata, and their anah--
can be classified into three
ooling and solidification o:'
Jcks are formed when the
ous rocks are formed whe:::.
md extrusive igneous roc~
;al, and biological processes
icles and soluble chemicals.
naterial is transported mvc.:
eathered materials is calle.:
eathered material accum c-
td grains that are deposit e.:
:arbonate ooze precipitate.:
material is buried and O\ - -
~d into solid rock. The ph y~­
ned into rock is called lir/::-
nples of the wide range c·
1entary rock provides clue':
pie, glaciers develop undc-
and soil produce sedime:::.-
tich contain a poorly son e~
zmorphic rocks are form e~
igh temperatures and prQ-
hen limestone is exposed z:
bedded layers of differe:::.
of plants and animals. ·-
~al processes so that the:
sils. Some researchers co:::.-
eas others contend that tt=
ns of extinct animals in sec-
tcient China it was thoug'-
as dragons. In Europe thes.::
had not been able to get : ~
od described in the Judec-
Lleontologists began to clas-
cs and their fossil cant e r.~
umed that strata that we:-:
~rlying strata. Thus, the fo-5-
forms of life than those ::.::
LIFE AND THE GEOLOGIC TIME SCALE 193
higher strata. This assumption is called the law of superposition. Second, they
assumed that the processes that weathered, eroded, and deposited rock in the
past were the same as those operative today. They therefore concluded that rates
of erosion and deposition in the past were no faster or slower than those
observed today. This assumption is called the law of uniformitarianism. From
observations of rock and fossil stratigraphy, mainly in Europe and North Amer-
ica, the geologic time scale was developed and refined during the nineteenth and
twentieth centuries.
Initially, geologists could only try to estimate the number of years that dif-
ferent strata represented and guess how old fossils were. Eighteenth- and nine-
teenth-century geologists such as James Hutton and Charles Lyell of Britain
realized that the earth must be very ancient in order for the rocks of so many dif-
ferent strata to be formed, weathered, eroded, deposited, and lithified. By the end
of the nineteenth century, geologists, paleontologists, and biogeographers con-
cluded that the earth must be several hundred million years old. During the twen-
tieth century, sophisticated means of dating rocks by using the radioactive decay
of unstable isotopes and elements have allowed more reliable estimates of the
age of the earth. Physicists have calculated how long it takes radioactive elements
to decay to their stable daughter elements. For example, it takes about 1.3 billion
years for half of the total amount of the radioactive element potassium 40 to
decay to calcium and argon. Physicists refer to the time needed for half of the
radioactive element present in a material to decay as the half-life of the element.
By examining the ratio of potassium 40 to calcium and argon in a mineral crystal,
the age of the crystal can be estimated. The oldest dated mineral crystals are zir-
cons, which are 4.1 billion years old. It is estimated that the earth is half a billion
years older than the age of these zircons. By using radiometric dating techniques,
we can affix ages to different strata and their fossils. The modern geologic time
scale is based on the classification of strata by their lithology, relative position,
and fossils, with ages provided by radiometric dating. The geologic time scale
encapsulates, in a very broad manner, the history of earth and its life.
The modern geologic time scale extends back over 4 billion years and is
subdivided into a hierarchy extending from eons to eras, periods, and epochs (Fig.
7.1). The names for the different periods can be quite confusing to the novice.
Some, such as the Pennsylvanian Period, are named for geographic regions where
diagnostic deposits of this age are found. Others such as the Tertiary Period are
drawn from other classification schemes that are no longer in use. Some, such as
the Silurian Period are even more obscure in origin. The Silurian is named after
an ancient tribe of Britain, the Silures.
We live in the Phanerozoic Eon, which began 570 million years ago. The first
era of the Phanerozoic is the Paleozoic, which is typified by the appearance of
complex forms of life. The appearance of complex, multicellular life appears to
have occurred during the Cambrian between 570 and 505 million years ago. The
rapid proliferation of multicellular exoskeleton (external hard covering) marine
species is called the Cambrian Explosion and peaked about 530 million years
ago. The ancestors of mollusks, crustaceans, and insects all appear in the oceans of
the Cambrian Period. The first marine vertebrate also appears in the Cambrian.
The Ordovician Period, extending from 505 to 438 million years ago, is noted for
the development of vertebrate saltwater and freshwater fish . The rise of primitive
land plants, land-dwelling invertebrates, and amphibians occurred during the
Silurian and Devonian periods about 438 to 360 million years ago. The oceans of
 "'
E
this time teem'
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c:: land plants wer

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s
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like plants wit!
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c:
<(
>-
<!) .g;"' gymnosperm tr
Silurian. By the
the land. By th(
of terrestrial v'
and Pennsylva1
.c ago, is characte
"%"' "' ancestor of the
~ ~ TheMes<
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~
.0
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Q; proliferation ar
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\
~ II> ern mammals <
.2:
Cii nance in the Tr
·c 'g
1ii
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~
~~ possessed somt
of dinosaurs, in
-o
0 ferns dominate
(.)

~ Cretaceous Per
·;;;; The curre
....."'0 by the extincti<
Q)

..,...,. + £ trial megafaun

C\11.0~
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0 One important
C\1 (")
c;; tiary Period an
(,)
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::-
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c:
Paleogene Peri
c."'
-o Neogene Perio
c:
"' adhere to the t
.!!2
The Quat
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and animals fo
c: c;;
....."' c: a:; the Holocene J
0

"'c.
Q) c"" tied by the pre:
c
....> ~
civilization and
c: c:
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tures of the H
"'0
::J c:: "'
·c: "'
·a. c: c::
c: c:: £
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'C
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c
our 10,000-yea
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~ i!="' "'c::c:: "' ·~ 'C
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(.)
--, 0.
& "'
~
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(,)
;: Charles I
"'
Q)
ai
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~ can be closely
E
(,)
·-0C l"'-
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record of clim<
Vl
--o He came to th
g 'i:0
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c Q)
.!:!~ .c c.
Q)
the general tir
·su~
= .2~
o= o- ii"'
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Q)
~

..
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c
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g=o
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~~
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periods of tim
However, duri
ll. r- Q)
w ~ widespread gt
cc&
:I ·- existed around
:l!OZOJ9U81.1d ":
u. .....
- 0
lion years age

194
 LIFE AND THE GEOLOGIC TIME SCALE 195

~ this time teemed with fish species and primitive sharks. The land plants likely
"'
a.
"0
started as semiaquatic algal seaweeds followed by primitive ferns. The first true
c:
land plants were only a few centimeters in height. By the end of the Silurian, tree-

=i (t
like plants with leaves over 10 em had long existed, and the ancestral form of
gymnosperm trees had arisen. Centipedes and spiders appeared on land in the
Silurian. By the middle Devonian, lobe-finned fish were making forays to feed on
the land. By the late Devonian, the first true amphibians appeared. Development
of terrestrial vegetation and land animals continued through the Mississippian
and Pennsylvanian periods. The Permian Period, some 286 to 245 million years
.s::; ago, is characterized by the appearance and development of true reptiles and the
~ "' ancestor of the crocodiles, caimans, and alligators.
~
.0
~
.0
Q) The Mesozoic Era began 245 million years ago and is best known for the

~
~
.!:
proliferation and dominance of dinosaur species. However, the ancestors of mod-
ern mammals also appeared at the same time that the dinosaurs rose to domi-
~
.E nance in the Triassic Period. Birds first appeared in the Jurassic Period between
;t

~
208 and 144 million years ago. A possible early ancestor of birds, Archaeopteryx,
~~ possessed some features of modern birds, such as feathers, but also had features
of dinosaurs, including its basic skeletal structure and teeth. Conifers, cycads, and
"0
0 ferns dominated the vegetation, but flowering plants also appeared by the early
u
~ Cretaceous Period, 144 million years ago.
·v; The current era is the Cenozoic. It began 65 million years ago and is typified
(/)

....0 by the extinction of the dinosaurs and the dominance of mammals in the terres-
<I>
+ £ trial megafauna. Flowering plants diversified and came to dominate the flora .
~
..,. ~
0
1'-
L()
0
0
co
....
0 One important group of flowering plants, the grasses, appeared in the mid-Ter-
"' ~
u tiary Period around 24 million years ago. We live in the Quaternary Period which
·a.
c began about 2 million years ago. In some revised versions of the geologic time
~
c:
scale, the Cenozoic, the Tertiary, and Quaternary periods are replaced by the
a."' Paleogene Period, which extends from 65 million to 24 million years ago, and the
"0
c:
Neogene Period, which extends from 24 million years ago to the present. We will
~~ ~ 2 1~1~1 2 1~1-t::
~ ~~~ ~ ~ j ~ ~ "'
:!!
adhere to the traditional Tertiary and Quaternary periods.
"'E The Quaternary is characterized by the appearance of the species of plants
c: and animals found living today, including our own species. We currently live in
."'0. =-
.::. the Holocene Epoch. The Holocene represents the past 10,000 years and is typi-
(/)
Q) fied by the presence of modern species of plants and animals. The rise of human
Cl.
c - civilization and the explosive growth of the human population are additional fea-
=-
Q)

c:
c: c:
£ tures of the Holocene. Despite the importance we place on human civilization,
"'
·.:: "'
·c::;
"'
~ £ our 10,000-year epoch represents but a brief moment in geologic time. Although
~
:i5
-~
"E
E "§ - we may think of the now extinct dinosaurs as failures, they did dominate the
0 "'
u
~
earth for a period of time that is 1000 times longer than the Holocene.
"'u
(/) ~

Charles Lyell recognized that since certain sedimentary rocks and fossils
Q)=
E-
·~ : can be closely related to past climatic conditions, it is possible to reconstruct a
u
"ii E
= record of climatic change over geologic time by carefully examining rock strata.
0
0Q) He came to the conclusion that the earth must have experienced a number of
c:
0: ~ large-scale swings in climate over its history. Continued work has documented
~
Q)
.!:.
1-
- the general timing and magnitude of such swings. On the broadest scale, long
E
~ .. -
,....
periods of time seem to have been dominated by generally warm conditions.
However, during certain intervals, there is evidence for cold temperatures and
II..
w - widespread glacial conditions. Within the Phanerozoic, such glacial climates
Cl:-
::l = existed around 450 million years ago during the late Ordovician, around 286 mil-
(!)
u:: lion years ago, at the boundary between the Pennsylvanian Period and the
-
196 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

Permian Period, and during the Pleistocene Epoch between 2 million and 10,00_
years ago. Within some of these periods of glaciation, we also find evidence c:
alternating warm climates and glacial conditions. The Pleistocene, for example. . :_
typified by long periods of glaciation in the high latitudes and mountains, a ~
other intervals when the climate is like that of today. What changes could ha\·=
occurred to the earth causing such dramatic shifts in climate over both long an.:
short time spans? This question will be explored in the next two sections of ths
chapter.

SHIFTING CONTINENTS

A casual look at a globe reveals a curious match between the eastern coastlin _
of North and South America with those of western Europe and Africa. It wouJ.:
seem that South America could fit nicely into the concave coastline of wester=
Africa, while northwestern Africa and Europe could be accommodated withi::.
the Caribbean Basin and adjacent portions of North America. This apparent :
between the Old and New World was noticed as early as 1598 by the cartograpbe:-
Abraham Ortelius who pointed out that such a fit between these coastlines su=-
gested a dislocation of the continents relative to each other. In 1858, the Amef.-
can Antonio Snider-Pelligrini published a detailed map showing how North an:.
South America may have at one time been joined to Europe and Africa. He p o~ FIGURE 7 .2 Th(
tulated that the biblical flood of Noah had somehow caused the continentc.. Wegener was the f
masses to be sundered apart. widely accepted dl
In addition to cartographic evidence, data from fossil finds, geological rna scientific expeditio
ping, and the distributions of living species also pointed to a closer connectio;::
between North America and Europe. For example, as early as the 1750s, tb.o
Compte de Buffon pointed out the similarity between fossils found in Irelan.: editions and pa]
and North America. Evidence of similarities in fossils and geology continued tc passing acquain1
accumulate. In 1872 another Frenchman, E. Reclus, argued for the existence of c. daughter of the •
closer connection between Europe and America on the basis of their similarity i;:: Wegener's
geological structure. In 1853 the biogeographer J.D. Hooker noted the similari ~ known as the tb
in plant species, genera, and families on the widely separated continents an" and 1929, he ou
islands of the southern hemisphere and concluded that the modern flora mus: First, he pointed
represent the remnants of a once continuous land vegetation that was broken u!= light rocks, or si;
by geological and climatic changes. By the early twentieth century, a number a:: dense basalts, or
geologists and paleontologists had speculated on the possibility that the conti- viscous mantle c
nental landmasses had been closer together in the geologic past. However, i: deposits in Braz
remained for the German astronomer, meteorologist and geophysicist, Alfre · cal areas were 01
Wegener, to marshal compelling evidence for continental movement and make c. formerly locatec
strong case for this phenomenon (Fig. 7.2). shape of con tine
Wegener warrants some description before we continue with our examina- For example, a F
tion of his ideas. He was born in Germany in 1880 and educated at the universi- is found in area~
ties of Heidelberg, Innsbruck, and Berlin. In 1906 he went to Greenland as an measurements c
expedition meteorologist for the Danish government. Following that, he taugb. 1923, and 1927 i1
meteorology and astronomy at Marburgh University in Germany. He wrote that m/yr. We now kn
his first ideas about continental movement arose in 1910 while he was looking at centimeters a ye
a geographic atlas and noticed the match between the Atlantic coastlines of the Based on t
Old and New Worlds. In 1912 he published his first account of his theory of the world's continen
Horizontal Displacement of the Continents. He would publish a number of book this landmass, wl
 SHIFTING CONTINENTS 197

:tween 2 million and 10,0()(

n, we also find evidence o:
Pleistocene, for example. is
itudes and mountains, an ·
·.What changes could ha\·c
climate over both long an ·
1e next two sections of th ~

veen the eastern coastlines

urope and Africa. It woul""
ncave coastline of wester::
I be accommodated withi;:
America. This apparent E:
1s 1598 by the cartograp h e~
tween these coastlines sug-
other. In 1858, the Amer.-
lp showing how North a n ~
~urope and Africa. He pos-
)W caused the continen ra.:
'ossil finds, geological rna
ted to a closer connectio;::
as early as the 1750s, the
~n fossils found in Irelan-=
. and geology continued tc
gued for the existence of .::
e basis of their similarity i=
:ooker noted the similarit'.
separated continents an "
1at the modern flora mus:
~tation that was broken u:
tieth century, a number o:
possibility that the conr:-
~eologic past. However. i:
t and geophysicist, Alfre ·
1tal movement and make c.
Jntinue with our examina-
educated at the universi-
went to Greenland as ar.
Following that, he taugb:
n Germany. He wrote tha:
10 while he was looking a:
Atlantic coastlines of th-
count of his theory of the
publish a number of boo'·
FIGURE 7.2 The German geophysicist and arctic explorer Alfred Wegener (1880-1930).
Wegener was the father of the modern theory of continental drift. His ideas were not
widely accepted during his lifetime. Wegener died on the Greenland ice cap during a
scientific expedition .
editions and papers on this topic between 1912 and 1929. He had a more than
passing acquaintance with contemporary biogeography: In 1913 he married the
daughter of the famous climatologist and biogeographer Koppen.
Wegener's ideas on the movement of the continents have come to be
known as the theory of continental drift. In a treatise published between 1912
and 1929, he outlined the following arguments in support of continental drift.
First, he pointed out that the crust of the continents was composed of relatively
light rocks, or sial. He suggested that the ocean floors were largely composed of
dense basalts, or sima. The lighter continental rocks could essentially float on the
viscous mantle of the earth and override oceanic crust. Second, Permian glacial
deposits in Brazil, South Africa, India, and Australia indicate that these subtropi-
cal areas were once glaciated, and this could only occur if those landmasses were
formerly located close to the polar regions (Fig. 7.3). Third, similarities in the
shape of continents, their geology, and fossils suggest closer proximity in the past.
For example, a Permian fossil flora that is typified by the fern genus Glossopteris
is found in areas of South America, Africa, and India (Fig. 7.4). Fourth, geodetic
measurements of the latitude and longitude of Greenland taken in 1823, 1870,
1923, and 1927 indicated a continuing westward movement of as much as 9 to 32
m/yr. We now know that this movement is real but occurs at a slower rate of a few
centimeters a year.
Based on the evidence he had marshaled, Wegener concluded that all of the
world's continents had once been joined in one huge landmass. He thought that
this landmass, which he called Pangaea , had existed until Permian-Pennsylvanian
198 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

times. The sout

0 Permo-Carboniferous Glaciati - Africa, Austral
(General Area)
while the nortl:
D Glossopteris distribution came to be call
Accordin.
Mesozoic. The ~
Africa that gre
had separated <
ory included Sf
absolute dates
edge about con
factory identifi1
on the mantle.'
coupled with o
tion, in drawin1
factual errors r•
Wegener'
FIGURE 7 .3 The distribution of Glossopteris fossils and the Permian-Carboniferous ter years were 1
glaciation as evident from tillites . The reconstructed configuration of South America, Afr icc criticized by th
Antarctica, Australia, New Zealand, India, and Madagascar during the late Permian is on continental
presented (after Windley, 1977). Geologists (A"
gists attacked 1
the South Afri·
and Gondwanc
the AAPG syn
theory for 25 )
matter for long
His body rema
Despite t
onward, new e'
continental dri
continents mm
1960s and 197
unearthed in J
India. The find
the past, and s
biogeographer
tions of many !
by a closer c<
Zealand in the
ioidea are fom
and New Zeal:
errors in geolo
tantly perhaps
to solve the rid
Much pr•
initiated by sul
FIGURE 7 .4 A fossil of a Glossopteris leaf from Permian-aged deposits in New South
the sea becam
Wales, Australia . Fossils of this land plant have been found in South America, Africa, India ,
sonar, which a
Australia, and Antarctica and provide evidence that all of these landmasses were joined as
to map the se
one continent in the Permian .

J Permo-Carboniferous Glaciatio-
(General Area)
~ Glossopteris distribution
\~\
.\\ .·
I I
,,/
he Permian-Carboniferous
ration of South America, Africc
luring the late Permian is
1ged deposits in New South
South America, Africa, ln dic
se landmasses were joined as
SHIFTING CONTINENTS 199
times. The southern portion of Pangaea, which included modern South America,
Africa, Australia, Antarctica and India, came to be known as Gondwanaland,
while the northern area comprised of North America, Europe, and most of Asia
came to be called Laurasia.
According to Wegener, the supercontinent of Pangaea broke apart in the
Mesozoic. The Atlantic Ocean formed as a great rift between South America and
Africa that grew northward. Wegener, thought that Europe and North America
had separated only in the last few million years. The problems with Wegener's the-
ory included sparse information on the geology of the deep oceans, a scarcity of
absolute dates and rates for many of the changes he envisioned, a lack of knowl-
edge about continental locations prior to the Pennsylvanian-Permian, and unsatis-
factory identification of the mechanism that caused the continents to move about
on the mantle. Wegener suggested that radioactive heating and flow of the mantle
coupled with other factors likely caused the continental blocks to move. In addi-
tion, in drawing together so many lines of evidence, Wegener made a number of
factual errors regarding geology and paleontology in his publications.
Wegener's theories were not greeted with universal acceptance, and his lat-
ter years were not happy ones. He and his theory of continental drift were harshly
criticized by the Anglo-American geological establishment. In 1926 a symposium
on continental drift was sponsored by the American Association of Petroleum
Geologists (AAPG), and a number of prominent American and British geolo-
gists attacked the theory. Wegener continued to have some supporters, such as
the South African geologist, Alexander du Toit, who coined the terms Laurasia
and Gondwana for the northern and southern continental masses. In many ways,
the AAPG symposium led to the almost complete disappearance of Wegener's
theory for 25 years. For Wegener himself, the shunning of his theory would not
matter for long. He died on the Greenland lee Cap during an expedition in 1930.
His body remains there still, entombed within the glacial ice.
Despite the widespread dismissal of Wegener and his theory, from the 1940s
onward, new evidence began to accumulate that would lead to the vindication of
continental drift. Some of the evidence simply supported the contention that the
continents must have been in closer proximity in the past. For example, in the
1960s and 1970s, fossils of an early Triassic reptile called Lystrosaurus were
unearthed in Antarctica. Fossils of this genus are also known from Africa and
India. The finds showed that Antarctica must have been considerably warmer in
the past, and suggest a closer connection to other continents. At the same time,
biogeographers such as P.J. Darlington were showing that the modern distribu-
tions of many genera and families of plants and animals could best be explained
by a closer connection between Africa, South America, Australia, and New
Zealand in the past. For example, freshwater fish species of the superfamily Galax-
ioidea are found only in southernmost South America, South Africa, Australia,
and New Zealand. Such information seemed to indicate that, despite his factual
errors in geology and paleontology, Wegener was on the right track. More impor-
tantly perhaps, geophysical evidence from the oceans and continents was helping
to solve the riddles of the history, rates, and causes of continental movement.
Much progress in oceanographic geophysics in the twentieth century was
initiated by submarine warfare in World War II. Conditions and features beneath
the sea became of great strategic interest. Work accelerated on devices such as
sonar, which allowed the detection of enemy submarines but could also be used
to map the sea floor. For example, the marine geologist H.H. Hess sailed on
200 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

American troopships across the Atlantic and used an echo sounder to chart '-= some regions, c
sea floor. Once maps of the ocean floor were completed, it was clear that the ::: gins. Deep-sea
between the submerged continental margins of areas such as South America a~ .: and most inten
Africa was even better than the fit that was seen when the modern coastlin:::. less than about
were compared. Over the 1940s and 1950s, it was discovered that a global syste= rocks, which ca
of midoceanic ridges bisected the South Pacific, Atlantic, and Indian oceans (Fi; ceanic ridges ar
7.5). These ridges are centers of volcanic activity and the emergence of la\·:o. Paleo mag
Oceanic trenches, several kilometers deep, are found near the margins of co n ~­ in the postwar }
nents. Volcanic activity and earthquakes are common near the oceanic trench :. When igneous
The famous Ring of Fire, the zone of intense volcanic and earthquake activi _ earth's magneti
along the edges of the Pacific Ocean, is associated with oceanic trenches arou - field. Once the
the Pacific. In addition, flat-topped mountains called guyots lie beneath the sea :.::. magnetization 1
that is formed t
much like a co
axis toward the
Eurasian
igneous rock, c~
Plate .) ure its remnant
ferent from th
movement of t
geometric anal:
tion of the lan<
have shown hu!
the past billion
lion years ago t
An impm
polarity. If the 1
compasses poir
they occur ove1
-Areas of
Plate live in a ,geom;
convergence 1--~;::,....._,
period began .
=Areas of
divergence
reversed most 1
Age (millions of years) (Ridges) and this period
2 1 0 1 2 of ocean basal1
ridges and that
Midoceanic ridge image of the ot
ing the Bruhne
earlier magneti
symmetrical on
Taken to!
---,r---
Mantle drag )
oceans over th<

I
nents are less d
/ \ Mantle
Mantle plume
evidence show~
during geologi<
Slab pull Fourth, the age
midoceanic rid
FIGURE 7. 5 The earth's plates, zones of spreading and collision (after Birkland and
continents had
Larson, 1989), and the Hawaiian Islands associated with the Pacific hot spot. Th e insert
anism that driv
shows a cross section of the earth at the mid-Atlantic Ridge . The symmetrical magnetic
band ing found on both sides of the ridge is indicated , as is the general paleomagnetic
The mod
stratigraphy for the past 160 million years. The factors contributing to plate movement tectonics. Geo1
are noted. mately driven

echo sounder to chart the
ed, it was clear that the fi:
uch as South America an
en the modern coastlines
vered that a global system
ic, and Indian oceans (Fig.
d the emergence of lava.
near the margins of conti-
near the oceanic trenche
c and earthquake activi .
1 oceanic trenches aroun ·
ryots lie beneath the sea i.;::_
Eu rasian
Plate .)
r-
-Areas of
convergence I 1vz:.,
=Areas of
divergence
(Ridges)
isi o n (after Bi rkland and
acific hot spot. The insert
he symm etrical magnetic
gene ral paleomagnetic
uting to plate movement
SHIFTING CONTINENTS 201
some regions, often being found at greater depths toward the continental mar-
gins. Deep-sea cores revealed that the oceans were indeed underlain by basalt,
and most interestingly, radiometric dating showed that this basalt was generally
less than about 150 million years old. This is quite young compared to continental
rocks, which can date back billions of years. In addition, basalts near the mido-
ceanic ridges are generally younger than those found in the oceanic trenches.
Paleomagnetism is an important geophysical technique that was developed
in the postwar period and has been applied to both marine and terrestrial geology.
When igneous rocks that contain elements such as iron or titanium cool, the
earth's magnetic field causes mineral grains to become oriented to the magnetic
field. Once the rocks have solidified, this orientation is locked in. This remnant
magnetization of the rocks can be measured in the laboratory. An igneous rock
that is formed by a volcanic eruption today will have remnant magnetization very
much like a compass needle, with the minerals preferentially oriented along an
axis toward the north magnetic pole. Geophysicists can collect a sample of ancient
igneous rock, carefully record its modern position and orientation, and then meas-
ure its remnant magnetic properties. If the magnetic orientation of the rock is dif-
ferent from the modern compass orientation, they know that there has been
movement of the land surface relative to the earth's magnetic poles. By careful
geometric analysis and radiometric dating of the igneous rocks, the past orienta-
tion of the landmass can be reconstructed over millions of years. These studies
have shown huge changes in the latitude and longitude of continental regions over
the past billion years. For example, paleomagnetic studies show that some 300 mil-
lion years ago the continent of North America was located at the equator.
An important feature of the earth's magnetic field is its capacity to change
polarity. If the magnetic field were to change polarity today, we would find all our
compasses pointing south instead of north. The cause of such shifts is unclear, but
they occur over periods of hundreds of thousands to millions of years. Today we
live in a geomagnetic period called the Bruhnes Normal Chron. This magnetic
period began about 780,000 years ago. The magnetic field of the earth was
reversed most of the time between 2.58 million years ago and 780,000 years ago,
and this period is called the Matuyama Reversed Chron. Paleomagnetic studies
of ocean basalts reveal that these rocks are banded parallel to the midoceanic
ridges and that the paleomagnetic banding on each side of the ridge is a mirror
image of the other side (Fig. 7.5). The younger rocks near the ridges formed dur-
ing the Bruhnes Normal Chron; older rocks away from the ridges formed during
earlier magnetic chrons. The increasing age of the rocks away from the ridges is
symmetrical on both sides of the ridge.
Taken together, the geophysical data collected from the continents and the
oceans over the past 60 years lead to four important conclusions. First, the conti-
nents are less dense than the rock underlying the oceans. Second, paleomagnetic
evidence shows that the continents have indeed moved very significant distances
during geologic time. Third, the oceans are much younger than the continents.
Fourth, the age of the oceanic basalts increases symmetrically outward from the
midoceanic ridges. These observations support Wegener's contentions that the
continents had indeed moved, and they also provide some evidence of the mech-
anism that drives such movements.
The modern theory of how continental drift occurs is referred to as plate
tectonics. Geophysicists now believe that the movement of the continents is ulti-
mately driven by the dynamics of the inner earth. To understand the forces
202 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

behind continental movement, we need to know a little about the composition -. relative to tht
the earth. At the center of our planet is a dense inner core surrounded by a 1 along the pla
km-thick outer core of molten liquid iron and nickel. This in turn is surround=-: ranges from t
by a 2900-km-thick mantle of extremely hot silica, oxygen, iron, magnesium, a;:: collision zone
other minerals. Above this is a thin layer of cool, solid rock that is 5 km thi · collision has 1
under the oceans and 60 km thick beneath the continents. This outer layer the world. Tht
called the crust. The upper 50 to 125 km of rock in the mantle and crust is rel=- called orogen
tively hard and brittle rock and is referred to as the lithosphere. Beneath the li - - continents, bt
osphere, the hot and plastic rock of the mantle is called the asthenosph e - earthquakes,;
Volcanic activity occurs when the molten rock of the asthenosphere penetra t~ With ra<
the crust. The plastic conditions of the asthenosphere extend some 300 km in- - plate tectonic
the mantle. The remainder of the mantle is called the mesosphere. The continen_ the continent:
are less dense than the mantle and therefore float on top of the lithosphere a.G: past 250 milli
asthenosphere. The fluid nature of the underlying asthenosphere allows rt:: physical geog
plates to move across the surface of the earth. Several forces appear to pro - ters, many asi
such movements. species are ex
Part of the force causing the continents to move comes from ridge pus During
This occurs when convective flow in the mantle causes molten rock to come i -. Gondwanalar
the surface in areas such as the midoceanic ridge zones. The magma cools an.: tralia, India, S
forms new solid basalt. This new material pushes older basalt outward away froc. Portions of G
the ridge zone (Fig. 7.5). Hence, the basalt near the midoceanic ridges is young :- tion during th
than the basalt near the continental margins, and symmetrical paleomagneti;: can and Eura~
banding occurs parallel to the ridges. Ridge push also explains why guyots furth e:- region. The pl
from the midoceanic ridges are older and more eroded than those close to the with the Afri<
ridges. Over millions of years, this process causes sea floors to expand, separatin= Laurasia. By 1
continental plates on either side of the midoceanic ridge. Lateral convective flo,\ led to the end
of the mantle occurs below the crust, and friction between the crust and the flo w- the Asian por
ing mantle also causes continental plates to move. This process, somewhat like a and North A1
conveyor belt, is referred to as mantle drag. Spreading due to ridge push an verged and bt:
mantle drag is not always confined to midoceanic ridges. Rift zones, such as th supercontiner
Red Sea Basin, are areas where spreading has occurred in continental areas. The and was surro
Atlantic Ocean began as a rift zone in the ancient supercontinent of Pangaea. Geologi
Along the continental margins, the lower density of the continental rocks allows began to split
the landmasses to ride over the expanding oceanic crust. Such areas are calle ceanic ridge o
subduction zones because the oceanic basalts are subducted beneath the conti- plates during
nents and remelted. The deep oceanic trenches are formed in the subductio Asia and Nor
zones. The process of subduction leads to much friction and fracturing of the Siberia. This L
crust. This causes earthquakes and volcanic activity in the vicinity of the subduc- Rifting also b
tion zone and explains why the Ring of Fire exists around the Pacific Ocean. The ago. By 150 rr
process of subduction also helps pull the remaining oceanic crust toward the sub- severed the la
duction zone. This process is called slab pulL land. During
Today we can identify 16 major tectonic plates that are bordered by spread- American Pla
ing zones, subduction zones, transform zones, and collision zones. Transform Ranges, Sierr:
zones and collision zones are areas of convergence where two plates move later- formed by thE
ally against each other or collide (Fig. 7.5). Volcanic activity and earthquakes are By the I
common near spreading zones and subduction zones. Earthquakes also occur separated No
near strike-slip fault zones. The San Andreas Fault of California represents the America itsell
strike-slip zone between the Pacific and North American plates. This region is Rifting and st
well known for earthquakes, including the 1906 quake that destroyed much of America apar
San Francisco. At the San Andreas Fault, the Pacific Plate is moving northward from each otl

~ about the composition o:
:ore surrounded by a 1700...
This in turn is surroundec
·gen, iron, magnesium, an
id rock that is 5 km thic.b:
inents. This outer layer is
e mantle and crust is rela-
wsphere. Beneath the lith-
called the asthenospher
asthenosphere penetrates
extend some 300 km intc
1esosphere. The continents
top of the lithosphere anc
asthenosphere allows thc-
:~1 forces appear to pro pe ~
e comes from ridge push
:s molten rock to come tc
1es. The magma cools an-'
basalt outward away froc::.
joceanic ridges is younge::-
•mmetrical paleomagneti.:
xplains why guyots furth e::-
~d than those close to th;:
oors to expand, separat in~
se. Lateral convective flo
~en the crust and the flo"' -
; process, somewhat like c.
ng due to ridge push anc
~es. Rift zones, such as th;:
j in continental areas. The
tpercontinent of Pangaec.
e continental rocks allows
ust. Such areas are callec
jucted beneath the cont!-
:ormed in the subductio::.
ion and fracturing of th;:
the vicinity of the subduc-
md the Pacific Ocean. Th;:
anic crust toward the sub-
tt are bordered by spreaC.-
ollision zones. Transforc::.
~re two plates move late::--
:ivity and earthquakes arc-
. Earthquakes also occu::-
California represents the
ican plates. This region is
e that destroyed much o:
'late is moving northwar-
SHIFTING CONTINENTS 203
relative to the North American Plate. Deformation of rock strata by movement
along the plate and volcanism is responsible for the development of mountain
ranges from the west coast to the Rocky Mountains. One of the most dramatic
collision zones occurs where the Indian Plate contacts the Eurasian Plate. This
collision has produced the Himalayan Mountains, the tallest mountain range in
the world. These major geologic disturbances which result in mountain ranges are
called orogenies. Thus, plate tectonics not only explains the past movement of
continents, but also helps us understand phenomena such as volcanic activity,
earthquakes, and some episodes of mountain building.
With radiometric dating of rocks, paleomagnetism, and an understanding of
plate tectonic mechanisms, it is possible to reconstruct the relative positions of
the continents over the past 400 million years. The plate tectonic history of the
past 250 million years is crucial (Fig. 7.6) to an understanding of the modern
physical geography and biogeography of the earth. As we will see in later chap-
ters, many aspects of the distributions of plant and animal families, genera, and
species are explainable in light of past continental movements.
During much of the early to mid-Permian (286-250 million years ago),
Gondwanaland, made up of the plates that comprise modern Antarctica, Aus-
tralia, India, South America, and Africa, was located in the southern hemisphere.
Portions of Gondwanaland nearest the South Pole experienced intensive glacia-
tion during this time. The Laurasian landmasses, consisting of the North Ameri-
can and Eurasian plates, were located in the northern hemisphere and equatorial
region. The plate that makes up modern North America had collided and joined
with the African Plate about 350 million years ago, linking Gondwanaland and
Laurasia. By the early Jurassic, the northward movement of Gondwanaland had
led to the end of glaciation and more complete fusion with Laurasia. In addition,
the Asian portions of Laurasia had fused together and joined with the European
and North American plates. In this manner, all of the continental masses con-
verged and become the supercontinent of Pangaea by 195 million years ago. The
supercontinent of Pangaea stretched from near the South Pole to the North Pole
and was surrounded by one large continuous ocean called Panthalassa.
Geologically speaking, the supercontinent of Pangaea was short-lived and
began to split apart 180 million years ago. A rift that would become the mido-
ceanic ridge of the Atlantic opened between the North American and European
plates during the Jurassic. By the late Cretaceous, about 75 million years ago,
Asia and North America became linked together near present-day Alaska and
Siberia. This land bridge between Eurasia and North America is called Beringia.
Rifting also began to separate Africa and South America by 180 million years
ago. By 150 million years ago, in the late Jurassic, continued sea floor spreading
severed the land connection between the North American Plate and Gondwana-
land. During the late Jurassic, the subduction of the Pacific Plate by the North
American Plate initiated the Cordilleran Orogeny. Mountains such as the Coast
Ranges, Sierra Nevada, and Rocky Mountains of western North America were
formed by the Cordilleran Orogeny.
By the late Cretaceous, some 90 to 100 million years ago, a large seaway
separated North America from the South American and African plates. North
America itself was divided into western and eastern landmasses by a shallow sea.
Rifting and subsequent sea floor spreading continued to drive Africa and South
America apart. In addition, India and Australia-Antarctica were now separated
from each other and the rest of Gondwanaland. By about 90 million years ago,
204 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

ago. By 2 to 3 r
oceans was essE
Prior tole
consider the sp
the most isolate
They have a cc
Hawaii northw
moved over a l
appears to have
lion years, pro-
magma. The rna
movement occt
-150 mya
islands formed l
rience volcanic
and is increasir
Oahu and Kau:
area through e
lies a chain of g
ots were once i~
and seamounts
low-sea reefs. E
sink hundreds
northwestward
ation of plate te
Cretaceous -100 mya Future -+250 my
tually, all of the
the waves. This
about 5 million
new island, callt

UATERNARY CLI

As described 2
repeated altern
today. Almost E
However, recog
FIGURE 7.6 The movement ofthe major plates over the past 250 million years and 250
tury. The conce
million years into the future (after Scotese and Baker, 1975; Scotese, 1988; PALEOMAP We b
today was devE
site http://www.scotese.com/earth.htm;Briggs, 1995).
working in the ,
existing alpine !
not only had Pangaea ceased to exist, but Laurasia and Gondwanaland had bee glaciers in low-1
split apart. be explained by
During the Tertiary Period, between 65 and 2 million years ago, the mod - sive than they a
ern configuration of our oceans and continents came into being. Northeastward of past ice ages
movement of Africa and India brought Africa in contact with the European began to look f,
Plate and India into contact with the Asian Plate by 14 million years ago. During formitarianist ~
the same period, Australia moved northeastward and separated from Antarc- occurrence of ~
tica. The northward movement of South America and formation of land by vol- today. Howeve1
canic activity led to the creation of the Central American Isthmus and the believed that th
joining of North America and South America by a land bridge 3.5 million years the result of rna

~50 my
.... UATERNARV CLIMATIC CHANGE
1st 250 million years and 250
;otese, 1988; PALEOMAP We:
j Gondwanaland had bee-
lillian years ago, the moe-
into being. Northeastwar:
Jntact with the Europea::
~ million years ago. Durie~
d separated from Antar.:-
1formation of land by vo:-
merican Isthmus and tt:-
:ld bridge 3.5 million ye a~
QUATERNARY CLIMATIC CHANGE 205
ago. By 2 to 3 million years ago, the configuration of the earth's continents and
oceans was essentially the same as it is today.
Prior to leaving the topic of crustal movement and plate tectonics, we might
consider the special geological history of the Hawaiian Islands. The islands are
the most isolated in the world and have never been part of any continental plate.
They have a core of oceanic basaltic rock. The island chain that extends from
Hawaii northwestward to Midway Island and was created as the Pacific Plate
moved over a hot spot extending into the mantle of the earth. This spot, which
appears to have remained relatively fixed in the central Pacific for at least 90 mil-
lion years, produces volcanic activity that leads to the buildup of extruded
magma. The magma eventually rises above sea level and forms islands. As crustal
movement occurs at a rate of approximately 4 em per century in this region, the
islands formed by the hot spot gradually move northwestward and cease to expe-
rience volcanic activity and growth. The big island of Hawaii lies near the hot spot
and is increasing in area due to active volcanism. Islands to the west, such as
Oahu and Kauai, are no longer near the hot spot and are decreasing in surface
area through erosion and subsidence. Northwestward, beyond Midway Island,
lies a chain of guyots and seamounts called the Emperor Seamounts. These guy-
ots were once islands much like the current Hawaiian chain. Cores of the guyots
and seamounts show that they once supported terrestrial environments and shal-
low-sea reefs. Erosion and crustal subsidence have caused these former islands to
sink hundreds of meters beneath the sea. Plate movement has carried them
northwestward thousands of kilometers. The Hawaiian Islands are both a cre-
ation of plate tectonics and striking evidence of the earth's crust in motion. Even-
tually, all of the current Hawaiian Islands will move westward and sink beneath
the waves. This process will take several million years. The oldest islands are
about 5 million years old. East of Hawaii, volcanic activity is already forming a
new island, called Loihi which will rise above sea level in 15,000 to 20,000 years.
As described above, a key feature of the Quaternary Period has been the
repeated alternation between cold glacial conditions and warm conditions like
today. Almost everyone knows that there have been past "ice ages" on earth.
However, recognition of these past glaciations only occured in the twentieth cen-
tury. The concept that glacial ice cover had once been much greater than it is
today was developed by the Swiss-American naturalist Louis Agassiz. While
working in the Alps, he noticed that new glacial deposits being formed there by
existing alpine glaciers were similar to older rock deposits found away from the
glaciers in low-lying areas. Agassiz concluded that these older deposits could only
be explained by the theory that the alpine glaciers were once much more exten-
sive than they are today and extended into the lowlands. He published his theory
of past ice ages in 1840. In 1848 he was appointed to Harvard University and
began to look for evidence of past glaciation in North and South America. Uni-
formitarianist geologists accepted Agassiz's theory because it explained the
occurrence of glacial deposits in areas that are too warm to support glaciers
today. However, others greeted the glacial hypothesis with skepticism. Some
believed that the tills and other glacial features that Agassiz identified were really
the result of massive water erosion and deposition caused by the flood of Noah.
206 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

Subsequent geological investigations in northern Europe and North Am ::-- into bubbles tr
ica confirmed the presence of glacial deposits and led to the development of_ tiny air bubble
fourfold model of glaciation. According to this model, there had been four majL provides impo1
glaciations in the Quaternary. Each time, glacial ice cover extended over all c· One imp1
Scandinavia, most of the British Isles, and the Alps. Almost all of Canada, po::-- of the 180 and
tions of the Great Lakes region, and New England were covered by ice, and tt~ heavier due to
glaciation extended as far south as portions of Kansas in the plains. The mo = the isotopes 181
also postulated that these glaciations were short-lived and that the interglaci=... the atmospher
climate of the Quaternary was similar to today's climate. This model remain-,:: and precipitatt
the accepted view on glaciation until the early 1970s. We now know that bo:= 160. The relati'
temporal and spatial patterns of glaciation were far more complex than the fo -- condensation a
fold model. and 160 in the
As the twentieth century progressed, new geological data and paleontol o~­ the high latitw
ical dating permitted a more complete picture of the extent of glaciation and i~ Antarctic ice C<
impact on the earth. In the late 1940s, the American physicist Willard Libr _ from the ice co
developed the radiocarbon technique for dating organic remains. Radiocarbo::. was as warm a~
dating is based on measuring the amount of the radioactive isotope carbon > declined by 6°
(1 4 C) found in plant and animal remains. Isotopes are forms of an element the.: years ago. The I
have the same atomic number but differ in the number of neutrons. Carbon h~ particularly lov
three isotopes: carbon 12 and 13, which are stable isotopes, and carbon 14, whi :. 10,000 years ag
is a radioactive isotope that decays over time to nitrogen. Carbon 14 is forme.: Globally,
naturally in the stratosphere when cosmic rays bombard nitrogen. Plants acquir;: present. In iee-e
14 C during normal photosynthesis and accumulate it in their tissue. When animalS
temperatures \1
eat plants, they accumulate 14C. When a plant or animal dies, it ceases to accume- regions, the difJ
late 14 C. The isotope decays to nitrogen at a rate of one half-life which equals to >5° C. In eq
about 5700 years. By measuring the amount of 14C in a plant or animal remain ,'"~ other areas, sue
can estimate the length of time that elapsed since the organism died. Radiocar- than today. Ice
bon dating of plant and animal remains provides good chronological control ove::- Scandinavia (Fi
events of the past 40,000 years. Thanks to abundant geological, geochemical, an · the northern l
paleontological evidence, coupled with many radiocarbon dates, we have a ver:
good understanding of conditions during the last glacial maximum (LGM), dur-
ing which global glacial ice reached its maximum extent about 20,000 years ago.
We can consider these conditions to be a very general approximation of what the
world was like during earlier Quaternary glaciations. In the classical fourfol ·
model of glaciation, the last glacial maximum is equivalent to the final period o:
the Wisconsin Glaciation in North America, the Devensian Glaciation in Britain.
and the Wtirm Glaciation in central and western Europe.
One of the most detailed records of climatic conditions during the las
glaciation comes from ice cores taken in Greenland and Antarctica. These core5
extend from the surface of the modern ice caps down to depths of 1500 to 3350 m. ..•
The upper portions of the cores contain annual layers of winter and summer sno'"
accumulation. Like tree-rings, these annual layers can be counted to provide
chronological control. Lower portions of the cores must be dated using the occur-
rence of volcanic ash layers, which have been radiometrically dated elsewhere, and
correlations with other paleoclimatic evidence. The Greenland ice cores provide
records dating back about 100,000 years ago, while the Antarctic cores date back
400,000 years. An important record of past atmospheric conditions can be cap-
tured from the ice cores. The fresh snow layers at the top of the ice contain much FIGURE 7. 7 Th
air space. Over time, as more snow accumulates, the older snow is compressed to exposed due to lo
firn (ice crystals) and finally clear ice. During this process, the air is compressed maximum (after R

n Europe and North Amer-
~d to the development of a
, there had been four maj o~
cover extended over all o:
Almost all of Canada, por-
·ere covered by ice, and the
as in the plains. The mode:
~d and that the interglaciC..:
nate. This model remaine(
'S. We now know that bor"
tore complex than the fou:-
gical data and paleontolo5-
extent of glaciation and i:s
an physicist Willard Libb:
anic remains. Radiocarbm::.
lioactive isotope carbon l -
e forms of an element the.:
'er of neutrons. Carbon he
:opes, and carbon 14, whic::
ogen. Carbon 14 is form ec.
1rd nitrogen. Plants acqu ir~
1 their tissue. When anim<Ls
al dies, it ceases to accumt.: -
one half-life which equ ~
plant or animal remain, w~
~ organism died. Radioca.1-
chronological control OYC
~ological, geochemical, anc.
rbon dates, we have a ver;
:ial maximum (LGM), du:-
;nt about 20,000 years age
approximation of what th.:
s. In the classical fourfolc.
'alent to the final period o:
nsian Glas;iation in Britaill..
,pe.
conditions during the las:
md Antarctica. These core:;
o depths of 1500 to 3350 rr:.
Jf winter and summer sno
an be counted to provic::
;;t be dated using the occu;--
rically dated elsewhere, anc.
reenland ice cores provic :-
; Antarctic cores date ba ·
~ric conditions can be cav
:op of the ice contain muc::
lder snow is compressed t.::
•cess, the air is compresse..::
QUATERNARY CLIMATIC CHANGE 207
into bubbles trapped in the ice. Geochemists and geophysicists can sample these
tiny air bubbles and extract the trapped atmospheric gases. Analysis of the gases
provides important clues about past climate and the age at the ice cores.
One important constituent of the ice core records is the relative abundance
of the 18 0 and 160 isotopes. The isotope 18 0 is chemically similar to l6Q but is
heavier due to the additional two neutrons. Variations in the relative amount of
the isotopes 180 and 16Q in ice cores appear to be related to the temperature of
the atmosphere at the time of precipitation. The heavy isotope, 18 0 , condenses
and precipitates more readily at high temperatures than does the light isotope
16 0. The relative abundance of 18Q increases with increasing temperature during
condensation and precipitation. Studying changes in the relative amounts of 18 0
and 16 0 in the ice cores allows scientists to estimate changes in temperature at
the high latitudes. Such estimates are generally consistent from Greenland and
Antarctic ice cores for the past 100,000 to 140,000 years. The temperature records
from the ice cores suggest that between 110,000 and 140,000 years ago, the earth
was as warm as it is today. By 100,000 years ago, high-latitude temperatures had
declined by 6° C and were variable but remained depressed until about 10,000
years ago. The last glacial maximum at 20,000 years ago coincides with a period of
particularly low temperatures during the long cool interval between 100,000 and
10,000 years ago.
Globally, climate during the last glacial maximum was much cooler than
present. In ice-covered areas such as Canada and Scandinavia, summer and winter
temperatures were as much as 20 to 28° C colder than today. Even in equatorial
regions, the difference between glacial and modern temperatures was generally 2
to >5° C. In equatorial regions, precipitation was significantly less than today. In
other areas, such as the Great Basin of the United States, precipitation was higher
than today. Ice sheets that were 2 to 4 km thick were centered over Canada and
Scandinavia (Fig. 7.7). These sheets extended far south into northern Europe and
the northern United States. A person standing at the location of present-day
D.~··
·.,
~-:
DIce sheet
cj'
0 Arctic sea ice
FIGURE 7. 7 The location of major ice sheets and major areas of continental shelf
exposed due to lower sea levels (eustatic sea-level changes) during the last glacial
maximum (after Roberts, 1989).
208 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

Winnipeg, Canada, or Stockholm, Sweden, 20,000 years ago would have expe:--
TECHNI
enced a climate and a view very similar to what someone standing in the mid · ~
of the Greenland Ice Cap would experience today. Interestingly, Alaska a::..::
Siberia were free of ice except in mountainous regions. It is likely that the glac:=-. Palynology
climate was cold in these regions, but it was too dry to develop ice sheets. Mo::-
tane ice caps developed over alpine regions such as the Rocky Mountains, Al .::. Conifers and
phytes and pr'
Andes, and Southern Alps of New Zealand, in both the northern and south :-
ferns and mm
hemispheres. Smaller alpine glacier complexes developed in mountains such ~ Pollen grains 2
the Sierra Nevada of California and the high peaks of east Africa and Ke 150 microns. 1
Guinea. flower to flow(
The growth of glacial ice had a profound impact on regional and global se_ ever, many pia
levels. At the regional level, the weight of the ice depressed the land surfa ~:: a
sal is relative
beneath it and caused the sea to inundate adjacent coastlines. However, in rna::_ of the same pi
areas, the ice sheets also caused more distant land surfaces to be displac.:: ering plants a1
upward. This caused land to rise above sea level. Such changes in sea level, cau ~ .:: lions of grains
by the depression and distortion of the earth's crust by glaciation, are called i the same speci
static sea-level changes. In addition, the growth of the ice sheets required gr ~ tree may prod1
Many wind-1
amounts of water. The growth of glaciers occurs when snow does not melt o\·=--
the ocean. M<
the summer and is buried by snowfall during the following year. Over rna;::;_ Although the I
years, the overlying snow builds up, and its weight causes compaction and de fo~­ the external w
mation of the snow crystals until solid ice results. Water evaporated from oceaGS. polymer called
lakes, rivers, and soil fell as snow on the ice sheets and remained frozen ther;: larly in anaerc
This water, therefore, was tied up in the growing ice and did not return to the s _ moss. Within s1
as runoff. As the ice sheets grew, more and more of the world's water was tied tr~ served indefin
in them. The result was a drop in the level of the world's oceans by some 80 to 1: recovered fron
m. Because all the oceans are connected, this drop in sea level was the sarr.::
everywhere. Such global changes in the level of the oceans are called eustatic se
level changes.
Eustatic sea-level changes during glacial periods caused some very imp o~­
tant changes in the geography of the earth (Fig. 7.7). The drop in sea level caus ~
the Bering Strait between Alaska and Siberia to disappear and produced a larg::
land bridge in Beringia that directly connected Asia and North America. Larg::
islands such as Sumatra, Java, and Borneo in Southeast Asia became linked wi .__
the Asian mainland. The islands of New Guinea and Tasmania were linked wi:..
Australia. Many other islands, peninsulas, and isthmuses were greatly expand -
in size.
In addition to drops in sea level, glaciation caused other significant hydro-
logical changes. In the Great Basin region of the United States, the glacial climar.:
was wetter and colder than the modern climate. The increased precipitation anc
decreased evaporation caused large lake systems to develop in what is now a::
extremely arid desert environment. Such water bodies, which appear with larg -
scale variations in climate, are called pluvial lakes. The modern Great Salt Lake
of Utah is a small remnant of a much larger pluvial lake called Lake Bonneville
One pluvial lake even occupied portions of Death Valley. The relict shoreline5
and sedimentary deposits from these lakes are found throughout the Great BasiL Fossil pollen gra
region of the United States. tundra zone of n
The altered climate and geography of glacial periods produced profounc The two large ki'
changes in the geographic distributions of the world's biomes. Studies of fossi: grain is about 1(
sediments of thi
pollen, also known as palynology, have been widely used to reconstruct past Qua-
the site. Biogeo~
ternary vegetation (Technical Box 7.1). Studies offossil pollen, along with analysis
 QUATERNARY CLIMATIC CHANGE 209

ars ago would have experi-

~one standing in the middle
'· Interestingly, Alaska an ·
1s. It is likely that the glaciC.:.
to develop ice sheets. Mon-
:he Rocky Mountains, Al- -
the northern and souther.::
loped in mountains such c.>
\:S of east Africa and Ne,-
ton regional and global se ~
depressed the land surfac.c
Jastlines. However, in m a n ~
d surfaces to be displace.:
changes in sea level, cause.:
by glaciation, are called is(}-
Je ice sheets required grec.-
~n snow does not melt o'·c~
following year. Over maL~
uses compaction and de fo~­
:er evaporated from oceans
and remained frozen ther.:
md did not return to the se.:.
1e world's water was tied c
i 's oceans by some 80 to 1::
in sea level was the sar:;::-
:eans are called eustatic se
TECHNICAL BOX 7.1
Palynology (Fossil Pollen and Spore Analysis)
Conifers and flowering plants produce pollen grains that contain male gameto-
phytes and protect them during transfer to receptive ovules. Lower plants such as
ferns and mosses produce spores that also aid in the dispersal of gametophytes.
Pollen grains and spores are very small, generally ranging in size from 10 microns to
150 microns. This is about the size of fine silt. Pollen grains are transferred from
flower to flower by many agents, including insects, birds, mammals, and water. How-
ever, many plants rely on wind to disperse pollen grains and spores. As wind disper-
sal is a relatively random process, most pollen grains will not reach a receptive ovule
of the same plant species and will not fulfill their reproductive function. Thus, flow-
ering plants and gymnosperms that use wind pollination must produce many mil-
lions of grains in order to ensure success in sexual reproduction with other plants of
the same species. For example, in one year a single lodgepole pine (Pinus contorta)
tree may produce over 25 billion pollen grains.
Many wind-dispersed pollen grains are deposited on soil and washed into lakes or
the ocean. Many others are directly deposited into lakes, oceans, or peat bogs.
Although the living cytoplasm in the interior of such grains quickly dies and decays,
the external wall of the grain, called the exine, is constructed of a relatively inert
polymer called sporopollenin. The sporopollenin is very resistant to decay, particu-
larly in anaerobic environments such as lake bottoms or within fast growing peat
moss. Within such environments, pollen grains and spores can be fossilized and pre-
served indefinitely. Spores that are more than 500 million years old have been
recovered from sedimentary rocks.

is caused some very im p o~­

rhe drop in sea level cause:
ppear and produced a l a r~ ::
and North America. La r~ ::
1st Asia became linked wi:.:
Tasmania were linked \\·i:.:
uses were greatly expand.::

sed other significant hydr;:-

ed States, the glacial clime.:::
increased precipitation ~­
develop in what is now :=.::
~s, which appear with la rg::-
he modern Great Salt Lai:::
:tke called Lake Bonnevi["'
Valley. The relict shore lin ::-
throughout the Great Bas= Fossil pollen grains recovered from the sediments of a small lake in the modern arctic
tundra zone of northern Canada. The pollen grains are approximately 5000 years old.
periods produced profoLC,_ The two large kidney-shaped pollen grains are from spruce trees (Picea). Each spruce
i 's biomes. Studies of fos,---:.. grain is about 100 microns long. The high numbers of spruce pollen grains in
sediments of this age from the lake provide evidence that spruce trees once grew near
sed to reconstruct past Q c -
the site. Biogeographers often use fossil pollen analysis to reconstruct past vegetation .
il pollen, along with ana l ~-,
21 0 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

much cooler aJ
Most biogeographers work with fossil pollen from the Quaternary. Biogeographers Packrat midde1
generally obtain sediment samples for pollen analysis from lakes, peatlands, and the modern Great
ocean. Treatments of acids and bases are used to digest other organic and inorganic
Some bio
material in the sediment samples and recover the fossil pollen and spores. After the
The Eurasian l
pollen grains and spores are recovered from the sediment samples, they are identified
and counted using transmitted light microscopy at 400x to lOOOx magnification. Pollen habitat. The w
samples also contain other material such as fossil charcoal, which can be used to restricted to s
reconstruct past fire regimes, and plant remains such as stomata or algal fragments. biome of Euro
Most Quaternary pollen grains and spores can be identified as to the genus or species formations, sig
of plant they belong to. Identifications are based on the shape, size, exine wall struc- species. In the
ture, openings in the exine (called apertures), and sculpturing on the exine. that are comr
By conducting many studies of the relationship between modern vegetation and rearrangement
the pollen and spores deposited in present-day lakes and peatlands, palynologists matic regimes 1
have demonstrated that a good relationship exists between the vegetation sur- the modern eli
rounding a site and the pollen and spores that are deposited. Since vegetation is world consiste1
often controlled by climatic conditions, good relationships have also been found
which do not 01
between pollen deposition and climate. Palynology is used to reconstruct both past
vegetation and climate for the Quaternary. Pollen records extending for thousands ice cores tells 1
to tens of thousands of years have been recovered and provide crucial insights into the last glacial
Quaternary environmental history. 100 parts per n
per million. De
photosynthesis.
greater numbe
of fossil wood, needles, and seeds, have allowed biogeographers to reconstru~ plants to exch
many aspects of the earth's vegetation during the last glacial maximum. Polle::. increase in stor
and other fossil plant remains are often recovered from lake sediments, pea·- sensitivity to w
ftood deposits, and wind-blown sediments. One of the most amazing repositori : tive to water st1
of fossil pollen and plant remains is the middens of packrats (Neotoma spp.). Tt.: today. The end
middens are collections of plant fragments that have been gathered by the pac - climate and atrr
rats and accumulated in small rocky hollows. The urine of the packrats cemen= not present tod
the plant remains together and protects them from deterioration. Many packn:.: During th1
middens dating from the last glacial maximum have been found in the dese:-: that were often
regions of western North America. evidence that gr
During the glacial maximum, much of the area occupied by arctic an- formed commm
alpine tundra today was covered by glacial ice. In turn, tundra and steppe vegetz- lared lemmings,
tion replaced the boreal forest biome in Siberia and Alaska, as well as the boreC:.: portions of the 1
and deciduous forests of Europe. Imagine the city of Paris, France, surround nia during the la
not by forests and fertile fields, but by tundra and cold steppe. In North Americc.. Identifyin2
boreal forest replaced the grasslands on much of the Great Plains. In westerr. and debate arne
North America, the temperate rainforest moved southward. Trees such as red- ages can be divi<
wood (Sequoia sempervirens) grew close to Los Angeles, while more northe.: glaciation or alt1
temperate rainforest trees such as sitka spruce (Picea stichensis) grew south o: Cenozoic Era. S
San Francisco. The tropics were much drier than today, and rainforest vegetatio.: ond, what cause1
was often restricted to areas along rivers and other favorable localities. Tropica: ditions? Let's ex
and subtropical savanna and desert expanded. Florida, for example, was domi- Some of tl
nated by sand dunes and dry scrub vegetation. Mountain vegetation in zones iL glaciation comes
the tropics and higher latitudes were depressed a thousand meters or more. In high latitudes ha
the Andes, vegetation zones may have dropped in altitude by close to 2000 m. The ago. These pebbl
Andean treeline occurred at about 2000 m in elevation during the last glacial to the midocean
maximum. Today the treeline is found at 4000 m elevation. Packrat midden geophysical evict
records show that the southwestern United States and adjacent Mexico were have been ice-cc

:)uaternary. Biogeographers
)ill lakes, peatlands, and the
other organic and inorganic
pollen and spores. After the
t samples, they are identified
'lOOOx magnification. Pollen
coal, which can be used to
stomata or algal fragments.
ed as to the genus or species
shape, size, exine wall struc-
ring on the exine.
~en modern vegetation and
nd peatlands, palynologists
:tween the vegetation sur-
)osited. Since vegetation is
hips have also been found
:ed to reconstruct both past
ds extending for thousands
)fOVide crucial insights into
geographers to reconstru.::
st glacial maximum. Polle:
'rom lake sediments, pea--
: most amazing repositoric
tckrats (Neotoma spp.) . Tt:-
been gathered by the pad :-
ne of the packrats ceme n ~
eterioration. Many packrc.:
~ been found in the des e ~
ea occupied by arctic an:
t, tundra and steppe vegetz-
\.laska, as well as the bore<:..
f Paris, France, surrounde:
j steppe. In North Ameri ~­
te Great Plains. In wester:
tthward. Trees such as red-
tgeles, while more northe:
~a stichensis) grew south o:"
y, and rainforest vegetatio;:.
avorable localities. Tropic<:..
da, for example, was dorni-
ttain vegetation in zones i:
ousand meters or more. I:.
:ude by close to 2000 m. Th::
tion during the last glaciC..:
elevation. Packrat midde:
and adjacent Mexico wert
QUATERNARY CLIMATIC CHANGE 211
much cooler and moister than present during the last glacial maximum (LGM).
Packrat middens provide evidence that coniferous trees grew in large areas of the
modern Great Basin, Mojave, Sonoran, and Chihuahuan deserts.
Some biomes, such as the boreal forest of Eurasia, virtually disappeared.
The Eurasian boreal trees grew as small populations in local areas of favorable
habitat. The woodlands of the Mediterranean biome in Europe also became
restricted to small fragments in protected locations. The deciduous forest
biome of Europe was similarly reduced to small fragments. In other vegetation
formations, significant changes took place in the relative dominance of plant
species. In the eastern deciduous forest of North America, many tree species
that are common today became extremely rare. The fragmentation and
rearrangement of major biomes suggest that some of the broad regional cli-
matic regimes that existed during the glacial maximum have no counterpart in
the modern climates of the world. The glacial climates in many parts of the
world consisted of combinations of seasonal temperatures and precipitation
which do not occur today. In addition, evidence from Greenland and Antarctic
ice cores tells us that C0 2 (carbon dioxide) levels in the atmosphere during
the last glacial maximum were only 180 to 200 parts per million. This is some
100 parts per million less than average Holocene concentrations of 290 parts
per million. Decreased C0 2 in the atmosphere can result in decreased rates of
photosynthesis. Under low amounts of atmospheric C0 2, many plants produce
greater numbers of stomata. The increased stomata may have allowed the
plants to exchange gases more freely with the atmosphere. However, the
increase in stomata also raises rates of transpiration and increases the plant's
sensitivity to water stress. Consequently, some plant species were more sensi-
tive to water stress and drought during the last glacial maximum than they are
today. The end result of these differences between glacial and nonglacial
climate and atmospheric C02 is that biomes existed during glaciations that are
not present today, and some modern biomes were absent.
During the last glacial maximum, it was not only vegetation communities
that were often greatly different from modern ones. Paleontologists have found
evidence that groups of animals that are not found living together today actually
formed communities during the glacial maximum. For example, species of col-
lared lemmings, shrews, and ground squirrels that today inhabit very different
portions of the North American continent were all living together in Pennsylva-
nia during the last glacial maximum (Fig. 7.8).
Identifying the cause of the ice ages has long been a topic of great interest
and debate among scientists. The question of what caused the Quaternary ice
ages can be divided into two main issues. First, there is no evidence for extensive
glaciation or alternating glacial and nonglacial conditions during the rest of the
Cenozoic Era. So, why did large-scale glaciation begin in the Quaternary? Sec-
ond, what caused the Quaternary climate to shift from glacial to nonglacial con-
ditions? Let's explore each of these issues in turn.
Some of the most important evidence for the first onset of widespread
glaciation comes from marine sediment cores. Some midoceanic cores from the
high latitudes have small pebbles in them starting between 2 and 3 million years
ago. These pebbles are called drop stones and could only have been transported
to the midocean by icebergs calving off of glaciers. Geologic, paleontological, and
geophysical evidence all suggest that areas such as Greenland and Antarctica
have been ice-covered throughout the Quaternary. In contrast, for much of the
212 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

Clearly, tl
This cooling of
had its ultimatt
masses, such as
around the nor1
at the South Pc
nental conditioJ
lation. In additi
oceans. The atm
dered by the de
10 million yean
between 60 anc
the east coast
Between 10 and
strong eastward
Warmer curren
cap began to fo
million years a~
form at the hig
cooling becaust
energy back intc
Plate tectc
• Sorex fumeus
help explain wh
land, the Cana
- Dicrostonyx hudsonius explain why the
tions, like the la
Spermophilus tridecemlineatus
plate movemen1
111111111 Dicrostonyx torquatus these large varia
the alternations
• Site where all four species found as fossils mine the freque1
It is diffic1
FIGURE 7.8 The modern distributions of eastern shrew (So rex fumens), eastern colla roc times major glac
lemming (Oicrostonyx hudsonius), prairie ground squirrel (Spermophilus tridecemlineatus at the start of a 1
and western collared lemming (Oicrostonyx torquatus), and a site in Pennsylvania where ice retreats, me
fossil evidence indicates that all four species coexisted during the last glacial maximum, deposits. As a re
although they clearly do not live together today (after Graham, 1986; Graham eta/., 1996;
covers evidence
Brown and Lomolino, 1998).
niques and more
classic fourfold 1
of glacial and n
Tertiary these areas were free of ice. Fossil evidence shows that 40 to 50 millio:: analysis, the terr
years ago, areas such as Greenland and the Canadian Arctic Islands supporte · for older glacial
swamp and deciduous forest. For example, the stumps, needles, and seeds of the nary glaciations
Tertiary tree Metasequoia occidentalis, a relative of the redwood, can be found o:: from the oceans
Axel Heiberg Island in Arctic Canada. Other fossil trees found in TertiaJ;o prone to erosion
deposits of the High Arctic include maidenhair tree (Gingko) and cypress ( Glyp- remain undistur
tostrobus). During the same time, tropical rainforest and dry subtropical fores: ment records thE
covered much of North America, southern Europe, and Siberia. By 25 millior: Several tyt
years ago, cooler temperate deciduous and boreal coniferous forests had sprea · struct past varia1
across northern North America, Europe, and Siberia. By 3 million years ago tun- world's water th
dra was developing in the arctic. using radiocarbc

JJ
Q
;orex fumens), eastern colla u;;:
permophilus tridecemlinea tus
3 site in Pennsylvania where
g the last glacial maximum ,
n, 1986; Graham et al., 1996;
shows that 40 to 50 millie::
m Arctic Islands supportc.:
)S, needles, and seeds of tL
e redwood, can be foun d o::
sil trees found in Terti<L'
Gingko) and cypress ( Gl_' ;-
: and dry subtropical fo rc,-
and Siberia. By 25 millie::
'niferous forests had spree..:
By 3 million years ago tn--
QUATERNARY CLIMATIC CHANGE 213
Clearly, the earth became cooler during the late Tertiary and Quaternary.
This cooling of the earth during the Late Cenozoic and the initiation of glaciation
had its ultimate roots in plate tectonics. As the Tertiary progressed, large land-
masses, such as northern North America, Europe, and Asia, became concentrated
around the north polar region. At the same time Antarctica moved to its position
at the South Pole. The positioning of land in the polar regions produced conti-
nental conditions with cold winters. The land also provides a base for ice accumu-
lation. In addition, the landmasses inhibit the northward transport of heat by the
oceans. The atmospheric transport of heat to the north polar regions was also hin-
dered by the development of the Himalayas and the Tibetan Plateau over the last
10 million years. Antarctica remained attached, or very close to South America,
between 60 and 10 million years ago, and warm oceanic currents moving along
the east coast of South America brought heat to the southern polar region.
Between 10 and 3 million years ago, the distance had increased to the point that a
strong eastward flow of oceanic currents developed between the two continents.
Warmer currents from the south could no longer reach Antartica, and so its ice
cap began to form. Finally, the development of the Isthmus of Panama about 3
million years ago further disrupted oceanic circulation. When large areas of ice
form at the high latitudes, they produce a positive feedback promoting further
cooling because their white surfaces reflect a considerable amount of solar
energy back into space.
Plate tectonic movements and associated orogenies in the Late Cenozoic
help explain why the earth cooled and ice caps developed in areas such as Green-
land, the Canadian Arctic, and Antarctica. Plate tectonics cannot, however,
explain why the Quaternary has experienced alternations between cold condi-
tions, like the late glacial maximum, and warm conditions like today. The rate of
plate movement and orogenic mountain formation is simply too slow to explain
these large variations in Quaternary climate. To understand what may have caused
the alternations between glacial and nonglacial conditions, we must first deter-
mine the frequency and timing of such climatic changes during the Quaternary.
It is difficult to use terrestrial geologic deposits to determine how many
times major glaciations have occurred in the Quaternary. As glacial ice advances
at the start of a glaciation, it erodes deposits left by prior ice advances. As glacial
ice retreats, meltwater also erodes tills or covers them with lake and stream
deposits. As a result, the advance and subsequent retreat of glaciers destroys and
covers evidence of prior glaciations. With the advent of radiometric dating tech-
niques and more sophisticated stratigraphic analysis, it has become clear that the
classic fourfold model of glaciation was based on an oversimplified classification
of glacial and nonglacial deposits. However, no matter how sophisticated the
analysis, the terrestrial record of Quaternary glaciation is problematic, especially
for older glacial episodes. Surprisingly, the best evidence of how often Quater-
nary glaciations have occurred, and how extensive these glaciations were, comes
from the oceans. Unlike deposits on land, deposits in deep ocean basins are not
prone to erosion. Sediments deposited in the deep ocean accumulate slowly but
remain undisturbed. Cores from the deep ocean can provide continuous sedi-
ment records that extend back millions of years.
Several types of evidence found in deep ocean cores can be used to recon-
struct past variations in ocean temperature and even the relative amount of the
world's water that is tied up in ice sheets. The sediment cores are often dated
using radiocarbon analysis and paleomagnetic dating. Sediment cores from the
214 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

deep ocean generally contain large numbers of shells from tiny planktonic pro- 2. 5
tozoans called foraminifera. These organisms live in the upper water column c: 9 11
5
the world's oceans and produce distinctive calcium carbonate shells. Mar;. 15
species of foraminifera and other plankton are sensitive to water temperatur :. 3. 5
and analysis of their fossil shells can be used to reconstruct past temperatur-:. ~
For example, glacial-aged sediments from the North Atlantic Ocean contai:::
-
high numbers of shells from the arctic-subarctic species Neogloboqadrir.
- 4. 5
pachyderma. It has also been found that the tropical species Pulleniantir. 14
810
obliquiloculata and Sphaeroidinella dehiscens are absent from equatorial wate:-:: 2 6 12 16
of the Atlantic and Caribbean during the last glacial maximum. It has been s u ~­ 5'5 '--''--''--''--'--1.---L-
gested that the presence of northern foraminifera and the absence of equatori::... 0.0 o.s.__
species indicate a drop in sea surface temperatures of as much as 7 to 8° C cor;:
pared to modern conditions. The silica shells of planktonic zooplankton, such c.=
radiolaria, and phytoplanktonic unicellular algae, such as diatoms, are aJs.;:
I
- 0' 5 r---,--r--,----,.-
found in deep-sea sediments and have also been used to infer past sea surfa ~
~ -0. 5 { \ 1\ !\
vv v \
temperatures. In general, such biological evidence points to a cooling in tt:c
world's oceans during the last glacial maximum, with decreases in temperatu:-:: ~ - 1.5-
ranging from 0.5 to 9.0° C. Reconstruction of long-term changes in sea surfa ~
temperatures from plankton assemblages indicate that there have been at leas- -2.5
six episodes of sea surface cooling similar to the last glacial maximum over tt::
past 500,000 years (Fig. 7.9). 19·0
Geochemical analysis, particularly of the calcium carbonate (CaC0 3 ) sheG - _1 5
of foraminifera, has proven an especially powerful tool for reconstructing tt::
timing and relative extent of Quaternary glaciations. When aquatic organis ~ _4 0
form calcium carbonate shells, they obtain oxygen from the water in which the.
26· 5 L..---'---'----'---
live to produce their shells. The oxygen atom of a water molecule (H 20) can ~ 0 50 100 150 200
either of the two isotopes 18 0 or 160, although 18 0 is much less abundant in tt:: Age in t~
oceans than 16 0. As ice sheets build up during a glaciation, they grow by the addi-
tion of snow. Most of the water in the atmosphere that forms the snow crysta.:.:: FIGURE 7.9 Var
comes from evaporation at the surface of the oceans. During this process, th:: Caribbean Sea ove
record of the 18Q c
light isotope, 16 0, evaporates more readily than the heavy isotope, 18 0. The war :-
that is tied up in the growing ice sheets does not flow back into the ocean rc
replenish the 16 0 that is preferentially evaporated. Over thousands of years, th::
relative amount of 18 0 to 160 in the oceans increases as the glaciers grow. Whe::. warm climate an
the glaciation ends and meltwater enters the oceans, the 160 is replenished anL is, instead, the e'
the relative amount of 18 0 decreases. As a rough approximation, the volume o:' It is clear
global glacial ice is reflected in the amount of 18 0/16 0 in sea water. The oxyge::. and warming ha·
isotope record for the past 3 million years suggests that there has been a genera: glacial and nong
increase in ice cover from about 3 million to 2.5 million years ago. This coincid 5 of the Quaterna
with the first appearance of drop stones in deep-sea cores. The record is marke · to this phenomE
with a number of swings in the relative abundance of 18 0 that suggest as many as orbital theory of
50 advances and retreats of glacial ice over the past 2.5 million years. However. orologist, Miluti1
very prolonged periods of large-scale glaciation, such as the last glacial maxi- tions occur in tht
mum, have only occurred in the past 800,000 years or so. Higher resolution analy- and latitudinal d
sis of deep-sea cores indicates that about 20 large glaciations have taken place iL tion trigger peric
the past 800,000 years. In addition, the isotopic records from the ocean indica! of the earth caus
that very warm periods similar to the Holocene Epoch are relatively rare in tb ~ Scottish natural
late Quaternary. Previous generations of geologists and biogeographers believec Milankovitch Th
that the ice ages were short-term anomalies in global climate and that warm con- orbit of the eartl
ditions, such as we experience at present, were the norm. We now know that the eccentricity incn

; from tiny planktonic pro- 2.5
the upper water column c:
m carbonate shells. Man'
tive to water temperature'-. 0 3.5
mstruct past temperature'-. ~
ch Atlantic Ocean contai::. ~
:0
species Neogloboqadri1:,; 4.5
)ical species Pulleniantil:,;
ent from equatorial wate::!
maximum. It has been s u~ ­
:1 the absence of equatoric_
f as much as 7 to so C cor:::.-
:tonic zooplankton, such "-'
such as diatoms, are als c +0 .5
~d to infer past sea surfae:o
~ -0.5
points to a cooling in tt:o
1 decreases in temperatu:-:: - - 1.5
~rm changes in sea surfac::
at there have been at leas· -2.5
glacial maximum over ti::::
1 carbonate (CaC03) she:..:C
tool for reconstructing r:::o
>. When aquatic organis- ·
)ill the water in which the
ter molecule (H20) can b:'
much less abundant in r:.::o
tion, they grow by the adc-
tat forms the snow crystL:
ts. During this process, r::=
~avy isotope, 18 0. The watt:
ow back into the ocean :-
ver thousands of years, t ~=
as the glaciers grow. Whe:
the 160 is replenished ac.:
)roximation, the volume c
) in sea water. The oxygc:
at there has been a gener::..
m years ago. This coincid ~
:ores. The record is marke-:
lSO that suggest as many ~
~.5 million years. HoweYe:-
:h as the last glacial ma'=-
;o. Higher resolution anal:.-
:iations have taken place :=
ds from the ocean indica:::
:h are relatively rare in tt::
td biogeographers belie\ c:
:limate and that warm co;::-
>rm. We now know that tt::
QUATERNARY CLIMATIC CHANGE 215
5
9 11
31 37 43
I I II 15
L
100
ll t 1 111'1 34
8 10 52
6
Q
12
w 20 Glacial Isotope Stages
16
2.0 2.5 3.0
.---.---r--r--.----.----.--.---.--.....---.-----l
l
150 200 250 300 350 400 450
Age in thousands of years
FIGURE 7.9 Variations in sea surface temperatures and 18 0 content values for the
Caribbean Sea over the past 450,000 years (after Imbrie et al., 1973) and a 3 million year
record of the 18 0 content of ocean waters from the equatorial Atlantic (after Raymo, 1992).
warm climate and resulting distribution of life that characterizes our world today
is, instead, the exception.
It is clear from the marine record that many episodes of climatic cooling
and warming have occurred over the Quaternary. With the number and timing of
glacial and nonglacial intervals documented, it is possible to search for the causes
of the Quaternary climate shifts. Although there are several theories pertaining
to this phenomenon, the most widely accepted hypothesis is the Milankovitch
orbital theory of glaciation elaborated in detail in the 1920s by the Serbian mete-
orologist, Milutin Milankovitch. He argued that three natural and periodic varia-
tions occur in the orbital geometry of the earth and cause changes in the seasonal
and latitudinal distribution of incoming solar radiation. These changes in insola-
tion trigger periodic glaciations. The concept that changes in the orbital geometry
of the earth caused ice ages is actually quite old and dates back to the work of the
Scottish naturalist, James Croll, in the 1860s and 1870s. According to the
Milankovitch Theory, periodic glaciations occur for three main reasons. First, the
orbit of the earth around the sun is not circular but eccentric, and the degree of
eccentricity increases and decreases on a -95,800-year cycle. The time of the year
216 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

when the earth is closest to the sun is called the perihelion; the time when it is fa;--
thest away is the aphelion. Today we experience an intermediate degree of ecce:;:- 100
tricity, and the perihelion occurs in January. This makes winter in the northe:-::
~
hemisphere slightly milder than it would be if the orbit was circular; second, tt:' ~
Ql
tilt of the earth's axis varies from 21.8° to 24.4° on a- 41 ,000-year cycle. If the · · -~ 50
"'0
is increased, the difference between winter and summer insolation is increase.: c:

This makes summers warmer and winters cooler. Today the tilt is 23.5°. Third, tt:' "'
-1

seasonal timing of the perihelion and aphelion (also called the precession of tt:' 0
0
equinoxes) varies on a 21,700-year cycle. Today the perihelion occurs in Januar:·
-1
eleven thousand years ago it occurred in June.
Milankovitch proposed that glaciation takes place when the orbital geom;-
try of the earth causes the high latitudes of the northern hemisphere to recei,-= 2: -2
1- -3
low amounts of summer insolation. Conditions in the northern hemisphere a;-~ en
en
crucial because this region offers the greatest amount of land area for new i :: -4
sheets to form. The growth of northern hemisphere ice sheets influences glob2... -5
cooling by reflecting large amounts of sunlight back into space and disturbini
ocean circulation. Changes in southern hemisphere insolation do not appear : - .----
be very important. Summer insolation is important because heating in the sue- 18
mer causes the ice and snow to melt. Winters in the high latitudes of the northe
hemisphere are always cold enough for snow to occur. If the summers are coc FIGURE 7 .10
enough, the winter snow does not melt and ice sheets form. Minimum radiatio::. predicted by thf
input to the northern hemisphere occurs when the eccentricity of the earth":; volume and sea
orbit is high, the tilt of the axis is low (21.8°) , and the aphelion occurs in the surc- Parrott, 1985). ~
mer. A comparison of glacial ice volume, global sea level, and changes in non - were developec
ern hemisphere insolation since the last glacial maximum shows a go -
relationship between minimum summer insolation 18,000 to 20,000 years ago an-
maximum ice volume (Fig. 7.10). The subsequent decay of the ice sheets ac.:
resulting infusion of meltwater in the world's oceans corresponds to increa s in~ UTURE CHANG!
summer insolation in the northern hemisphere during the late Pleistocene an.: :ONTINENTS AI'
early Holocene. The timing of earlier Pleistocene glaciations correlates very w _
with past variations in insolation. Detailed analysis of sedimentalogical recor · We have seer
of earlier glaciations, such as the Permian, also suggests that Milankovitch cy cl~ and climate c
produced variations in glaciation during those times. With an undt
The Milankovitch Theory also casts some light on large-scale climati: possible to p
changes during the Holocene. There is much evidence from a number o: plates (Fig. 7
regions that summer temperatures were a few degrees warmer than prese : and the rangt
during the early to mid-Holocene. The timing of this warm period, known as th:: cold today wi
altithermal, or climatic optimum, varies regionally but ranges from about 80 · Antarctica ar
to 4000 years ago. For example, well-preserved tree stumps are found across t h ~ and collided ,
arctic tundra of northern Eurasia and show that boreal forest grew right up tc exist. Indeed.
the present Arctic coastline between 8000 and 4000 years ago. The timing of till:; Eventually, tt
maximum Holocene warming lags behind the timing of maximum summ : north as mod'
insolation, which occurred between 12,000 and 9000 years ago. It is likely tha: about the san
the presence of large areas of decaying ice in northern North America and the tinentalland1
relatively slow warming of the oceans kept summer temperatures cool until the Pangaea will
mid-Holocene. By 7000 to 6000 years ago, the ice had melted, and the sea sur- In the s
face had warmed. The cooling that has been experienced in the past 4000 yea ~ tions of incre
is probably the result of decreasing summer insolation caused by the changin ~ ical evidence
orbital geometry of the earth (Fig. 7.10). This cooling is entirely consistent wit t about 4000 yt
the Milankovitch Theory. should enter
 FUTURE CHANGES IN CONTINENTS AND CLIMATE 217

~lion; the time when it is far- 8

termediate degree of ecce;:;- 100 IIIIIIIIIIIIIIIUt,,,,,,

.
tkes winter in the norther::.
bit was circular; second, tt::
~
~
,,,~
...... ?f!.

41,000-year cycle. If the t:..::

Q)
.>2 50
'0
........
''..,,,,Ice 4 -;
.Q
Oi
mer insolation is increa s e ~ c ,,,,
ay the tilt is 23.5°. Third, tt::
C1l
_J ,,,,
..,,,,,,,,,,,,, ~
called the precession of tt:: 0 0 ~
(/)
0 ,/
erihelion occurs in Januar - I
-1 ,/ z
.S
1ce when the orbital geomc- 2: -2 /
.·ssT -4 g'
Q)

hern hemisphere to recei,·:: ,I C1l

1e northern hemisphere a:-::

tn
(/)
-3 .s:::
()

nt of land area for new ic:: -4

ice sheets influences gl o b~
;: into space and disturbiL.;
[nsolation do not appear : -
Jecause heating in the suc.-
gh latitudes of the northe::-::.
ur. If the summers are coc
.s form. Minimum radiatic-
eccentricity of the ean t .
aphelion occurs in the s
level, and changes in n or~ -
maximum shows a goc_
,000 to 20,000 years ago a::_
lecay of the ice sheets a;:,_
s corresponds to increasiL.;
ng the late Pleistocene a::.:.
~iations correlates very we
::Jf sedimentalogical recor.:_
sts that Milankovitch eye:::-
~ht on large-scale climL_
[dence from a number ,::
;rees warmer than prese::;
warm period, known as r:::.=
IUt ranges from about 8C·:
:tumps are found across t:::c=
real forest grew right up :
1ears ago. The timing of t'-
ting of maximum sumn:.::--
) years ago. It is likely tL
~rn North America and ::::
temperatures cool until r::_
ad melted, and the sea sc -
nced in the past 4000 y e c...~
[on caused by the changi==
g is entirely consistent \\-:,:.:
-5 -8
18 15 12 9 6 3 0
Thousands of years ago
FIGURE 7.10 Changes in summer and winter insolation for the northern hemisphere as
predicted by the Milankovitch Theory for the past 18,000 years. Changes in global glacial ice
volume and sea surface temperatures (SSTs) are also shown (after Kutzbach and Street-
Parrott, 1985). Modern conditions of global sea surface temperatures and glacial ice extent
were developed by 6000 to 7000 years ago.
TURE CHANGES IN
:ONTINENTS AND CLIMATE
We have seen that the past has experienced profound changes in the geography
and climate of the earth. It is natural to wonder what is in store for the future.
With an understanding of the patterns and mechanisms of plate tectonics, it is
possible to predict the future movement and configuration of the continental
plates (Fig. 7.6) . As the continents change position, global climate will change,
and the ranges of species will be split or brought together. Some areas that are
cold today will warm and other regions will cool. In 150 million years from now,
Antarctica and Australia will have merged again. Africa will have moved north
and collided with the European landmass. The Mediterranean Sea will no longer
exist. Indeed, the shrinkage of the Mediterranean Sea is already commencing.
Eventually, the northern coast of Africa will lie at 60° north latitude. That is as far
north as modern Scandinavia. In contrast, Siberia will have swung south and lie at
about the same latitude as modern southern China. In 250 million years, the con-
tinental landmasses will have rejoined each other or will lie very close together.
Pangaea will exist again.
In the shorter term, the changes in the earth's orbit have produced condi-
tions of increasingly low summer insolation in the northern hemisphere. Geolog-
ical evidence indicates that mountain glaciers around the world began to expand
about 4000 years ago. However, orbital calculations do not indicate that the earth
should enter another glacial period for several tens of thousands of years.
218 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

Aside from shifting continents, changes in the orbital geometry of the ean
and other natural influences, a new and potent force is exerting its influence on t t~
earth's climate. This new force is human activity. The ultimate mechanism behi ::.
anthropogenic climate change is increasing amounts of greenhouse gases in tt:::
atmosphere. Greenhouse gases, are molecules such as C02 , methane (CH4), an.:
nitrous oxide (N20) which absorb long-wave radiation (heat) in the atmosphe:-=
rather than allowing it to escape into space. Almost all of the absorption of h ~·
energy in the atmosphere takes place within the lower 4 km of the tropospher-
The most common of the greenhouse gases listed above is C0 2, which makes t:~
only about 360 parts per million of the troposphere today. Methane constitute
only 1725 parts per billion by volume, and nitrous oxide accounts for 311 parts pe:
billion. Despite their tiny concentrations, the absorption and reradiation of tt=
escaping heat energy by greenhouse gases keep the atmosphere and surface of t =
earth significantly warmer than they would be without the presence of such gas :.
Today, the average temperature at the surface of the earth is 15°C. If our atm :
phere did not contain greenhouse gases, the average temperature would b::
- 18°C. Without greenhouse gases, life would not exist on our planet.
Greenhouse gases such as C02, CH4, and N20 are natural constituents c:
the atmosphere. Carbon dioxide is released in volcanic eruptions, the weatheri.G;
of calcareous rocks, by respiration, organic decay, and burning of biomass b.
wildfires. Methane is mainly produced by anaerobic decay in soils, wetlands, an.:
peats. Insects, such as termites, and animals, such as cows, also produce significar.:
amounts of CH4 in digestion. Volcanic activity, biomass burning, respiration, an.:
decomposition of organic matter are important sources of N20. In the nature..
carbon cycle, photosynthesis by terrestrial and aquatic plants removes C02 fr or::
the atmosphere and incorporates it into plants and, eventually, animal tiss u ~
Increased levels of atmospheric C02 serve as a fertilizer promoting increase.:
plant growth. The resulting increase in green biomass and photosynthesis c~ have also studieci
then serve to reduce the C02 in the atmosphere. This negative feedback helps - bubbles from Gn
regulate the concentration of C02 in the earth's atmosphere. Natural feedbac -, pheric C02 incre<
within biogeochemical cycles have maintained relatively stable concentrations o: million observed i
C02 in the atmosphere during the Holocene. However, the amount of C0 2 in th::: cate that late Hoi
earth's atmosphere has varied greatly during the Pleistocene and over long ~ prior to the indus
geologic time. that at no time ov
Several different human activities have recently led to increasing amoun:: been as high as tr
of C0 2 and other greenhouse gases in the atmosphere. Fossil fuel, such as coa.._ alization, defores
oil, and natural gas, are formed from the biomass of organisms that died millio ' rapid rise in atmo
of years ago. The carbon in these remains has remained stored underground an.: ing of coal accow
has not been released back into the atmosphere. In burning fossil fuels, \\-::: use accounts for
release this stored carbon into the atmosphere at a greatly enhanced rate Deforestation anc
Destruction of vegetation can also release carbon into the atmosphere an-' and its replaceme1
decrease the rate at which carbon is taken up by photosynthesis. With increasi n~ increase in C02.
world population, industrialization, and land-use change starting about 200 yea_r:; increased in a ve
ago, there have been great increases in the amount of fossil fuels burned eact human populatior
year to produce energy. The rate of the destruction of natural vegetation has alsc atmospheric cone
increased. Has this activity caused changes in C02and other greenhouse gase ' tinue to increase i
Direct measurements of C02 in the atmosphere at remote regions such as the to~ will produce a dm
of the Mauna Loa volcano in Hawaii, Point Barrow, Alaska, and the Vostok Sta- Although it i
tion,Antarctica, show that since the late 1950s, C0 2 has increased throughout the in the amount of
world from 315 parts per million to 360 parts per million (Fig.7.11). Scientists how such increas1
 FUTURE CHANGES IN CONTINENTS AND CLIMATE 219

·bital geometry of the eartt. 360 i~ I I I I I I I I I I I I I I I

exerting its influence on tb::-
340 Anthropogenic C0 2 increase
ultimate mechanism behinC. from 270 ppm to 360 ppm
of greenhouse gases in tb:e between -1800 and 2000 AD
s COz, methane (CH4), an.: 320
m (heat) in the atmosphere
til of the absorption of hec.: ~ 300
E
~r 4 km of the troposphere 0.
0.
~ 280
we is C02 , which makes e;-
~
today. Methane constitu t~
le accounts for 311 parts pc:- ~ 260
c:

tion and reradiation of tt::
nosphere and surface of tt::
t the presence of such gasa.
earth is 15°C. If our atm
1ge temperature would
on our planet.
are natural constituents c~
ic eruptions, the weatheriL~
md burning of biomass c:
1ecay in soils, wetlands, ar::
>ws, also produce significa::-
tss burning, respiration, ac:
rces of NzO. In the natur.::..
c plants removes C02 fr o;:::
l, eventually, animal tisst.:::
tilizer promoting increase:
1ss and photosynthesis cc.-
negative feedback helps :
osphere. Natural feedb ac'·
ely stable concentrations c·-
:r, the amount of C02 in tt::
leistocene and over lon g ~-
;; led to increasing amou r::.
re. Fossil fuel, such as coc._
>rganisms that died millio=-
ed stored underground ar..:
In burning fossil fuels, \\ ::
a greatly enhanced ra t~
into the atmosphere ac
tosynthesis. With increas i.r::.~
1ge starting about 200 yea.::
of fossil fuels burned eac::.
·natural vegetation has als:
td other greenhouse gas e ~
note regions such as the tc~
:\laska, and the Vostok St.:.-
lS increased throughout tt::
nillion (Fig.7.11). Scientis:...
8~
240
220
200
180 C02 Vostok ice core
0 20 40 60 80 100 120 140 160
Age
(Thousands of years)
FIGURE 7 .11 Changes in atmospheric C0 2 concentrations for the past 140,000 years as
evident in the Vostok Ice Core from Antarctica (after Jouzel et al., 1993). Recent increases in
atmospheric C0 2 are also indicated.
have also studied the concentration of C02 and other greenhouse gases in air
bubbles from Greenland and Antarctic ice cores. They have found that atmos-
pheric C0 2 increased from 280 parts per million prior to 1800 to the 360 parts per
million observed in recent times (Fig. 7.11). Analysis of long ice core records indi-
cate that late Holocene levels of C0 2 were around 270 to 280 parts per million
prior to the industrial revolution. It is apparent from these long ice core records
that at no time over the past 100,000 years have levels of C0 2 in the atmosphere
been as high as they are today. Accelerating human population growth, industri-
alization, deforestation, and land-use change have produced an increasingly
rapid rise in atmospheric C0 2 over the past century. It is estimated that the burn-
ing of coal accounts for 31% of the unprecedented increase in COz. Petroleum
use accounts for an additional 31%, and natural gas burning contributes 13%.
Deforestation and land-use changes that add carbon by the burning of vegetation
and its replacement by nonvegetated surfaces is responsible for about 23% of the
increase in C02. Other greenhouse gases such as CH4 and N 20 have also
increased in a very similar manner over the last 200 years. Most analysis of
human population growth, fossil fuel demands, and land-use patterns suggest that
atmospheric concetrations of C02 and many other greenhouse gases will con-
tinue to increase in the future. Even a 2% annual increase in greenhouse gasses
will produce a doubling of C0 2 within the next 100 years.
Although it is widely accepted that human activity has led to a great increase
in the amount of greenhouse gases in the atmosphere, it is less well understood
how such increases will impact world climate. It has long been suggested that
220 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

increasing greenhouse gas concentrations will lead to the increased retention

heat energy in the atmosphere. This would produce higher temperatures in t::.::
troposphere and at the earth's surface. In 1861, the British scientist Joy Tynd<=-.
first suggested that changes in atmospheric C02 could produce climatic chang :.
In the 1890s, Svante Arrhenius concluded that a doubling of C0 2 could lead t o~
5° C warming of the atmosphere. In the 1970s and 1980s, American climat o lc~
gists Stephen Schneider and others provided fresh evidence and warnings abo:_
the potential warming of the earth due to increasing greenhouse gases. Since t ::
1980s, intense efforts have been made to estimate the impact of increasing C _
and other greenhouse gases on climate. This difficult problem has been tackJ -:
mainly through the use of sophisticated models of the earth's climate that
on supercomputers. A number of such models are in use today. These Gene

2.5 FIGURE 7.13 T

Alberta. The glaci<
62.0 and small lake in f
2.-
Q)
century. Such retn
0>
climate warming.
lij 1.5
.r::.
()

l!?
~ 1.0
Qi Circulation Moe
0.
E of C02 will lead
~ 0.5
century. Warmin
it is likely that r:
0
The GCMs prov
1850 likely that some
Year (AD) ence decreases :
Despite uncertai
widely forecast 1
(Fig. 7.13) and tl
level of 20 to 14(
be very problem
most of Venice, l
as large portions
tiona! dikes. Sucl
country like Ban
prone to inunda
could simply cea:
Is there an
gases has taken .
ture records frorr
global temperatt
FIGURE 7.12 The historical instrumental record of variations in global temperature an d (Fig. 7.12). As wt
climate model estimates of increases in annual mean temperature (°C) that could be occur in the higl::
experienced by the year 2050 due to greenhouse warming. The map shows the increased records of then
warming that might be experienced in different regions (°C) (after Hadley Center, 1995; provides some e'
Bryant, 1997). ever, such obser

J the increased retention o:
higher temperatures in th=
British scientist Joy Tynda=..
d produce climatic changes.
Jling of C02 could lead to.=.
1980s, American climatolc-
lidence and warnings abo::-
greenhouse gases. Since tt::
.e impact of increasing C _
t problem has been tackle::
:he earth's climate that n:::
n use today. These Gener::...
/
2050
ons in global temperature ar:
rature (°C) that could be
rhe map shows the increas e
(after Hadley Center, 1995;
FUTURE CHANGES IN CONTINENTS AND CLIMATE 221
FIGURE 7.13 The front of the Athabasca Glacier in the Canadian Rocky Mountains,
Alberta. The glacial front has been retreating rapidly over the past 100 years. The rocky area
and small lake in front of the glacier were covered by ice at the start of the twentieth
century. Such retreat is evident for a number of alpine glaciers and may be a result of global
climate warming .
Circulation Models, also known as GCMs, consistently suggest that the doubling
of C02 will lead to an increase in global temperature of 1.5 to 4.5° C in the 21st
century. Warming will not be even across the whole earth (Fig. 7.12). For example,
it is likely that particularly strong temperature increases will occur in the arctic.
The GCMs provide less consistent results in terms of changes in precipitation. It is
likely that some regions, such as the Great Plains of North America, will experi-
ence decreases in precipitation, whereas other areas will experience increases.
Despite uncertainties in predicting future precipitation, one important change is
widely forecast for the hydrosphere. It is estimated that the melting of glaciers
(Fig. 7.13) and thermal expansion of water in the ocean will produce a rise in sea
level of 20 to 140 em. This may seem relatively minor, but such an increase would
be very problematic for major coastal cities throughout the world. For example,
most of Venice, Italy would become uninhabitable. Some agricultural areas, such
as large portions of the Netherlands that lie below sea level, would require addi-
tional dikes. Such mitigation measures are available to rich developed nations. A
country like Bangladesh, which has much of its land near sea level and is already
prone to inundation, would be devastated. The nation of the Maldives Islands
could simply cease to exist.
Is there any evidence that global warming due to increasing greenhouse
gases has taken place already? Analysis and synthesis of instrumental tempera-
ture records from meteorological stations around the world show that the average
global temperature increased by a half degree or so over the twentieth century
(Fig. 7.12). As we discussed earlier, some of the greatest warming is expected to
occur in the high latitudes of the northern hemisphere. Analysis of temperature
records of thermometer readings from scattered northern weather stations
provides some evidence of a warming trend during the twentieth century. How-
ever, such observations are very sparse for the arctic and many other regions.
222 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES

TECHNICAL BOX 7.2

counting, eros
struct tree-rin
Dendroclimatology (Tree-Ring Analysis) dendroclimatc
wood density.
We have already seen how the analysis of tree-rings can be used to reconstruct the these samples
history of forest stands and fires (Chapter 5). In addition, tree-rings are one of the amalgamation
most important tools used by biogeographers to reconstruct recent past climati in the ring rec
changes. In many parts of the world, instrumental climate records from meteorolog- competition ~
ical stations extend back in time 100 years or less. This span is too short to allow rings are then
analysis of natural climate variability or to detect the impact of increasing green- recorded at n
house gases. In some areas, coniferous trees can be found that are 1000 to over 4000 mathematical
years old. In addition, it is sometimes possible to find preserved wood from trees climate. The v:
that lived and died thousands of years ago. By analyzing the rings of ancient living fer function rr
trees and preserved wood, it is often possible to construct long records of past cli- number of sta
mate that extend for hundreds to thousands of years. mate are relia
The climate of the past is captured in tree-ring records because trees in cold species and m;
regions or arid areas are often limited in terms of growth by temperature or precip- arctic treeline
itation. For example, a ponderosa pine (Pinus ponderosa) growing on a rocky ridge east Asia. Tree
in the western interior of the United States will grow more vigorously during a wet bristlecone pil
year than during a dry year. When growth is vigorous, the size of the annual ring pro- Holocene. A r
duced by the tree is large. During a dry and stressful year the size of the ring is small. evidence that
Tree-ring samples are usually obtained using a very fine coring device that pen- warming than
etrates the tree but does not injure it. Dead wood is sampled by sawing a cross sec-
tion of the trunk. The samples are mounted finely and then sanded for counting
using a stereomicroscope and special computerized measuring table. By carefully

Biogeographers
changes in temp
matic technique
(Technical Box
America or Sibe
years. The growt
ularly during the
tend to be wide;
and measuring e
struct annual ch
and other paleo•
around the wor
more sustained J
years. Evidence
nounced in the h
records that are ;
earth is warmin!
due to human ac

The author collects tree-ring samples from a larch tree (Larix dahurica) in northern
Siberia . A small boat anchored on the lake is being used as a platform to recover EY WORDS AND .
sediment cores for pollen analysis. Analysis of arctic tree-ring cores from Siberia and
elsewhere provides evidence that the twentieth century has experienced unusually :=:_sta tic sea-level change
warm temperatures compared to the past 400 to 1000 years . -:;_:sils
::;:ologic time scale

, be used to reconstruct the
n, tree-rings are one of the
tstruct recent past climatic
~records from meteorolog-
, span is too short to allow
mpact of increasing green-
j that are 1000 to over 4000
Jreserved wood from trees
~ the rings of ancient living
1ct long records of past eli-
ords because trees in cold
1 by temperature or precip-
z) growing on a rocky ridge
Jre vigorously during a wet
size of the annual ring pro-
the size of the ring is small.
ine coring device that pen-
tpled by sawing a cross sec-
t then sanded for counting
:asuring table. By carefully
1rix dahurica) in northern
ts a platform to recover
ring cores from Siberia and
as experienced unusually
IrS.
KEY WORDS AND TERMS 2 23
counting, cross-dating, and measuring the rings, dendroclimatologists can con-
struct tree-ring records that extend far into the past. In addition to optical counts,
dendroclimatologists also use X rays of wood samples to examine variations in
wood density. Generally, many trees are sampled at each site, and the counts from
these samples are combined into an indexed record for that location. Prior to
amalgamation, mathematical detrending techniques are used to remove variations
in the ring records that might arise due to biological factors such as tree-aging or
competition with other trees. The resulting detrended annual variations in the
rings are then statistically compared to annual variations in climate that have been
recorded at nearby meteorological stations. From this statistical comparison a
mathematical model is developed to link changes in ring widths to variations in
climate. The variations in the rings are then used as input in a mathematical trans-
fer function model that estimates past climate on the basis of the tree-rings. A
number of statistical tests are performed to make sure the estimates of past cli-
mate are reliable. Tree-ring records of past climate are available from many tree
species and many parts of the world, ranging from larches (Larix dahurica) at the
arctic treeline in Siberia to teak (Tectona grandis) trees at the equator in South-
east Asia. Tree-ring records from living and preserved oaks in Europe and the
bristlecone pines (Pinus aristata) in California extend back through the entire
Holocene. A number of tree-ring studies from the arctic and elsewhere provide
evidence that the twentieth century experienced longer and more prolonged
warming than any time over the past 400 to 1000 years.
Biogeographers have turned to paleoclimatological techniques to reconstruct past
changes in temperature. Tree-ring analysis is one of the most important paleocli-
matic techniques used to reconstruct recent changes in the northern climate
(Technical Box 7.2). Boreal trees such as white spruce (Picea glauca) in North
America or Siberian larches (Larix sibirica and L. dahurica) can live for over 400
years. The growth of these trees is often limited by cold climatic conditions, partic-
ularly during the summer. During warm summers, the annual rings of the trees
tend to be wide; during cold summers, the rings are narrow. By counting the rings
and measuring changes in their width, dendroclimatologists are able to recon-
struct annual changes in temperature back hundreds of years. Tree-ring records
and other paleoclimatic data from ice cores and lake sediments from many sites
around the world show that the twentieth century experienced greater and/or
more sustained high temperatures than any other time over the past 400 to 1000
years. Evidence from the arctic shows that climate warming has been most pro-
nounced in the high latitudes of the northern hemisphere. The more paleoclimatic
records that are acquired, the clearer the evidence becomes that the climate of the
earth is warming. A likely cause for this warming is increased greenhouse gases
due to human activity.
EY WORDS AND T ERMS
:=:...:static sea-level change Glaciation Law of superposition
::;_"S.Sils Greenhouse gases and effect Law of uniformitarianism
-:eo logic time scale Isostatic sea-level change
224 CHAPTER 7 CHANGING CONTINENTS AND CLIMATES
Milankovitch orbital theory of Orogeny
glaciation Plate tectonics
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A PTER 8
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which were onc'
the farmers livi
and their famili'
they could and
tral valley of Cc
hardships of the
as John Steinbe
was, it illustrate
mental change <
isms to alter tl
species, is in rna
mental change.
such changes in
As we ha\
have occurred i1
can be either nE
ing climate may
pitable. At the
Species escape t:
advantage of ne
to disperse eithE
For many l
to the establish!
marram grass (£
stem fragment c
animal species a
population. The1
birds, a single fE
Snail species car
lation. Some tis
changes in gend
In view of this d
refer to the stagE
~ht,
~
H. E. Jr., and Barnowsk ~
Quaternary Environments
versity of Minnesota,
The appearance and disap·
•r vegetation assemblages:
tiona! dynamics in eastern
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CHAPTER 8
DISPERSAL, COLONIZATION,
AND INVASION
During the 1930s, a series of extremely severe droughts affected the Great Plains
of the United States. These droughts destroyed crops and vegetation over large
areas of the Great Plains. Lack of vegetative cover allowed the top soil to be
removed by the wind. Fields were rendered infertile due to lack of water, loss of
soil, or deep accumulations of dust. Areas of Oklahoma and adjacent states,
which were once productive farmland, became known as the Dust Bowl. Many of
the farmers living in the Dust Bowl region could no longer support themselves
and their families. A number of such displaced farmers packed what possessions
they could and moved westward to work as field hands in areas such as the cen-
tral valley of California which were not as deeply impacted by the drought. The
hardships of those times are recounted in family histories, songs, and novels such
as John Steinbeck's Grapes ofWrath. As personally tragic as the 1930s drought
was, it illustrates that all species, including humans, are vulnerable to environ-
mental change and that one of the ways of surviving such changes is for organ-
isms to alter their geographic distributions. The history of plant and animal
species, is in many ways a story of changing distributions in the face of environ-
mental change. In this chapter, we will explore how plants and animals facilitate
such changes in geographic distributions.
As we have seen, changes in the geography and environment of the earth
have occurred in the past and will continue into the distant future. Such changes
can be either negative or beneficial to individual organisms and species. Chang-
ing climate may make the current geographic area occupied by a species inhos-
pitable. At the same time, new areas may become available for occupation.
Species escape extinction due to unfavorable environmental conditions and take
advantage of new areas that are available for occupation, by having the capacity
to disperse either themselves or their progeny to new geographic locations.
For many plant species, a single seed that germinates and matures can lead
to the establishment of a viable population. In the case of some plants, such as
marram grass (Ammophila arenaria) which is common on beach dunes, a single
stem fragment can grow vegetatively and establish a new population. For most
animal species at least one male and one female are required to establish a viable
population. There are, however, some exceptions. For many species of insects and
birds, a single female carrying fertilized eggs can establish a viable population.
Snail species can self-fertilize, and a single individual can produce a viable popu-
lation. Some fish species have been shown to actually undergo spontaneous
changes in gender, allowing two females to establish a reproductive population.
In view of this diversity of strategies, biogeographers use the term propagule to
refer to the stage in the life cycle (a plant seed, for example), part of the organism
227
228 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

(a piece of marram grass that can develop a new set of roots and grow into a ~ away from its p
sized grass plant), or group of organisms (a male and female rabbit of mat~ range of the spt
age) that is required to establish a new reproducing population. Upon arrival a: : and is especial!:
new geographic location, the propagule must then be able to establish a via - the species avo
reproducing population in order to survive. In many cases, propagules are ~ most interest tc
persed to locations where they cannot establish a viable population due to t~ Organism
existing physical or biological conditions. In other cases, the establishment of "' sal mechanism~
new species causes dramatic changes in the physical and biological environme- force to move 1
of the newly occupied area. In many cases, propagules disperse and establis:: water, or it can
within the current geographic ranges of the species. Such processes are crucial Plants cannot r
the survival of a species within its range. In other cases, dispersal and establis~ dispersers. Act
ment occur at sites that are beyond the current geographic range of the speci : :. Species that pn
Such events that lead to range expansions are called colonizations or invasio ns. to cross 3000 k
The term invasion is often used to describe geographic range extensions that a:-:: ibis) (Fig. 8.1).
caused by the introduction of exotic species by humans. The changes in ran== crossed the Atl
boundaries that occur due to dispersal and establishment are an important cor:;::,- South America
ponent of understanding the geographic distributions of species. In this chapte: km. Even large
we will examine the processes and patterns of dispersal, colonization, and rang:: distances of 10
expansion events. We will also look at some examples of biological invasions a - sive dispersers.
their impacts. seeds. One exa:
a parasite on 1<
The dwarf mist]
DISPERSAL can eject the ri
dispersers. For '
Although dispersal can be defined simply as the movement of an organism awe~
from its point of origin, this definition is too vague to be very useful. From the pe:--
spective of the biogeographer and the ecologist, there are two main categories o:
dispersal events. The first category is intra-range dispersal or ecological dispersa.:
which results in the movement of a propagule from its place of origin to a new sit::
within the current geographic range of the species. Intra-range dispersal :S
extremely important in maintaining species populations within their geographi -
ranges. In some instances, the physical environment of a plant or animal's place of
origin may not be suitable for survival of its progeny. Many tree species provid
good examples of this. Seedlings of light-demanding genera, such as pines, haY
low photosynthetic rates when shaded. These seedlings cannot survive when they
germinate directly under the canopy of their parents. Dispersal away from the par-
ent tree is crucial for the survival of progeny. The study of seedling distributions
for trees and shrubs often shows a lack of regeneration directly beneath the par-
ent. In many cases, dispersal of progeny decreases competition with parents and
siblings for light, water, food, or nesting sites. In some instances, dispersal also
decreases rates of predation upon offspring. Many seed predators such as rodents
and squirrels concentrate their feeding in areas close to parent plants where seeds
are most plentiful. In such cases, seeds that are dispersed away from the parent
have a higher survival rate. Finally, changes in physical and biological conditions.
caused by disturbance and subsequent succession, can make sites uninhabitable
for species. Many early successional herb species cannot survive shading and com-
FIGURE 8 . 1 A
petition by shrubs and trees that come later. In order to survive, the early succes- the nineteenth ce
sional species must be able to disperse seeds onto newly disturbed sites. spread througho1
The second general form of dispersal is extra-range dispersal or biogeo- 1960s have now I
graphical dispersal. This type of event results in the movement of the propagule be attributed to i r
 DISPERSAL 229

:roots and grow into a fu U- away from its place of origin to a new site that lies outside the current geographic
d female rabbit of mating range of the species. Extra-range dispersal allows species to colonize new regions
>pulation. Upon arrival at and is especially crucial when an environment becomes unfavorable since it helps
~ able to establish a viabl the species avoid extinction. Obviously, these types of dispersal events are of the
cases, propagules are dis- most interest to biogeographers.
ible population due to th- Organisms disperse their propagules in a number of different ways. Disper-
ses, the establishment of c. sal mechanisms may be passive or active. Passive dispersal requires an outside
md biological environmen: force to move the propagule. The force can be a physical one, such as wind and
ties disperse and establist water, or it can be in the form of biological agents, such as birds and mammals.
ch processes are crucial fo~ Plants cannot move under their own power and are almost all obligate passive
es, dispersal and establish- dispersers. Active dispersal relies on the propagule itself to provide motion.
·aphic range of the species.. Species that propel themselves, such as birds, bats, and insects, have been known
colonizations or invasions. to cross 3000 km of open ocean. A recent example is the cattle egret (Bubulcus
c range extensions that ar~ ibis) (Fig. 8.1). Either a breeding pair of egrets or a female with fertile eggs
tans. The changes in ran g ~ crossed the Atlantic Ocean from Africa and established a population of egrets in
tent are an important con:- South America in the late 1800s. The distance flown in this case was over 2000
of species. In this chapte::- km. Even large terrestrial mammals, such as elephants, have been shown to swim
;al, colonization, and rang~ distances of 10 km and can disperse to nearby islands. In general, plants are pas-
of biological invasions ar:.: sive dispersers. However, there are some interesting species that actively disperse
seeds. One example is the dwarf mistletoe (Areuthobium americanum), which is
a parasite on lodgepole pine (Pinus contorta) and jack pine (Pinus banksiana).
The dwarf mistletoe carries its seed in its fleshy base from which pressure changes
can eject the ripe seed to distances of several meters. Not all animals are active
dispersers. For example, the larvae of many marine invertebrates, including many
:ment of an organism awe.:
~ very useful. From the pe::--
are two main categories c:
rsal or ecological disp ers,_;_
place of origin to a new si:::
s. Intra-range dispersal _
ms within their geograpL :
a plant or animal's place
Many tree species prO\ic::
genera, such as pines, ha\ ::
s cannot survive when t h::~
ispersal away from the pc.::--
jy of seedling distributio::.
n directly beneath the pc.::--
npetition with parents ~:
1e instances, dispersal als
j predators such as rodec
) parent plants where see-
rsed away from the pare-
1 and biological conditio::..
1 make sites uninhabita2.=
>t survive shading and co=-
FIGURE 8.1 A cattle egret (Bubulcus ibis) flying over a waterway in southern Florida . In
o survive, the early succe-~
the nineteenth century, the cattle egret spread from Africa to South America, and it has now
ly disturbed sites. spread throughout much of the New World. Cattle egrets that spread to New Zealand in the
range dispersal or biogr: 1960s have now become established the re. Part of the recent spread of the cattle egret can
wvement of the propag::...= be attributed to increased areas of pasture land throughout the world .
230 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

gastropds, bivalves, worms, and crustaceans, exist as free-floating plankton tba:

are moved by water currents. Some planktonic invertebrate larvae found in th::
Pacific Ocean can survive three to six months and travel distances of 2000 to 4
km to new shallow-shore habitats. Such dispersal is responsible for the rich inve -
tebrate fauna associated with reefs on even the most remote Pacific islands su ~
as Hawaii. Mature barnacles, such as the common goosefoot barnacle (Lepas fru-
cicularis), adhere to marine organisms such as turtles and whales, as well as ships.
and are passively dispersed.
Passively dispersing plants and animals are often classified according to tb;:-
modes by which their propagules are dispersed:
Anemochores are dispersed by the wind. For many plants, the wind di:-
perses their seeds which have specialized shapes and structures to aid in dispe -
sal. The seeds of maple trees (Acer spp.) and many pines (Pinus spp.) posse -
winglike structures of thin, hard tissue attached to the seed coat. Seeds of this
type are referred to as samara seeds. The name comes from the winged helme '
worn by ancient Japanese warriors. When samara seeds fall, the wing causes the
seed to rotate much like a helicopter blade. The spinning imparts a small amoun:
of lift and decreases the settling velocity of the propagule. This allows the seed tc
remain suspended in the air and be transported away from the parent tree. Othe:-
seeds, such as the common dandelion (Taraxacum officina/e) and aspen trees
(Populus tremuloides), possess small tufts of hairlike material that catches the
wind and helps keep the seeds aloft. Interestingly, some plants, such as birche
that live in environments with winter snow cover release a proportion of their
seeds in winter when the seeds can be blown large distances across the smoot t FIGURE 8.2 C
snow surface. Tumbleweeds exhibit another interesting form of anemochory. widespread distr
Adult tumbleweeds die and are rolled along by the wind. During this movemen r. the ability of pair
the tumbleweed releases seeds. The Russian tumbleweed (Salsola iberica), origi- addition, Polyne:
nally from Eurasia but now found widely in North America, disperses some many remote is!;

20,000 to 50,000 seeds over great distances through this strategy.

Anemochory is not restricted to plants. For example, young black widow
spiders (Latrodectus mactans) spin long strands of web, which catch the wind an must be able
can transport the small spiders many kilometers. Even without such an adapta- plants, contac1
tion, wind can carry insects very great distances. Insect fauna captured in air sam- plants, like tht:
pling traps over the central Pacific Ocean include spiders, mites, flies, butterflie water. A stud)
moths, beetles, and other insect groups in roughly the same proportion as these Ocean showec
invertebrates are represented in the fauna of Hawaii. Ancestors of many of the eight weeks of
native spiders, mites, and insects found on Hawaii likely made the 3000- to 4000- Anemoh
km trip from Asia, Australia, or North America as anemochores. For these of aspen trees
insects, and in many cases for birds and bats, long-distance dispersal is caused useful for floo1
mainly by the organism being wafted along by strong winds rather than by the Zoochor
navigational efforts of the organism itself. adapted to clir
Hydrochores are dispersed by water. The adults, larvae, and eggs of many exo-zoochory.
aquatic organisms are hydrochores. Even relatively large organisms such as crabs spikes that ho
and starfish may have a juvenile stage that is dispersed as plankton by water cur- pha) produce
rents. One of the most striking hydrochores among terrestrial plants is the seeds as the t1
coconut palm (Cocos nucifera). The thick husk and shell of the coconut keep the colorful garde
seed afloat and protected from salt water for very long periods (Fig. 8.2). When adhere to pass
deposited by storm waves on suitable beaches, the coconuts germinate and grow. Dispersa
In this manner, coconuts have been able to spread to many tropical islands. of zoochory. B
Organisms dispersed by sea water are also called thalassochores. Such propagules often carry an
 DISPERSAL 231

ree-floating plankton tha·

:brate larvae found in the
1 distances of 2000 to 400C
Jonsible for the rich inver-
emote Pacific islands suet
efoot barnacle (Lepas f a.< -
ad whales, as well as ship

classified according to the

tarry plants, the wind d i ~­

tructures to aid in disper-
Jines (Pinus spp.) possess
e seed coat Seeds of this
from the winged helmets
s fall , the wing causes the
tg imparts a small amoun:
tle. This allows the seed tc
om the parent tree. Oth e~
Fficinale) and aspen trees
material that catches th.::
1e plants, such as birches.
ase a proportion of thei:-
:tances across the smoot
ng form of anemochon
d. During this movemen
ed (Sa/sola iberica), origJ-
America, disperses som;
. strategy.
nple, young black wido\•
which catch the wind an -
1 without such an adapta-
·auna captured in air sam-
~rs, mites, flies, butterflies..
same proportion as these
<\ncestors of many of the
vmade the 3000- to 4000-
anemochores. For these
tance dispersal is cause<::
winds rather than by the
larvae, and eggs of ma n ~
e organisms such as crabs
as plankton by water cur-
terrestrial plants is th-
l of the coconut keep th-
periods (Fig. 8.2). Whe
nuts germinate and gro\\
o many tropical islands.
ochores. Such propagules
FIGURE 8.2 Coconut palm (Cocos nucifera) seeds germinating on a tropical beach . The
widespread distribution of the palms on tropical coastal areas and islands owes much to
the ability of palm seeds to remain viable after long transport on ocean currents. In
addition, Polynesians brought coconut palms with them and introduced the species to
many remote islands such as the Hawaiian Islands.
must be able to withstand submergence in salt water. For the seeds of many
plants, contact with salt water causes mortality. However, the seeds of other
plants, like the coconut, can tolerate relatively long periods of contact with sea
water. A study of seeds from plants growing on the Aldabra Atoll in the Indian
Ocean showed that 81% of the native species had seeds that could survive up to
eight weeks of submergence in sea water.
Anemohydrochores are dispersed by wind or water. The tiny plumed seeds
of aspen trees can be dispersed by both wind and water. This ability is particularly
useful for floodplain species and plants that exist on islands.
Zoochores are dispersed by animals. Many plants have seeds that are
adapted to cling to the coats of passing mammals. This form of transport is called
exo-zoochory. For example, ragweeds (Ambrosia spp.) produce seeds with small
spikes that hook into fur or clothing. Plants like burclover (Medicago polymor-
pha) produce barbed pods that fasten to fur or clothes and disperse a number of
seeds as the transporting animal or person moves on. Other plants, such as the
colorful garden vines of the genus Plumbago, produce seeds that are sticky and
adhere to passing animals.
Dispersal of seeds through the feeding habits of animals is a common form
of zoochory. Birds and squirrels, for example, feed on acorns from oak trees and
often carry and store the acorns in the ground for later use. Some of the acorns
232 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

are not retrieved by the animals and grow into new trees. Jays can bury as rna= seeds (Fagus 1
as 4600 acorns in a single season and often transport caches of five seeds over l_ ( Ectopistes mig
m from parent trees. Other plants, such as fig trees, produce nutritious fruit tha _ tury hinders be
eaten by birds and other animals. The seeds contained within the fruit p ~­ In terms
through the digestive track of the animal and are eventually excreted. Some ' chores and zoo
these seeds come to rest at sites where they can germinate and grow. This form compare the a
transport is called endo-zoochory. (Eucalyptus re~
Anthropochores are zoochores that are dispersed by humans. Aside fro:: pean jay (Gan
agricultural crops such as corn (Zea maize), which have been specifically b r ~: seeds are depo ~
and cannot effectively disperse seeds without human intervention, most of ~= transported me
anthropochores are dispersed by a number of different animals in addition :. ual oak seeds l
humans. The common weed, ribwort (Plantago lanceolata), is a good exam 1::. 100 m. Howeve
The ribwort is native to Eurasia but is now found growing throughout the wor::. carry these mm
Its small brown seeds are sticky and are easily carried on clothing and fur. : of seeds are of
spread wherever Europeans settled and introduced horses and cattle graz :. persed distance
Native Americans sometimes referred to it as "white man's footprint." plant establishr
Anemochory and zoochory are the two main dispersal strategies for plan:::. species, that b<
What are the relative advantages and disadvantages? A prime adaptation f . persed hundrec
wind dispersal is lightness. Wind-dispersed seeds contain very little endosperm ::
support the seedling after germination. The resistance to death by desiccation "
by shading of germinating seedlings of plants such as oaks and chestnuts, whi ·-
possess large seeds and much endosperm material, is often greater than f _
species such as birch which have small light seeds (Fig. 8.3). In addition, the dis-
tribution of anemochorous seeds is relatively random depending on the win -
Such seeds can as easily be deposited on inhospitable sites as favorable sites.
Many zoochorous seeds are transported from the parent plant to similar sit ~
that are capable of supporting the plant species. However, each of these larg::
seeds costs the parent plant more resources than is required to produce a sm ~
anemochorous seed. One result is that fewer seeds are produced for dispers ·
The dispersal of zoochorous species is also dependent on the animal that trans-
ports the seeds. It has been suggested that the dispersal of North American be ec~

• Betula lenta
5
• Rhus glabra "0 80
•
Betula populifolia
Q)
t:
0
4 0.
(/)
• Pinus strobus c::
60
Q)
Acer rubrum : Liriodendron tulipifera ~
~3 (/)~

.J::. -g~
til
Q)
Q)
(/) 40
02 "iii
Gleditsia triacanthos • ::;)
"0
·:;:
'0 20
Quercus rubra • E
Castanea mollissima 0
o+--,--~~--~-r--r-~~~~
-10 0.5 0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 0
Log mean weight seed reserve (mg)
FIGURE 8.3 The relationship between seed size and mortality by shading for a number FIGURE 8.4 SE
of tree species found in North American and European deciduous forests (after Grime and seeds of a zoocho
Jeffery, 1965). 1965; Began et al.
 DISPERSAL 233

rees. Jays can bury as many
aches of five seeds over l OC
)duce nutritious fruit that is
tined within the fruit pas_
entually excreted. Some o:"
nate and grow. This form o:
:ed by humans. Aside frorr:
1ave been specifically brec
1 intervention, most of th-
·ent animals in addition tc
~olata), is a good exampl
.ving throughout the worlc
ied on clothing and fur. I;
horses and cattle graze
nan 's footprint. "
.persal strategies for plam -
;? A prime adaptation fo:;
tin very little endosperm tc
, to death by desiccation o~
oaks and chestnuts, whict
is often greater than fo~
g. 8.3). In addition, the dis-
seeds (Fagus grandifolia) was in part dependent on the passenger pigeon
(Ectopistes migratorius), and the extinction of this bird in the early twentieth cen-
tury hinders beech dispersal today.
In terms of actual transport distance, the differences between anemo-
chores and zoochores are sometimes not as great as might be expected. We can
compare the average transport distances for the small seeds of a eucalyptus
(Eucalyptus regnans) and the zoochrous acorns of oaks dispersed by the Euro-
pean jay (Garrulus glandarius) (Fig. 8.4) . The vast majority of the eucalyptus
seeds are deposited within 50 m of the parent stands, and only a tiny portion are
transported more than 100m away. The same is true for the transport of individ-
ual oak seeds by jays. In general, jays do not transport single seeds more than
100m. However, when the jays transport groups of two to five seeds, they often
carry these more than 100m. For both anemochores and zoochores, the majority
of seeds are often dispersed very close to the parent plant and a few are dis-
persed distances of hundreds to thousands of meters. We know from evidence of
plant establishment on isolated islands, or the spread of newly introduced plant
species, that both anemochorous and zoochorous seeds are occasionally dis-
persed hundreds to thousands of kilometers.

~
n depending on the wine Anemochore
lle sites as favorable site-
lrent plant to similar sit e ~
wever, each of these large
.c~

"0 "'
Q)"O
- c:
·u; ca
0 "'
a.:::l
400i
\ (Eucalyptus)
by wind

Q) 0
~quired to produce a sma.:. : "0 ~ 200
"'~
.re produced for dispersal. "0
Q)
Q)
t on the animal that trans- en
[ of North American beec·
0
0 50 100
Meters

"0
2 80
0
a.
"'c: 60 Zoochore
~ (Quereus)
"'~ by Jays
"0~
Q)g__
Q)
40
"'
Cii
:::l
"0
s 20
i3
E

ity by shading for a number
ous forests (after Grime and
0
0 50 100
Meters
FIGURE 8.4 Seed dispersal curves for an anemochore (Eucalyptus) and the individual
seeds of a zoochore (Quercus) carried by jays (Garrulus glandarius) (data from Cramer,
1965; Begon et al., 1996).
234 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

If we look more closely at plots of the distance that propagules are dis- establish perrr
persed from parents, we see that the resulting dispersal curve for most species fol- we should con
lows a generally similar pattern. High numbers of individuals are dispersed close sal--<:olonizati<
to the parent, and an exponential or normal curve of declining numbers of indi- episodic irrup1
viduals are dispersed futher away (Fig. 8.4). An exponential dispersal curve, witt Seasonal
a very high proportion of offspring dispersing close to the parent, appears to b ularly occupie·
more common for passive anemochores, while a normal curve, with less offspring ditions, feedin
establishing very close to parents, is often observed for animals. The difference they move in <
may be because animals can actively choose to disperse further, thereby avoiding follow the sarr
competition, while passive dispersal by wind is essentially a physical proce ~ the summer is
dependent on winds and settling velocities of the propagules. area occupied
We have seen that dispersal is important for species to escape inhospitabl taken repeate<
conditions and reach new habitats. However, a propensity for distant dispers al Seasonal
can also be disadvantageous. The fact that an organism occurs in a specific place the eastern U
often indicates that conditions at that site are favorable for individuals of tha during the win
species to survive and grow to reproductive maturity. Dispersal away from th as golden plov
site may bring the organism to a location in which conditions are inhospitable. dra biome of ·
For example, anemochorous seeds from island-dwelling plants are likely to b coastal region:
blown into the ocean and die. Similarly, flying insects and birds could potentially (Fig. 8.5). Inse
venture too far from the island and be lost. Interestingly, some plant species thai
have evolved on islands lack plumed or winged seeds that are typical for related
species living on the mainland. Similarly, some island species of birds, flies, and
beetles have lost the ability to fly. The dodo bird (Raphus cucullatus), an extinc:
relative of the pigeons that lived on Mauritius Island in the Indian Ocean, is a
good example. The ancestors of the dodo had dispersed to the islands by flying.
but the dodo birds were incapable of flight. The typical distribution patterns o ·
both anemochorous and zoochorous seeds, with the majority of propagules
falling close to the parent and a few being transported great distances, naturally
increases the chance of species survival. This form of dispersal curve means that
the majority of propagules found close to the parent can take advantage of the
probability that sites there will be hospitable. However, the smaller number of
propagules that are dispersed to great distances may help to advance the range
and long-term survivability of the species by taking advantage of potentially
favorable conditions at new sites. Some plants, such as the telegraph weed (Hel-
erotheca latifolia), produce both plumed seeds for air dispersal and nonplumed
seeds that fall close to the parent plant. In general, the ability to disperse propag-
ules great distances seems favored by organisms living in frequently disturbed
environments, while it is not so favored by species in very stable environments or
on islands.

COLONIZATION, SEASONAL
MIGRATIONS, AND IRRUPTIONS

Colonization occurs when a propagule arrives in an area previously unoccupied

by the species and establishes a reproducing population, thereby expanding the
geographic range of the species. Being adept at dispersal does not benefit a FIGURE 8.5 E
species if it is unable to colonize its new location. Failure to establish a permanent migration by diff
population is common in widely dispersed organisms. Each year, about 100 (1992). Irruption'
species of birds from Asia and Europe are seen in North America, yet do not breeding areas ir
 COLONIZATION, SEASONAL MIGRATIONS, AND IRRUPTIONS 235

e that propagules are dis- establish permanent populations. Before we discuss the process of colonization,
curve for most species fo l- we should consider two interesting phenomena that do not represent true disper-
viduals are dispersed close sal-colonization events. These two phenomena are seasonal migrations and
declining numbers of indi- episodic irruptions.
ential dispersal curve, wit Seasonal migrations are the annual movements of organisms from one reg-
> the parent, appears to be ularly occupied geographic region to another for purposes of avoiding harsh con-
:tl curve, with less offspring ditions, feeding, and mating. Many species exhibit such behavior. Not only do
or animals. The differenc they move in a regular annual fashion from one region to another, but they often
e further, thereby avoidin g follow the same routes between regions. The area occupied by the species during
'ntially a physical process the summer is often referred to as its summer range to differentiate it from the
agules. area occupied in the winter, which is called its winter range. The regular paths
::ies to escape inhospitabl taken repeatedly between winter and summer ranges are called migration routes.
msity for distant dispersa: Seasonal migrations are very common for bird species. Many song birds in
n occurs in a specific place the eastern United States migrate to Mexico and Central and South America
ble for individuals of tha: during the winter and return north to breed in the summer. Some shorebirds such
. Dispersal away from the as golden plover (Pluviialis dominica) and knots (Calidris spp.) breed in the tun-
mditions are inhospitable. dra biome of Europe, Asia, or North America and spend the winter months in
mg plants are likely to be coastal regions of southern Africa, South America, Australia, and New Zealand
md birds could potentiaU,- (Fig. 8.5) . Insects may also undertake considerable seasonal migrations. One of
~ly, some plant species tha:
that are typical for relatec
species of birds, flies, an ·
hus cucullatus), an extinc:
in the Indian Ocean, is c.
~ d to the islands by flying.
a! distribution patterns o::
e majority of propagules
1 great distances, naturalh-
Espersal curve means th a:
can take advantage of the
er, the smaller number o::
help to advance the rang-
advantage of potenti al! ~
. the telegraph weed (He:-
dispersal and nonplume ·
ability to disperse propag-
tg in frequently disturbe -
:ry stable environments o;-
0 • Knot migration
\ D Locust irruption
~
(}
·ea previously unoccupie - ng
)n, thereby expanding the
Jersal does not benefit c. FIGURE 8.5 Examples of migration and irruption . Remarkable long-distance annual
e to establish a permanen: migration by different species of Eurasian knots (Ca/idris) (after Piersma and Davidson
1s. Each year, about l OC (1992). Irruption expansion of the desert locust (Schistocerca gregaria) swarms from
orth America, yet do no: breeding areas in central Africa (after Begon et al., 1996).
236 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

the most celebrated instances of this pattern is the travel of the monarch butte:-- Mod
fiy (Danaus plexippus), which migrates annually from a summer range in tl:::
northern United States and southern Canada to a winter range in coastal Califo:--
nia and southern Mexico. Monarch butterflies that have been tagged in Ontaric
Canada, have been found over 2600 km to the south in their Mexican wintc:
range. Wildebeests ( Connochaetes spp. ), zebra (Equus spp. ), gazelle (Gaz e/.'..:

I!
spp.), and other grazing animals of the Serengeti Plain in Africa also travers::
hundreds of kilometers each year from feeding and breeding grounds in tl::::
south to feeding grounds in the north. Many marine animals also display season;o_
migratory behavior. The Pacific gray whale (Rhachianectes glaucus) spends tl:::
summer in the North Pacific off the coast of Alaska and the winter off the coas- ~
ofMexico.
What triggers animals to begin their seasonal migration and what guid :
them along the same routes and to the same seasonal ranges has long been ~
topic of research and debate. Recently, some migratory bird species have be : : .
shown to have unusual concentrations of metal in their brains. These birds me.
be guided, in part, by the earth's magnetic field.
Irruptions are episodic explosions in the population size and geographi.: Beetles

ranges of insects or animals. Locusts are the best known example of species tha:
experience dramatic irruptions. The desert locust (Schistocerca gregaria), Africa:.
locust (Locusta migratoria), and red locust (Nomadacris septemfasciata) all haw
core areas in central Africa where they live and reproduce. Increases in populc:-
tion, coupled with low food supplies in their home range, cause the locusts r
swarm and disperse over huge areas of Africa and adjacent Asia (Fig. 8.5). Som"'
African locust swarms have even crossed the Atlantic to South America. How-
ever, the locusts cannot reproduce and permanently occupy this extended range
The range of the species eventually collapses back down to a core area. Th""
desert locust has peaks in such irruptions roughly every 15 years.
During a true colonization event, a species arrives in a new area and esta
lishes a reproducing population. Population biologists have long been intereste ·
in the demography of such events. If an organism were to colonize a new area.
which possessed unlimited resources to support the species, we would expe :
population growth to be exponential (Fig. 8.6). Imagine that a male and fema l~
finch are blown to a large island and reproduce. The next year the original breed-
ing pair may have produced two male and two female offspring. The three pairs
of finches that now exist on the island in turn produce a total of 12 more off-
spring, bringing the total population to 18 in a relatively short span of time. If w
were to plot the growth of the finch population against time, we would observe ar.
exponential curve (Fig. 8.6). The rate of exponential population growth can b
described by the simple equation
FIGURE 8.6 Ex
dN/dt= rN population growtt
Crombie, 1945). A
where dN =the change in population size (N) , dt equals the time increment (one (Picea) colonizing
generation in our example) , and r is the per capita rate of population growth. (the The population si;
number of finches that each parent produces minus the number that die prior to spruce pollen dep
reproduction in our example). The rapidity of growth that is possible due to
exponential increases in population explain why it is often so difficult to control
invading species.
If we ponder the example of the colonizing finches a bit more, we must con-
sider that the island that they have invaded does not really offer infinite resources
 COLONIZATION, SEASONAL MIGRATIONS, AND IRRUPTIONS 237

tvel of the monarch butter- Model

iffi a summer range in the
ter range in coastal Califor-
ve been tagged in Ontario. dN
h in their Mexican wint e~ dt
tus spp.), gazelle (Gazellfl
ain in Africa also traver~e
l breeding grounds in the
timals also display seasona:
nectes glaucus) spends the
nd the winter off the coa~:

nigration and what guid e~

II ~~
~e<c'
«,:+-«0~,../
,~ "'"' 'c
\..0"'
al ranges has long been ::
>ry bird species have bee=
~ir brains. These birds rna\
Time (t)
ation size and geographi c Beetles Spruce trees
5.0 . . . . - - - - - - - - - - - - , 1 0 0
Nn example of species th e.:
!stocerca gregaria), Africa= 4.5
I %Total Picea
~400
ris septemfasciata) all ha\·;e ~
oduce. Increases in populc.- (IJ
~
4.0 0
·ange, cause the locusts rc ]l 200
acent Asia (Fig. 8.5). Some c:o ~ 3.5
~ to South America. H m\ - 'U
;;, 3.0 oQ.
ccupy this extended range ')' co
:::J
down to a core area. Th-o 100 200 300 "0
a: § 2.5 ([)

y 15 years. Days if (IJ

c C'([)l
'(ii :::J
!S in a new area and estab- 0, 2.0 a.=.
have long been intereste-' c 3
~
re to colonize a new area. & 1.5
species, we would expe, .
1e that a male and fem ale 1.0
:xt year the original breec-
• offspring. The three palli 0.5
ce a total of 12 more oE-
0 1 ~
I I I I 0
ly short span of time. If wE 11.2 10.5 9.9 9.3
time, we would observe a= Age
14
population growth can b-e ( C yr. BP (x 103))

Is the time increment (onE
of population growth. (t h-o
e number that die prior tc
:h that is possible due tc
•ften so difficult to cont ro~
FIGURE 8.6 Exponential and logistic population growth curves. An example of logistic
population growth observed in the laboratory for beetles (Rhizopertha dominca) (data from
Crombie, 1945) . An example of logistic population growth for lodgepole spruce tree species
(Picea) colonizing areas of western Canada during the Holocene (after MacDonald, 1993).
The population size of the spruce trees in the study region is inferred from the amount of
spruce pollen deposited in lake sediments.

:s a bit more, we must cou-

llly offer infinite resource~
238 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

for population growth. Eventually, all of the sites on the island which can suppor TECHNI
finch nesting and feeding will be occupied, and the population cannot continue -
grow. Our hypothetical island may only be able to support 100 finches. We rna:
recall from earlier chapters that the environmentally controlled limit to the por- Reconstrm
ulation size of a species in a particular area is called the carrying capacity. In oc;
Fossil pollen :
example, the carrying capacity of our island for finches is 100. As population siz:: reconstruct tl
approaches carrying capacity, intraspecific competition increases for resourc :. North Ameri•
For example, two pairs of finches that live near each other may compete for nes:- tocene. The tc
ing sites and food. In the end, only one pair might survive and reproduce. Th:..> in the vicinit)
increasing competition between offspring causes the rate of population growth r- pollen in sedi1
slow as carrying capacity is approached. The population growth of a colonizin~ increases in t
species in a setting with finite resources has the S-shaped form of logistic popula- deposited wil
tion growth (Fig. 8.6). We can mathematically describe the logistic populatio;: bon dating is
growth of a colonizing species as stratigraphic 1
ing the increa
nologists coui
dN/dt=rN (-K -N)
-
K colonized the
of species and
where K is the carrying capacity. The equation simply expresses that as TheAmeri<
approaches K the rate of population increase will slow and there is no gro the postglacia
after N = K. Since all environments can only support a limited number of ind:- sil pollen site~
viduals of any species, the logistic model is a better general approximation of tb-- fully radiocar
growth of colonizing species than is the exponential model. colonization <
Reconstructio
One convenient way of comparing the growth rates for different popula-
ofNorthAme
tions that are increasing either exponentially or logistically is to calculate the Pollen anal'
population doubling time. This is simply the number of years it takes for the po tion dynamics
ulation to double in size. Once the per capita rate of population growth rate (r) is because many
calculated, the population doubling time (td) is calculated from the equation to thousands <
biogeographic
td = ln2/r
imperfect one
The In 2 approximately equals 0.70, and this rounded value is typically used in tb pollen that cm
equation. that lay far be
The logistic equation for the population growth of colonizing species bas firs, produce ~
records from ~
been tested in many laboratory experiments on population growth of single-celled
relationship b
organisms such as yeast cells, insects such as fruit flies (Drosophila melanogaster). lakes and bog~
water fleas (Daphnia magna), beetles (Tribolium and Rhizopertha), and simp! records are be
plants such as aquatic duckweed (Spirodela oligorrhiza). There is also evidence
that natural populations grow in a logistic manner. For example, a study of the pri-
mary succession on the volcanic island of Surtsey near Iceland showed that the
colonization of bare rock by moss followed a clear logistic pattern. Fossil pollen from studies of 1
evidence of the population growth of some, but not all, tree species colonizing the Bering Sea.
deglaciated landscapes in North America, Asia, and Australia (Technical Box 8.1) Island in order 1
at the start of the Holocene also reveals logistic growth (Fig. 8.6). The population of the populatic
doubling times for different tree taxa ranged from less than 30 years to over 1000 2000 reindeer. '
(Table 8.1). growth rate, the
In addition to the smooth s-shaped pattern of simple logistic growth, some remained in 19:
colonizing populations show evidence of a phenomenon that can be called carry- very low. The de
ing capacity overshoot. In these cases, the initial growth of the population contin- deer and sharp
ues beyond the average carrying capacity of the area, and the population experiences rapid populatio1
a catastrophic decline to the carrying capacity, or below it. One example comes entire island's (

~ island which can support
ulation cannot continue to
'port 100 finches. We may
ontrolled limit to the pop·
e carrying capacity. In our
is 100. As population size
n increases for resources.
1er may compete for nest-
rvive and reproduce. Th is
te of population growth to
)n growth of a colonizing
~d form of logistic popula·
'e the logistic population
tply expresses that as S
w and there is no growth
a limited number of indi-
eral approximation of th
ldel.
1tes for different popula-
tically is to calculate tbe
years it takes for the po
pulation growth rate (r) is
~d from the equation
lue is typically used in the
of colonizing species has
on growth of single-cellec
)rosophila melanogaster).
Rhizopertha), and simple
z). There is also evidenc
xample, a study of the pri-
Iceland showed that th ~
stic pattern. Fossil polle
ll, tree species colonizin§
tralia (Technical Box 8.1
(Fig. 8.6). The populatior.
han 30 years to over 100C
pie logistic growth, som
1 that can be called earn--
of the population contin-
1e population experiences
"' it. One example comes
COLONIZATION, SEASONAL MIGRATIONS, AND IRRUPTIONS 239
TECHNICAL BOX 8.1
Reconstructing Plant Colonization Using Fossil Pollen
Fossil pollen grains found in lake sediments and peatlands have often been used to
reconstruct the history of plant colonization, particularly the northward spread of
North American and European tree species that occurred at the end of the Pleis-
tocene. The technique is based on the premise that when a plant species first arrives
in the vicinity of a lake or peatland, it will be represented by very low amounts of
pollen in sediments deposited at that time. As the population of the colonizing plant
increases in the vicinity of the site, the amount of pollen from that species being
deposited will increase in proportion to the size of the plant population. Radiocar-
bon dating is usually used to derive a time scale for the lake sediment and peatland
stratigraphic records. The British palynologist, Keith Bennett, showed that by study-
ing the increase in the accumulation rates of tree pollen over time at one site, paly-
nologists could reconstruct the population growth rate for the tree species as it
colonized the area around the site. Such studies have been carried out for a number
of species and genera in England, Europe, Australia, and North America.
The American palynologist, Margaret Davis, pioneered the approach of mapping
the postglacial geographic expansion of tree species by using networks of many fos-
sil pollen sites. By collecting pollen stratigraphies from a number of sites and care-
fully radiocarbon dating those stratigraphies, it is possible to trace the northward
colonization of tree species that occurred following the close of the Pleistocene.
Reconstructions of postglacial plant migrations have been carried out for a number
of North American and European tree species and genera.
Pollen analysis provides one of the only ways in which we can study the popula-
tion dynamics and geographic patterns of naturally colonizing plant species. This is
because many of these events occurred thousands of years ago and took hundreds
to thousands of years to complete. Written records do not exist for these important
biogeographical events. However, it is recognized that the fossil pollen record is an
imperfect one. For example, some trees, such as pines, produce great amounts of
pollen that can be carried far from the parent tree. Pine pollen can be found at sites
that lay far beyond the range limits of the trees. Alternatively, some trees, such as
firs, produce small amounts of heavy pollen and may not be detected in pollen
records from sites around which the trees are present. Finally, it is not known if the
relationship between tree population size and the amount of pollen deposited in
lakes and bogs is linear. Thus, reconstructions of population growth based on pollen
records are best considered approximations.
from studies of reindeer (Rangifer tarandus) populations on the Pribilof Islands of
the Bering Sea. In 1911,4 male and 21 female reindeer were introduced to St. Paul
Island in order to develop herds for food, clothing, and export. The initial growth
of the population was exponential, and by 1938 the island supported more than
2000 reindeer. Then, with no prior evidence of logistic decline in population
growth rate, there was a massive and continued decline until only eight reindeer
remained in 1950. The carrying capacity of reindeer on the islands has remained
very low. The decline appears to have been prompted by overgrazing by the rein-
deer and sharp declines in lichens and plants. In a sense, the introduction and
rapid population growth of the reindeer caused a catastrophic disruption of the
entire island's ecosystem. Such exponential growth followed by a precipitous
240 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

TABLE 8 . 1 The Postglacial Population Growth which have tl

Rates (as Expressed as Doubling slower rates o
lime) of Some Tree Genera at
Various Sites in North America 8
I FFUSION VERS
Tree Genus Doubling Time (Years)

Fir (Abies) 408 In many case~

Pine (Pinus) 80-1100 adjacent area
Douglas fir (Pseudotsuga) 52-365 form of range
Hemlock (Tsuga) 31-239 be more prop
Beech (Fagus) 124-444 perse seeds cl
each generati1
8 Estimates are calculated from fossil pollen data.
Source: MacDonald, 1993. provided by
United States
decline down to carrying capacity has been shown in a number of laboratory anc maximum, me
field-based studies of colonizing organisms. The phenomenon seems prevalent i.:: in what is nov
species that have significant overlaps in generation and where breeding aged indi- the retreat of
viduals can survive the initial onset of resource decline. northward. u~
Many factors can work to impede colonization. In some instances, unfavor- cal Box 8.1),
able physical environmental conditions cause mortality or reproductive failure. canadensis), d
For example, coconut palms cannot survive frost. Although on occasion coconuts years ago (Fi~
may be carried to high-latitude beaches, they will not germinate, or the seedlin= perate tree sp
will be killed at the first frost. The same fate awaits tropical birds or insects the.: and range frm
are blown to high-latitude locations. Coral larvae are widely dispersed as plank- diffusion was
ton in oceanic currents; yet most corals do not grow in waters that fall belo\\- sian were like
180 C in temperature, and many of these larvae end up in waters in which the\· dispersal.
cannot survive. Biological factors can also work against successful colonization. In anim<
The arriving species may be confronted by a superior competitor and not be able banded armad
to survive. Many laboratory studies show that rates of population growth declin the southwest
significantly in colonizing populations in the presence of a competitor. Plant- taken about 11
eating insects are often highly stenophagous and may not survive because thei.:' The diffusion '
food plant is not present. Finally, some species may face predators or pathogellS During 1
against which they have no defense. short generati
What determines the general probability that an organism will be able to The Europear
successfully colonize a new area? Some organisms seem particularly well suitec 1980, the speci
for rapid dispersal and successful colonization. These species are sometimes Alaska. The a
referred to as supertramps. The characteristics that enhance the ability of species 40,000 m per
to disperse and colonize new habitat include widely dispersed propagules, rapid species that "'
population growth rates, and ability to survive and reproduce in a wide range o:: century. It spr'
environmental conditions and resources (in other words possessing a generalis: to that of the ~
niche). Many supertramp species also appear adept at colonizing disturbed sites. In some
The Eurasian dandelion (Taraxacum officinale) is an excellent example of a expansions al1
supertramp. This annual plant has wide climatic tolerance, thrives in a number of lished thousar
soil conditions, matures and sets seeds in one growing season, and has small events are kn<
plumed seeds that are easily carried by wind or affixed to fur and clothing. It is perse in this rr
often inadvertently transported by humans. Thanks in part to human dispersal. importance ar
the dandelion is now found on every continent except Antarctica. Surprisingly. the plants anc
the propagules of many supertramps are spread by passive dispersal rather than organism spre
active dispersal. An analysis of dispersal and colonization rates, compared to small wind-blc
body size and maximum speed of active dispersal, showed that large animals. by flying orga

DIFFUSION VERSUS J U MP DISPERSAL
number of laboratory anc
menon seems prevalent iL
where breeding aged indi-
1 some instances, unfavor-
ty or reproductive failur e.
mgh on occasion coconuts
germinate, or the seedlin ~
)pi cal birds or insects tha-
Nidely dispersed as plank-
in waters that fall belo"'
1p in waters in which th e ~
st successful colonization
ompetitor and not be able
Jopulation growth declin"'
:e of a competitor. Plant -
not survive because thei:-
:e predators or pathog e ~
1 organism will be able tc
m particularly well suite ·
;e species are sometim e ~
tance the ability of spe ci e ~
ispersed propagules, rapi ·
,roduce in a wide range o:
·ds possessing a generalis:
colonizing disturbed site
1 excellent example of c:
1ce, thrives in a number o:
ng season, and has smat.:
d to fur and clothing. It ~
. part to human dispers al.
t Antarctica. Surprising]\.
;sive dispersal rather thar:
~ation rates, compared tc
wwed that large animals.
DIFFUSION VERSUS JUMP DISPERSAL 241
which have the potential for active dispersal over large distances, often have
slower rates of dispersal and colonization than small passive dispersers.
In many cases, species expand their geographic ranges by dispersing into closely
adjacent areas. Each generation expands the range of the species slightly. This
form of range adjustment is sometimes referred to as slow penetration but should
be more properly thought of as diffusion. Diffusion is typical of plants that dis-
perse seeds close to the parent and rely on the germination and reproduction of
each generation in order to expand into favorable new habitat. One example is
provided by the northward expansion of tree species into the northeastern
United States and adjacent Canada during the Holocene. During the last glacial
maximum, most of the tree species typical of the northern deciduous forest lived
in what is now the southeastern United States. With the warming of climate and
the retreat of ice at the end of the Pleistocene, these species expanded their range
northward. Using fossil pollen to reconstruct their northward expansion (Techni-
cal Box 8.1), we can show that some species, such as eastern hemlock (Tsuga
canadensis), did not reach their northwestern range limits until a few hundred
years ago (Fig. 8.7). The rates of range expansion exhibited by boreal and tem-
perate tree species in eastern North America during the Holocene varied greatly
and range from <100m per year to 400 m per year (Table 8.2). It is clear that this
diffusion was slow in some instances and quick in others. The rates of tree diffu-
sion were likely controlled by rates of Holocene climatic change and propagule
dispersal.
In animals, an excellent example of slow diffusion is provided by the nine-
banded armadillo (Dasypus novemcinctus). Spreading naturally from Mexico into
the southwestern United States and introduced into Florida, the armadillo has
taken about 100 years to expand its range from the Mexican border to Arkansas.
The diffusion of the armadillo is an example of slow penetration diffusion.
During diffusion events, animal species with high dispersal abilities and
short generational times can expand their geographic ranges extremely quickly.
The European starling (Sturnus vulgaris) was introduced in New York in 1891. By
1980, the species had expanded its range in North America to the Pacific Coast of
Alaska. The average rate of range expansion for the starling was greater than
40,000 m per year. The house sparrow (Passer domesticus) is also a nonnative
species that was introduced to North America from Europe in the nineteenth
century. It spread across North America between 1852 and 1910 at a rate similar
to that of the starling.
In some cases, dispersal and colonization events do not occur as gradual
expansions along the range limits of a species. New populations can be estab-
lished thousands of kilometers away from the range limits of the species. Such
events are known as jump dispersals. If species did not possess the ability to dis-
perse in this manner, remote islands such as Hawaii would be devoid of life. The
importance and frequency of successful jump dispersal events are evident from
the plants and animals that are found on remote islands. The probability of an
organism spreading by jump dispersal is generally high for plant species with
small wind-blown seeds and for flying animal species. Propagules that are carried
by flying organisms are also prone to jump dispersal events. It is not surprising,
 then, that re
mals except
Atlantic Oce
over a very g
• the Atlantic
• westward in
expanded it
1960s. TheN
that the catt
New Zealan
part attribu
Europeans.
domesticate
Arece
riers by plan
toa followin
1883. The isl
eruption, de
A recent an
that sea-disf
nized the is
Present range of rates, and ne
Tsuga canadensis eruption. M:
12 14
C yr. BP (x 103 ) date of that grow ir
Tsuga canadensis arrival anemochon
• Pollen sample sites colonize. Zc
are still arri'
FIGURE 8. 7 Example of apparent diffusion dispersal on a continental scale. The natural enced by th(
spread of eastern hemlock (Tsuga canadensis) in North America during the postglacial the island f,
(after MacDonald, 1993, adapted from Davis et al., 1986). such as vegE
Their
chores, and
nizing speci1
TABLE 8.2 The Apparent Postglacial Range of colonizat
Expansion Rates for Tree Genera propagules
in Eastern North America 8
sequent arri
Tree Genus Migration Rate (meters per year) plants that 1
probability
Fir (Abies) 159-200 chance eve1
Larch (Larix) 189 species prol
Spruce (Picea) 141-250 populations
Pine (Pinus) 81-400 been able tc
Hemlock (Tsuga) 200-250 colonizers. r

Maple (Acer) 126-200 events requ

Birch (Betula) 212 cal envirom
Beech (Fagus) 169-200 Jump
Oak (Quercus) 126-350 expanses o
Elm (Ulmus) 134-250 large areas
• The estimates are calculated from fossil pollen data.
propagules
Source: MacDonald, 1993. population
 DIFFUSION VERSUS JUMP DISPERSAL 243

then, that remote islands such as New Zealand or Hawaii have no native mam-
mals except for bats. The dispersal of the cattle egret (Bubulcus ibis) across the
Atlantic Ocean from Africa to South America is an example of a jump dispersal
over a very great distance across a barrier. Interestingly, the jump dispersal across
the Atlantic was followed by the diffusion dispersal of the egret northward and
westward into South and North America. More recently, the cattle egret has also
expanded its range to New Zealand, becoming established there in the early
1960s. The New Zealand populations arrived from Australia. Although it appears
that the cattle egret crossed from Africa to the New World and from Australia to
New Zealand on its own, its successful establishment in these new regions is in
part attributable to land clearance and the introduction of cattle and sheep by
Europeans. Cattle egrets feed on insects typically found in fields that support
domesticated livestock.
A recent example of the efficiency and pattern of jump dispersal across bar-
riers by plants and animals is provided by the revegetation of the island of Kraka-
toa following the destruction of all life there by a massive volcanic eruption in
1883. The island lies about 40 km from other islands. Yet, within 60 years after the
eruption, close to 300 plant species and 30 bird species had colonized Krakatau.
A recent analysis of plant dispersal strategies for the plants on Krakatau shows
that sea-dispersed plants had the highest initial colonization rates and fully colo-
nized the island within about 40 years. Anemochores had slower colonization
Present range of rates, and new species were still being added to the island flora a century after the
Tsuga canadensis eruption. Many of the first anemochores were species of herbs, grasses, and ferns
12 14
C yr. BP (x 103) date o that grow in open conditions. As forest cover has formed on the island, other
Tsuga canadensis arriVE anemochores such as orchids, which require moist and shady conditions, began to
• Pollen sample sites colonize. Zoochores had the slowest initial colonization rates, and new species
are still arriving on the island. The slow rate of colonization by zoochores is influ-
continental scale. The natu re enced by the fact that the birds and bats that carry these seeds will likely not visit
·ica during the postglacial the island for any appreciable amount of time until environmental conditions,
such as vegetation cover, provide suitable habitat.
The increasing species diversity of the island for thalassochores, anemo-
chores, and zoochores follows a logistic pattern with rapid initial increase in colo-
nizing species and then a flattening out of colonization events. The logistic pattern
of colonization probably reflects two factors. First, species with readily dispersed
propagules and populations close to Krakatoa dispersed there very quickly. Sub-
sequent arrivees were likely representative of less efficiently dispersed species or
plants that had parent populations located further away. Such species had a lower
probability of reaching Krakatoa, and greater amounts of time elapsed before a
chance event carried their seeds to the island. Second, the early-arriving plant
species probably faced little competition and were able to establish reproducing
populations relatively easily. Some of the species that arrived later may not have
been able to establish populations on the island because of competition from early
colonizers. The revegetation of Krakatoa shows us that successful jump dispersal
events require both the arrival of the propagule and suitable physical and biologi-
cal environmental conditions for establishment.
Jump dispersal events are not always restricted to species crossing large
expanses of water or other barriers. Sometimes rapid range expansion over
large areas of suitable habitat can occur via jump dispersal. In these cases,
propagules are dispersed over areas of suitable habitat and establish disjunct
populations beyond the main diffusion front. Such events are important for
244 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

colonization or invasions because these small disjunct populations serve as

inoculation centers from which the colonizing or invading species can ther:
expand. One example of continental jump dispersal comes from the early
Holocene spread of beech (Fagus grandifolia) in eastern North America. A:
the end of the last glacial maximum, beech spread northward from the south-
ern United States. A detailed examination of many fossil pollen records sug-
gests that beech spread rapidly across eastern North America between 10,00C
and 8000 years ago through the establishment of small, disjunct population
The size of these populations grew, and they coalesced as beech became a
dominant tree in the deciduous forest during the middle Holocene. Beech is a
zoochore, and its rapid and extensive jump dispersal in the early Holocene
may have been facilitated by the transport of beech nuts by the passenge:-
pigeon. A similar pattern of postglacial jump dispersal is evident for lodgepole
pine (Pinus contorta) in western North America.
The difference between diffusion and jump dispersal in environments tha: 5(

do not contain large barriers can often be more a matter of scale than anything
else. A detailed analysis of the spread of many invading species generally show: ~4(
E
that small disjunct populations are often established in advance of the main dif- ~

fusion front. A good example of this comes from the spread of the house finch ~ 3(
c
(Carpodacus mexicanus) in eastern North America. The birds are native to west- "'
u;
~ 2(
ern North America. The first house finches in the east were released on Long
Island in the 1940s. By the mid-1970s, the species had spread from Massachusetts
"'
'0
a:"' 1c
to North Carolina. A careful and spatially detailed analysis of bird count records
shows how the spread of the finch was often facilitated by jump dispersal and the
establishment of small disjunct populations in advance of the main diffusion
front (Fig. 8.8).
In analyzing the diffusion and continental jump dispersal for a number of
FIGURE 8.8 T
different species, some general patterns can be identified. In particular, when the
diffusion in easte
geographic range expansion of a colonizing species is graphed relative to time. Kawasaki, 1997).
the form of the relationship is often a logistic curve. This can be seen for both ani-
mals and plants and for diffusion and jump dispersals. The northwestward expan-
sion of the American muskrat (Ondatra zibethica) following its accidental
introduction near Prague, Czech Republic, provides an excellent example of slow
initial spread followed by rapid expansion and then gradual slowing. Following its animals, high c
introduction in 1905, the range expansion of the muskrat in Europe was very slow dispersal as i
diffusion. The rate of expansion increased between 1920 and the 1950s and then instances, the i
slowed greatly as the species approached its climatically controlled range limits for the specie~
in 1963. In addition, increased trapping programs likely contributed to the decreas- when the spec
ing rate of expansion. Cheat grass (Bromus tectorum) was accidentally intro- reproduction.
duced to western North America in the late nineteenth century. Through a series fact that Texa
of jump dispersal events, aided by the transport of its seeds in feed and with live- Rates of range
stock, it spread throughout the West by the 1930s. Although it continues to fill in the species m
portions of its western range, cheat grass intially expanded its range in a logistic decline as th(
manner between 1900 and 1930 (Fig. 8.9). approached. T
The logistic pattern of range expansion has three phases. First, the initial tions near the
rate of range expansion is often relatively slow because the small initial popu- growth and pr
lation size of the colonizing species limits propagule production and dispersal. be lower, an1
As the size of the colonizing population increases, propagule numbers climb decreases. The
and expansion rates increase. The initial slow spread of the house finch in the Canada and tl
vicinity of New York City likely reflects this principle. In actively dispersing these environr
 DIFFUSION VERSUS JUMP DISPERSAL 245

met populations serve c:s

nvading species can th e::.
;al comes from the earl'.
1stern North America. A:
orthward from the soutt -
fossil pollen records sug-
t America between 10,00:
nall, disjunct populations.
=seed as beech became ~
ddle Holocene. Beech is ~
;al in the early Holocec::
ch nuts by the passenge::-
al is evident for lodgepo:::

~
-=-2. 1973
ersal in environments the.: 500 r - - - - - - - . ,
tter of scale than anythir:.; 10
ng species generally show-o ~400
E U)

n advance of the main cL:- 6 ·c:Ql

0
spread of the house fin e::. ~ 300 0l.)
~ ..~
1e birds are native to wes::- U)
2 5
'6 200
lSt were released on Lor.;: Cii
'6 0
"'
U)

;pread from Massachuse r..! "' ci

a: 100 z

1lysis of bird count recorc
I by jump dispersal and t:::::
nee of the main diffusic::.
dispersal for a number c:
1ed. In particular, when t::.:'
; graphed relative to tilL =
is can be seen for both a~­
The northwestward expa=-
) following its accid e n ~
t excellent example of s!o-
tdual slowing. Following ==
at in Europe was very s!o-
120 and the 1950s and the::.
tlly controlled range Iilli=
contributed to the decree5-
n) was accidentally intr;:-
h. century. Through a sef. =-:.
;eeds in feed and with liY::-
lough it continues to fill =
nded its range in a logis:::
ee phases. First, the initi.:....
use the small initial por :.-
production and disperse
propagule numbers cl i.c:
I of the house finch in r::::
pie. In actively dispersi=.;
II nrrt~l~b
~
0 • r- 1 1 1 0 r 1 I '
1950 1960 1970 1980 0 100 200 300 400
Years Distance from primary range (km)
FIGURE 8.8 The spread of the house finch (Carpodacus mexicanus) by jump dispersal and
diffusion in eastern North America (after Mundinger and Hope, 1982, and Shigesada and
Kawasaki, 1997). The mean jump distance for establishment of disjunct populations is 115 km.
animals, high densities at the point of introduction may further promote active
dispersal as individuals move to avoid intraspecific competition. In some
instances, the initial point of colonization may not be the optimal environment
for the species. Rates of population growth and range expansion can increase
when the species reaches areas that provide better conditions for growth and
reproduction. The rapid spread of the armadillo across Texas may reflect the
fact that Texas provided better habitat than the desert of northern Mexico.
Rates of range expansion generally increase after population size builds up and
the species moves into favorable habitat. In many cases, rates of expansion
decline as the environmentally controlled range limits of the species are
approached. This decline probably reflects unfavorable environmental condi-
tions near the range limits of the species that cause slower rates of population
growth and propagule production. In addition, survival rates of offspring may
be lower, and the chance of successful establishment and colonization
decreases. The slow rate of range expansion for the starling in northwestern
Canada and the armadillo in northeastern Texas likely reflects the fact that
these environments are not optimal for these species.
246 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION BARF

ians. Most speci

with salt water.
and colonize i~
species will thri
species (Salmo
fish and do ver~
insurmountablE
(Lepomis spp.)
These genera w
mainland Nortl
of these fish trm
well. Isolated isl
amphibian spec
to cross. The frc
those found on
(f)
Land area
100 Many marine s
% of eventual range
Even small lane
75
Isthmus of Pan
species are restr
50
the physical ba
25 platurus, which
eastern Pacific 1
Caribbean Com
1910 1920 193 Environm
FIG U RE 8.9 The logistic expansion of invading species spreading by either diffusion o· ation can also p
jump dispersal as displayed by cheat grass (Bromus tectorum) expansion in western Nor;:- barrier for the 1

America (after Mack, 1981; Shigesada and Kawasaki, 1997). Sahara Desert
ranean region a1
of the Mediterr
Saharan Africa.
BA RRIERS, CORRIDORS, FILTERS, ST EPPING separated from
STONES, AND SWEEPSTAKES America by the
found in Califor
Biogeographers often classify geographic features in terms of their impact on d~ such as Douglas
persal and colonization. Geographic features that block dispersal and colonizc.- ern North Arne
tion are called biogeographic barriers. In contrast, geographic features th-· posed by the dr
promote dispersal and colonization are called biogeographic corridors. As w~ cold winter dim
will see, the terms barrier and corridor are very relative in nature. For exampk the north.
large water bodies may serve as barriers to land organisms, but may also act cs Within the
corridors that allow the dispersal of aquatic species. When the term barrier ~ rise above the 1

used in a biogeographic sense, it must be related to specific species or groups c:' small mammal f
species. A barrier for one species is often a corridor for another species! munks that cann
Large expanses of water are a barrier to the dispersal of most terrestric... of these mounta
organisms. Very few reptile or mammal species native to Europe and Asia ar.: meters serve as
native to North America, despite similarities in the physical environments avaL.- montane and air
able on both continents. The tree species found in North America are also diffe-- during glacial p
ent from those found in Asia and Europe. Finally, even among land birds anc cooler than it is
bats, relatively few species are native to both Eurasia and North America. OceaiE Mountains
also serve as important barriers to the dispersal of freshwater fish and amphib- tions have cool1

(f)
00 .
% of eventual range
75
50
25
1910 1920
reading by either diffusion
7) expansion in western N o r~-
....:II
erms of their impact on ~
JCk dispersal and coloniz.c.
, geographic features L::.:.
!ographic corridors. As ~:
ive in nature. For exam c ~~
:misms, but may also ac. _
. When the term barrier _
pecific species or groups
)r another species!
ispersal of most terrestr:
ve to Europe and Asia c.:-:
hysical environments a,·c._
rth America are also dit:;:-
ven among land birds c::__
md North America. Ocec.=
·eshwater fish and ampl-.·-
BARRIERS, CORRIDORS, FILTERS, STEPPING STONES, AND SWEEPSTAKES 24 7
ians. Most species adapted to life in freshwater cannot tolerate prolonged contact
with salt water. Few freshwater fish species ever disperse across ocean barriers
and colonize islands and other continents. We know that in many cases, fish
species will thrive in new regions if introduced by humans. North American trout
species (Salmo spp.) , for example, have been introduced to New Zealand as game
fish and do very well there. However, even small expanses of salt water can pose
insurmountable barriers to freshwater fish. Bass (Micropterus spp.) and sunfish
(Lepomis spp.) are extremely abundant in lakes and streams in North America.
These genera were not, however, able to cross the relatively short distances from
mainland North America to the islands of the Caribbean Sea. Again, populations
of these fish transplanted to the Caribbean Islands by humans have survived very
well. Isolated islands, such as New Zealand and Hawaii commonly have no native
amphibian species. Even short expanses of salt water are difficult for amphibians
to cross. The frog species found on islands in Indonesia are far less diverse than
those found on nearby mainland Asia.
Land areas form barriers for the dispersal of marine and freshwater species.
Many marine species found in the Pacific Ocean do not occur in the Atlantic.
Even small land areas can pose insurmountable barriers to aquatic animals. The
Isthmus of Panama is less than 100 km wide in some places, yet many marine
species are restricted to either the Pacific Ocean or the Caribbean Sea because of
the physical barrier of the isthmus. Even the poisonous marine snake Pelamis
platurus, which can survive for short periods out of water, is found only on the
eastern Pacific Coast and has not been able to disperse overland to the nearby
Caribbean Coast.
193 Environmental conditions vary across landmasses and oceans, and this vari-
ation can also pose a barrier to dispersal. For example, expanses of desert pose a
barrier for the dispersal of mesic plants and moisture-demanding animals. The
Sahara Desert in Africa is an important barrier between the circum-Mediter-
ranean region and central and south Africa. The northern African flora and fauna
of the Mediterranean coast have great affinity to Europe and much less to sub-
Saharan Africa. The relatively moist areas of coastal and montane California are
separated from the mesic habitat of the Rocky Mountains and eastern North
America by the Mojave and Great Basin deserts. Over 1500 species of plants
found in California are not found in areas east of the state. Western tree species,
such as Douglas fir (Pseudotsuga menziesii) grow very well when planted in east-
ern North America, but their populations cannot naturally cross the barriers
posed by the dry deserts and grasslands in central North America, or the very
cold winter climate and short summers of the boreal forest and tundra biomes to
the north.
Within the Great Basin of the United States, a number of mountain ranges
rise above the desert and provide mesic montane and alpine conditions. The
small mammal fauna of these ranges includes species such as squirrels and chip-
munks that cannot survive desert conditions. Analysis of the small mammal fauna
of these mountains shows that desert distances of a few tens to hundreds of kilo-
meters serve as a barrier to the dispersal of these species. It is thought that the
montane and alpine mammal species dispersed to these isolated mountain ranges
during glacial periods when the climate of the Great Basin was moister and
cooler than it is today.
Mountains are a particularly interesting form of barrier. High-altitude loca-
tions have cooler temperatures than adjacent low-elevation sites. In addition,
248 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION BAR I

mountainous regions are often moister than nearby lowlands. Since many spe ·:: between North
of plants and animals are extremely sensitive to cold temperatures, high mo --- sal biogeograpl
tain ranges can be effective barriers to migration and dispersal. However, -.. = Between
degree to which mountains serve as a barrier is also related to latitude. In hi: -- South America
latitude locations, winter temperatures are cold in both the mountains and on ·- = Isthmus of Pan
adjacent lowlands. Consequently, the flora and fauna at the higher latitudes m of about 150 m
be adapted to withstand cold conditions regardless of whether they live in ·-:: tinents preclud
mountains or the lowlands. In the tropics, however, there is much less seasOI:.:.. and flightless b
variation in temperature, and lowlands are generally always warm. High e]e\: ity along the P'
tions may warm up slightly during the summer but in general remain relatiYe_ Cocos Plate to
cool throughout the year. Tropical lowland flora and fauna are often incapable continued rise<
withstanding cold temperatures, and even in the summer the mountains may r: a continuous Ia
be warm enough to allow survival of lowland organisms. Mountains in the tropi the Isthmus of
therefore, are greater barriers than high-latitude mountain ranges. and terrestrial ·
In concluding our consideration of barriers, it is important to note that bc:- trial fauna that
riers to dispersal and migration can be biological as well as physical. Stenophago_ the Great Arne
predators cannot survive in areas where their prey species are lacking, even if ~= For them<
physical conditions are satisfactory for the predator. Intense competition fr = from other con
other species can also serve as a barrier to colonization. Animal behavior -- and colonizers J
present yet another barrier to dispersal. A number of tropical bird genera incl :- facilitated by th
ing cotingas, toucans, and ant birds have been shown to avoid crossing even \. :- subsequent elm
short distances of open water, despite the presence of suitable habitat only a f American man
hundred meters away. dominated by n
As mentioned earlier, areas that promote dispersal and colonization a:-:: exchange of rna
called corridors. A river joining two lakes would be a corridor for the dispersal -- mals such as lla
aquatic organisms, while an isthmus joining two continents would serve a si~ the modern So
role for terrestrial plants and animals. According to its strict biogeographical de:: North America.
nition, a corridor allows the unrestricted movement of all taxa from either side of =-- the south. Asidt
Over time, dispersal and colonization along such a corridor lead to the harmoniz.c.- (mammals that
tion of the flora and fauna on both sides. Biogeographical harmonization mea::: cophaga) and g
that similar species of flora and fauna are found on both sides of the corrid :- that stood abou
According to this definition, land corridors are continuous areas of similar clima:=: appears that N•
and vegetation. Thus, some extensive biomes, such as the boreal forest of northe::::. South America
North America or northern Eurasia, form corridors. In the case of the North Arne:-- ward. It may h
ican boreal forest, the flora and fauna found in Alaska are indeed very similar superior to the ~
the flora and fauna found across the continent in eastern Canada. Mountain rang :: When the
can serve as corridors for alpine species. Many of the same species of plants ar:: and open savan
animals found in the southern Rocky Mountains of Canada occur as far south c by many differe
the Rocky Mountains of Colorado. developed in 1<
Obviously, in many instances geographic connections, such as rivers or is - allowed dispers
muses, link two regions, but because of physical or biological conditions, all speci : tion of North 1
cannot disperse with equal ease. Avenues of dispersal and colonization which ar~ tropical forest c
not equally favorable for all species are called filters. When filters exist, they allo Tropical forest
some species to cross and restrict others. Physical environmental conditions, sue: desert species c;
as temperature, may restrict some species. In addition, biological factors along th• northern portio
filter route, such as competition with other species, may also serve to restrict th- northward mov
dispersal and colonization of some species. The end result is that the flora anc the dispersal of
fauna on both sides remain distinctive. The Isthmus of Panama, linking North an - and Eurasia, re
South America, is an excellent example of a modern biogeographic filter. The for- been unable to
mation of the Isthmus of Panama and the biogeographic impact of this landbridge and colonize Sc

viands. Since many specie ~
temperatures, high moun·
d dispersal. However, the
~lated to latitude. In high-
the mountains and on the
t the higher latitudes mu
f whether they live in the
tere is much less season ~
always warm. High eleva-
generalremain relative! ~
una are often incapable o:
er the mountains may no:
;. Mountains in the tropics.
tain ranges.
mportant to note that bar-
as physical. StenophagotE
:ies are lacking, even if th-
Intense competition frorr:
ion. Animal behavior ca.:.
:apical bird genera indue-
) avoid crossing even ve r:
mitable habitat only a fe
Tsal and colonization ar.:
)rridor for the dispersal o:
ents would serve a similc..:-
;trict biogeographical def-
1taxa from either side of i:..
dor lead to the harmonizz-
tical harmonization mea~
Joth sides of the corrido:
ms areas of similar climat.:
e boreal forest of norther::.
1e case of the NorthAme:-
are indeed very similar · -
L Canada. Mountain rang~
;arne species of plants an::
nada occur as far south c
ions, such as rivers or istt-
gical conditions, all spe ci~
md colonization which ar.:
hen filters exist, they allov
·onmental conditions, sue:::.
'iological factors along th.:
y also serve to restrict th.:
esult is that the flora an;:
>anama, linking North an;:
)geographic filter. The fo:-
c impact of this landbridg.:
BARRIERS, CORRIDORS, FILTERS, STEPPING STONES, AND SWEEPSTAKES 249
between North and South America is one of the most interesting topics in disper-
sal biogeography and is well worth our detailed consideration.
Between 16 million and 3 million years ago, the continents of North and
South America lay at about their modern geographic positions. In place of the
Isthmus of Panama, there was a series of islands set upon a relatively shallow sea
of about 150m in depth. The lack of a continuous land bridge joining the two con-
tinents precluded the exchange of most terrestrial fauna, particularly mammals
and flightless birds. The islands were formed by displacement and volcanic activ-
ity along the point of contact between the Caribbean Plate to the east and the
Cocos Plate to the west. The ongoing collision between the two plates led to the
continued rise of land in the area of the Isthmus until 2.4 million years ago when
a continuous landbridge was formed between North and South America. Once
the Isthmus of Panama was in place, it facilitated the dispersal of land animals
and terrestrial plants north and south along its length. The movement of terres-
trial fauna that occurred following the establishment of the isthmus is known as
the Great American Exchange.
For the most part, mammal species in South America had evolved in isolation
from other continents, while those in North America represented native families
and colonizers from Eurasia. Exchange between North America and Eurasia was
facilitated by the late separation of northern Europe and North America and the
subsequent close proximity of Asia and North America at Beringia. The North
American mammals were placental, whereas South America's mammals were
dominated by marsupials. When the Isthmus was formed, there was a considerable
exchange of mammals between the two continents (Table 8.3). Interestingly, mam-
mals such as llamas and jaguars, which we typically think of as representatives of
the modern South American fauna, are actually derived from colonizers from
North America. The opossum, common in North America today, is a colonizer from
the south. Aside from the opossum, there were many other interesting marsupials
(mammals that carry their young in a pouch), such as giant anteaters (Myrme-
cophaga) and giant sloths (Eremotherium), and a giant carnivorous bird (Titanis)
that stood about 4 m tall that came northward across the landbridge. However, it
appears that North American mammals were far more successful in colonizing
South America than South American mammals and birds were in moving north-
ward. It may be that the North American-Eurasian species were competitively
superior to the South American species, which had evolved in relative isolation.
When the isthmus first developed, it contained a landscape of both forest
and open savanna and functioned, more or less, as a corridor allowing dispersal
by many different species. Howder, during the Pleistocene, dense tropical forest
developed in lowland Central America, and the isthmus became a filter that
allowed dispersal by tropical forest species but restricted the southward migra-
tion of North American grazing fauna such as bison and antelope. The dense
tropical forest of the modern southern Isthmus still serves as a filter to dispersal.
Tropical forest animals can traverse it, but temperate, grassland, savanna, and
desert species cannot. The more xeric savanna and desert conditions found in the
northern portions of Central America and Mexico in turn serve as barriers to the
northward movement of tropical species. The Isthmus also serves as a filter for
the dispersal of plants. Pines, which are widely distributed across North America
and Eurasia, reach their modern range limits in Nicaragua and appear to have
been unable to cross the lowland tropical environment of the southern isthmus
and colonize South America. Indeed, even humans are daunted by this tropical
250 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION BAR

TABLE 8.3 Some of the Most Commonly Known Mam- gs 30

mal Families of the Great American Faunal ·c::;
Exchange between North and South America 2i
C/)
25
via the Isthmus of Panama ~ 20
0,
Origin Family Common Name ~ 15
::2
North America 0 10
Cii
Leporidae Rabitts -g 5
Sciuridae Squirrels ::l

Cricetidae Field mice

z 0-+---
0
Felidae Cats

~
Mustelidae Skunks and otters
Canidae Foxes
Ursidae Bears
Equidae Horses
Camelidae Camels and ilamas
NewG1
Cervidae Deer
South America
Didelphidae Opossums
Dasypodiadae Armadillos
Megatheriidae Giant ground sloths FIGURE 8.10
Bradypodidae Three-toed sloth South Pacific (da'
Myrmecophagidae Anteaters colonize the mos
Cebidae Monkeys
Erethizontidae Porcupines
Caviidae Guinea pigs Adjacent
Source: Webb, 1997. dispersal of me
tains of the Un
over 2000 m pr,
Lower elevatio
forest barrier-there is still no paved road connecting South and North Ameri tane species. 11
due to the difficulty of traversing the Darien Gap in southern Panama and adj~­ gradually decli:
cent Colombia. northward. The
Chains of closely distributed islands form a special type of dispersal rou t~C desert and cor
known as stepping stones. Stepping stones are always filters mainly because no: mammal specie
all species of flora and fauna can cross the gaps between them. The islands o:": Some dis1
Indonesia and the Philippines provide such stepping stones for the dispersal o:' ules in air or Wl
flora and fauna from Asia. A number of bird species from Asia are found in the colonization of
Philippines or eastern Indonesia. The amphibian and reptile fauna originating i:: Most will perisl
Asia, however, is very poorly represented away from the mainland and adjacen: colonization. 11
islands. Similarly, the proportion of reptile and insect fauna from New Guinec dispersal, but a.
and Australia decreases sharply westward toward the Asian mainland. Ne..- new colonizabl
Guinea and the Melanesian Islands provide stepping stones for the dispersal a: ful, they are sor
Asian and Australian fauna eastward into the Pacific. The distribution of man- tant for the col
grove tree species in New Guinea and Melanesia shows the filter effect of this Waif dispersal i
stepping stone route (Fig. 8.10). A steady and marked decrease in the number a: small anemoch
mangrove taxa is encountered westward from New Guinea. Clearly, due either to ters, for not all
differences in dispersal and colonization abilities, or due to chance, some species Thus, the Haw;
have been able to disperse across the Pacific island stepping stones while others smaller insect f:
have not. lie closer to the
 BARRIERS, CORRIDORS, FILTERS, STEPPING STONES, AND SWEEPSTAKES 251

rn Mam- ~ 30
'(3
.c;(t ' ."· - ..

:r
1Faunal .

1 America
~ 25
en
~ 20
0,
1on Name ai 15
t '
.
:::2:
0 10
~
.0
5
E
::::l
liS
z 0
1ice I
0 800 1,600 2,400 3,200 4,000 4,600 5,200

\
km
; and otters

' Solomon
'\:,"'

; and ilamas
,, "" Islands
"'· ~\ 1
"" Samoa
Islands
"'~

Jms
illos
1round sloths
:oed sloth
'lrs
ys
ines
, pigs
~ South and North Ameri:::
;outhern Panama and ad':=-
cial type of dispersal ro c:.::
s filters mainly because r..
ween them. The islands -
stones for the dispersal ::c
from Asia are found in G::c::
reptile fauna originating = Most will perish, and a very small proportion may end up on a beach suitable for
the mainland and adjacc
;t fauna from New Guin.:_
the Asian mainland. t\e
. stones for the dispersal ~·
::. The distribution of mc..:_-
ows the filter effect of (
I decrease in the number c
uinea. Clearly, due either :-
lue to chance, some spec ;:-.
:epping stones while otbe.
~
0 Fiji
FIGURE 8 . 10 The impact of an island stepping stone filter on mangrove species in the
South Pacific (data from Woodroffe, 1987). The numbe r of species able to disperse to and
colonize the most distant islands is a fraction of the genera found in New Guinea.
Adjacent mountain ranges can also serve as stepping stone filters for the
dispersal of montane and alpine flora and fauna. In the southern Rocky Moun-
tains of the United States and the Sierra Madre Occidental of Mexico, elevations
over 2000 m provide relatively cool and moist montane and alpine environments.
Lower elevations are dominated by desert, which forms a barrier for the mon-
tane species. The proportion of northern forest and woodland mammal species
gradually declines southward, while the proportion of southern species declines
northward. The filter effect probably results from difficulties dispersing across the
desert and competition from the already established northern and southern
mammal species on the mountains.
Some dispersal routes are typified by the long-distance transport of propag-
ules in air or water currents, which provide a relatively small chance of successful
colonization of new regions. An example is the release of coconuts into the sea.
colonization. The planktonic larvae of coral reef fauna and flora depend on such
dispersal, but also face the probability that most will not succeed in dispersing to
new colonizable environments. When such long-distance dispersals are success-
ful, they are sometimes called waif dispersal events. Waif dispersal is very impor-
tant for the colonization of coral atolls and other shallow marine environments.
Waif dispersal is also important for the colonization of islands, such as Hawaii, by
small anemochorous insects and some plants. Waif dispersal routes are also fil-
ters, for not all organisms can survive the transport or colonize the new regions.
Thus, the Hawaiian Islands have much smaller coral reef fauna and flora and
smaller insect fauna than mainland areas of Asia or the Americas, or islands that
lie closer to those continents. Such relatively impoverished biota are sometimes
252 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

called waifflora or waif fauna due to the impact that isolation and dispersal co--
straints have on them.
Some routes of dispersal and migration only rarely allow successful dispe:--
sal and colonization. Crossing such routes occurs by chance and has a very 10'
probability. Such dispersal pathways are called sweepstakes routes. The chan ~
of an organism successfully dispersing by this route and colonizing a new area _
similar to the odds of a person winning a lottery or sweepstakes. However, O\·::
thousands to millions of years, some organisms will disperse across sweepstak=--
routes by unlikely events and be able to colonize new areas. The dispersal of ·--
cattle egret across the Atlantic to North America is an example of a sweepsta ·::-
route. The fact that even extremely remote islands, such as Hawaii in the Pac·--
and the Azores Islands in the Atlantic, have flora and fauna derived from a nu=-
0:100/145
ber of different families, not all of which are known waif dispersers, shows th--
despite the low probability of success, sweepstakes routes are important mod::-
of dispersal and colonization.

RECENT INVASIONS BY EXOTIC SPECIES

As we discussed earlier, the term biological invader is often used to refer :: FIGURE 8 . 1 1
exotic species that are introduced to new areas by humans. In some instanCA..~ offshore islands o
such introductions are made deliberately as in the case of agricultural crops, ga:--
den plants, and game species, but in most cases they are purely accidental. 11::.~
problem of invading plant and animal species became acute with the widesprea::
European exploration and colonization of the late fifteenth century. Prior to the.: is perhaps eve1
time, Eurasia and North America were relatively isolated from each other. c.:: plant species b1
were temperate parts of North and South America. Australia was isolated fro= invasions are n1
Eurasia, Africa, and the New World. The introduction of exotic species oft e~ species found 1
causes significant damage to native ecosystems. These invasive species can aL- other continen1
cause considerable problems for agriculture, forestry, and other resources us ::_ are invaders, w
by humans. Invading species often spread at low population levels and are difE- been introduce'
cult to initially detect. Once established, they then increase in a logistic manne: The impa'
making them difficult to detect and eradicate before their population grov. ._ ecosystem funct
accelerates and it is too late to stop the invasion. and its impact
The magnitude of biogeographic invasions over the past 500 years is st a ~:­ (Opuntia stricat,
gering. In most parts of the world, the modern flora and fauna contains large form hedges. Th
numbers of introduced species (Fig. 8.11). Some 2000 to 3000 species of exo ti- ble for humans
plants are known to grow in the continental United States and Canada. 'I1lis Americas, lacke
represents at least 10% to 25% of the flora. Approximately 50% to 70% of the prickly pear wa~
so-called weedy species that cause damage to agricultural crops in North Amer- free of predator
ica are introduced Eurasian plants. Central Europe is host to 100 invasi\·e cactus increased
species of woody plants alone. These trees and shrubs were brought there fro n: it was recognize
the Americas and Asia. The entire European continent is home to over 150C fields. By 1900, t
nonnative plant species. New Zealand supports 1500 species of alien vascul a: eastern Australi
plants, which now make up about 50% of its total flora. Within the Unitec to 243,000 sq k1
States, California is host to 1025 nonnative plant species. This is about 17% o: catastrophe. It v.
the total California flora. Plants commonly associated with the California land- ually. In additim
scape, such as the eucalyptus (Australia), Mexican fan palm (Mexico), Canary brings short-ter
Island pine (Canary Island), giant pampas grass (Argentina), and bamboo remain viable in
(China), are all introduced alien species. The situation in the Hawaiian Islands discovered that

isolation and dispersal con-
ely allow successful disper-
chance and has a very low
•stakes routes. The chances;
nd colonizing a new area is
weepstakes. However, over
.isperse across sweepstak e~
' areas. The dispersal of the
1 example of a sweepstakes
tch as Hawaii in the Pacifi-
fauna derived from anum-
waif dispersers, shows tha-
Jutes are important modes
' is often used to refer tc
mmans. In some instances.
e of agricultural crops, ga:--
are purely accidental.
acute with the widesprea;:
eenth century. Prior to thE:
Jlated from each other. ES
'\.ustralia was isolated froc
on of exotic species ofte=
e invasive species can alsc
, and other resources use::
ulation levels and are diff-
;rease in a logistic mann e~
e their population gro\,-·-
· the past 500 years is st a~­
t and fauna contains larg ::
0 to 3000 species of ex o r:~
j States and Canada. Th:.!
mately 50% to 70% of tb::
:ural crops in North Am e~ ­
'e is host to 100 invasi \·;:
s were brought there fr o=
tent is home to over 150;:
) species of alien vascu l ~
flora. Within the Unite.:
~cies. This is about 17% c:
d with the California lan" -
m palm (Mexico), Cana .
Argentina), and bambcx:
min the Hawaiian Islan-'-
RECENT INVASIONS BY EXOTIC SPECIES 253
··,
0:100/145
0 = Oceanic Islands
=
C Continents
S = Continental Islands
Mammals/Birds
FIGURE 8.11 Introduced mammal/bird species for the continents, coastal islands, and
offshore islands ofthe earth (after Ebenhard, 1988).
is perhaps even more striking. The islands have a native flora of about 1996
plant species but host about 800 additional species of alien plants. Large-scale
invasions are not just limited to plants. In California, 35% of the freshwater fish
species found today are introduced from other parts of North America and
other continents. In Hawaii, 40% of the bird species now found on the islands
are invaders, while 94% of the mammal species and 100% of the reptiles have
been introduced by humans.
The impact of introduced plants can be extremely disruptive to natural
ecosystem functioning and to human land use. A classic example of plant invasion
and its impact comes from Australia. In 1893 a species of prickly pear cactus
(Opuntia stricata) was brought from the United States to Australia and planted to
form hedges. The species forms dense thickets that are over 1 m high and impossi-
ble for humans or large animals to pass through. Cacti, which are native to the
Americas, lacked natural predators and parasites in Australia. In addition, the
prickly pear was well adapted to thrive in the semiarid climate of Australia. Being
free of predators and competitively superior to Australian plants, the prickly pear
cactus increased dramatically in numbers and geographic distribution. In 40 years,
it was recognized as a major pest that displaced native flora and destroyed grazing
fields. By 1900, the prickly pear had spread over an area of 40,000 sq km in south-
eastern Australia. By 1925, the area infested with prickly pear cactus had increased
to 243,000 sq km. The invasion of fields by prickly pear was a major economic
catastrophe. It was not economically feasible to poison or remove the plants man-
ually. In addition to the prohibitive cost of removing the cacti, such a strategy only
brings short-term relief. Prickly pear can resprout from its roots, and its seeds
remain viable in the soil for some 15 years. Fortunately, Australian entomologists
discovered that a moth species from Argentina called Cactoblastis cactorum
254 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

produced larvae that feed exclusively on the cacti. The larvae burrow into the ca -
tus, facilitating the entry of fungi and other pathogens. Starting in 1925, Cacra-
blastis cactorum were imported to Australia and introduced into areas where
prickly pear were widespread. By 1940 the prickly pear population had plum-
meted, and the species survives in Australia today only as scattered individua_
plants.
Australia has, in a way, repaid the United States for the damage caused b:
the prickly pear cactus. During the late nineteenth and early twentieth centurie-
the Australian eucalyptus (Eucalyptus spp.) was introduced and widely planted i;:.
California. At the same time, the Australian melaleuca tree (Melaleuca spp.) \Va5
introduced to Florida. The two genera were imported and planted as garde;:
specimens, windbreaks, and timber trees. In both states these introduced tree
genera lacked serious predators and pathogens. Wildfire is common in Australia
and both trees are adapted to frequent burning. They resprout easily from roo '
and grow quickly following fires. Melaleuca trees that survive fires produce ·
lions of seeds, and the young trees can grow as much as 1 m in a single year. Th
dense shade, thick leaf and bark litter, and oily residue found beneath eucalypt tc
and melaleuca stands often kill all understory plants. Eucalyptus and melaleu
have escaped from cultivation and become established widely. In California.
eucalyptus groves are particularly problematic because they replace oak-gra " FIGURE 8.12
woodlands and some coastal forest and shrubland communities. Little grO\\"S the post-fire ve~
beneath the eucalyptus canopy. Melaleuca has become a particular problem ir: dries out in the '
native species o
the Everglades National Park of Florida. The tree was introduced in the region as
a timber species and has become impossible to control. The trees have a pro-
found impact on native vegetation. Areas covered by melaleuca typically suppor:
only two or three native plant species. Similar areas, which do not have melaleu
stands, generally support 60 to 80 different native plant species. Both th Another
melaleuca and eucalyptus are particularly serious fire hazards. The trees contain (Tamarix pent
high amounts of volatile oils and burn quickly at very high intensities. A 1992 fir produces copi
in the eucalyptus-covered hills of Oakland, California, covered 650 ha, destroye introduced to·
over 3000 homes, and killed 25 people. In 1985 a melaleuca fire near Miami. all southwest ,
Florida, consumed over 3500 ha in a 35-day period. west as the Me
Perhaps the most insidious terrestrial plant invading North America at the been greater t
present time is cheat grass (Bromus tectorum). Some of the first reports of thi tation, each m
Eurasian plant in North America came from British Columbia, Canada, in 1889. has been estin
By 1930, cheat grass was reportedly growing as far south as Utah, Nevada, and soils as is used
California (Fig. 8.9). By the 1990s, cheat grass had expanded into southern Ari- sible to contro
zona and occupied a total of some 40 million ha. It is estimated that cheat grass i Invasive
still expanding its density of coverage within its range in western North America world. The An
at a rate of over 1800 ha per-year. Cheat grass is spread by fire and helps promote as an orname1
burning. In the summer at the end of its growing season, the plant dries and pro- and purple flo
duces a continuous cover of fine fuel. Following burning, cheat grass seeds into Victoria is the
burned areas vigorously. In areas formerly dominated by semiarid shrublands. people. By 19 1
the continuous cover of cheat grass promotes frequent and widespread fires. important fisl
which cause a decline in areas dominated by native shrubs such as sage brush entry to harbc
(Artemisia spp.) and the replacement of species-rich shrubland by cheat grass- tlements and J
dominated grassland (Fig. 8.12). The establishment of such grasslands in turn of huge propo
leads to a decrease in the species diversity of insects, birds, mammals, and reptiles. In Nortl
In addition to its negative impact on native flora , cheat grass is a common and introduced frc
destructive weed in wheat fields. in the 1950s. 1
 RECENT INVASIONS BY EXOTIC SPECIES 255

trvae burrow into the cac-

'· Starting in 1925, Cacto-
Jduced into areas wher
ar population had plum-
ly as scattered individual

·o r the damage caused b~

early twentieth centurie
ced and widely planted i;:;
tree (Melaleuca spp.) was
j and planted as garde
es these introduced tree
e is common in Australia_
esprout easily from roon:
;urvive fires produce
; 1 m in a single year. The
'ound beneath eucalyptl:S
:ucalyptus and melaleuc::
ed widely. In California.
e they replace oak-grass
)mmunities. Little grO\\ "S
~ a particular problem iL
ttroduced in the region as
Jl. The trees have a pro-
elaleuca typically suppor:
ch do not have melaleucc.
plant species. Both th""
tazards. The trees contai-
gh intensities. A 1992 fir;:-
:overed 650 ha, destroye -=
laleuca fire near Mia
ing North America at t
)f the first reports of this
>lumbia, Canada, in 1889
tth as Utah, Nevada, anc
anded into southern Ar: -
imated that cheat grass is
1 western North Americe.
Jy fire and helps prom or-
, the plant dries and pro-
lg, cheat grass seeds intc
by semiarid shrubland-
nt and widespread fir es.
1rubs such as sage brus
hrubland by cheat grass-
such grasslands in tur:;.
is, mammals, and reptiles.
t grass is a common an ·
FIG URE 8 . 12 Cheat grass (Bromus tectorum), an invasive plant from Eurasia, dominates
the post-fire vegetation in a recently burned site in southwestern Utah . Cheat grass dies and
dries out in the summer, providing fuel for frequent fires that eventually exclude many
native species of trees, shrubs, herbs, and grasses.
Another problematic invader in the western desert regions is the salt cedar
(Tamarix pentrandra). Salt cedar is an Asian desert shrub that grows quickly and
produces copious amounts of seeds that are dispersed by wind and water. It was
introduced to Texas in 1877. In less than 100 years, this invasive plant had spread to
all southwest desert floodplains, sweeping as far north as central Utah and as far
west as the Mojave Desert of California. The rate of expansion for the salt cedar has
been greater than 20,000 m per year. Aside from displacing native floodplain vege-
tation, each mature salt cedar can take 800 liters of water out of the soil each day. It
has been estimated that salt cedars now remove as much water from southwestern
soils as is used by all of the cities in southern California. The plant has proven impos-
sible to control and is found in even the most remote canyons of the Southwest.
Invasive water plants have caused significant problems in many parts of the
world. The Amazonian water hyacinth (Eichornia crasssipes) was taken to Africa
as an ornamental plant in the nineteenth century. It has beautiful floating foliage
and purple flowers. In 1990, escaped hyacinths were discovered in Lake Victoria.
Victoria is the second largest lake in the world, and its fishery supports millions of
people. By 1996, water hyacinth occupied 90% of the lake's shoreline, destroying
important fish-breeding areas. The plant grows thick enough to choke off the
entry to harbors for fishing boats. In addition, it clogs water intake pipes for set-
tlements and power plants. A collapse of the lake's fishery would mean a famine
of huge proportions. Efforts are underway to fight this invader.
In North America, the aquatic species hydrilla (Hydrilla verticilatta) was
introduced from Southeast Asia as an aquarium plant and escaped into the wild
in the 1950s. This plant can grow as much as 10 em per day and spreads through
256 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

the dispersal of leaves and stems. It is now a very damaging weed in both the One of 1
southern United States and along the West Coast. By 1990, approximately 40'}c America is tl
of Florida's waterways were infested by hydrilla. Hydrilla grows into dense mats bivalve is nati
that choke off light and cover submerged surfaces. The decay of the mats deplete_ million eggs a
oxygen from the water and can kill fish and other organisms. Aside from displac- perse readily ·
ing native plants and disrupting aquatic ecosystems, hydrilla is a major culprit iL the zebra mm
clogging water intake valves, irrigation systems, and natural waterways. spread as far
Insects, mollusks, mammals, and birds have also been very successfU:. barrier to stoj
invaders. Many common pest insects encountered in North America are alie appears that
invaders. These include the Argentine ant, the fire ant, the European earwig, anci released that
the gypsy moth. In turn, North and South American insects have been introducec mussel. By 19
throughout the world. For example, the Colorado beetle is an agricultural pes: terns and into
that was accidentally introduced to France in the early twentieth century and has Zebra rr
now spread as far east as Russia. At the present time, the most infamous of thes terns and bur
insect invaders in North America is the Africanized honey bee. predators of 1=
The European honey bee (Apis mellifera lingustica) was introduced to square meter
North and South America with the first European colonists. In addition to pro- 8.13). They ex
viding honey, these bees are extremely important for pollinating orchard trees declines in na
and some other crops. The African subspecies of honey bee (Apis mellifera sattel- species of nat
lata) is very similar in form and habit to its European counterpart. However, tb mussels. Wate
African subspecies is extremely aggressive in protecting its hives. Swarms of tb- Zebra musseh
bees will pursue animals or people that disturb their hives. Often, the intruders phytoplankto1
receive a lethal number of stings. Bee keeping for honey production and pollina- and permits g
tion is much more difficult with African bees than with the European subspecie- decreasing th
The African subspecies was brought to Brazil for breeding experiments to thought that z
develop a hybrid that would exhibit the African subspecies' ability to surviYe
tropical conditions and the European bee's more docile behavior. In 1956-19--
African bees escaped and established breeding populations in Brazil. The African
bee can aggressively drive out European bees. However, in many cases, th
African and European bees mate to produce hybrids. Unfortunately, the hybrid
bees display the aggressive behavior of the African subspecies. Since hybrids are
now so common, the aggressive strain of bees that is presently invading South
and North America is referred to as the Africanized honey bee.
The Africanized bees have been dispersing and colonizing South, Central.
and North America at a rate of roughly 110 km per year. They now occupy most
temperate and tropical portions of South and Central America. The northward
spread of the Africanized bee slowed when they reached the drier portions of
northern Mexico in the late 1980s. However, by the mid-1990s they were estab-
lished throughout the southwestern United States. By 1999 the Africanized
bees had spread to the Pacific Coast of California and caused their first human
fatalities in that state. It is thought that all wild populations of honey bee had
become Africanized within about two years of their appearance in southern
California.
Alien species of mollusks have also become major pests in North America.
The large brown garden snail (Helix aspersa) that commonly destroys agricul-
tural and ornamental plants in North America was introduced from Europe.
Polynesians introduced at least two species of land snails to Hawaii. In recent FIGURE 8.13
years, the giant African land snail (Achatina fulica) has been accidentally intro- station on Lake
duced to a number of Pacific islands. This large mollusk causes damage to native late 1980s, zebn
causing econon
flora and competes with species of native snails.

lamaging weed in both the
y 1990, approximately 40 ~c
lrilla grows into dense mats
~ decay of the mats deplete ~
anisms. Aside from displac-
lydrilla is a major culprit iL
atural waterways.
also been very successfu.:
1 North America are ali e
:, the European earwig, anc
tsects have been introduce -
~etle is an agricultural p e ~:
y twentieth century and h ~
the most infamous of thes;
oney bee.
;ustica) was introduced tc'
)lonists. In addition to pro-
,r pollinating orchard treQ
y bee (Apis mellifera sew er-
counterpart. However, t h~
ing its hives. Swarms of th~
hives. Often, the intru d e ~
tey production and pollinc.-
h the European subspecies.
breeding experiments · -
)species' ability to survi\ .o
;ile behavior. In 1956-195 -
ttions in Brazil. The Africa=.
wever, in many cases, tl:.::
. Unfortunately, the hybri-
tbspecies. Since hybrids ar::
> presently invading Sout:
oney bee.
colonizing South, Centra:
~ar. They now occupy m o~­
tl America. The northwar.::
ched the drier portions c:
1id-1990s they were estat-
By 1999 the Africanize.::
d caused their first huma::
ulations of honey bee ha.:
r appearance in souther:.
or pests in North Americc..
Jmmonly destroys agricU:-
introduced from Europe
:nails to Hawaii. In recer::
as been accidentally int ro-
:k causes damage to nati\-;
RECENT INVASIONS BY EXOTIC SPECIES 257
One of the most destructive alien mollusks in western Europe and North
America is the zebra mussel (Dreisena polymorpha). This small freshwater
bivalve is native to the Caspian Sea of central Asia. A single female can release 5
million eggs annually, and the larvae of the species are free-swimming and dis-
perse readily in freshwater. The building of shipping canals in the 1700s allowed
the zebra mussel to invade eastern Europe. By the 1830s, the zebra mussel had
spread as far west as England. The saltwater of the Atlantic Ocean produced a
barrier to stop the further spread of the mussel until around 1986-1988 when it
appears that a freight ship that had taken on freshwater ballast in Europe
released that ballast in Lake Erie. In that ballast were the larvae of the zebra
mussel. By 1996, the zebra mussel had spread throughout the Great Lakes sys-
tems and into the Mississippi River.
Zebra mussels are a particularly damaging invader to both natural ecosys-
tems and human activities. In North America, where they have few effective
predators of parasites, they reach densities of up to 400,000 individual adults per
square meter on any firm substrate from rocks to boat hulls to water pipes (Fig.
8.13). They exclude other mollusks and plants from these areas and have caused
declines in native shellfish populations. It~!las been suggested that 140 different
species of native North American shellfish might face extinction due to zebra
mussels. Water intake and outlet pipes quickly become plugged by the mussels.
Zebra mussels are extremely effective filter feeders and remove large amounts of
phytoplankton from the water, which, in turn, causes changes in the food chain
and permits greater amounts of sunlight to penetrate lake and river waters. By
decreasing the amount of plankton available to invertebrates and fish, it is
thought that zebra mussels can limit the amount of fish that lakes can support. It
FIGURE 8.13 Zebra mussels (Dreissena po/ymorpha) clog the intake pipes to a power
station on Lake Erie. Following an accidental introduction into the lower Great Lakes in the
late 1980s, zebra mussels have rapidly spread throughout much of eastern North America,
causing economic damage and the loss of benthic habitat along many lakes and rivers.
258 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION

has been suggested that the zebra mussel has cost commercial fishermen on Lak:- generally hav1
Erie as much as $400 million in decreased catch per year. As destructive as t :- Third, the inv:
zebra mussel is, it is even more sobering to realize that it is only one of 141 alic::. the invading s
species of fish, invertebrates, and plants to invade the Great Lakes system in t :- the new en vir<
past 200 years. the invaders. ~
Several mammal and bird species have become notorious invaders. Tl ~ and lack effe<
European rabbit (Orryctalagus cuniculus) was brought to the temperate regio;:: such as habit:
of southern Australia in the 1860s. By 1900, it occupied the southeastern quan e:- species, aids ir
of the continent and expanded almost to the northern coastline by 1980. Rabbi_ Can we
have proven extremely destructive to native Australian herbs and shrubs. Tt:- of informatio
black rat (Rattus rattus), which infests almost all of temperate North America . .::. and the plan
a native of Eurasia that has been introduced around the world. Interestino ~ information i
extremely stringent control measures at its borders, combined with a very co ~:. Brunswick, C
climate, have kept the black rat from invading the province of Alberta, Canad2.. potential suc<
The Polynesian rat (Rattus exulans) was introduced to Hawaii by Polynesian s :- range of the i
tiers and poses a major threat to native mollusks, insects, and some birds. Tt:o species' pote1
native birds of Hawaii face an additional challenge in the form of large populc.- occupy large
tions of the alien bird species myna (Acridotheres tristis), Japanese white- e~-= the right con
(Zosterops japonica), and red-billed leiothrix (Leiothrix lute a) which compete f _ breadth to s
food and nesting sites. In Europe, large flocks of Canada geese (Branta canader;- humans. Ho\\
sis) have become nuisances in parks and ornamental waterways, and drive o ·- invaders. For
native waterfowl from other habitat. range in sma
Amphibians and reptiles have also been introduced to many regions b. forestry speci
human activity and have proven to be destructive invaders. The cane toad (B ufc pine has spre
marinus) was brought to Australia to control insect pests, but the voracious aduJ - general, it hm
and tadpoles consume native insects, reptiles, and even small mammals at a:: sive invaders
alarming rate. It has proven impossible to eradicate the species. One of the mos For bett
insidious reptile invasions occurred on the Pacific island of Guam. Beginning i= phere by intrc
the 1960s, scientists noticed significant declines in the native bird population o: rivers, and sea
the island. During the 1980s, six of the eleven native bird species of Guam·: tions; others I
forests went extinct. Surviving individuals of other species were captured an - eluded that th1
taken to breeding facilities to be kept alive in captivity. Scientists puzzled ove: introduced by
the cause of the population declines and extinctions. After much study, it was are probably 1
determined that the culprit was a bird-eating snake called Boiga irregularis th2: do not contair
was native to the Solomon Islands. It appears that the snake had been acciden-
tally introduced to the island sometime during the 1940s. The snake approaches
bird nests at night and eats adult birds, chicks, and eggs with proficiency. Due to c:.
variety of prey, and lack of serious predators, the snake population grew to unna t-
ural densities, and the native birds of Guam paid the ultimate price for the intro- EY WORDS ANI
duction of this alien species.
Destructive invading species need not be large organisms. Fungi, micro- -.aemochores
scopic parasites, and bacteria have also been introduced to new regions b ~­ -.aemohydrochores
human activity with disastrous results. The North American deciduous forest has ~ t h ropochore

experienced catastrophic declines in elm trees (Ulmus spp.) and chestnut trees 3arrier
(Castanea dentata) due to Eurasian pathogens. Colonization
Corridor
The success of invading species often reflects several common factors. First.
:)iffusion
these species have been introduced to environments that provide physical condi- :)ispersal- active and
tions that are well suited for the survival of the invaders. Second, the invaders passive

1mercial fishermen on Lake
year. As destructive as the
1t it is only one of 141 a!i e;:;
• Great Lakes system in the
1e notorious invaders.
ht to the temperate regi o ~
:d the southeastern quam:~
1 coastline by 1980. Rab b i~
!ian herbs and shrubs. 11:..:
mperate North America . ~
td the world. Interestingl:
combined with a very co!.:
ovince of Alberta, Canadc._
J Hawaii by Polynesian se:-
tsects, and some birds. Th.:
1 the form of large populc.-
·ristis ), Japanese white-ey.:
ix lutea) which compete fc:
tda geese (Branta canader.-
l waterways, and drive o::-
heed to many regions _
aders. The cane toad (B u_-·
;ts, but the voracious ad ub
ven small mammals at c.::.
1e species. One of the mO!-
md of Guam. Beginning c
: native bird population c:
re bird species of Gua ~ -~
:pecies were captured a1::
ity. Scientists puzzled OYe:
. After much study, it w~
1lled Boiga irregularis the.·
e snake had been accide:::-
40s. The snake approach ~
; with proficiency. Due to.:.
population grew to un na:-
,ltimate price for the int rc-
: organisms. Fungi, micrc-
duced to new regions _
:rican deciduous forest h ~
S spp.) and chestnut tre~
eral common factors. Fin:·
tat provide physical cond:-
ders. Second, the invade__
RECENT INVASIONS BY EXOTIC SPECIES 259
generally have short generation times and produce large numbers of offspring.
Third, the invading plants or animals have readily dispersed propagules. Fourth,
the invading species have no serious natural predators, parasites, or pathogens in
the new environment. Fifth, native competitors, if they exist, are less efficient than
the invaders. Sixth, in the case of predatory invaders, prey species are abundant
and lack effective defenses against the invader. In many cases, human activity,
such as habitat disturbance or the hunting of native competitors or predator
species, aids in the success of the invaders.
Can we determine which species have the potential to be invasive if a lot
of information is available on the autecology and synecology of the invader
and the plant and animal communities being invaded? In most cases, such
information is in very short supply. A recent study of invasive plants in New
Brunswick, Canada, produced an interesting result in terms of predicting the
potential success of an invader. In that study, it was found that the size of the
range of the invading plant in its native habitat was the best predictor of the
species' potential to invade new territory. It appears that species which can
occupy large geographic ranges in their native areas are often likely to have
the right combination of reproductive traits, competitive ability, and niche
breadth to successfully invade large areas when they are introduced by
humans. However, organisms with small natural ranges are not always poor
invaders. For example, the Monterey pine (Pinus radiata) has a tiny native
range in small portions of coastal California but is successfully grown as a
forestry species in many parts of the world. In New Zealand, the Monterey
pine has spread from forestry plantations and invaded native vegetation. In
general, it has proven very difficult to predict which organisms will be aggres-
sive invaders and which ecosystems will be most prone to invasion.
For better, but mainly for worse, we have dramatically changed the bios-
phere by introducing thousands of alien species to new islands, continents, lakes,
rivers, and seas. Some environments have been radically altered by these introduc-
tions; others have suffered relatively little disruption. However, it can be con-
cluded that there are few, if any, large ecosystems left that do not host alien species
introduced by humans in the past few centuries. Aside from the high arctic, there
are probably no national parks or other nature preserves in North America that
do not contain significant numbers of alien species of plants and animals.
EY WORDS AND TERMS
-.ilemochores Exponential population Logistic population growth
emohydrochores growth Propagule
thropochore Filter Samara seeds
.;m-ier Great American Exchange Seasonal migration
Colonization Harmonization Stepping stones
Corridor Hydrochore Supertramp
::J'J fusion Invasion Sweepstakes routes
Jispersal-active and Irruption Waif dispersal events
passive Jump dispersal Zoochore
260 CHAPTER 8 DISPERSAL, COLONIZATION, AND INVASION
REFERENCES AND FURTHER READING
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Billings, W. D. (1990). The mountain forests of
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Elton, C. S. (1958). The Ecology of Invasions b\
=-in 64 (supplement), 187-
Animals and Plants. Methuen, London.
::.2.:llmen, D. (1996). Song OJ
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3 !ogeography in an Age Oj
(1999). Predicting invasiveness of plant
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Conservation Biology 13, 422--426.
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Gorman, M. L. (1979) . Island Ecology. Chapma:. Oxford.
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Ecology 70, 339-347. __-:s of Geographic Patternil
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ondary dispersal of tree seeds on snow. Jo ur- : ge, J. A. (1987). Extinctic
nal of Ecology 85, 329-340. :es avifauna by an introdu
Grime, J.P., and Jeffrey, D. W. (1965). Seedling '660-668.
establishment in vertical gradients of sunligh:.. ·eltema, R. S. (1986) . Long
Journal of Ecology 53, 621-642. 2y planktonic larvae of she
Grimm, W. C. (1967). Familiar Trees of America. · vertebrates among centr
Harper and Row, New York. Bulletin of Marine Science
Harper, J. L. (1977). Population Biology of
Plants. Academic Press, New York.
Heywood, V. H. (1995). Global Biodiversity, Assess-
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Higgins, S. 1., and Richardson, D. M. (1996). A
review of models of alien plant spread. Eco-
logical Modelling 87, 249-265 .
Huggett, R. J. (1998). Fundamentals of Biogeog-
raphy. Routledge, London.
Huntley, B. and Webb, T., III (1989) . Migration:
Species' response to climatic variations
caused by changes in the earth's orbit. Journ a!
of Biogeography 16,5-19.
Krebs, C. J. (1994). Ecology, 4th edition. Harper-
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MacDonald, G. M. (1993). Reconstructing plant
invasions using fossil pollen analysis.
Advances in Ecological Research 24, 67-110.
MacDonald, G. M., and Cwynar, L. C. (1985). A
fossil pollen based reconstruction of the late
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Quaternary history of lodgepole pine (Pinus Shigesada, N., and Kawasaki, K. (1997). Biologi-
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Delcourt, H . R. (1987). Long- of Canada. Canadian Journal of Forestry University Press, Oxford.
!amics of the Temperate Zon Research 15, 1039-1044. Silvertown, J. W. (1982). Introduction to Plant
Late-Quaternary Forests in _.fack, R.N. (1981). Invasion of Bromus tectorum Population Ecology. Longman, New York.
merica. Springer-Verlag, Ne" L. into western North America: An ecological Silvertown, J. W. , and Doust, J. L. (1993). Intro-
chronical. Agroecosystems 7, 145-165. duction to Plant Population Biology. Black-
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Rejmanek, M ., and geography. Van Nostrand Reinhold, New York. Simpson , G. G. (1940) . Mammals and land
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ve. J Wiley, New York. of the winter range of the house finch . Ameri- of Science 30, 137-163.
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68, 660-668. Zacharin, R. F. (1978). Emigrant Eucalypts: Gum
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i) . Global Biodiversity, A ssess-
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:hardson, D. M . (1996) . A
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Fundamentals of Biogeog-
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, T. , III (1989). Migration
to climatic variations
in the earth's orbit. Jou rna.
16,5-19.
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blishers, New York.
993). Reconstructing pl an;
ssil pollen analysis.
ogical Research 24, 67-11 0.
d Cwynar, L. C. (1985). A
reconstruction of the la te
 CHAPTER 9 terms cladogen
are caused by<
to the creation
lutionary · dive1
EVOLUTION, SPECIATION, descendant spe
anomie orders.
AND EXTINCTION more detail, w
relate to sexual

Some Basi'
It is impossible to consider the great variety of plant and animal species that e ··_.. The basic phys
today and not ask the question, how did this diversity of life arise? We have see:. cal structures c
that plants and animals possess extraordinary anatomical and physiological fec.- parents to offs
tures that allow them to exist in many different environments. How do such adar- deoxyribonuclE
tations develop? Even before the time of Aristotle, people contemplated the ~ binations of su:
fundamental questions. Indeed, the origin of plant and animals species rema ~ pounds consisti
among the most vigorously debated issues in theology, philosophy, and science. The DNA stru
In addition to the plants and animals we see today, we are aware from ge0- on the sides an
logical and historical records that there are many species that lived in the past b ': entist Francis C
do not exist today. Everyone is familiar with the dinosaurs, but our informatio;:. basic structure
on these animals comes to us only from fossils. The last living dinosaur disa 1940s. The desc
peared from the face of the earth some 65 million years ago. Not a single dinosau: scientific disco'
exists today, and yet for for over 100 million years (from the Triassic to the end o: Within th
the Cretaceous) huge numbers of dinosaurs lived on the land, in the air, and ~ like structures
the sea. Why did all the hundreds of species of dinosaurs cease to exist some 6:: chromosome i~
million years ago? numbers of ch
What causes species to appear, adapt to certain environments, and disa genes located a
pear is a fundamental question for biogeographers. In this chapter we will con- somes arrangec
sider how geography and environmental change may influence the developmen: chromosome f
of new species. We will then look at the role of geography and environmenta... only 4. Plants 2
change in the elimination of species. Finally, we will consider the positive role as each of thei1
that the elimination of some species plays in promoting the development of othe:- some pairs are
new species. animals, contai
as being haplo
haploid female
EVOLUTION AND SPECIATION bine. This resul
ducing plant 01
In order to study the way in which geography and environmental change haYe parent and hal
contributed to the great diversity of species, we need to consider two relate Many the
processes-evolution and speciation. The term evolution refers to genetica l!~­ chromosomes.
controlled changes in physiology, anatomy, and behavior that occur to a species cause differenc
over time. Evolutionary change within an individual species or population is Such genetica
called microevolution, whereas evolutionary change within larger taxonomi referred to as I
units such as families is termed macroevolution. In this chapter we will concern are called aile)
ourselves mainly with microevolution. Speciation refers to the development of be heterozygo1
two or more genetically differentiable species from a single common ancestor an important n
species. Speciation results from evolutionary change, but not all evolutionary The natu
change leads to the development of two or more species from a common ances- reproducing 01
tor. The different species that arise from the same ancestor are called a clade. The implications. F

262

'N,
nd animal species that ey:--
of life arise? We have see:.
nical and physiological fec.-
nments. How do such ada:::-
people contemplated thes.:c
1d animals species remai;:.
1, philosophy, and science.
1
ay, we are aware from gec-
ies that lived in the past bL
)Saurs, but our informa tio-
last living dinosaur disa;-
s ago. Not a single dinosa
om the Triassic to the end c:
the land, in the air, and :.=
mrs cease to exist some c:
1 environments, and disa_::-
n this chapter we will co:::-
influence the developme;:;-
graphy and environmen:2...
consider the positive ro: ~
g the development of ot b:c-
nvironmental change ha· ~
d to consider two rel ate.:
11tion refers to genetica:.:
rior that occur to a spec>
I species or population _
within larger taxono- -
s chapter we will conce:-
ers to the developmenr c
single common ancesr-
but not all evolutiona..-o
es from a common an c~
star are called a clade. T.:~
EVOLUTION AND SPECIATION 263
terms cladogenesis and speciation can be used synonymously. Speciation events
are caused by evolutionary changes, but not all evolutionary developments lead
to the creation of additional species lines. When speciation occurs, continued evo-
lutionary divergence can lead to greater differences between ancestral and
descendant species and the development of new genera, families, and higher tax-
onomic orders. Before we consider the processes of evolution and speciation in
more detail, we should review some aspects of genetics and heredity as they
relate to sexually reproducing species.
Some Basic Genetics
The basic physiology, anatomy, and behavior of species are controlled by chemi-
cal structures called genes, which are found in living cells and are passed from
parents to offspring. The genes of plants and animals consist of molecules of
deoxyribonucleic acid (DNA). The DNA molecules are made up of various com-
binations of sugars and phosphates that are joined together by nitrogenous com-
pounds consisting of adenine, thymine, cytosine, and guanine bound by hydrogen.
The DNA structure resembles a twisted ladder with the sugars and phosphates
on the sides and the nitrogenous compounds forming the rungs. The English sci-
entist Francis Crick and his American colleague James Watson first described the
basic structure of this crucial DNA molecule at Cambridge University in the late
1940s. The description of the DNA structure stands as one of the most important
scientific discoveries of all time.
Within the cells of an organism the genes are arranged along paired, thread-
like structures called chromosomes. The point at which a gene is located on a
chromosome is called a locus (the plural is loci). Different species have different
numbers of chromosomes, different numbers of genes, and different types of
genes located at the loci of the chromosomes. Humans have a total of 46 chromo-
somes arranged in 23 pairs. The American black bear (Ursus americanus) has 37
chromosome pairs, while the common fruit fly (Drosophila melanogaster) has
only 4. Plants and animals typically have the same number of chromosome pairs
as each of their parents. Organisms and cells that have complete sets of chromo-
some pairs are said to be diploid. Certain reproductive cells, such as sperm from
animals, contain only one-half of each chromosome pair. These cells are described
as being haploid. During sexual reproduction a haploid male cell fuses with a
haploid female cell, and the two sets of chromosomes and associated genes com-
bine. This results in the formation of a diploid embryo. Thus, each sexually repro-
ducing plant or animal receives half of its genes and chromosomes from its male
parent and half from its female parent.
Many thousands of different gene forms can be arranged along individual
chromosomes. Differences in the specific chemical form of the gene at a locus
cause differences such as hair coloring, skin coloring, or eye coloring in humans.
Such genetically controlled variation in appearance within a population is
referred to as polymorphism. The different gene forms that exist for a given locus
are called alleles. When a locus has different alleles associated with it, it is said to
be heterozygous. The degree of heterozygosity within a species or population is
an important measure of genetic diversity.
The nature of genes and chromosomes, and the way in which sexually
reproducing organisms pass along their genes to their progeny have important
implications. First, the genes can best be thought of as particles of information
264 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

that themselves do not change or blend during reproduction. The genes rema:=
stable from generation to generation. Second, if the chromosomes and loci of tLo 100

male and female do not match perfectly, reproduction generally cannot occe.::
This blocks interbreeding between different species. Third, because chromoson::-
numbers, and the genes at each loci, must match, when species reproduce th _
produce offspring that are genetically similar to the parents. Therefore, the ge;:-
eral physiology, anatomy, and behavior of the offspring will be like those of t :-
parent. Fourth, since each offspring is a product of the recombination of chrom
somes and genes from the male and female parent, the alleles found at the loci c:
the offspring will not exactly match either of the parents. Thus, the offspring
differ from the parents in traits for which there are different alleles. This allo\,_
for individual variability in traits, such as eye color, within the same species. Ge -
der is also controlled by chromosomes and associated genes. The complete rang::
of genes present in a species is called the genome. The human genome consists c:'
more than 25,000 different genes. The wide variety in human appearance is on:-
product of this genetic diversity.
Differences in the physiology, anatomy, or behavior of different species ~
Ic:
0
individuals of the same species are called phenotypic variations. These obsen ·- ·~

able variations are often the result of genetic differences referred to as genotypi s;
[i
variations. For example, all white spruce trees have thin needles, whereas willO\\~
have broader leaves. These phenotypic differences between spruces and will O\\~ West A
are genotypic and are inherited by the two plant species from their parents. No: FIGURE 9 . 1 A
all phenotypic differences are genotypic. The environmental conditions to whict California. The hi
an organism is exposed can also influence the growth or form of the individua: from both high a
plant or animal. For example, the seed of white spruce trees (Picea glauca persist. This sho1
planted in relatively warm forested sites can grow into a tree that is 30 m tall. I: altitudes represe
the same seed is planted at a cold and unprotected site near the treeline, the
spruce will grow as a very low shrub with sparse foliage. The difference betweer:
the tall spruce of the forest and the stunted spruce of the treeline has nothing to transplanted tc
do with genetic differences; it is caused by differences in the environment. Thu uals that still ~
not all phenotypic differences are genetic. genetically em
Because both genetics and environmental conditions can influence the follL allows them t(
or behavior of an organism, it is sometimes difficult to determine to what degre protected and
two individuals are genetically distinct. Two individuals may appear to be differen distinct memb
simply because of the environmental conditions to which they have been exposed. to as geograpl
This problem is particularly important for biogeographers. We frequently find tha such as the hei
plants and animals of the same species living in different geographic regions dis- Today, s<
play slightly different features or behavioral patterns. One way to assess if there is exact genetic c
a genetic control behind such phenotypic differences in plant species is to conduc often found th
transplant experiments. In such experiments, the seeds from one region are trans- differences in
planted into a different region and the development of the transplanted seedlings example, lodg
is observed. If the transplanted seedlings develop features that are typical of United States
plants in their native habitat, but distinctive from the plants found in the trans- tions found in
plant area, we can assume that the phenotypic differences we observe in plant Yukon a few t
from the different regions are indeed genetically controlled. subset of the t
A classic transplant experiment was conducted on yarrow plants (Achillea the eastern U1
lanulosa) from California (Fig. 9.1). The yarrow plants grow from near sea level of marine and
to alpine tundra at elevations of over 3600 m. The yarrows growing at low eleva- black sea bass
tions typically develop flower stalks that are up to 70 em in height, while those row (Ammod1
found at the highest elevations grow stalks that are less than 20 em tall. When from populati

juction. The genes remai
romosomes and loci of th
on generally cannot occu r.
1ird, because chromosom
~n species reproduce the ~·
arents. Therefore, the gen-
tg will be like those of the
recombination of chromo-
alleles found at the loci o:·
Its. Thus, the offspring cac
fferent alleles. This allO\\·s
hin the same species. Gen-
genes. The complete range
human genome consists o:
human appearance is o n ~
·ior of different species o:-
variations. These obsen ··
=s referred to as genotyp i
1 needles, whereas willows
ween spruces and willows
es from their parents. I\·o:
1ental conditions to whic::
or form of the individua:
ruce trees (Pice a glauca
• a tree that is 30 m tall. I:'
;ite near the treeline, th.o
e. The difference betwee::
he treeline has nothing t c
in the environment. Thn<
)ns can influence the fo ~
determine to what degre;:
nay appear to be differec.:
h they have been expose -
:rs. We frequently find the.:
rrt geographic regions dis-
ne way to assess if there :S
plant species is to cond uc:
:r om one region are trans-
the transplanted seedlin=-
ttures that are typical c::
Jlants found in the trans-
lees we observe in plan:s
ied.
n yarrow plants (AchillE.:.
grow from near sea ]e\·;:_
WS growing at low e]eY<:-
cm in height, while thos::
ss than 20 em tall. Whe::
EVOLUTION AND SPECIATION 265
100
E'
.:=..
E
Cl
"(i)
..c 50
cctl
a:
Ic
0
~
w
West A East B
FIGURE 9.1 A cline in yarrow (Achillea lanu/osa) races found at different elevations in
California. The high elevation varieties are shorter than the low elevation ones. When plants
from both high and low elevations are grown together near sea level the differences in height
persist. This shows that the height differences are genotypic and the plants found at different
altitudes represent genetically distinct races (after Clausen et al., 1948; Krebs, 1994).
transplanted to low elevation, the seeds from the alpine plants produced individ-
uals that still grew to only 20 em. Thus, the difference in size of the yarrows is a
genetically controlled feature. It is likely that the low stature of the alpine plants
allows them to survive at high elevations because it keeps the flowering heads
protected and closer to the warm ground surface. Genetically and phenotypically
distinct members of the same species that occur in different regions are referred
to as geographic races. A geographic gradient in a genetically controlled trait,
such as the height of the yarrow plants, is called a cline.
Today, sophisticated biomolecular techniques can be used to examine the
exact genetic differences that occur across clines of plant and animals species. It is
often found that within the geographic range of any species there are significant
differences in the genetic composition of different geographic populations. For
example, lodgepole pine (Pinus contorta) trees in the Rocky Mountains of the
United States have significantly higher numbers of different alleles than popula-
tions found in the Yukon Territories of Canada. The species only arrived in the
Yukon a few thousand years ago, and these northern populations contain only a
subset of the total genetic diversity present in the older southern populations. In
the eastern United States, the Gulf of Mexico populations of widespread species
of marine and shoreline animals such as the ribbed mussel (Geukensia demissa),
black sea bass (Centropristis stiata), tiger beetle (Cicindela dorsalis), seaside spar-
row (Ammodramus maritimus), and many other species are genetically distinct
from populations of the same species that live along the Atlantic coastline. It is
266 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

thought that these Atlantic-Gulf distinctions arise because the marine and shore- Historical I
line populations in the two regions have long been separated from each other b: We have seen
the Florida Peninsula. spring. We ha
Many times, as in the case of the California yarrow plants, genetic diffe:-- mutations. He
ences within species cause the populations found in different geographic regia ' instantaneous
to have significant differences in morphology or physiology. For example, popula- tions that allc
tions of the Douglas fir species (Pseudotsuga menziesii) can be found growiL~ species evolve
from central British Columbia southward along the Rocky Mountains to th:: history of this
mountains of northern Mexico and along the coast to northern California. Trans- The idea
plant experiments show that there are a number of different genotypes withi::. in Greece at l•
this large geographic range. One important manifestation of this geographi: was at odds w
diversity in genotypes is that trees from coastal California are killed by temper~­ the great philc
tures of less than -16°C, while northern and high-elevation populations are muc::: continued to l
more frost resistant. Bacon suggest
The discussion of differences in the heights of yarrows or frost resistance i:: be transmuted
Douglas firs raises an interesting question. Can offspring of an organism inheri: tion. In 1766 tl
phenotypic traits that are acquired due to environmental conditions acting upo;: animal familie
the organism? The answer is no. We now know that such acquired traits do no: species prodw
affect the alleles and chromosomes of the organism and are not passed down tc over a long pe
the offspring. If, for example, a bull elephant loses a tusk in a fight, the elephant·: degenerate sp
offspring will still develop two tusks. However, as late as the middle twentietl:: naeus suggest<
century, there were scientists who argued that acquired traits could indeed be two different :
inherited. progeny are cc:
We can see that genetics and heredity act to maintain the physiologicaL ing the charac
anatomical, and behavioral attributes of species from one generation to the nex force for creat
How then do new genes and genetically controlled traits appear in specie s ~ Lamarck (174·
Genes and chromosomes are subject to random mutations that create new allele5 on earth as sin
and chromosomal structures. Such mutations can give rise to new traits. Poi1 : thought the in
mutations occur when individual genes experience changes in their chemica acquiring ada
structure. Mutations also arise when chromosomes break into two or more example, a gin
smaller chromosomes and alter the chromosomal number of the organism. Fo branches. As a
example, some populations of the intertidal snail Thais lappillus have 26 chromo- With each gen
somes, while others have 36. In some cases, the broken chromosome fragments to Lamarck, o
fuse in new or inverted configurations, and the gene loci are rearranged along the developed ne\
chromosome. In plants, it is relatively common for some offspring to have twic species evolve
the chromosome number of either parent. These plants are called polyploids. Lamarck's the
Finally, during recombination, some chromosome chains may not line up exactly. organism's pre
and some genes may be duplicated or deleted. All of these genetic mutations can The biol•
give rise to new phenotypic traits. In most cases, however, major genetic muta· of Lamarck, a
tions are lethal or do not allow successful reproduction by the mutant organism. teenth-centuq
However, some mutations may be benign or beneficial to the organism. In some word "evolutic
cases, the mutant organism cannot mate with nonmutant members of the same ary change an
species but can mate with siblings that share the mutation, or reproduce asexu- geography. Th
ally. In this case, instantaneous speciation occurs. evolution. Ou
The genetic composition of a population can change over time as new genes remarkable E
arise via mutation and other genes are lost through chance processes. Such ran- win (Fig. 9.2)
dom changes are referred to as genetic drift. Over time, genetic drift can lead to Charles
the development of new species, particularly if the population is small and geo- educated in m
graphically isolated from other members of the same species. versity. By all

:a use the marine and shore-
parated from each other b~
trrow plants, genetic diffe:--
lifferent geographic regi o ~
ology. For example, populc-
esii) can be found growiL§:
~ Rocky Mountains to th:-
northern California. Trans-
different genotypes with:::
:station of this geograp lll:
lrnia are killed by temperc:-
ation populations are muc:.
trrows or frost resistance ·-
ring of an organism inher:·
ntal conditions acting upo::
mch acquired traits do nc·
.nd are not passed down :-
tsk in a fight, the elephan: _
te as the middle twentie::.:.
red traits could indeed c·:-
naintain the physiologic~
one generation to the nex-_
traits appear in species
ions that create new allele<_
'e rise to new traits. P 01r
changes in their chemiC?.
break into two or mo:-:-
mber of the organism. f,~ ­
! lappillus have 26 chromc-
~n chromosome fragm en:..
:i are rearranged along tt::
ne offspring to have twic::
mts are called polyploids.
ns may not line up exactl:
1ese genetic mutations <:2:
ever, major genetic m u r ~­
n by the mutant organis
I to the organism. In soc::::
ant members of the sac::::
ttion, or reproduce ase'-• -
tge over time as new gen.:::
1ance processes. Such ra;:-
e, genetic drift can lead :-
pulation is small and gec-
pecies.
EVOLUTION AND SPECIATION 267
Historical Development of Evolutionary Theory
We have seen how genetically controlled traits are passed from parent to off-
spring. We have also seen that new traits can arise in species through genetic
mutations. However, the capacity for the random generation of new traits or
instantaneous speciation does not explain how species evolve specific adapta-
tions that allow them to survive in new environments. The problem of how
species evolve has baffled scientists from ancient times. Let's briefly consider the
history of this scientific inquiry.
The idea that species could be transformed into new species was introduced
in Greece at least a century before Aristotle (384-322 B.c.). This idea, however,
was at odds with Aristotle's view that species and genera were immutable, and
the great philosopher argued against it. Aristotle's belief in the stability of species
continued to be dominant until the time of the European Renaissance. Francis
Bacon suggested in 1631 that there were some cases in which species appeared to
be transmuted into other species and that this phenomenon warranted investiga-
tion. In 1766 the French scientist, Comte de Buffon suggested that the number of
animal families or genera of the earth represented the original number of animal
species produced by divine creation. Under the influence of the environment,
over a long period of time, these original species had subdivided into a number of
degenerate species that now inhabit the planet. During the same period, Lin-
naeus suggested that new species could arise through interspecific mating. When
two different species mate, the process is called hybridization, and the resulting
progeny are called hybrids. Linnaeus saw hybridization as a means of both chang-
ing the characteristics of existing species and producing new species. A different
force for creating new species was proposed by the French biologist Jean Baptiste
Lamarck (1744-1829). In the early 1800s, Lamarck postulated that life first arose
on earth as simple primitive organisms that developed in the primordial mud. He
thought the impact of the environment on individual organisms resulted in their
acquiring adaptive traits that they could then pass on to their offspring. For
example, a giraffe feeding on savanna trees would stretch its neck to reach higher
branches. As a result, the progeny of that giraffes would be born with a long neck.
With each generation the necks of the giraffes would become longer. According
to Lamarck, organisms that were exposed to different environmental conditions
developed new adaptations and passed these down to their offspring. Thus, new
species evolved to take advantage of different environments. At the root of
Lamarck's theory was the hypothesis that acquired traits are passed down to the
organism's progeny. We now know this is not possible.
The biological use of the term evolution appears to have begun at the time
of Lamarck, although he never referred to his ideas using this term. Other nine-
teenth-century scientists, such as the Scottish geologist Charles Lyell, did use the
word "evolution" in reference to Lamarck's ideas. Thus, the concept of evolution-
ary change and speciation was introduced into the language of biology and bio-
geography. The stage was now set for the development of the current theory of
evolution. Our current ideas about evolution owe much to the work of two
remarkable English naturalists and consummate biogeographers, Charles Dar-
win (Fig. 9.2) and Alfred Russel Wallace (Fig. 9.3).
Charles Darwin (1809-1882) came from a relatively affluent family and was
educated in medicine at Edinburgh University and divinity at Cambridge Uni-
versity. By all accounts, he was not a particularly dedicated or gifted student in
268 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

either subject. •
British naval s
stations to aid
whose role waE
and provide ap
captain of the ~
ous and energ<
read works by
Charles Lyell. J
to environmen
evolution thro1
grandfather, th
during the VO)
much on Darw
During h
influence his tl
remains and nc
animal species.
common over .
geographic reg
islands such as
bled those on
FIGURE 9.2 The young Charles Darwin (1809-1882). Darwin was only in his 20s wh en - that the specie~
embarked on the HMS Beagle for his scientific voyage around the world.
to represent di
ago. He would l
toises, finches, ::
When he
by publishing ·
also worked 01
acquired on tt
answer the ride
his thoughts ar
(1766-1834) ar
man and econ•
poverty in a tre
was capable o:
agricultural pre
human popu!a1
This would lea<
the hoarding
nations. The re
and the strong
last provided h
evolution.
What wa:
species have m
for example, so
FIGURE 9.3 Alfred Russel Wa llace (1823-1913) as a young man. Wallace began his have slightly lo
scientific career as a f ree-lance collector of exotic biologica l specimens and began his horses have !01
travels to places such as South America and the Malaysian Archipelago while in his 20s. ulations of pla1

in was only in his 20s whe n
:l the world.
man. Wallace began his
>ecimens and began his
chipelago while in his 20s .
EVOLUTION AND SPECIATION 269
either subject. In 1831 he set out upon a five-year voyage on the HMS Beagle. The
British naval ship's mission was to chart the coast of South America and set up
stations to aid navigation in the South Pacific. Darwin was an unpaid naturalist
whose role was to make geological and biological observations, collect specimens,
and provide appropriate company and intellectual stimulation for the upper-class
captain of the ship. The young Charles Darwin turned out to be a very adventur-
ous and energetic member of the expedition. Before and during the voyage, he
read works by the biogeographer Alexander von Humboldt and the geologist
Charles Lyell. Darwin accepted Lyell's ideas that the earth was ancient and prone
to environmental change. Young Darwin was also familiar with incipient ideas of
evolution through the writings of Lamarck, and those of Charles Darwin's own
grandfather, the physician-philosopher-biologist, Erasmus Darwin. It is clear that
during the voyage the question of where and how different species arose was
much on Darwin's mind.
During his explorations, Darwin observed several phenomena that would
influence his thinking about how different species arose. He found many fossil
remains and noted that some of these closely, but not exactly, resembled modern
animal species. He noticed that in many cases bird or animal species would be
common over a large area, but when one moved from that area into a different
geographic region, the species were quickly replaced by other species. He visited
islands such as the Galapagos and noted that some species of animals there resem-
'= bled those on the mainland, while others were quite distinct. He also observed
that the species of birds and tortoises on some of the Galapagos Islands appeared
to represent different varieties from those found on other islands in the archipel-
ago. He would later learn from experts in Britain that the different varieties of tor-
toises, finches, and mockingbirds he collected were actually different species.
When he returned to England, Darwin established his scientific reputation
by publishing the geological observations he had made during the voyage. He
also worked on the huge body of zoological and botanical information he had
acquired on the expedition. In secret, he also began to use his data to try to
answer the riddle of how species evolved. In this effort, he kept extensive notes of
his thoughts and observations. In 1838 he read the writings of Thomas Malthus
(1766-1834) and was greatly impressed by them. Malthus was an English clergy-
man and economist who set forth his views on human population growth and
poverty in a treatise published in 1789. Malthus proposed that human population
was capable of expanding in an exponential manner, while resources, such as
agricultural produce, could only increase in a linear manner. Thus, over time the
human population would grow beyond the ability of the resources to support it.
This would lead to increasingly fierce competition, such as fights between people,
the hoarding of resources by certain segments of society, or wars between
nations. The result of such competition was that the weaker would be destroyed
and the stronger would persist. Darwin later wrote that the ideas of Malthus at
last provided him with "a theory by which to work" on his own ideas of natural
evolution.
What was the concept that Darwin formulated? Darwin observed that all
species have natural variability in physiology, anatomy, and behavior. In humans,
for example, some people have black hair and some have blond hair. Some birds
have slightly longer beaks than other individuals of the same species, while some
horses have longer legs than other horses, and so on. He also observed that pop-
ulations of plants and animals can grow rapidly to great sizes, but this growth is
270 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

limited by intensive intraspecific and interspecific competition. If an individ u ~ to work as as

possesses a trait that gives it an advantage in competition, say, for example. _ exotic insects, ·
longer beak that aids in hunting worms, that individual will be more likely to ha\ = and could be s
sufficient energy to survive and produce healthy offspring. The trait that gives tt= two books on I
individual a reproductive advantage over its competitors is in turn inherited _ spent the next
its offspring. These offspring are more numerous and more likely to survive tha::. conditions we1
the offspring of competitors. Alternatively, individual birds of the same species malaria.
that have slightly shorter beaks would be at a disadvantage in capturing prey aL.: In his letl
less likely to survive and breed. Over time, the long-beaked variety of bird \\ to the pressun
increase due to its success in feeding and reproduction, while the short-beak :.: favor some va:
variety will disappear. Darwin referred to this process as natural selection. Trai_ ments with ex;
that provide an advantage in reproduction are selected for, whereas disadvan - the various is!~
geous traits are selected against. that the geogr
Natural selection provides a means for evolutionary change within a speci - new species fr
How does natural selection produce new species? As we will see, this is still a vex- were extreme!
ing problem. Darwin thought that new species could develop when individu ~ that a young, ~
moved into new habitat and when natural selection within that new environm e~­ the wilds of M;
acted to promote certain traits and depress others. Darwin also considered tt= Darwin. The ol
establishment of geographically separated populations by the dispersal of propa=-- the luxury of ~
ules to islands or distant continents to be an important force in allowing the deve:- Darwin could I
opment of new species. ory. Thus, he we
Modern genetics had not been developed by Darwin 's time, and he did n0: by being the fir
know about genes and how they pass along traits from parent to offsprin; action would r
Despite his lack of knowledge of genetics, Darwin did not believe in Lamarck-: confided his dil
ideas about acquired traits being inherited. It is amazing in some ways that D a;-- arrange for a jc
win developed his theory without knowing how new inheritable traits could ari_E naean Society <
as genetic mutations or how the recombination of genes during reproductio:: gle article enti
allows variability in offspring. If we wish to consider natural selection in light o:"" perpetuation o
modern genetics, we might say that Darwinian evolution works to select for gen : Darwin expanc
that impart traits that lead to reproductive success. These genes are preserved ii: of Natural Selt
the population, while genes associated with traits that limit reproductive succe ' Life. This pivo·
are winnowed out. remains in prin
Darwin first developed his theory in 1844 and 1845. At that point, he share Darwin a
his ideas with a few close colleagues, such as the noted botanist Joseph Hooke both scientific <
Darwin's ideas, that species could change over time and give rise to new specie human species.
were in some ways heretical to the teaching of the Church of England and th tion to the thee
religious beliefs prevalent at that time. He knew his theory would generate con- feared, D arwin
troversy, and he was very hesitant to publish it. Darwin enjoyed independen- attacks increase
wealth, so he could continue to spend time refining his ideas and collecting eYi- evolved from p
dence to support his theory of natural selection. Then, on June 18, 1858, he to Sex. Howevt:
received an alarming letter from Malaysia that changed his plans. had embraced
The letter that caused Darwin so much distress came from a young English accorded many
naturalist named Alfred Russel Wallace (1823-1913). Rather than being an Parliament ace<
enemy, Wallace was a great admirer of Darwin's work in geology and biology_ some of Englan
Before we consider the contents of Wallace's letter, it is interesting to compar scientific career
him to Darwin. Unlike Darwin, Wallace came from a relatively poor background including an ho
and never attended university. Indeed, he had only six years of formal schooling the shadow of
and acquired his knowledge of biology largely though self-training. For a number medallion at We
of years, Wallace worked as a land surveyor in Great Britain. In 1848 he Even thot:
embarked on a voyage to the Amazon Basin with the entomologist, Henry Bate . lication, and a

>mpetition. If an individuc...
•etition, say, for example. =.
l will be more likely to ha,·:-
ring. The trait that gives tt:-
itors is in turn inherited
more likely to survive the.=
1 birds of the same specie-:
ntage in capturing prey a=:
beaked variety of bird ,,-c_
on, while the short-beakc-:
; as natural selection. Tra;:.:
~d for, whereas disadva n ~-
ary change within a specin.
ve will see, this is still a \ e~-
1 develop when individuc......
ithin that new environme;:.
)arwin also considered r::-.::
. by the dispersal of propa;-
force in allowing the de,·e.-
trwin 's time, and he did r.
from parent to offspri.L.;
d not believe in Lamarc:.;. .
ing in some ways that Dc.:--
lheritable traits could ar..s::
~enes during reproductic-
latural selection in ligh
m works to select for ger.::-
tese genes are preserved --
limit reproductive succes
45. At that point, he shar.e:
d botanist Joseph Hook.o:
1d give rise to new sp e cie~
:hurch of England and t~::
1eory would generate co;:-
·win enjoyed indepen de::.
is ideas and collecting e· --
len, on June 18, 1858. :':,:o
d his plans.
arne from a young Engl:s-
3). Rather than being 2.-
·k in geology and biolo;-
: is interesting to compc.:-::
elatively poor backgrou:-..:
years of formal schooli::..=
;elf-training. For a nu m b~
freat Britain. In 1848 ::.::
:ntomologist, Henry Bat e:..
EVOLUTION AND SPECIATION 271
to work as a self-employed collector of natural history specimens. Examples of
exotic insects, birds, and flowers were very much in demand in Victorian England
and could be sold at a profit. Wallace spent four years in the Amazon and wrote
two books on his travels. In 1854, he departed on a collecting trip to Malaysia. He
spent the next eight years collecting specimens and writing scientific articles. The
conditions were incredibly difficult, and his challenges included severe bouts of
malaria.
In his letter to Darwin, Wallace outlined a theory on how species evolve due
to the pressure of competition, predation, and other environmental factors that
favor some varieties and are deleterious to others. Wallace backed up his argu-
ments with examples from the species of animals and insects he encountered on
the various islands of Southeast Asia. He was particularly interested in the effect
that the geographic separation of islands had on the eventual development of
new species from common ancestors. Darwin saw at once that Wallace's ideas
were extremely similar to his own theory of natural selection. How incredible
that a young, poor, and relatively uneducated adventurer, working and living in
the wilds of Malaysia, should so quickly develop the same theory of evolution as
Darwin. The older scientist was the product of the best British education and had
the luxury of spending over 20 years at home to work on his ideas. Of course,
Darwin could have put Wallace's letter aside and quickly published his own the-
ory. Thus, he would bring his life's work to fruition and secure his place in science
by being the first to publish on the concept of natural selection. However, such an
action would not have been in keeping with the character of the man. Darwin
confided his dilemma to his close friends Charles Lyell and Hooker. They helped
arrange for a joint reading of separate papers by Darwin and Wallace at the Lin-
naean Society of London on July 1, 1858. The papers were then published as a sin-
gle article entitled "On the tendency of species to form varieties; and on the
perpetuation of varieties and species by means of natural selection." In 1859,
D arwin expanded his ideas in a book entitled On the Origin of Species by Means
of Natural Selection, or the Preservation of Favoured Races in the Struggle for
Life. This pivotal volume went through several revisions in Darwin's time and
remains in print today.
Darwin and Wallace 's concepts of evolution caused great controversy in
both scientific and civil society. Bishops debated botanists over the origins of the
human species. Famous geologists such as Louis Agassiz (1807-1873) took excep-
tion to the theory, while others such as Charles Lyell were supporters. As he had
feared, Darwin was the object of some personal attacks. The tempo of these
attacks increased in 1871 when Darwin published his views that humans likely
evolved from primates in the book, The Descent of Man, and Selection in Relation
to Sex. However, by the end of his life the majority of the scientific community
had embraced Darwin's views on evolution by natural selection. He was
accorded many great honors, and when he died in 1882 a petition to the British
Parliament accorded him burial in Westminster Abbey. This is the resting place of
some of England's most famous figures. Wallace also had a long and distinguished
scientific career. In fact, Wallace lived for 91 years and was granted many awards,
including an honorary doctorate from Oxford University. He was, and remains, in
the shadow of Darwin, however. It is telling that he was accorded a memorial
medallion at Westminster Abbey but not burial there.
Even though almost 150 years have passed since Darwin and Wallace's pub-
lication, and a huge body of research has been conducted during that time, an
272 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

incredible amount of controversy continues to surround evolution. Much scie=- to a family (D1
tific debate has centered on five key questions: (1) Can new species develop if "'-: that the Hawai
founding populations are not somehow first separated geographically? (2) D. of finch that sc
evolutionary changes occur gradually or rapidly? (3) Is some general directi ~ than 15 to 20 n
toward increasing physiological or morphological complexity present in evoL- different specit
tionary history? (4) Does natural selection produce species that are perfec- _ events such the
adapted to their environments? (5) Has evolution produced a continuo (Fagus) trees fc
increase in species diversity over geologic time? All of these questions are hio share a commo
relevant for biogeographers. Let's tackle them individually and briefly review arated about 3
current state of debate. Ocean and Ar
plates separate
Isolation and Speciation Although
honeycreepers
Darwin's insights and writings on evolution via natural selection are genera[_ how many diff
clear and detailed. However, on the question of how natural selection leads to rt:- islands. We als
splitting of ancestral species into two or more new descendant species, Dan,-=: from one com
was uncertain, and his writings reflect this doubt. He clearly thought that evol.:- comes from tht
tionary changes caused by natural selection resulted ultimately in speciation, b:_ ferent species c
he did not deeply delve into how such splits might occur. We should not be t - renee of such
hard on Darwin, for the question of how evolution results in speciation is sri_ requires that n
hotly debated today. some instance~
Here is the problem we face when we try to explain how evolution ~ mental change:
changes lead to speciation. In order for new biological species to be formed, the:-:: However, such
must be some barrier to reproduction between the newly formed species and tt:: lids. In additi01
members of the original species. Evolutionary scientists refer to this barrier E! we find on con
reproductive isolation. Without isolation, the evolutionary changes that are dev _- large-scale ge<
oped within a population will be passed along throughout the entire species po.- America founc
ulation, and although new traits may arise and spread in this manner, new speci : barriers from c
are not created. For speciation to occur, new genotypic traits must develop, and tt:: The deve
individuals and populations that possess those traits must become reproductivet sympatric sped
isolated from populations of the original species. Once isolation occurs, the twc the same geogr
species can undergo further differences in their genetic composition as furtbe:- lation. New tra
natural selection or genetic drift occurs. population. It i~
In some cases, mutations can cause instant reproductive isolation and spe :- lated new spec
ation because the mutant individuals are genetically blocked from mating wi· ·- occur. For exan
members of the nonmutant population. Many plant species have arisen becaus.: tory of cichlid J
polyploidy occurred and produced new individuals with double the chromosom e.:: species must ha
number of the parent species. The mismatch in chromosome number between the are finding tha1
original species and the polyploid offspring produces instant reproductive isola- occur between
tion. How can reproductive isolation occur when there is no mutationally derivec in a number of
genetic barrier to reproduction to start with? The most obvious answer is geo- Differenc
graphic separation of the populations. Geographic barriers, including distanc . cause reproduc
prevent new traits that evolve in one population from being introduced to othe::- grasses that arE
populations. In time, geographically separated populations may develop geneti different time t
traits that preclude interbreeding. Darwin himself was led to the conclusion tha: soils. The differ·
geographic separation was an important means for speciation to occur. The for - toxic soils fror
mation of new species by geographic isolation is called allopatric speciation. adapted to gro'
Many phenomena can cause populations of species to be isolated geo- cal plants, parti
graphically. A chance dispersal event may cause a population to become estab- in their floweri
lished on a distant island or continent. The honeycreeper birds of Hawaii belong tion and maint<

und evolution. Much scien-
1 new species develop if th.:
ed geographically? (2) De
) Is some general directio::.
•mplexity present in evolt:.-
' species that are perfect!:
n produced a continuoc..;:
f these questions are highi:
ually and briefly review tL:-
tral selection are general._
atural selection leads to tt:e
lescendant species, Darwi::.
clearly thought that evok-
tltimately in speciation. be
::cur. We should not be to-=
results in speciation is st:..::.
explain how evolutiona_:-:
species to be formed, th e:-::
wly formed species and rt::
ists refer to this barrier .::..c
tary changes that are de\·e>
tout the entire species po;-
in this manner, new speci~
traits must develop, and tl:.:e
mst become reproductin · __
:e isolation occurs, the t\\ -
:tic composition as furth::-
ductive isolation and spec-
blocked from mating \\i · -
pecies have arisen becal!S::
h double the chromosoJ::J::.:'-.
•some number between L:;:e
instant reproductive i s o~­
is no mutationally deri\·.::.
Jst obvious answer is gec-
arriers, including distanc::
being introduced to ott::-
tions may develop gen e:::
; led to the conclusion tt.=..
oeciation to occur. The fo::--
1 allopatric speciation.
Jecies to be isolated gec-
pulation to become esra:--
Jer birds of Hawaii bela::.::
EVOLUTION AND SPECIATION 273
to a family (Drepanididae) that occurs nowhere else in the world. It is thought
that the Hawaiian honeycreeper species evolved from a single ancestral species
of finch that somehow reached these remote islands in the distant past but less
than 15 to 20 million years ago. The Hawaiian descendants of the finch are now
different species and genera from the mainland finches. In other cases, geologic
events such the splitting of continents can cause isolation. For example, the beech
(Fagus) trees found in Europe and North America belong to the same genera and
share a common ancestor, but they evolved into different species after being sep-
arated about 30 million years ago by the increasing size of the North Atlantic
Ocean and Arctic Ocean that occurred as the North American and Eurasian
plates separated.
Although large-scale geographic separation can explain why Hawaiian
honeycreepers are different species from continental finches, it cannot explain
how many different species of honeycreepers may have evolved on individual
islands. We also see many similar instances of a number of species evolving
from one common ancestor in aquatic habitats. The most striking example
comes from the cichlid fish in Africa. Lake Victoria alone supports over 300 dif-
ferent species of cichlids that likely arose from a common ancestor. The occur-
rence of such speciation on small islands or in isolated small waterbodies
requires that new species arise in the absence of large geographic barriers. In
some instances, cichlid species may have evolved allopatrically after environ-
mental changes caused lake levels to drop and populations to become isolated.
However, such events cannot explain all of the species diversity in African cich-
lids. In addition, it appears unlikely that all of the species of plants and animals
we find on continents could have evolved and speciated allopatrically through
large-scale geographic isolation. An analysis of mammal species in North
America found that only about 45% of species are separated by geographical
barriers from closely related species of the same genus.
The development of new species within the same geographic area is called
sympatric speciation. The challenge of sympatric speciation is that individuals in
the same geographic region have no physical barriers to cause reproductive iso-
lation. New traits that do arise through mutations can be passed throughout the
population. It is, therefore, difficult for genetically distinct and reproductively iso-
lated new species to arise. Yet, evidence exists that sympatric speciation does
occur. For example, detailed study of the DNA structure and environmental his-
tory of cichlid fish populations in African crater lakes suggests that a number of
species must have evolved sympatrically within the lakes. Evolutionary ecologists
are finding that the reproductive isolation required for sympatric speciation can
occur between individuals of the same species within the same geographic region
in a number of different ways.
Differences in life cycle timing, particularly the timing of reproduction, can
cause reproductive isolation. For example, it has been shown that certain races of
grasses that are genetically adapted to withstand toxic soil conditions flower at a
different time than other races of the same species which grow nearby on normal
soils. The difference in the timing of flowering prevents the species adapted to the
toxic soils from interbreeding with the more numerous individuals that are
adapted to growth on nontoxic soils. Similarly, many interfertile species of tropi-
cal plants, particularly orchids, remain reproductively isolated due to differences
in their flowering season. Differences in flowering times prevent cross-fertiliza-
tion and maintain separate species. Since the time of flowering is often controlled
274 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

by day length or temperature, it is easy to see how populations of plants distriJ::.-.

uted along long latitudinal gradients or across large elevational gradients wi....
flower at different times of the year and thus be reproductively isolated fr oc
other populations. One could imagine how California yarrow plants at low elevc-
tions would flower and be reproductively receptive to pollination early in tho::
spring, while high-elevation plants would not flower until later in the summe::-
Temporal patterns can also reproductively isolate animal populations. One re -
son why two closely related species of North American flies (Drosoph i~
pseudoobscura and D. persimilis) do not interbreed is because D. pseudoobscuru.
is sexually active in the evening and D. persimilis is sexually active at night. -~
similar situation occurs for the orange- and white-colored races of sulfur butter- [
flies (Colias eurytheme). The orange-colored butterflies are active in the mid-day.
while the white ones are active in the morning and late afternoon.
Mate choice in animals can be a complex process, and small differences C.
coloring or behavior may cause individuals to mate with one individual and no:
others. Barriers to breeding that are caused by behavior, particularly mate choi
are referred to as ethological isolation. There are many interfertile species an ·
races of cichlid fish in the lakes of Africa. It has been observed that small differ-
ences in male coloring or courtship behavior are critical for female mate choi fCold)
and this keeps the cichlid races and species reproductively isolated and allows fo:-
sympatric evolution. Somewhat similar mate choice behavior appears responsi- N
ble for sympatric speciation among stickleback fish (Gasterosteus) in lakes i.;: l(warm)
British Columbia. The males of many tropical bird species have very distincti\-- ~w~
(Dry
plumage, while the females have plumage that is somewhat similar to that o: 8
other species. It is thought that mate choice by females is, in part, responsible fo:-
the reproductive isolation that has led to such high rates of speciation among t
these birds. In groups such as the birds of paradise found in New Guinea, the FIGURE 9.4
males must possess the proper plumage and engage in specific elabora t land masses.
courtship behaviors in order to entice the female to mate. In North America, the
mallard duck (Anas platyrhynchos) and pintail duck (A. acuta) can physically
interbreed but do not because the females do not recognize the courtship beha\-
iors of males of the other species. these offsprin1
Finally, the difference between what we define as allopatric speciation an and usually pe
sympatric speciation may be largely a problem of scale. We might consider a larg the inability ol
field of shrubs to be one geographic unit. However, for stenophagous insects tha: the evolutionc
spend their entire life on one individual plant, such a field is really an ocean o:' same geograpl
isolated islands. Relatively little gene flow may occur between the insect popula- likely at the cc
tions on different plants. For example, at least 27 different species of weevils have
evolved on the 6-km sq island of Rapa in the South Pacific. This tiny island pro- The Temp(J
vides these small insects with a checkerboard of different habitat types and geo-
graphic locales for adaptation and speciation. For many species, even a fairly Darwin clear]
small area presents a patchwork of many different habitats. By restricting their occurred main
activities to certain habitat types, animal populations become isolated from other traits that inf,
populations. Genetic drift and natural selection for traits that impart reproduc- become domir
tive success in the specific habitat occupied by the population lead to genetic dif- as phyletic gn
ferentiation and eventually to speciation. Habitat differences within a given area through a nun
can also promote reproductive isolation by influencing mortality rates among off- tion to genen
spring. For example, two closely related species of iris live in the Mississippi Delta should be pres
region. Iris fulva grows on drier river banks, while Iris giganticaerulea grows in years provides
damp marshlands. The species are interfertile and can produce hybrids. However. sils. Over tim(
 EVOLUTION AND SPECIATION 275

pulations of plants distri Present Future

elevational gradients wi[ Tectonic
Jroductively isolated fra n:: ..__
Movement
~
Allopatric (Vicariance)

yarrow plants at low eleva-

to pollination early in tb.:
until later in the summe~
mal populations. One rea-
8EJB
..__ ~

Jump Dispersal
nerican flies (Drosoph il::. Allopatric (Dispersal and Founder Event)
~
because D. pseudoobscur::.
;;exually active at night. _:.._
>red races of sulfur butt e~­
s are active in the mid-d a\
EJD EJB
J
Peripat~ic~SpB
~ afternoon.
,_ -t
;s, and small differences i:.
ith one individual and no:
r, particularly mate choice:
I= SpA SpA

- ~~spc
ny interfertile species an.: ~

:>bserved that small diffe ~­ Sympatric (Parapatric)

:al for female mate choice. fCold) ~
'ely isolated and allows fo-
,ehavior appears respons:-
(Gasterosteus) in lakes i:.
N
I(Warm)-
~
ecies have very distincti\·;: .-w~
S (Dry) E(--..W
newhat similar to that e:
; is, in part, responsible fc~
+ ot)

rates of speciation am o c~
ound in New Guinea, tt:-
'lge in specific elabora::-
ate. In North America, c~ -
(A. acuta) can physicaL_
gnize the courtship beha'.-
s allopatric speciation ar..:
We might consider a larg:-
stenophagous insects tb--
field is really an ocean e·
1etween the insect populc.-
~nt species of weevils ha' :-
'lcific. This tiny island prc-
ent habitat types and gee-
any species, even a fair..
bitats. By restricting the::-
~come isolated from othc
1its that impart reprod uc-
Lilation lead to genetic cii:-
rences within a given ar e~
mortality rates among o:=-
ve in the Mississippi D e l~
s giganticaerulea grows i:::-
,roduce hybrids. Howew :
FIGURE 9.4 Different possible modes of geographic speciation. The rectangles represent
land masses.
these offspring are not successful in growing on either the dry or very wet sites
and usually perish. Thus, genetic mixing between the two species is restricted by
the inability of the hybrids to survive. This form of speciation, which is caused by
the evolutionary divergence of populations that occupy different habitat in the
same geographic area, is called disruptive selection, or parapatric speciation. It is
likely at the core of sympatric speciation through natural selection (Fig. 9.4).
The Temporal Pattern of Evolution
Darwin clearly thought that evolution and speciation via natural selection
occurred mainly as a slow and gradual process. New traits arise by mutations, and
traits that infer greater reproductive success are selected for and eventually
become dominant over many generations. Evolution in this manner is referred to
as phyletic gradualism. If evolution proceeds in this manner, species should go
through a number of transitional forms as natural selection works from genera-
tion to generation to favor certain adaptations. The earlier transitional forms
should be preserved as fossils. The fossil history of horses over the past 55 million
years provides evidence of such gradual evolution in the form of transtional fos-
sils. Over time, the size and shape of the horses ' long bones, skulls, and teeth
276 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

changed as the group developed from a very small ancestor and speciated in: ferent taxa. Pa
the clade that include horses, zebras, and donkeys today. However, even in Da:- associated witl
win's time, critics of natural selection pointed out that such transitional forn:... the speciation
were rare in the fossil records of evolving species. Instead, the fossil record oft ~ Gould's cancer
shows that species persist for long periods of time with little outward chan g ~ that species pt
Some paleontologists use the term stasis to refer to these long periods of time ::. occurs. These r•
which fossil lineages seem to undergo little change. The arrival of new specie- stability of the
appears to occur very rapidly, often with no evidence of transitional forms. Tt.= So, which
fossil records of shellfish, for example, often include long periods of stasis aiL: equilibrium? V
rapid speciation events. Interestingly, such periods of stasis often occur at tt= believed that b
same time for many different species living within the same region. These episod tion. This is pr
are called coordinated stasis. Darwin and many other evolutionists explained l = mon for organi
lack of transitional forms by stating that the fossil record is often imperfect, ari: areas of simila
transitional forms might be missing because either they had not been fossilir ..: small bays tha
due to geologic conditions or their fossils had not yet been discovered by paleo:.- peripheral envi
tologists. However, by the middle of the twentieth century, after much paleon gence. The pun
logical research had been conducted, it became clear that some speciation eve :.. our understanc
appear to have occurred very rapidly and there is little evidence for transitio n~ from larger an•
forms existing in certain cases. cept explains v
In the 1940s the great biogeographer, G. G. Simpson introduced the conce;:: replaced witho
of quantum evolution to explain the apparent rapid appearance of new speci · lineages, such ;:
He proposed that many evolutionary changes arise from very small populatio;:_ gradual evoluti
that lie at the periphery of the main ranges of the species, or are geographica.:... ism and punctu
isolated. Simpson argued that evolutionary changes were more likely to occur c mation of hom
such small populations, but due to the small numbers of individuals involved, fos- millions of ye<
sil evidence of such changes would be rare. Eventually, traits developed in su . _ cotherium pass
small populations could, if they were particularly beneficial, spread rapi ·_ possessed by m
throughout the larger population. The fossil record would capture the arrival : forms with red1
these new forms, but not their development in small, isolated populations. cotherium and i
Simpson's ideas were developed and refined by the noted paleontologis:: the transitions
Steven J. Gould and Neals Eldredge. Gould is famous for his popular writings o:. idly. Punctuate<
evolution, natural history, and the history of science. In the 1970s Eldredge ar;: evolution and :
Gould proposed that most new species arise from small peripheral or isolat - events. Phyletic
populations. Such small populations are prone to relatively rapid speciation . I:. most clearly ap
addition, many such peripheral and isolated habitats present different enviro::.- tion encompas~
mental conditions and selective pressures than are found in the main geograp : It might t
range of the species. Due to natural selection related to their distinctive enviro:.- can evolutiona
ments, and genetic drift, the peripheral populations will evolve traits that are ct:- actual number ·
ferent from those possessed by the main ancestral population of the species. Tt:: highly variable
evolutionary changes that occur in such small populations are not likely to ::. severity of sele
well represented in the fossil record. were accidenta
Eldredge and Gould (1977) postulated that environmental change als - increase in win
plays a role in the timing of large-scale speciation events. If the environme;:- have been seer
changed and produced conditions in the main range of the species that we-::: become isolate
more typical of those experienced by the peripheral races or species, the ance:>-- to be exception
tral species would be replaced by the peripheral population. They felt that tt::
rapid speciation events apparent in the fossil records of many species actuaL._ D irection in
represented periods of environmental change that allowed peripheral races aL:
species to replace the main ancestral species. For example, the rapid speciatio:. Is there a gener
events for shellfish in Lake Turkana, Africa, occur at the same time for many dif- tion leads to m<

.cestor and speciated in tc
ty. However, even in Da:--
tt such transitional forrn5
ad, the fossil record ofte::.
ith little outward change
se long periods of time :-
he arrival of new species
of transitional forms. Th::
ong periods of stasis aL::
stasis often occur at tt::
me region. These episode:-
volutionists explained t2::
•rd is often imperfect, ac.:
:y had not been fossilize::
:en discovered by paleo::.-
:ury, after much paleontc--
at some speciation ever::.c
: evidence for transiti o c~
Jn introduced the con c e~
Jpearance of new specin
•m very small populatio:::...
;ies, or are geographica:.:.
:re more likely to occur :::
f individuals involved, f~
r, traits developed in su.--
Jeneficial, spread rapi -
•uld capture the arrival c
)lated populations.
the noted paleontologis:..
or his popular writings c:
n the 1970s Eldredge a::.:
tall peripheral or isolat::-.:
tively rapid speciation. ::.:;
Jresent different enviro::.
1d in the main geograpL
> their distinctive enviro::.
l evolve traits that are ·
ulation of the species. T::.:
tions are not likely to :--:
tvironmental change a3
vents. If the environmc
of the species that \W ::-:
1ces or species, the a n c~
1lation. They felt that L:.:
of many species actuL
•wed peripheral races c:;,_
nple, the rapid speciat:.--
e same time for many ·
EVOLUTION AND SPECIATION 277
ferent taxa. Paleoenvironmental analysis shows that these speciation events are
associated with environmental changes caused by shifts in lake levels. Between
the speciation events there are long periods of coordinated stasis. Eldredge and
Gould's concept of evolution is referred to as punctuated equilibria. They suggest
that species persist for long periods of stasis when little evolutionary change
occurs. These relatively stable species are then replaced by new species when the
stability of the environment is punctuated by environmental changes.
So, which mode of evolution is correct, phyletic gradualism or punctuated
equilibrium? When Simpson developed the concept of quantum evolution, he
believed that both quantum evolution and gradualism operated in driving evolu-
tion. This is probably correct. Punctuated equilibrium seems particularly com-
mon for organisms such as shellfish, which occupy lakes possessing large central
areas of similar habitat but also contain peripheral areas of shallow water or
small bays that offer different environmental conditions. It is in these small
peripheral environments that the small populations undergo evolutionary diver-
gence. The punctuated equilibrium concept is attractive because it conforms to
our understanding of how small populations can evolve differently and diverge
from larger ancestral populations. In addition, the punctuated equilibrium con-
cept explains why many fossil species seem to persist for so long and then are
replaced without evidence of transitional forms. However, fossil evidence from
lineages, such as the horse, includes transitional forms that allow us to trace the
gradual evolution of certain traits. Perhaps the conflict between phyletic gradual-
ism and punctuated equilibrium is also a matter of scale. When we look at the for-
mation of hooves in horses, we can see the transition that occurred over tens of
millions of years between the four separate toes that the early genus Hyra-
cotherium possessed, the three toes of intermediate genera, and the single hoof
possessed by modern horses. However, we have little evidence of the transitional
forms with reduced fourth toes that might have existed between four-toed Hyra -
cotherium and its three-toed descendants. In addition, the fossil record shows that
the transitions between four-toed , three-toed, and hooved horses occurred rap-
idly. Punctuated equilibrium provides insights on the pattern and mechanism of
evolution and speciation that might occur at the scale of individual speciation
events. Phyletic gradualism, identified by distinctive transitional forms, is perhaps
most clearly apparent when we step back and look at the long histories of evolu-
tion encompassing many ancestor and descendant species.
It might be asked, how fast, in absolute terms, such as years or generations,
can evolutionary change and speciation occur due to natural selection? The
actual number of years it takes for a species to evolve and speciate appears to be
highly variable and dependent on both the genetics of the organism and the
severity of selection pressure. Studies of fruit flies (Drosophila subobscura) that
were accidentally introduced to California show that a genetically controlled
increase in wing size developed in less than 20 years. Similarly, rapid changes
have been seen in increasing wing sizes in Amazon bird populations that have
become isolated in forest fragments. However, these very rapid changes appear
to be exceptional.
Direction in Evolution
Is there a general direction in evolutionary development? We might ask if evolu-
tion leads to more complex organisms? Many nineteenth- and twentieth-century
278 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

scientists studying evolution felt that there was a general direction in evoluti ~ Some ev
that leads to organisms that are more complex in form and function. Darwin w::... tion through 1
strongly influenced by the concept that there was a progression in evolutiona;- change that a
development from simple to more complex forms. He used terms such as lowe- million years.
and higher forms of life to rank the evolutionary development of plants and a ·- time scales of
mals. Darwin considered humans to be the highest form of life and single-cell e~ important not
organisms, such as bacteria to be the lowest forms. He saw evolution as the pr quently, a tyr:
gressive development from lower to higher life forms. 20,000-, 40,00(
Most modern evolutionists do not subscribe to the idea that there is a g ;::-. ticular climati
eral pattern of evolutionary development from simple to complex. In fact , it .::. subsequent ch
often quite difficult to define what is meant by the terms simple and complex. =..: continuous di
you look at the flower of a grass plant, you will see a very tiny and sim p~:: species to ace
arrangement of reproductive organs. The grass flower lacks petals, bright colo::.. tions through
and nectar. In contrast, the flowers of the magnolia are large, have bright pet rus. that for the C
and possess specialized organs for producing sweet-smelling nectar. From an e\·c- rna tic change.
lutionary perspective, however, the grasses evolved fairly recently, in the Terti a,~ of the boreal J
In contrast, magnolias are an ancient genus of plants that developed their form ~.:­ forest plants a
flower in the Cretaceous, tens of millions of years before the grasses. The simp:...- east coast wht:
fied flowers of the grasses are evolutionary reductions that evolved from see - in the modern
ingly more complex forms. There are many cases from the fossil records of la.:::c: boreal climate
and marine plants and animals where morphologically complex species ha :: glaciation and
given way to reduced forms over time. ticity to survi
One interesting general pattern that occurs in the evolutionary history L - found in thest
many species is a trend toward larger size as the lineage evolves. Horses, f zone but may
example started as a small dog-sized mammal and experienced a progres i :: must be careft
increase in size as they evolved into the modern genus Equus. This general pa:- ent environmc
tern of increasing size is referred to as Cope's Rule. There are, however, mar.. ern environmt;
exceptions to this pattern.
Speciation through evolutionary development is driven by natural selecti =
to specific environmental conditions and by chance factors, such as mutations ar:~ Increasing
genetic drift. Natural selection works to promote reproductive success above a_
We might ask i
else. Greater complexity in form or function does not necessarily translate ;
animal species
greater reproductive success. Thus, one cannot assume a general direction in eYo-
restrial or mar
lution from complex to simple.
species, gener2
families of ver
Perfection in Evolution past 2 million
Can evolution and speciation be considered to produce perfect optimal adap ta- million years ;
tions to environmental conditions? The degree of morphological and physi vertebrates. It
logical change that can occur within a species is governed by genetic constrain ' life first began
that limit the degree of change that can occur in the physiology and morphol- years, the fossi
ogy of a species through evolution. In a classic discourse, Gould used the exam- lies is not as cl
ple of the panda's thumb to illustrate this principle. Pandas must strip larg"' understanding
amounts of leaves from bamboo plants in order to sustain themselves. The pa past few millio
of the panda appears to have a set of digits and a thumb, which allows the ani - more distant
mals to grab the bamboo and strip the leaves. However, pandas have evolve · increase in the
from carnivorous, clawed ancestors that did not possess a thumblike articulate · years are mo1
digit. Upon closer inspection, the thumb of the panda is actually an elongate · increasing dive
wristbone that serves as an imperfect replacement for a real thumb provided b\- ent have chan1
a flexible digit. increased signi

~raldirection in evolutio::
and function . Darwin was
·ogression in evolutionar:
used terms such as lowe:-
.opment of plants and mi.-
m of life and single-celle.:
saw evolution as the pro-
1e idea that there is a ge::-
~ to complex. In fact , it ~
ms simple and complex. ::.:
e a very tiny and simp:::-
lacks petals, bright color""
~ large, have bright petE
=lling nectar. From an e\ ·c~
Jy recently, in the Tertia;:
at developed their form o:
•re the grasses. The simp:..:-
: that evolved from see ~­
L the fossil records of laL.:.
lly complex species ha\::
1e evolutionary historY
teage evolves. Horses. : --
!xperienced a progress:· =
s Equus. This general pc.:-
There are, however, rna:.
jriven by natural select:.::=
cors, such as mutations a::..:.
oductive success above 2.-
)t necessarily translate :
a general direction in e\ --
;e perfect optimal adap:__
torphological and phy s :~
ned by genetic constra.C-
physiology and morpL
rse, Gould used the ex · -
. Pandas must strip Ia:-.:-
stain themselves. The p~
.mb, which allows the c=
ver, pandas have evol\ ::-_
ss a thumblike articul2:::-_
a is actually an elong2:::-_
a real thumb providec
EVOLUTION AND SPECIATION 279
Some evolutionary scientists have also pointed out that the rate of adapta-
tion through natural selection may be slower than the rate of environmental
change that a species encounters. For example, many species persist for over 1
million years. In contrast, the Milankovitch cycles of climatic change occur on
time scales of 100,000 years and less. These climatic cycles appear to have been
important not just in the Quaternary, but earlier in geologic time as well. Conse-
quently, a typical species must survive environmental changes that occur on
20,000-, 40,000-, and 100,000-year cycles. If a species is highly adapted to a par-
ticular climatic or environmental condition, it may not be able to survive the
subsequent change in environment brought on by the Milankovitch cycles. The
continuous disruptions caused by climatic changes may restrict the ability of
species to accumulate very fine adaptations to particular environmental condi-
tions through natural selection. Some evolutionary scientists have concluded
that for the Quaternary period in particular, species are often adapted to cli-
matic change. For example, during the last glacial maximum, the present location
of the boreal forest biome in North America was largely covered by ice. Boreal
forest plants and animals lived in southern areas such as the central plains and
east coast where seasonal differences in hours of daylight are less extreme that
in the modern boreal zone and where the climate was different than the modern
boreal climate. Plant and animal species that have survived through the cycles of
glaciation and nonglaciation must have enough phenotypic and genotypic plas-
ticity to survive this wide range of environmental conditions. Certain traits
found in these species may not be optimal for survival in the modern boreal
zone but may have been important for survival during glacial periods. Thus, we
must be careful in assuming that all species are optimally adapted to their pres-
ent environments or that traits we observe today represent adaptations to mod-
ern environmental conditions.
Increasing Species Diversity
We might ask if there is a continuing growth trend in the total number of plant and
animal species on earth over geologic time? At first glance, the fossil record of ter-
restrial or marine plants and animals appears to show a clear trend of increasing
species, genera, and family numbers over geologic time. For example, almost 300
families of vertebrate terrestrial animals are known from the fossil record of the
past 2 million years, while fewer than 100 are known from the Triassic some 200
million years ago. Similar trends are apparent for terrestrial plants and marine
vertebrates. It is obvious that there are more species today than there were when
life first began in the Precambrian. However, during the past 100 or 200 million
years, the fossil evidence for ever increasing numbers of species, genera, and fami-
lies is not as clear as it first seems. We have many more fossils and a much better
understanding of the taxonomy of recent plants and animals that lived over the
past few million years than we have for the plants and animals that existed in the
more distant geologic past. Some paleontologists believe that the apparent
increase in the number of species, genera, or families over the past 200 million
years are more an artifact of the fossil record than a true representation of
increasing diversity over recent geologic time. Although the species that are pres-
ent have changed over time, it is not completely clear that species numbers have
increased significantly or uninterrupted during the past few hundred million years.
280 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

GEOGRAPHY AND EVOLUTION: (Tanysiptera) i

FOUNDER EFFECTS, BOTTLENECKS, contain at leas1
VICARIANCE EVENTS, ADAPTIVE RADIATION, of kingfishers. '
AND EVOLUTIONARY CONVERGENCE main island an
species of king
Now that we have reviewed some basics, let's examine a little more closely ho Tanysiptera gal
geography may impact on processes and patterns of evolution and speciation. : a combination
is thought that rates of evolutionary change and speciation are enhanced wh - populations of
very small populations are geographically separated from their parent popu ~­ contributed to
tion. In such cases, the smaller population often has a different genetic camp o ~ island kingfishE
tion than is average for the larger populations. For example, imagine a smL Many un
group of tourists that become stranded on a remote island. The group might o · islands of the F
contain individuals with genes for blond hair and blue eyes. This is a subsample : tlenecks that c
the entire wide range of human genes for hair and eye color. Although, G:::o colonizing pop
stranded tourists may reproduce and increase in population size, their offsp~ islands and ur
will have less genetic diversity than is evident in the entire human populati =.. development c
The populations that develop following such founder events appear to be pan :::- selection. Thrc
ularly prone to speciation because they are already genetically different from :: develop traits
ancestral population and new alleles that may develop via mutations rema= island. Many SI
within the isolated populations. In addition, the isolated population is cut o=: Zealand, are fl
from alleles that may develop in the ancestral population. The idea that su - fliers. Hawaii, f,
events promote rapid allopatric speciation in geographically isolated populatio=- closely related
(Fig. 9.4) is called the founder principle. proportion of
Consider another scenario in which half of the stranded tourists sail aw~ _ species of islan•
and half remain. Some of the genetic diversity that had existed on the island \\-:_ have reduced <
have been lost because of the emigration of the people who sailed away. R :c growing on the
decreased genetic diversity that results from a decreased population size and as island because
ciated decrease in genetic diversity is called a bottleneck. Genetic bottlenec ·:: large predaton
generally occur when environmental factors, such as a rapid change in clima ~ escape predatic
cause the population of a species to be greatly reduced in size. During such ca ~­ Unfortun.
strophic events, it is likely that certain genes will be lost from the species becaus= and alien preda
all individuals carrying those genes perish. The small population that survive ::. duction of pred
bottleneck may produce offspring and increase in numbers. However, the poS":- itous decline i1
bottleneck population does not contain the range of genetic diversity found in tb:o species went ex
original ancestral population. Over time, mutations lead to the introduction : gle remaining •
new alleles and the development of new traits. Some of these traits may be advar.- shorter wings tl
tageous and are selected for by natural selection. Others become common a ::. Even with mod
result of genetic drift. In any case, because the population is small, these allel : of extinction. B
spread rapidly among it. If the population is isolated from the original ancestr2... today due to pr
population, these traits do not spread back to the original population. In some c
Because of the founder effect, speciation is often more likely to occur at tht- logic events su•
edges of species range than in the central portions of their geographic distribe- ranges of speci•
tions. Small isolated populations at the edges of species geographic distributionS" events (Fig. 9.{
often have lower genetic diversity than is found in more central populations.. speciation can <
Events that cause such peripheral populations to become geographically isolat · trees into sepal
from the main population accelerate genetic divergence and speciation throu~ formation of th
the founder effect. This geographic pattern of speciation is sometimes referred tc can also produ
as peripatric speciation (Fig. 9.4). A classic example of divergence and speciatio species and the
due to the founder effect acting on geographically peripheral populations is pro- Ice Complex SI
vided by the distribution of different races and species of kingfisher birdS" portions. Gene

IN,
e a little more closely ho'"'
volution and speciation.:-
:iation are enhanced whc=
from their parent popul;;.-
different genetic compos-
example, imagine a smC.::.
and. The group might O IL~
eyes. This is a subsample c:
eye color. Although ,
1lation size, their offsp ri.::.~
entire human populatio=--.
;vents appear to be pani.::-
letically different from C::=
lop via mutations rema:=
ated population is cut o::
lation. The idea that su.::.::.
tically isolated populatio=:
;tranded tourists sail aw::.
j existed on the island w:_
>ple who sailed away. 1'.:=
d population size and ass..:::-
neck. Genetic bottlenect..
a rapid change in clima:=:
i in size. During such ca ~­
.t from the species becau..<.::
population that survives _
mbers. However, the pos:-
netic diversity found in ~=
;ad to the introduction ;:_
these traits may be adva::::-
ters become common as :
1tion is small, these aile!.:::
'rom the original ancestr.::....
1al population.
more likely to occur at -~­
their geographic distrib:..-
~s geographic distributio=:
more central populatio- -
ne geographically isolate
ce and speciation throu: ·
,n is sometimes referred ::
divergence and speciati0:::
tpheral populations is prc-
>ecies of kingfisher bir -'
GEOGRAPHY AND EVOLUTION 281
(Tanysiptera) in New Guinea. The outer coastline and islands of New Guinea
contain at least two different species and eight morphologically distinctive races
of kingfishers. The main species, Tanysiptera galatea, is found at the center of the
main island and on the largest of the surrounding islands. It is likely that the
species of kingfishers on the small islands and coastal areas have diverged from
Tanysiptera galatea into new races and one new species. The divergence is due to
a combination of genetic drift and natural selection acting on small peripheral
populations of birds. Rising sea levels at the close of the Pleistocene may have
contributed to the geographic isolation and peripatric speciation of the outer
island kingfisher populations by creating and isolating islands.
Many unique species of birds, invertebrates, and plants found on remote
islands of the Pacific Ocean are the result of founder effects and population bot-
tlenecks that occurred following long-distance dispersal and colonization. The
colonizing populations of animals and plants arrived from the mainland or other
islands and underwent evolution and allopatric speciation. The evolutionary
development of an isolated species can be caused by genetic drift or natural
selection. Through natural selection, a species on an isolated island might
develop traits that are related to increased survival and reproduction on the
island. Many species of island birds, such as the kiwi (Apteryx australis) in New
Zealand, are flightless. The continental relatives of these birds are often good
fliers. Hawaii, for example, had at least eight species of flightless geese that were
closely related to the migratory Canada goose. Darwin himself noted the high
proportion of flightless birds on islands. Flightlessness is also seen in many
species of island insects. In addition, the seeds of a number of island plant species
have reduced capacity for wind dispersal when compared with close relatives
growing on the mainland. The inability to fly might be selectively favored on an
island because it keeps individuals from getting blown out to sea. In many cases,
large predators are absent on islands, so being able to fly is not as crucial to
escape predation as it might be on the mainland.
Unfortunately, flightless bird species are extremely vulnerable when humans
and alien predators arrive on islands. The arrival of human hunters and the intro-
duction of predators such as cats, rats, and dogs to Hawaii contributed to a precip-
itous decline in the population of Hawaiian geese. All of the flightless geese
species went extinct after the arrival of Polynesians about 1500 years ago. The sin-
gle remaining species of Hawaiian goose, the nene (Branta sandvicensis), has
shorter wings than its mainland relatives but can fly moderate distances (Fig. 9.5).
Even with moderate flying abilities the nene has not been excluded from the risk
of extinction. By 1951 only 30 nene remained on the islands. The species survives
today due to protection and propagation using captive populations.
In some cases, large populations become geographically separated by geo-
logic events such as the splitting of continents. Such changes, which divide the
ranges of species into geographically isolated distributions, are called vicariance
events (Fig. 9.4). Once the populations are split by geographic barriers, allopatric
speciation can occur. The divergence of North American and Eurasian deciduous
trees into separate species is the result of such a vicariance event caused by the
formation of the Atlantic Ocean and Arctic Ocean. Large-scale climatic changes
can also produce vicariance events, which lead to the geographic isolation of
species and the evolution of new species. During glacial periods, the Laurentide
Ice Complex split the North American Boreal biome into western and eastern
portions. Genetic analysis of jack pine (Pinus banksiana), common to eastern
282 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

FIGU RE 9 .5 The last surviving species of Hawaiian goose, the nene (Branta
sandvicensis). Like a number of species of bird and insect species that have evolved on
remote islands, all the other species of Hawaiian goose have lost their ability to fly. Thos e
species are now extinct. The nene can still fly moderate distances.

FIG URE 9.6

North America, and lodgepole pine (Pinus contorta), common to western No r~ (Pinus contortc;
America, suggests that these two species developed from a common ance s t r~ development o
population that was split apart by an advance of the Laurentide Ice Complex an~ the ancestral pi
geographically isolated some 500,000 years ago. During glacial periods the ance5-
tors of jack pine grew in the eastern United States, while the ancestors of lodge-
pole pine were restricted to the Rocky Mountains westward. During the perio · · In the c
of separation caused by continental glaciation, the single ancestral speci-: ariant specie1
diverged allopatrically into the two modern species (Fig. 9.6). terfiy, bird, a
The large populations created by continental scale vicariance events cor:- created when
tain large amounts of genetic diversity relative to the total diversity of the ances- ranges into t'
tral population. This contrasts with the lower amounts of genetic diversi _ separate spec
contained in small founder event populations. In theory, evolutionary divergen ': some 10 to L
and speciation will occur more slowly within these large vicariance populatior:s shows that ev
and more quickly in small founding event populations. There is some eviden ;:- the lowlands
that species differentiation in large vicariance event populations does indeec Founde
occur relatively slowly. For example, the general morphology, favored clima · · gence of om
conditions, and growth habitat typical of the European beech tree (Fagus sylva:- more than c
ica) and the North American beech species (Fagus grandifolia) are extremely found on the
similar. A very close similarity in the niches of the beeches persists, despite the- developed fr
fact that the vicariance speciation event that led to the creation of these two water bodies.
species was the separation of North America from Eurasia some 30 million years results from <
ago. Many Eurasian species of plants, such as sycamores (Platanus) not only haH· species evolv
similar niches with North American relatives, but reproduce easily with thei.:" colonized re~
North American representatives despite a separation of some 30 million year a species is sc
Jack and lodgepole pine also hybridize when they grow together today (Fig. 9.6). ent honeycre

he nene (Branta
:ies that have evolved on
lSt their ability to fly. Tho se
;es.
:ommon to western Nor...:
'rom a common ance st::-~
urentide Ice Complex a;::.:
~ glacial periods the ancQ-
ile the ancestors of lodg.:-
tward. During the peri<X._
single ancestral speci::-
g. 9.6).
1le vicariance events co::-
)tal diversity of the ancQ-
unts of genetic diversi~
y, evolutionary divergen.::::
ge vicariance populatio::_
s. There is some eviden.::::
populations does inde.:.:
phology, favored climae::
1 beech tree (Fagus sy/n;.:-
randifolia) are extreme._
eches persists, despite tL·
:he creation of these t\\ -
asia some 30 million ye a:-
s (Platanus) not only ha'::
produce easily with the
of some 30 million ye<L5.
' together today (Fig. 9.
GEOGRAPHY AND EVOLUTION 283
0 Laurentide and • Lodgepole Pine
Cordilleran Ice
0 Jack Pine
• Lodgepole Pine
Hybrids
Ill Jack Pine
(General Distributions )
FIGURE 9.6 The modern distribution of two closely related tree species, lodgepole pine
(Pinus contorta) and jack pine (Pinus banksiana). It is thought that divergence and the
development of these two species occurred allopatrically when Pleistocene glaciations split
the ancestral pine distribution into separate western and eastern populations.
In the case of animals, a similar close correspondence in the niches of vic-
ariant species was shown in an examination of 37 different pairs of upland but-
terfly, bird, and mammal species in southern Mexico. These species pairs were
created when the formation of the Isthmus of Tehuantepec Lowlands split their
ranges into two separate areas. Allopatric speciation then led to the evolution of
separate species north and south of the lowlands. This vicariance event occurred
some 10 to 2.4 million years ago. Yet a careful analysis of the species' niches
shows that evolutionary divergence between the species to the north and south of
the lowlands has not been very great.
Founder events and vicariance events can explain the evolutionary diver-
gence of one species into two. However, in many cases speciation produces
more than one new species. The many species of honeycreepers originally
found on the Hawaiian Islands or cichlid fish species found in African lakes
developed from the arrival of a single founding species at isolated islands or
waterbodies. The development of many species from a single species often
results from a process called adaptive radiation. During adaptive radiation, new
species evolve from a common ancestor to fill all of the available niches in the
colonized region. The niche space that is occupied by the adaptive radiation of
a species is sometimes referred to as the adaptive zone. For example, the differ-
ent honeycreepers on the Hawaiian Islands have a wide range of beaks that
284 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

allow different species to take advantage of different foods. Some species - have evolved
honeycreepers have short, thick beaks adapted for eating fruit and seeds (E; period of les5
9.7). Others have beaks that are adapted for prying bark apart to find insec-..:. lizards of the
Finally, some species possess long and slender beaks with which they ob ta_ gence might
nectar from flowers. Cichlid fish in African lakes display a similar wide varie. were experirr
of morphological adaptations to food and habitat that have developed throu_:- covers. The is
adaptive radiation (Fig. 9.7). By evolving traits that allow individuals to tal.:= of years. The
advantage of different foods and habitats, and thus decrease niche overlc..:;: size after a pE
competition is decreased between the species and carrying capacity _ The pro
increased. Among other forms, some cichlids have large mouths adapted iL great variety
eating fish, others have small mouths adapted for consuming zooplankton, a::_:: Since natura
still others possess sharp, pointed mouths adapted to picking invertebrates c= expect that si
of aquatic vegetation and the lake bottom. different spec
In comparison with the apparently slow rate of species divergence seen : : : . conditions. 11
the large vicariance events, evolutionary divergence and adaptive radiation · unrelated spe
small founder populations often occur rapidly. The cichlid fish in Lake Vict o:-~ evolution. Th
of Australia <
and Eurasia.
deserts, savan
has been sep:
native mamn
,o that carry the
tinue to matu
Hawaii possum (Dea
Telespyza cantans Pseudonestor xanthophyrys comprised of
external sack
ing variety of
behavior to I
tralia. These i
version of a k
tion to fill thE
Loxops virens Hemignathus procerus were selected
tion produce,
their placent<
Conver1
that evolved i
to the cacti th
course, why 1
look similar 1
Docimodus johnstoni Haplochromis polydon continent to <
Africa
cial in produc
mate that is a
One int
the very high
mon ancestOJ
Labidochromis vellicans Rhamphochromis macrophthalmus evolution is u
lations derive
logically sim
FIGURE 9. 7 Two examples of adaptive radiation: Honeycreepers from Hawaii (after Hispaniola, J
Raikow, 1977) and cichlid fish from Africa (after Fryer and lies, 1972). species of an<

~nt foods. Some species c:
eating fruit and seeds (Fig
, bark apart to find insecG.
ks with which they obtai::
>play a similar wide vari e ..
at have developed throug._
t allow individuals to t a k~
1s decrease niche overla::
and carrying capacity :s
large mouths adapted fc::-
'nsuming zooplankton, ar::.
o picking invertebrates o""
species divergence seen :.:.
and adaptive radiation .
ichlid fish in Lake Victor_:
,o
D that carry their young in sacks called marsupium where the young feed and con-
~?0
Hawaii
{)
~
rs
epers from Hawaii (after
1972).
GEOGRAPHY AND EVOLUTION 285
have evolved into hundreds of new species with extremely different niches in a
period of less than 12,000 years. Experiments have been conducted using Anole
lizards of the species Anolis sagrei to try to assess how quickly evolutionary diver-
gence might occur for small founding populations. A small number of anoles
were experimentally introduced to several islands that have different vegetation
covers. The islands were revisited, and the lizards were examined over a number
of years. The anoles displayed significant evolutionary divergence in shape and
size after a period of only 10 to 14 years.
The process of adaptive radiation is fundamental to the development of the
great variety of plant and animal species that exist today and existed in the past.
Since natural selection is a response to environmental conditions, we might
expect that similar morphological and physiological traits will be selected for in
different species that evolve in different regions that have similar environmental
conditions. The development of similar morphological or physiological traits in
unrelated species living in geographically separated regions is called convergent
evolution. The classic examples of convergent evolution are the marsupial fauna
of Australia and Tasmania, and the placental mammal fauna of North America
and Eurasia. The continent of Australia has a host of environments, including
deserts, savanna, tropical rainforests, temperate forests, and grasslands. Australia
has been separated from all other continents for some 50 million years, and its
native mammal fauna has been dominated by marsupial mammals (mammals
tinue to mature). Aside from some marsupial species in the Americas, such as the
possum (Dedelphis virginiana), the mammal fauna of the rest of the world is
comprised of placentals (mammals such as humans that do not carry young in
external sacks). When Europeans first arrived in Australia, they found an amaz-
ing variety of marsupial species that were close counterparts in morphology and
behavior to placental mammals found in similar environments outside of Aus-
tralia. These included a marsupial "cat," a "wolf," a "mole," and even a marsupial
version of a kangaroo rat (Fig. 9.8). These marsupials evolved via adaptive radia-
tion to fill the many niches offered by the Australian environment. Similar traits
were selected for in Australia as on the other continents, and convergent evolu-
tion produced similarities between the species of marsupials in Australia and
their placental counterparts that lived elsewhere.
Convergent evolution also occurs in plants. The succulent euphorb plants
that evolved in the deserts of Africa and Eurasia share gross similarities in form
to the cacti that evolved in the Americas. The convergent evolution of plants is, of
course, why the tropical forests in Africa, Southeast Asia, and South America
look similar but are largely comprised of very different species of plants from
continent to continent. The prevalence of convergent evolution in plants is cru-
cial in producing the close linkage between general vegetation structure and cli-
mate that is apparent in the world 's biomes.
One interesting aspect of adaptive radiation and convergent evolution is
the very high degree of similarity that develops in the descendant species of com-
mon ancestors that evolve in geographically separate areas. The term parallel
evolution is used to describe these instances where geographically isolated popu-
lations derived from the same ancestor evolve into morphologically and physio-
logically similar descendant species. For example, the islands of Cuba,
Hispaniola, Jamaica, and Puerto Rico each support a number of distinctive
species of anole lizards. Although the species that are present differ from island
286 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

A BLE 9.1 Parallel E

.;.ab itat Anole

--ae crown Large bo<

Marsupials

_ :Joer trunk Large toe

- ·.i gs Short boc

,. t runk Long fore

_: .ver trunk Stocky be

_ und/bush Slender t

FIGURE 9 .8 An example of convergent evolution: mammals and ecologically similar 3::urce:Losos and de Queiroz, 11
marsupials (after Baker and Garland, 1982).

interactions.
to island, all of these species appear to have evolved from the same commo;:: between plar
ancestor. On each island, species have evolved that have remarkably similar mor- only when tb
phological adaptations and utilize similar habitats. For example, each island has 2 physical trait
species with a large body and large toe pads that lives in tree crowns. Three of th= they feed fn
islands also have a species with very slender bodies and long tails that lives on the beaks and th
ground. In fact, each of the islands possesses its own set of unique anole liza rd The shapes o
species that are adapted for life either in tree canopies, upper trunks, twig< land finches <
midtrunks, lower trunks, or the ground (Table 9.1).1t appears that on each islan tion between
competition caused the anoles to evolve into specialists that take advantage o:
the different habitats available in the forests of the islands. The close similarity
between the canopy-dwelling species or lower trunk dwelling species on the dif- EXTIN CTION
ferent islands reflects the fact that all of these geographically separate specie::
evolved from a common colonizing ancestral species and share a common set o: Extinction is
ancestral genes. Similar evidence of parallel evolution is seen in a number o: genus, family
other organisms, including stickleback fish from isolated lakes in British Colum- nomic order
bia and fruit flies found on different Hawaiian Islands. regions. The
A final pattern of evolutionary development that is tightly tied by geog- local extincti
raphy occurs when two unrelated species evolve traits that are tied to their bison) could
 EXTINCTION 287

TABLE 9.1 Parallel Evolution in Anole (Anolis} Lizard Species in the Caribbean

Anole Morphology Island Species

---
ee crown Large body, large toe pads Cuba A. equestris
Hispaniola A. ricodii
Jamaica A. garmani
Puerto Rico A. cuvieri
pper trunk Large toe pads, change color Cuba A. porcatus
Hispaniola A. chlorcynus
Jamaica A. grahami
Puerto Rico A. evermanni
--.vi gs Short body, slender legs Cuba A. angusticeps

~~tive"cat"
Hispaniola A. insolitus
Jamaica A. valenciennii
~(! ";-.ruasyurus) Puerto Rico A. occultus
.'i dtrunk Long forelimbs, flat body Cuba A. loysiana
Hispaniola A. distichus
Jamaica None
Puerto Rico None
w er trunk Stocky body, long hind limbs Cuba A. sagrei
Hispaniola A. cybotes
A. lineatopusi

~
Jamaica
'
. Puerto Rico A. gundlachi
. . ::·o und/bush Slender body, long tail Cuba A. alutaceus
Hispaniola A. olssani
arsupial "mole" Jamaica None
(Notoryctes)
Puerto Rico A. pulche/lus

Is and ecologically similar 5::urce:Losos and de Oueiroz, 1998.

interactions. This pattern is called coevolution. Mutualistic associations

j from the same commC'::.
7 e remarkably similar mo:-
example, each island has :
n tree crowns. Three of L:.,::
I long tails that lives on r.::.::
set of unique anole liza::
>pies, upper trunks, t\\i_
ppears that on each islar::.
sts that take advantage -
:lands. The close similar. .
welling species on the d::-
aphically separate sp e~e­
md share a common set -
m is seen in a number c
ed lakes in British Colw---
tat is tightly tied by g e o~­
:tits that are tied to the::
between plant and animal species are a result of coevolution and can develop
only when the species are in close proximity. One example of coevolution in
physical traits is provided by nectar-eating honeycreepers and the flowers that
they feed from, and thus pollinate. The length and curvature of the bird's
beaks and the floral tubes that they preferentially feed from are very similar.
The shapes of the beaks and the flower tubes are different from those of main-
land finches and close relatives of the Hawaiian flowers; this suggests coevolu-
tion between the two species on the islands.
EXTINCTION
Extinction is the loss of all individuals in the population of a given species,
genus, family, or order. Local extinction occurs when a species or higher taxo-
nomic order disappears in one or more geographic areas but persists in other
regions. The history of the buffalo in North America provides an example of
local extinction. In the late eighteenth century, the American Buffalo (Bison
bison) could be found across a continuous range that extended south from the
288 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

northern plains of Canada to northern Mexico and east from the Rocky Moun-
tains to New York (Fig. 9.9) . By 1888, hunting pressure from European settle~
coupled with land-use changes had led to the local extinction of buffalo from a....
but a few tiny isolated areas of the interior plains. By 1888, the total number o::'
buffalo was likely less than 1000 animals. Captive breeding and protection of th::
little remaining buffalo habitat have led to a resurgence in their numbers OY ~
the past hundred years.
Global extinction refers to the loss of a species, or higher taxonomic ordc
over its entire range. Once global extinction occurs, the unique genome that was
that species is lost forever. Global extinction for a species or genus is similar tc
death for an individual organism; it is final. The mammoth (Mammuthus spp.
provides an example of global extinction. Mammoths were relatives of the ele-
phant and were extremely abundant during the Pleistocene. Prior to 12,000 ye ~
ago, species of mammoths were found across a huge range, extending from wes:-
ern Europe to northern Asia and across the unglaciated portions of North Amer-
ica. By about 10,000 years ago, all species of mammoth became extin :
throughout almost all of their range on all three continents. The last mammoths
probably lived on Wrangle Island in northeastern Siberia until about 4000 ye a~ -~
Original
ago. The global extinction of the mammoth has long been a topic of interest. It is range
likely that a combination of environmental change and hunting by huma -
caused their extinction. Today there are ongoing attempts to re-create this los:
genus by recovering DNA from mammoth tissue that is frozen in the permafro_:
of Siberia. In theory, cloning procedures could be used to produce a new mam-
moth from well-preserved DNA. However, the technical hurdles to such effom
are considerable. For the present, globally extinct species such as the mammoths
are lost forever.
The extinction of one species can have a profound impact on the entir ~
ecosystem in which it lives. For example, the loss of an important prey species ca.;:;
cause further extinctions because of the loss of food for higher predators. Su e~
events are called trophic cascades. The loss of the mammoths may have cause ·
such a trophic cascade of global and local extinction in North America abou:
10,000 years ago. The sabertooth cat (Smilodon spp.) was a very large and rela-
tively common carnivore that existed in North America during the Pleistocen
and likely fed upon mammoths and other large mammals. The sabertooth ca:
went extinct at about the same period as the mammoths. The extinction of th 1880
·~
sabertooth cats was global, and we only know of these predators from their fos i.
remains. Trophic cascades can also impact on scavenger species. The Califorrna
condor (Gymnogyps californianus) exists today only in highly protected moun-
tain areas in southern California. It is the largest bird in North America and cah
have a wing span of 3 m. Fossil evidence shows that at the end of the Pleistocen
the California condor had a much more extensive geographic range, living as far FIGURE 9.9
east as New York State. It is thought that this large scavenger subsisted mainly on throughout the
the carcasses of large mammals such as the mammoth. The extinction of th
mammoth and other large North American mammals likely led to the local
extinction of the California condor over most of its range. The small populatior. If death
in southern California may have survived in part by feeding on the carcasses of species? Frorr
whales and other sea mammals that wash up on Pacific Coast beaches. Since species have <
European settlement and land-use changes, condor populations have undergone However, it is
a dramatic decline. The species survives today only because of intensive captive actually repre
breeding programs and protection of its remaining habitat. many cases, S
 EXTINCTION 289

ast from the Rocky Mour:;-

ue from European settle._
:tinction of buffalo from ~
y 1888, the total number o:
eding and protection of th.o
~nee in their numbers O\·e::-

or higher taxonomic orde

1e unique genome that WG5
1ecies or genus is similar :.::
mmoth (Mammuthus spp
s were relatives of the elc-
ocene. Prior to 12,000 yea.-;;
ange, extending from wes:-
~d portions of North A me:--
nammoth became extin.:-·
inents. The last mammot'- ·
'eria until about 4000 yea:_
1een a topic of interest. It --
~ and hunting by hum a::::
~mpts to re-create this l o~
is frozen in the permafro>
~d to produce a new m a~::­
ical hurdles to such effo. ~
:;ies such as the mammotl:.::

Jund impact on the ent iJ=

important prey species c~
for higher predators. Sue.: •• . 10
1mmoths may have cause.: 200 .
1 in North America abo_ •26

was a very large and relc.- . . 20

"ica during the Pleistocec::

• • 25

nmals. The sabertooth c"'

oths. The extinction of tl:.::
1880 1888
predators from their fo s~
ger species. The Californ;._:_
in highly protected mou=.-
in North America and c~
the end of the Pleistocec::
graphic range, living as i2: FIGURE 9.9 The widespread local extinction of the North American buffalo (Bison bison)
venger subsisted mainly o- throughout the interior of North America (after Grant, 1991 ).
oth. The extinction of t:c-:
als likely led to the Joe=-.
mge. The small populatic= If death is the fate of all individuals, is global extinction the fate of all
eeding on the carcasses .::_ species? From the fossil record , we can estimate that over geologic time far more
cific Coast beaches. Sine:: species have evolved and gone extinct in the past than exist on the earth today.
>pulations have undergo=.:: However, it is not always the case that species that go extinct in the fossil record
~cause of intensive capti' -: actually represent evolutionary lineages and genomes that have truly died out. In
)itat. many cases, species disappear from the fossil record because they evolved into
290 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

crab and the cc

biloba) is an e;
genus develop'
The fossi
First, for plan1
time. For exm
extinct at an a·
term base-lev(
background eJ
extinction rate
rates include t
lion years ago)
million years
episodes of ve
extinctions. Pe1
mian, and the
FIGURE 9.10 A true living fossil, the horseshoe crab (Limulus). These relatives of spid e ~ and marine ar
have existed in forms similar to their modern one for over 400 million years. the existing ar
mass extinctio
land and marir
new species. This is called phyletic extinction, and many of the genes of the origi- tion that signiJ
nal species persist in the descendant species. For example, 55 million years age Cretaceous le•
there was a small herbivore mammal called Hyracotherium that browsed o included man~
shrubs and small trees. The members of this genus stood only 25 to 50 em tat_ enced by mari
From the fossil record we can see that Hyracotherium went extinct at least 40 mil- during that tin
lion years ago. However, further analysis of the fossil record tells us that Hyra- It is belie
cotherium is actually the ancestor of modern horses and their relatives in the events such as
genus Equus. Although apparently extinct, Hyracotherium evolved through vari- of the earth. T
ous stages to become the modern genus Equus. In this case, many of the gen ~ appear to hav(
possessed by Hyracotherium have been preserved and passed down. When 2 Yucatan Penin
species disappears without any descendant species, this event is called a true tion remains '
extinction. Unlike the modern horses, there are no descendant species fro rr: strophic coolin
mammoths or sabertooth cats. These lineages are the victims of true extinctions. Others believe
When we further examine the fossil record for extinctions (both true anc mate. Instead,
phyletic), we see that for commonly found organisms, such as marine invertebrate dioxide balanc
the average rate of extinction over the past 500 million years has been two com- ducing widesp
plete families or organisms each million years. The vast majority of mammal genera dence that mas
have only lasted 10 million years or less. The majority of primate mammal genera. The extinction
the group that includes humans, only persist for 5 million years or less. In general. was likely caus
individual species persist for fairly short periods, generally well less than 1 to 2 mil- nary glacial cy
lion years for complex animals. There are some very interesting plants and animals the extinction I
that appear to have unusually long persistence, one of the most notable being gests that then
horseshoe crabs (Limulus spp.). These relatives of the spider have escaped extinc- When co
tion and have persisted with very little evolutionary change for the past 400 to 500 what attribute
million years (Fig. 9.10). Another interesting example of a long-lived species is the to persist for
coelacanth fish (Latimeria chalumnae), which appears to have persisted relatively extinctions? Ir
unchanged for about 70 million years. Indeed, the coelacanths were thought to adapt to long-
have gone extinct about 70 million years ago in the Late Cretaceous, until a fishing drift and dim:
vessel landed a living one in 1938. A number of other coelacanths have been caught events. For ex<
since that time. Ancient taxa that persist to the present day, such as the horseshoe the dispersal r

us). These relatives of sp ide:
million years.
r of the genes of the o ri ~­
aple, 55 million years ag -
therium that browsed c::.
>od only 25 to 50 em tL
rent extinct at least 40 m:.-
record tells us that Hyr~­
and their relatives in t'::::
;um evolved through va.-:-
s case, many of the gee::_
1d passed down. When _
1is event is called a tn..:::
descendant species fro=
ctims of true extinctior:s.
:xtinctions (both true a;:::
;has marine invertebrate.
years has been two co-
tajority of mammal gene__
·primate mammal gen e~
n years or less. In generc_
ly well less than 1 to 2 m=::-
resting plants and animi:.
f the most notable bee~
>ider have escaped ex tiL.:-
1ge for the past 400 to .5 ~•
a long-lived species is C:::::
J have persisted relatiYe_
lacanths were thought : -
Cretaceous, until a fislffi:;
acanths have been cau ~
ay, such as the horsesb0::
EXTINCTION 291
crab and the coelacanth fish, are often called living fossils. The ginkgo tree (Ginkgo
biloba) is an example of a living fossil plant. Fossil leaves of ginkgo show that this
genus developed over 200 million years ago in the Permian.
The fossil record of extinctions over time shows two interesting features.
First, for plants or animals there is continuous extinction throughout geologic
time. For example, over the past 400 million years, plant families have gone
extinct at an average base-level rate of about 1 every 4 million years. These long-
term base-level rates of extinction are sometimes called normal extinctions or
background extinction rates. In addition, there are clearly time periods where
extinction rates become very high. For plants, these periods of high extinction
rates include the Late Devonian (360 million years ago), the Permian (250 mil-
lion years ago), the Late Triassic (200 million years ago), the Late Cretaceous (65
million years ago), and the Late Tertiary (2 million years ago). Such discrete
episodes of very high extinction rates are called catastrophic extinctions or mass
extinctions. Periods of catastrophic extinction in the Late Devonian, the Late Per-
mian, and the Late Cretaceous are clearly apparent in the fossil records of land
and marine animals and plants. During the Late Permian, for example, 50% of
the existing animal families became extinct. Others think that the late Permian
mass extinction may have been even more severe, with perhaps 90 to 96% of all
land and marine species going extinct. Interestingly, this was the only mass extinc-
tion that significantly impacted on insects. The mass extinction at the end of the
Cretaceous led to the annihilation of the dinosaurs. Late Tertiary extinctions
included many marine organisms. Extinction rates of 50 to 75% were experi-
enced by marine bivalves in portions of the North Atlantic and Mediterranean
during that time.
It is believed that most periods of mass extinctions coincide with catastrophic
events such as rapid climatic change or the impact of large meteors on the surface
of the earth. The late Cretaceous extinctions, including the loss of the dinosaurs,
appear to have been triggered by the impact of an asteroid in the vicinity of the
Yucatan Peninsula (Technical Box 9.1). The cause of the Late Permian mass extinc-
tion remains debated. Some scientists believe it may have resulted from cata-
strophic cooling due to climatic change related to the positions of the continents.
Others believe that the extinction event occurred too rapidly to be explained by cli-
mate. Instead, some have postulated that rapid changes in the oxygen and carbon
dioxide balance of the oceans may have poisoned the waters and atmosphere, pro-
ducing widespread marine and terrestrial extinctions. Finally, there is some evi-
dence that massive volcanic eruptions may have led to a cooling in the atmosphere.
The extinction of plant families and marine organisms at the close of the Tertiary
was likely caused by the onset of climatic cooling that ultimately led to the Quater-
nary glacial cycles. How frequent are such mass extinction events? An analysis of
the extinction histories of 9773 marine animals over the past 300 million years sug-
gests that there is a mass extinction every 26 million years.
When considering normal extinctions, biogeographers have long pondered
what attributes cause some species to be prone to extinction while others are able
to persist for many millions of years. We might first ask, what causes normal
extinctions? In some cases, species may simply not have the genetic diversity to
adapt to long-term changes in their physical environment caused by continental
drift and climatic change. Some other normal extinctions may relate to chance
events. For example, species may evolve on an isolated island and may not have
the dispersal mechanisms needed to migrate away if the island is destroyed or
292 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

TECHNICAL BOX 9.1 that a globa

impact of tb
humans fac•
Extinction of the Dinosaurs
would also:
Finding the cause of the late Cretaceous extinctions of the dionsaurs is one of the great phe is more
detective stories in the history of geology, paleontology, and biogeography. Much of the the impact<
credit for the evidence and arguments supporting this theory can go to the father and
son team of Luis and Walter Alvarez. The geologic boundary between the Cretaceous
Period (the time during which the last dinosaurs lived) and the Tertiary Period is called
experiences c
the KT Boundary and is dated at about 65 million years ago. In many geologic sections,
may evolve a
this boundary is relatively sharp and suggests a very rapid transition between the time
of the dinosaurs and the time of the mammals. Although the span of time encompassed
important ro
by the KT Boundary is difficult to date precisely the apparent sharpness of the bound- extinction h~
ary in many sections led some paleontologists to suggest that some catastrophic event Hypothesis d
had led to the extinction of the dinosaurs and many other organisms at the end of the comes from~
Cretaceous. Luis Alvarez thought that by studying the concentration of a rare element the book the
called iridium he might be able to affix a firm age to the time that elapsed over the KT same place." '
Boundary and determine whether the extinction of the dinosaurs really was a rapid ing and becOJ
catastrophic event. Iridium found at the surface of the earth largely comes from small keep pace wi
extraterrestrial particles and meteors and is deposited at a relatively constant rate. interspecific '
Alvarez reasoned that if the KT Boundary occurred very quickly, it would have low
always prate•
concentrations of iridium. If the boundary encompassed a large amount of time, it
highly adapt<
should have higher amounts of iridium. Much to Alvarez's surprise, the samples of the
KT Boundary that he studied contained almost 100 times more iridium than he had remote lake.
estimated. He concluded that the only explanation for such high levels of iridium was adapt to new
the impact of a meteor or large comet on the surface of the earth during the time of the as an evoluti(
KT Boundary. Evidence of high concentrations of iridium has subsequently been can fix speci•
found in many other sites that possess the KT Boundary. In addition, geological and Such evoluti<
geophysical evidence shows that a very large asteroid impacted the earth in the area of likely for higl
the modern Yucatan Peninsula of Mexico at the time of the KT Boundary. This huge tlenecks. The
geologic feature is called the Chicxulub Crater. the species h.
How would a large comet or meteor in Mexico cause global extinctions of terres- Anum
trial and marine life? The initial crater was likely some 40 km deep and several hun-
tions have b<
dred kilometers across. The heat and shock waves would have killed everything within
Complexity,
several thousand kilometers of the impact. Huge tsunami waves would have inun-
dated coastal areas and caused more deaths of plants and animals. Geological and
with the life
paleontological evidence of such destruction by tsunamis has been found at sites and planktOJ
along the Gulf of Mexico. Dust from the impact and fine charcoal from resulting fires contrast, rna
would have darkened the atmosphere and caused global temperatures to plummet for thought that
several months. This would have been lethal to many plants and animals. Precipitation phological ac
would have been extremely acidic due to increased levels of sulfur from the impact ment in whic
zone and the formation of nitric acid caused by the heating of the atmosphere. The extinction th
acidic precipitation would have killed both terrestrial and aquatic organisms. Once require grea
the fine dust had settled, increased amounts of carbon dioxide and water vapor in the environment
air caused by the impact would have created extreme greenhouse warming that would take advant~
kill other organisms. Perhaps what is surprising about the KT Boundary is not that the
take advanta
dinosaurs went extinct, but rather how any organisms were able to survive.
Could such a catastrophe occur again and cause our extinction? It is likely that
to survive. P1
other large comets or meteors will eventually hit the earth. In 1994, a relatively large resources du
comet impacted Jupiter and was observed by astronomers on earth. Perhaps such an sumers. Thm
event will not occur to the earth for many millions of years. Maybe we will develop Specialist sp•
defenses against such an incident. What is more immediately troubling is the fact generalist sp
environment

tionsaurs is one of the great
biogeography. Much of the
•ry can go to the father and
ry between the Cretaceous
the Tertiary Period is called
·. In many geologic sections.
ransition between the time
~ span of time encompassed
:nt sharpness of the bound-
at some catastrophic event
Jrganisms at the end of the
entration of a rare element
e that elapsed over the KT
nosaurs really was a rapid
b. largely comes from small
a relatively constant rate.
quickly, it would have low
a large amount of time, it
;urprise, the samples of the
more iridium than he had
high levels of iridium was
:arth during the time of the
m has subsequently been
n addition, geological and
ted the earth in the area of
e KT Boundary. This huge
obal extinctions of terres-
km deep and several hun-
ve killed everything within
waves would have inun-
animals. Geological and
; has been found at sites
.arcoal from resulting fires
1peratures to plummet for
and animals. Precipitation
of sulfur from the impact
tg of the atmosphere. The
aquatic organisms. Once
de and water vapor in the
1ouse warming that would
Boundary is not that the
able to survive.
tinction? It is likely that
In 1994, a relatively large
Jn earth. Perhaps such an
. Maybe we will develop
tely troubling is the fact
EXTINCTION 293
that a global nuclear war would cause many of the same atmospheric events as the
impact of the Chicxulub asteroid 65 million years ago. Not only would any surviving
humans face a prolonged period of intense cooling, called a nuclear winter, but they
would also face dangerous levels of radiation. Perhaps this made-at-home catastro-
phe is more likely in the immediate future and hopefully is more preventable than
the impact of comet or meteor.
experiences climatic changes. In other cases, a virulent pathogen, such as a virus,
may evolve and destroy the species. It is also likely that competition may play an
important role in generating extinctions. The potential role of competition in
extinction has been elegantly summarized in what is called the Red Queen
Hypothesis developed by Leigh Van Valen in 1973. The name of the hypothesis
comes from a line in Lewis Carroll's classic book Through the Looking Glass. In
the book the Red Queen says, "it takes all the running you can do to stay in the
same place." The point is that all of a species' competitors are continually evolv-
ing and becoming more competitive. If a species cannot evolve quickly enough to
keep pace with the evolution of competing species, it will go extinct because of
interspecific competition. However, evolutionary change and adaptation do not
always protect a species against extinction. In some cases, species may become
highly adapted for life in a very isolated geographic area, such as an island or a
remote lake. If the environment changes, such species may have lost the ability to
adapt to new conditions. Such overspecialization in an isolated site is referred to
as an evolutionary trap. In addition, the loss of genetic diversity during evolution
can fix species into modes of evolutionary development which become lethal.
Such evolutionary paths are called blind alleys. This phenomenon is particularly
likely for highly specialized species or those that develop after strong genetic bot-
tlenecks. The loss of genetic diversity over time will be most problematic when
the species has to contend with pressures brought on by environmental change.
A number of specific factors related to individual organisms and popula-
tions have been identified as important controls on the likelihood of extinction.
Complexity, in terms of behavior or form, appears to be negatively correlated
with the life span of species. Simple species, such as marine bivalves, mollusks,
and plankton, have average durations of 10 million years before extinction. In
contrast, mammal species rarely persist for more than 3 million years. It is
thought that species which employ complex behavioral, physiological, or mor-
phological adaptations in order to survive are less likely to persist if the environ-
ment in which they have evolved changes. Large animals appear more prone to
extinction than small ones. The reason for this difference is that large animals
require greater resources to support them and exist in a more coarse-grained
environment. Should the environment change, a smaller animal may be able to
take advantage of favorable microsites and survive. The larger animals cannot
take advantage of such microsites or cannot obtain enough resources from them
to survive. Predators at the top of the food chain are more sensitive to decreasing
resources due to chance events and to trophic cascades than are primary con-
sumers. Thus, animals at the higher trophic levels are more prone to extinction.
Specialist species with narrow niches also appear more prone to extinction than
generalist species. Species with narrow niche breadth are less likely to survive
environmental changes.
294 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

graphic distrib
Birth/Death= 100
10,000 Birth/Death = 10 with poor disr
Birth/Death = 2 quickly enougl
always ensure
1,000 released well-c
bivalves that d
c:
0
t5c: 100
~
Q)
HE RELATIONSH
.9
~ 10 Birth/Death = 1
:VOLUTION AND
<IS
~
Although the <
very negative c
evolution of li:
ancestral speci
force in the de
10 100 1,000
Population size
adapted to the
removal of om
FIGURE 9.11 Mathematical modeling results suggest how the duration of a species pric- vacant niche sr
to extinction is influenced by both population size and the birth rate/death ratio (after lutionary adap
MacArthur and Wilson, 1967). aspects of extir
In recen
Population size appears very important for species persistence. Species tha: extinction and
have very small population sizes are prone to extinction due to chance even - lies that typica
such as severe drought, whereas if a species has a large population size there i c ciation rates. 1
greater probability that some individuals will survive. Studies of the local extin - broad scale the
tion of tropical birds in Amazon forest patches over several decades indicate tha: related with tl
populations with 50 individuals or more are about five times less likely to go marine bivalve
extinct than those with 5 or less. Species that have rapid generation times an - these bivalves
high birth/death ratios can produce rapid population regrowth after chan c ~ persist for less
events and are more likely to evade extinction. Mathematical models have bee found at low p
used to explore the sensitivity of extinction rates to population size and bin to speciate allc
rates/death rates. Such models show that species with very high birth rates rela- lations are alsc
tive to death rates can persist at relatively low population sizes for long periods Catastror
of time. Species with low birth rates relative to death rates are prone to rapid When a catastr
extinction even when population sizes are high (Fig. 9.11) . environment in
Geography also appears very important in determining the probabl typically experi
longevity of a species. In general, species that have large geographic distribu- occupy the nov
tions are less likely to go extinct than species with small ranges. This is becaus by animals ove
a chance event, such as a particularly severe winter or drought, is not likely to extinction and
severely impact a large region as it is a small region. Thus, even though such extinction even
chance events may cause local extinctions, they will not cause global extinc- ceous in which
tions. Studies of the local extinction of tropical birds on forest patches in th e lowed by simila
Amazon show that species living on 250-ha patches have local extinction rates As we clc
that are almost half the rate for species living on 24-ha patches. Large geo- of mammals t(
graphic distributions also appear to be negatively correlated with extinctions demise of the c
during catastrophic events. For example, widely distributed marine inverte- extinction caus
brates showed lower rates of extinction during the late Cretaceous extinction species diversi
than invertebrates with small geographic distributions. Finally, species with astrophic extin
high dispersal capability may be more likely to escape extinction than those tive radiation
with restricted abilities to disperse. Good dispersers can change their geo- not have evolv

THE RELATIONSHIP BETWEEN
EVOLUTION AND EXTINCTION
:he duration of a species pri
1 rate/death ratio (after
:s persistence. Species the.:
ion due to chance even ~
population size there is :.
ltudies of the local extinc-
'eral decades indicate thc.-
ve times less likely to g-
pid generation times ar::.:
n regrowth after chanc:e
natical models have bee:.
population size and bin ·-
very high birth rates relc.-
ion sizes for long perio-'-
rates are prone to ra-·-
1).
~termining the proba u"::
1rge geographic distribt: -
lll ranges. This is becaus:e
r drought, is not likely tc
. Thus, even though sue:.
not cause global extin.:--
on forest patches in tt:e
we local extinction r a t ~
~-ha patches. Large geo-
rrelated with extincti o~
ributed marine inven e-
te Cretaceous extinctio:.
•ns. Finally, species wiL::.
)e extinction than th os:e
; can change their geo-
THE RELATIONSHIP BETWEEN EVOLUTION AND EXTINCTION 295
graphic distributions quickly during periods of environmental change. Species
with poor dispersal capabilities may not be able to change their distributions
quickly enough to escape extinction. Interestingly, dispersal capability does not
always ensure survival. It has been found that marine bivalve species that
released well-dispersed planktonic larvae had just as high an extinction rate as
bivalves that did not have planktonic larvae.
Although the extinction of a species is often compared to death, and thus has a
very negative connotation, extinction does play a very positive role in the overall
evolution of life. Phyletic extinction, after all, represents the replacement of an
ancestral species by its own descendant. If natural selection has been a strong
force in the development of the new species, it can be assumed that it is better
adapted to the environment in which it lives. When true extinction occurs, the
removal of one species provides habitat and resources for competing species. The
vacant niche space that is created by the extinction of one species can lead to evo-
lutionary adaptation by other species to fill that niche. Darwin saw these positive
aspects of extinction when he formulated his ideas on evolution.
In recent years, evolutionists have looked at the relationship between
extinction and evolution in more detail. It has been found that genera and fami-
lies that typically have high rates of species extinction also tend to have high spe-
ciation rates. In his classic investigations, Steven Stanley documented that on a
broad scale the speciation rates for animals and higher plants are negatively cor-
related with the average length of time that species persist. Thus, species of
marine bivalves often persist for 10 million years or more. The speciation rate for
these bivalves is about one-quarter the rate for mammal species, which typically
persist for less than 2 million years. It has been suggested that species that are
found at low population sizes, with fragmented distributions, are the most likely
to speciate allopatrically via founder effects. However, such small, isolated popu-
lations are also the most likely to go extinct.
Catastrophic extinctions play a very crucial role in generating speciation.
When a catastrophic extinction event occurs, the loss of so many species creates an
environment in which many potential niches are unfilled. The surviving species then
typically experience high rates of adaptive radiation as they evolve and speciate to
occupy the now vacant niches. A comparison of extinction rates to speciation rates
by animals over the past 500 million years shows this relationship between mass
extinction and rapid adaptive radiation and speciation (Fig. 9.12). Catastrophic
extinction events such as those at the end of the Permian or the end of the Creta-
ceous in which as many as 50% of the living animal families become extinct are fol-
lowed by similar or greater magnitude pulses in the evolution of new families.
As we close this chapter, we might reflect on the fact that the great diversity
of mammals to which we belong occurred as adaptive radiation following the
demise of the dinosaurs in the late Cretaceous. Even though the Late Cretaceous
extinction caused many mammal species to go extinct, within 10 million years the
species diversity of mammals had increased to its modern level. Without the cat-
astrophic extinction event that destroyed the dinosaurs and the subsequent adap-
tive radiation of mammals which it triggered, it is probable that humans would
not have evolved, and you and I would not exist today.
296 CHAPTER 9 EVOLUTION, SPECIATION, AND EXTINCTION

?:Jlaeogeography, Palaeocl
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Terborgh, J., and Winter, B. (1980). Some causes savanna regions
of extinction. In Conservation Biology. An a single native s
Evolutionary-Ecological Perspective (ed. M . ::.. faunas of some ;
Soule and B. A. Wilcox) , pp.119-133. Sinau e~
Why groul
Associates, Sunderland, MA.
van Noort, S., and Compton, S. G. (1996). Conve -
lar regions and <
gent evolution of Agaonine and Sycoecine raphy. The expla
(Agaonidae, Chalcidoidea) head shape in continents and i
response to the constraints of host fig mor- occupy them. Ir
phology. Journal of Biogeography 23, 415-4_.! earth into biog(
Van Valen, L. (1973). A new evolutionary law. plant and anim
Evolutionary Theory 1, 1-33. together in one
Wallace, A. R. (1858). On the tendency of vari- earth history, ev1
eties to depart indefinitely from the original modern biogeo1
type. Journal of the Proceedings of the Lin-
graphic regions
nean Society (Zoology) 3, 45-62.
Williamson, P. G. (1981). Paleontological docu-
mentation of speciation in Cenozoic molluscs
from Turkana Basin. Nature 293,437-443. :J EFINING BIOGEO
EGIONS, AND Pfi
When wemapc
cover some ver~
 tural populations of guppies
'ata). Science 275, 1934-1936.
J2) . The Meaning of Evolutior
hicago Press, Chicago.
CHAPTER 10
The Darwin Reader. W. W.
Jrk. REALMS, REGIONS,
leehler, B. M. (1990). Patterns
Indian birds. Journal of Bio-
,39-648.
AND PROVINCES:
limon, W. D. (1990). Causes o_:
!leontological Perspective, p.
THE BIOGEOGRAPHIC
of Chicago Press, Chicago.
Allopatric speciation: ded uce.: SUBDIVISIONS OF
j_ Journal of Biogeography 17.

44). Tempo and Mode in Eva-

a University Press, New York
53). Major Features in Evolu-
University Press, New York.
55). The Geography of E volu-
lOk Co., Philadelphia.
~X tinction. Scientific Am eri ~ e ­
~k.
').Macroevolution: Pattern m::.
'r eeman, San Francisco.
1). The general correlation
speciation and rate of extin· -
~ausallinkages. In Causes o.
leontological Perspective (ed.
W. D. Allmon) , pp.103-127.
1icago Press, Chicago.
inter, B. (1980) . Some causes
Conservation Biology. An
~ological Perspective (ed. M.S.
Wilcox), pp.119-133. Sina u e~
lerland, MA.
:ompton, S. G. (1996). Con \·e-
f Agaonine and Sycoecine
1lcidoidea) head shape in
~onstraints of host fig mar-
of Biogeography 23, 415--4 _-
).A new evolutionary law.
eory 1, 1-33.
-l). On the tendency of vari-
!definitely from the original
•he Proceedings of the Lin-
lOlogy) 3, 45-62.
981) . Paleontological docu-
:ciation in Cenozoic moll us__
1sin. Nature 293,437--443.
THE EARTH
Whether you climb to the alpine treeline in the mountains of northern Europe,
northern Asia, or North America, you will encounter trees that belong to the same
genera: pine (Pinus) , spruce (Picea), fir (Abies), or larch (Larix). The coniferous
trees found in the Alps are closely related to those found in the Rocky Mountains
of Canada or the Ural Mountains of Russia. In contrast, if you were to venture to
the mountains in New Zealand, Australia, or the southern tip of South America,
you would find the alpine treeline occupied by species of southern beech (Nothofa-
gus). You would not find a single native species of pine, spruce, fir, or larch. Simi-
larly, not a single species of southern beech is native to North America, Europe, or
Asia. Descending from the high elevations and walking across the savanna regions
of Africa, you will encounter vast herds of gazelles but not a single kangaroo. The
savanna regions of Australia support several different species of kangaroos but not
a single native species of gazelle. Why are the species that comprise the floras and
faunas of some areas very similar and very different from other areas?
Why groups of closely related plants or animals are found together in particu-
lar regions and are absent from others has long been a central question in biogeog-
raphy. The explanations of such patterns generally lie in both the geologic history of
continents and islands, and the evolutionary history of the plants and animals that
occupy them. In this chapter we will consider how biogeographers subdivide the
earth into biogeographic regions on the basis of the geographic distributions of
plant and animals species. We will examine why certain groups of species occur
together in one region and are absent in other areas. We will see how environment,
earth history, evolution, and extinction are all key forces behind the development of
modern biogeographic regions. Finally, we will briefly review the major biogeo-
graphic regions of the planet to consider their unique fauna, flora, and histories.
DEFINING BIOGEOGRAPHIC REALMS,
REGIONS, AND PROVINCES

When we map out the global geographic distributions of mammal families, we dis-
cover some very interesting facts. Excluding bats and marine mammals, there are

299
300 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

about 100 known families of mammals. Of these families, most are restricted :.:: arctic treeline
one or two continents, and no family is native to all of the continents of the wor - level of famil:
We find broadly similar results when we study the global geographic distributior:: if we compar
of birds, freshwater fish, or plant genera. When a species, genera, or family - reason for thi
restricted to one or a few geographic regions, it is referred to as being endemi and genera of
The term endemic is very relative. Lodgepole pine (Pinus contorta) is endemic : - tively differer
western North America, but within this region it grows from Alaska to southe:::. Several
California and from the Pacific Coast to the Rocky Mountains. In contrast, soo::: cosmopolitan.
species are narrow endemics that occur in only a very small area. For examp> These species
Santa Catalina mahagony (Cercocarpus traskiae) has a global distribution consis:- family to occu
ing of only six individuals that grow in Wild Boar Gully on Santa Catalina Islac might be muc
off the coast of southern California. Widely distributed families, genera, or speciP-5 ties or chance
are said to be cosmopolitan. Owing in part to human transport, several mammc._ one species oJ
families are indeed truly cosmopolitan and are found today on all continen:.: enced by the
except for Antarctica. These include the Muridae (rats and mice), the Leporid~:: and family. Th
(hares and rabbits), and the Canidae (wolves and dogs). The degree of enderni species evolvt
in plants and animals increases with taxonomic level. Individual species are oft = lies. Therefon
far more endemic than the genera or families to which they belong. The reaso - family and ge
for the generally wider distributions of genera and families compared to speci : bution. Fourtl
will be discussed later in this chapter. In general, endemism is the common sta·:: nate individu;
for land plants and animals, while true cosmopolitans are rare. genus might e
Recognizing that many families, genera, or species of plants and animals ~:: Thus, an indi'
endemic, biogeographers have sought to divide the earth into geographic regia " might persist ·
that have similar species, genera, or families within each region. How can or.:: If we ex<
objectively determine whether the species, genera, or families present in two <K- and the coeff
ferent areas are similar or distinct? Biogeographers often use mathemati .:..: interesting pa
indices to provide quantitative estimates of the floral or faunal similariti : are not found
between regions. These indices are referred to as coefficients of similarity. Two o: the NorthAm
the most commonly applied coefficients of similarity are the Jaccard and tt:: native to Afri'
Simpson. The Jaccard coefficent has the form: ern Eurasia. 1
America and ,
c tralia shares n
N1 +N2-C the continent<:
exists betwee
where Cis the number of families, genera, or species found in both regions, N 1 :S
faunas. For ex
the number of families, genera, or species found in region 1, and N 2 is the numbe:-
of families, genera, or species found in region 2. The Simpson has a simpler fo
c TABLE 10.1
N1
For both equations, a value of 1 indicates that the flora or fauna of both regions Country
are identical. Biogeographers would say that the flora or fauna of regions are ii: Canada
harmony. A value of 0 indicates that there is no similarity in the flora and faun a o~
the regions. By multiplying the coefficients by 100, we can transform them into
percentages of similarity that range in value from 0% for regions that do no:
share any of the same families, genera, or species to 100% for areas that are com- Russia
pletely harmonized.
When comparisons of biogeographic similarity are carried out, we fin ·
that the higher the taxonomic order being examined, the higher the biogeo- Simpson coeffic
graphic similarity. For example, if we compare the coniferous trees along the

, most are restricted to
:ontinents of the worlcl
eographic distributions
~s, genera, or family is
d to as being endemic.
contorta) is endemic to
om Alaska to southe
tains. In contrast, sam
nall area. For exampL.
bal distribution consist-
n Santa Catalina Islan ·
!lilies, genera, or species
1sport, several mamma:
oday on all continents
td mice), the Leporidae
he degree of endemisr:::.
vidual species are ofte:::.
tey belong. The reasons
es compared to species
;m is the common stat::
rare.
f plants and animals are
into geographic regia ~
h region. How can one
tilies present in two dif-
•ften use mathematica:
l or faunal similarities
~nts of similarity. Two o:
re the Jaccard and the
!ld in both regions, N 1 '-
1, and Nz is the numbe ~
>son has a simpler form.
>r fauna of both regi a~
fauna of regions are i.;:
n the flora and fauna o:"
an transform them in tc
or regions that do no:
for areas that are com-
e carried out, we fi n ·
he higher the biogeo-
iferous trees along the
DEFINING BIOGEOGRAPHIC REALMS, REGIONS, AND PROVINCES 301
arctic treeline of Canada with those of northern Russia , we will find that at the
level of family or genus the biogeographic similarity coefficient is 1. However,
if we compare the two regions at the level of species, the coefficient is 0. The
reason for this difference is that both regions are occupied by the same families
and genera of coniferous trees, but those genera are represented by the distinc-
tively different species in Canada and Russia (Table 10.1).
Several reasons can be cited as to why higher taxonomic levels are more
cosmopolitan. First, families and genera often include many different species.
These species might have greatly different niches, which allows the genera or
family to occupy a wide variety of environments, although each individual species
might be much more restricted in its distribution. Second, better dispersal abili-
ties or chance events might lead to long-distance dispersal and colonization by
one species of a genera or family but not by others. The range expansion experi-
enced by the individual species creates a larger geographic range for the genus
and family. Third, because evolution works directly at the level of the species, new
species evolve and diverge much more quickly than do entire genera and fami-
lies. Therefore, new species may evolve and exist as narrow endemics, while the
family and genus to which they belong remain relatively cosmopolitan in distri-
bution. Fourth, extinction due to either normal or catastrophic events may elimi-
nate individual species of a genus or family, but other species of the family or
genus might escape due to differences in their niches or geographic distributions.
Thus, an individual species may go extinct in a region, but the family or genus
might persist through the survival of other member species.
If we examine the global geographic distributions of plant and animal species
and the coefficients of similarity between different continents, we notice some
interesting patterns. Each continent possesses families of mammals or plants that
are not found on other continents. Thirteen different families of mammals occupy
the North American continent at present. Interestingly, none of those 13 families is
native to Africa. However, five of the North American families are found in north-
ern Eurasia. As a result, the Simpson index of similarity is 0% between North
America and Africa and 38% between North America and northern Eurasia. Aus-
tralia shares no common mammal families with any other continent. If we consider
the continental distribution of flowering plants, we find that much greater similarity
exists between the floras of the different continents compared to the mammal
faunas. For example, the Simpson index of similarity between the flowering plant
TABLE 10.1 Biogeographic Similarity between Treeline
Conifers in Canada and Northern Russia
Country Family Genus Species
Canada Pinaceae Larix laricina
Pinaceae Pice a glauca
Pinaceae Pice a mariana
Pinaceae Pinus banksiana
Russia Pina cea e Larix sibirica
Pinaceae Picea obovata
Pinaceae Pinus sylvestris
Simpson coefficient 1.0 1.0 0.0
302 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

families of North America and northern Eurasia is 47%. The similarity between th:: of vertebrates,
plant families of Australia and those of South America, Africa, and southern A ;~ nonflying man
ranges from 69% to 63%. Neither the mammal families nor the flowering plar.· regions is deer
families are completely harmonized between any of the continents. Each contine · regions recogn
contains families of mammals and plants that are unique. We see a similar patter:; Sclater, althou1
when we examine birds, reptiles, or freshwater fish. nal regions stil
Our brief examination of the worldwide distributions of mammals an:: systems. The S
flowering plants raises three interesting questions. First, why do different con - into six biogeo~
nents contain not just individual families, but groups of families that are not pre5-
ent on other continents? Second, why does Australia have no mammal families ir.
common with other continents, while Eurasia and North America share so man_
families with each other and other regions? Third, why are the flowering plar.:
families more similar from continent to continent while the mammal families ar::
so different? Such questions have always intrigued biogeographers.
The tendency for different regions to possess unique species, genera, or fam:-
lies is called biogeographical provincialism. The modern scientific study of biogeo-
graphic provincialism can be traced back to the work of nineteenth-centur:
European phytogeographers such as F. Schouw and A. de Candolle, while the cu:--
rent concept of subdividing the earth into biogeographic regions is attributable
Philip Sclater, a well-known British ornithologist responsible for the scientific ciis
covery and description of over 1000 different bird species from all over the worl ·
Sclater was a friend of Charles Darwin and very active in the Royal Geographi ~
Society of London. He was very well informed regarding the biological and geo-
graphical discoveries of his time. Based on his data regarding the global geographi:
distributions of birds, plants, and animals, Sclater argued that previously propose<:
biogeographic subdivisions of the earth were based on latitude, longitude, and c
some degree climate, and did not take into account the provincialism observed i::.
the geographic distributions of flora and fauna. He observed that, even thou~'­
places such as tropical Africa and South America have similar climates, they ar:·
occupied by very different species of plants and animals. Sclater proposed that th~
earth should instead be subdivided on the basis of the geographic distributions o:
related species, genera, or families of organisms rather than on geographic positio
environmental conditions, or vegetation structure. Sclater's approach is, of course.
quite different from the biome approach for subdividing the biosphere (see Chapte:-
6). He believed that areas where a species is found today likely represents the geo-
graphic area in which the species first developed. Therefore, Sclater reasoned tha:
divisions of the earth based on species distributions would provide insights into bio-
logical history. Sclater developed his subdivision using passerine birds (perchin=
birds such as sparrows), for this was the group he knew best, and he presented his
results in 1857 before the Linnaean Society of London. Interestingly, Sclater's pres-
entation occurred one year before Darwin and Wallace read their paper on evol -
tion to the same society. The system proposed by Sclater was called the Scheme
Avium Distributionis Geographicae. This hierarchical scheme divided the earth into
two super regions: the New World (North and South America) and the Old Worl ·
The landmasses were then divided into six biogeographic regions based on the sim-
ilarity in the bird species within each region and the dissimilarity from the birc
species found in other regions.
Sclater's concept of biogeographical regions was further developed by th
great evolutionist and biogeographer, Alfred Wallace. In 1876 Wallace presentee FIGURE 10.1 1
a subdivision of the earth into biogeographical regions based on the distributiom fauna biogeograp
 DEFINING BIOGEOGRAPHIC REALMS, REGIONS, AND PROVINCES 303

. The similarity between the of vertebrates, with particular reference to nonflying mammals. By focusing on
, Africa, and southern AsiE nonflying mammals, the impact of long-distance dispersal between different
ies nor the flowering plan: regions is decreased and the boundaries between regions should be clearer. The
continents. Each continen: regions recognized by Wallace correspond relatively closely to those proposed by
1e. We see a similar patter::: Sclater, although some of the names differed. The Sclater-Wallace system of fau-
nal regions still remains widely used today and forms the basis of more recent
.butions of mammals a- - systems. The Sclater-Wallace system divides the major landmasses of the earth
st, why do different conu- into six biogeographic faunal regions (Fig. lO.l). These six faunal regions as typically
: families that are not pres-
we no mammal families i.::
th America share so rna--
h.y are the flowering pla::-
e the mammal families a:::'
geographers.
ue species, genera, or fam:-
1 scientific study of biogec-
Jrk of nineteenth-cenhh
je Candolle, while the cu.:--
ic regions is attributable : -
asible for the scientific <fu-.
es from all over the wor:C..
: in the Royal Geograplll:
ng the biological and g~
·ding the global geograph::.:
j that previously propo~::
latitude, longitude, and · -
provincialism observed :=
bserved that, even thou=--
: similar climates, they a:-::
:. Sclater proposed that rt::
~eographic distributions c-
tan on geographic positio;::.
er's approach is, of course.
the biosphere (see Chapte_
:y likely represents the gee-
fore, Sclater reasoned th=..·
d provide insights into b:c-
passerine birds (percbi- ~
best, and he presented
nterestingly, Sclater's pre:;
read their paper on evo!::-
:er was called the Scher ;-
erne divided the earth in: _
1erica) and the Old Wor~.::..
; regions based on the sir.::.-
lissimilarity from the be.:

further developed by r:::.::

n 1876 Wallace present.::.: FIGURE 10.1 The world's faunal regions, floral regions, and Pielou's combined floral and
based on the distributio::.. fauna biogeographic regions (from a number of sources) .
304 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

recognized today are the Nearctic, Palearctic, Neotropical, Ethiopian or Africar species. Within
Oriental, and Australian. of local endemi
The success of the Sclater-Wallace system can be assessed by looking at tht In the late
relationship between modern terrestrial mammal distributions and the propose C. Pielou, propc
faunal regions. Of the 90 most common mammal families, 11 have relatively co~­ regions based c
mopolitan distributions and are native to all continents except Australia an-= 10.2). The gener
Antarctica. These families are the Soricids (shrews), Sciurids (squirrels and chip- for his faunal n
munks), Cricetids (hamsters and lemmings), Cervids (deer), Ursids (bears lace also differ
Felids (cats), Mustelids (weasels and badgers), Bovids (cows), Murids (rats treatment of oc1
Canids (dogs), and Leporids (rabbits). These families are called the 11 wandering ary between bic
families because of their relatively cosmopolitan distributions. Fifty-one (57%) o:
the 90 mammal families are endemic to only one faunal region. Most families ar ~
restricted to one or two of the recognized regions, and this supports the gener~ :> ETERMINING THI
applicability of the scheme. In addition, one can examine the degree of edemici . BETWEEN REGION
within each region to see how unique each region's mammal fauna is. The degret
of edemicity ranges from 91% for the Australian region to 3% for the Palearcti As we have see:
Therefore, the Australian region is the most clearly differentiatable on the ba s~ tinctive ftowerin
of mammal fauna, while the Palearctic is hard to differentiate from the adjace~:: ous differences
Oriental, African, and N earctic regions. to determine th
A number of efforts have been undertaken to subdivide the earth into bio- tal regions. Sud
geographic regions based on plant distributions. One of the earliest attempts w~ these boundarie
by Engler, who distinguished four floral realms in the ninteenth century. A mor; his Australian a
recent scheme developed by the Soviet botanist A. Takhtajan in the twentieth cer- the west of Wal
tury subdivides the earth into six floral regions (Fig. 10.2) on the basis of flowerin~
plant distributions. The currently recognized floral regions include the Holarcti-
N eo tropical, Paleotropical, Cape, Australian, and Antarctic. It is sometimes difficul:
to define clear biogeographic regions on the basis of plant families. Only abot::
18% of the world's plant families are endemic to one floral province. In some cases.
such as the Holarctic, the floral regions encompass more than one faunal region. l:::.
contrast, the Cape floral region of Africa is a tiny area of endemic plants compare -
to the African faunal region. Despite its small size, the unusual flora of the Cape o:
Good Hope area of South Africa contains over 500 different species of plan '
many of which are narrow endemics found only in the Cape region.
In theory, biogeographic regions should follow a hierarchy with biogeo-
graphic realms that incorporate several continents and large landmasses being
the largest, followed by the biogeographic regions which subdivide the earth r:
the continental level followed by biogeographic provinces which subdivict
smaller areas. Finally, provinces are sometimes further divided into subdivisio "
It must be kept in mind that biogeographic realms, regions, and provinces art
based on the genetic relationships between plant and animals species within ea ~
region and are thus very different from the biomes, which are based on the sim:-
larity of vegetation structure and climate within each biome.
The division of regions into provinces relies on the careful identificatio::;
and mapping of endemic species within each region. For example, the Nearcri:
region, which consists of North America, can be divided into broad fto rC..:.
provinces that represent the arctic, the boreal-temperate forests and grasslands.
the western mountains and deserts, and the Mexican-Caribbean region. Califor- FIGURE 10.2 Tt
nia, because of its high degree of plant endemism, is generally recognized as ~ biogeographic line
separate subprovince, or province. Within California, a number of biogeographi: regions. The north1
subdivisions are recognizable on the basis of high numbers of endemic plac eucalyptus, are als1
 DETERMINING THE BOUNDARIES BETWEEN REGIONS 305

ical, Ethiopian or Africar. species. Within these subdivisions, smaller areas with particularly high numbers
of local endemics can also be recognized.
assessed by looking at thO" In the late twentieth century, the Canadian biogeographer and ecologist, E.
ibutions and the propose:: C. Pielou, proposed a scheme to divide the world into a number of biogeographic
lies, 11 have relatively co~ regions based on the combination of both faunal and floral distributions (Fig.
~nts except Australia ar::: 10.2). The ge~eral regionalization proposed by Pielou is similar to that of Wallace
;iurids (squirrels and chi;-- for his faunal regions. Pielou adds a separate Antarctic region. Pielou and Wal-
ls (deer), Ursids (bearsc lace also differ regarding the locations of specific regional boundaries and the
[ds (cows), Murids (rats treatment of oceanic islands. As we shall soon discuss, affixing the precise bound-
re called the 11 wanderic: ary between biogeographic regions can be difficult.
mtions. Fifty-one (57 %) c:
1l region. Most familie s a:-::
l this supports the gener.::... DETERMINING THE BOUNDARIES
ne the degree of edemic::: BETWEEN REGIONS
mmal fauna is. The degrc-:-
n to 3% for the Palearcr:: As we have seen, Australia has very different mammal fauna and somewhat dis-
fferentiatable on the b ~-:_ tinctive flowering plant flora when compared to mainland Asia. Despite the obvi-
rentiate from the adjace= ous differences in the fauna and flora of Australia and Thailand, it is not so easy
to determine the exact geographic boundary between the Australian and Orien-
bdivide the earth into bic- tal regions. Such boundaries are called biogeographic lines. The most famous of
f the earliest attempts w.::... these boundaries is Wallace's Line, which Wallace himself proposed to separate
ninteenth century. A mo::-:- his Australian and Oriental regions (Fig. 10.2). In general, plants and animals to
ttajan in the twentieth ce:::.- the west of Wallace's Line are dominated by Oriental species, while Australian
~) on the basis of flower'..;:;
[ons include the Holarc:::.:..
tic. It is sometimes diffi
,•
plant families. Only abo _
ral province. In some cas~
~than one faunal regio n .~­
f endemic plants comparc.-
nusual flora of the Cape c
jifferent species of piaL _
:ape region.
a hierarchy with biog
td large landmasses be:.::.=
ich subdivide the earth ::
rovinces which subdi \ ~.::=
divided into subdivisi o=.::..
·egions, and provinces c.:-::
nimals species within e2.:-
tich are based on the si.c::;_-
liome.
the careful identificatiL
For example, the NeaL ~­
jivided into broad fl o._
ate forests and grasslan "' -
:=aribbean region. Califc-- FIGURE 10.2 The Wallacea region of southeastern Asia and Australia and several of the
generally recognized as _ biogeographic lines that have been proposed to separate the Australasian and Oriental
number of biogeograpt.:._ regions. The northwestern range limits of the Australian groups, the marsupials and
umbers of endemic pic:: eucalyptus, are also shown (after Zacharin , 1978 and other sources).
306 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

species dominate to the east. However, the exact location of the bounda:-: it is perhaps b
between Australasia and the Oriental region is hard to fix because many grou;- will tackle this
of animals and plants have different geographic limits in the region. Con_c-
quently many different biogeographic lines have been proposed to delineate t '-::
Australian and Oriental regions (Fig. 10.2). For example, marsupials are the o,· :-- -ACTORS BEHINI
whelmingly dominant mammal in Australia and Tasmania but are not found 31 0GEOGRAPHIC
mainland Asia. The geographic range limits of marsupials extend northwestwa::
to Sulawesi but do not extend to the Philippines or the islands west of Tim - As we have sc
However, some lines would include some of these smaller nonmarsupial islan- plants and ani
within the Australian region, while other proposed lines would not. Eucalyp _ species found
trees are a dominant element of Australian vegetation, comprising 95% of c:: understanding
forest trees on that continent. Within the Australian region, eucalyptus are or:.:_ this question v
found as far northwest as New Guinea, Sulawesi, and the southern Philippine __ behind the ex
number of smaller islands east of Wallace's Line that do not contain native eu -- present locati•
lyptus are still grouped within the Australian region, while the Philippines a::: plates, and (3)
not. Some more recent lines would also exclude Sulawesi and the Philippin - look at each o
others would not. As we c
The region of Wallace's Line is a narrow archipelago of islands separated graphic regiOI
relatively deep water. The channels between the islands provide barriers to t plants and ani
dispersal of land plants and animals and should help to produce a relatively cl ~ and Atlantic c
boundary between the Oriental and Australian biogeographic regions. As the western ar
have seen, even here the exact location of the line between the biogeograp : to terrestrial c
regions is not so clear cut. It is even more difficult to differentiate biogeograp : boundaries be
lines when they cross continuous land areas. One example of this is the line se p ~­ tions of the Sc
rating the Palearctic and Ethiopian regions. At least 18 different biogeograp h:.: flora between
lines have been proposed for this region. The locations of these lines extend fro;:: highlands and
the Mediterranean Coast southward to the southern edge of the Sahara Dese:- the Oriental r•
In the case of the Palearctic and Ethiopian regions, the Sahara provides a barri ~ cal and biolo!
to dispersal by some plant and animal species but not to others. A similar dif£- bridge betwet:
culty is encountered in delineating the boundary between the Nearctic a .: the southern E
Neotropical regions. Based on the northern range limits of South Americ<:c animal comm1
mammal families and the southern range limits of North American mammal far;:- temperate, de
ilies, it would be possible to suggest a number of different biogeographic lines northern Mex
separate the regions. The lines could be drawn as far north as central Mexico a : blocks the pas
as far south as Panama. Of COUTS•
Although large barriers, such as the Atlantic and Pacific oceans, cleaL: the biogeogra1
delineate regions such as the Neotropical from the Ethiopian, determining sh ~ as the moden
boundaries and distinct biogeographic lines between regions which are clos e:~ Alaska and Sil
adjacent or share continental areas is much more difficult. Such less distill - ods, the eustat
boundary areas are referred to as biogeographic transition zones. Some author:- and Nearctic r
ties consider the area between the Malaysian Peninsula and Australia to be :::. mate and tree
transition zone between the Australian and Oriental regions. They call this tran_i- the dispersal o
tion zone Wallacea in honor of Wallace. The Nearctic and the Neotropical, th"' duced land cm
Palearctic and the Oriental, and the Ethiopian and Palearctic regions might all - the islands in 1
considered to be separated by transition zones. islands such as
It is not surprising that the exact boundaries between biogeographi.: 120m and rem
regions are not sharply definable. Different species from each region will ha,·- the ocean serv•
different environmental requirements, dispersal and colonization abilities, an · during glacial]
histories. All of these factors contribute to the blurring of the boundaries betwee;: As we sc
realms, regions, and provinces. Instead of asking why boundaries are not distinc- changed over

location of the boundac
to fix because many group:
lffiits in the region. Conse-
n proposed to delineate t t~
ple, marsupials are the ove:--
mania but are not foun d :.:.
pials extend northwestwar:.
· the islands west of Timo:
naller nonmarsupial islan-' ·
lines would not. Eucalyp n::s
ion, comprising 95% of r:::~
region, eucalyptus are oC.:_
the southern Philippines..-.
do not contain native euCE.-
1, while the Philippines a:~
ilawesi and the PhilippinQ
:lago of islands separated ·
mds provide barriers to r::. ~
to produce a relatively cl e ~
,geographic regions. As .. _
between the biogeograpl:::
differentiate biogeograpL:
mple of this is the line sep::-
18 different biogeograpl::.::
ts of these lines extend fro=
edge of the Sahara Dese:--_
te Sahara provides a barrie:
ot to others. A similar diE-
between the Nearctic ac:.
limits of South Amer i ~--
rth American mammal fac-
~rent biogeographic lines :-
1orth as central Mexico a.::.:
and Pacific oceans, clear:_
thiopian, determining sha.,.--::
1 regions which are close::
difficult. Such less dist in.:-
~sition zones. Some autho:-:-
tsula and Australia to be .:.
·egions. They call this trallS-
ic and the Neotropical, tt::
learctic regions might all to::
es between biogeograpbi:
from each region will h a\·~
l colonization abilities, ac:
~ of the boundaries betwee::
boundaries are not distinc-_
FACTORS BEHIND THE MODERN BIOGEOGRAPHICAL REGIONS 307
it is perhaps better to ask why biogeographic regions exist in the first place. We
will tackle this question in the next section.
FACTORS BEHIND THE MODERN
BIOGEOGRAPHICAL REGIONS
As we have seen, the question as to why regions of the earth possess groups of
plants and animals that differ, often dramatically, from the families, genera, and
species found in other regions has long intrigued biogeographers. Prior to an
understanding of evolution and plate tectonics, a complete scientific answer to
this question was impossible. We now know that the three most important factors
behind the existence of clearly definable faunal and floral regions are: (1) the
present locations of biogeographic barriers, (2) the history of the continental
plates, and (3) the evolutionary history of modern animal and plant families. Let's
look at each of these factors in turn.
As we discussed earlier, the boundaries between the present biogeo-
graphic regions are formed by physical barriers to the dispersal of terrestrial
plants and animals. The sea, both in the form of large bodies such as the Pacific
and Atlantic oceans, or smaller bodies of water such as the channels between
the western and eastern islands of the Malaysian Archipelago, is a major barrier
to terrestrial organisms. Deserts and mountains also form barriers that serve as
boundaries between biogeographic regions. The hot and extremely arid condi-
tions of the Sahara Desert and Arabian Desert restrict exchanges of fauna and
flora between the Palearctic and Ethiopian regions. The cold conditions of the
highlands and mountains of the Tibetan Plateau and the Himalayas help isolate
the Oriental region from the Palearctic. Some boundaries are formed by physi-
cal and biological barriers. Although the Isthmus of Panama provides a land
bridge between the Nearctic and Neotropical regions, the tropical climate of
the southern end and competition from the associated tropical vegetation and
animal communities preclude the northward or southward migration of most
temperate, desert, or tundra species. Similarly, the arid desert conditions of
northern Mexico and the adjacent United States produce a barrier that also
blocks the passage of most temperate, tropical, or tundra organisms.
Of course, the physical and biological conditions of the boundaries between
the biogeographic regions have been subject to changes. The Bering Strait serves
as the modern boundary between the Nearctic and Palearctic regions. Here,
Alaska and Siberia are divided by less than 100 km of ocean. During glacial peri-
ods, the eustatic sea level dropped by 80 to 100 m- enough that the Palearctic
and Nearctic regions were joined by a large landbridge. However, the arctic cli-
mate and treeless tundra environment of the region was far too harsh to allow
the dispersal of many species of plants and animals. Drops in sea level also pro-
duced land connections between the mainland of Asia or Australia and many of
the islands in the Malaysian Archipelago. However, the main channels between
islands such as Borneo and Sulawesi in the Malay Archipelago are deeper than
120m and remained inundated by the sea even during glacial periods. As a result,
the ocean served as a barrier between the Australasian and Oriental regions even
during glacial periods.
As we saw in Chapter 7, the geographic positions of the continents have
changed over time. Thus, some continents, such as Europe and Asia, have been
308 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

joined as a continuous landmass for at least 50 million years. Others such a:; pattern is tha
South and North America have become linked by a landbridge only in the pas: the many vac
few million years. Australia has been isolated from all other continents, includin; However, rec
Antarctica, for over 50 million years. It is not surprising, therefore, that Euro - that many of
and northern Asia form one modern biogeographical region, the Palearctic. It ~ earlier in the
also not surprising that the Australasian region is the most distinct of all the b i~ structure of tl
geographical regions. have begun 1
The biogeographic regions we are considering here are based on the mo ·_ were in the p
ern distributions of mammal and flowering plant species. The evolutionary his- America was
tory of these groups explains why the mammal fauna of the differer;: sea level rose
biogeographic regions is more distinctive than the plant flora, why marsupi ~ based reconst
dominate the native mammal fauna of Australia, and why no southern bee :::. speciation cau
trees are native to the northern hemisphere. Let's briefly consider the evolutior:.- opment of epc
ary history of the mammals and flowering plants as it is known today. modern mam
mammal fossi
number and b
Evolution of the Mammals
Cretaceous ex
It may be surprising to learn that the first primitive mammals appeared at abo - One oft
the same time as the first dinosaurs. Fossils suggest that the first mammals a ;: development <
dinosaurs appeared during the Triassic some 220 million years ago. Bo:.:::. are known as
dinosaurs and mammals evolved from reptiles. Mammals descended fro m ~ evolved first , 1
group of reptiles called the therapsids, which appeared in the Permian about r · evolved in No
million years ago. Hair, which typifies mammals, likely evolved from scales. OD.:c 100 million ye.
of the first known mammals was a very small, shrewlike creature called t h~ eastward to A
Megaostrodon. These earliest mammals likely laid eggs. The only surviving e&,- early marsupi<
laying animals are the monotremes, which include the duckbill platypt.:_;; to be membei
(Ornithorhynchus anatinus) and echidnas (Tachyglossus and Zaglossus) found i.;: American opo
Australia, Tasmania, and New Guinea. Fossilized teeth are by far the most con:- also represent'
mon remains found of the earliest mammals. These teeth indicate that some ear · must have star
mammal species fed on plants, others fed on insects, and still others were omni- Today, m
vores. The omnivore group, called the pantotheres, are likely the ancestors o: in North Ame1
modern mammals. Unlike the dinosaurs, most mammals remained small in sir remain domin:
and their populations remained small during the next 155 million years. Durin; once far more
this time, the dinosaurs grew in physical size, abundance, and range of adaptc:- marsupials toe
tions, and came to be the dominant large animals on land, sea, and air. This woulc mals. An epeir
all change with the close of the Cretaceous about 65 million years ago. kept the mars1
The close of the Cretaceous Period represents one of the greatest care- also separated
strophic extinction events that the earth has experienced (see Chapter 7). The nized these re
dinosaurs, which had been so dominant, disappeared along with many other t e ~­ shortly after tl
restrial and marine animals and plants. It seems indisputable that this event was cental mamma
caused by the impact of a large meteor or comet near the present-day Yucata:. which were pri
Peninsula of Mexico. Fires, tidal waves, acidic precipitation, changes in the ozone tavora). By 50 •
layer, and later climatic cooling caused by the extraterrestrial strike destroye-' America, Afric
the dinosaurs directly and through trophic cascades events. Some mammals, how- (includes monl
ever, survived this catastrophe, even though many species apparently did gc morpha (inclw
extinct at this time. The small size of the mammals, along with their warm- mals were not,
blooded physiology, likely contributed to their survival. separated fror
When we look at the fossil evidence for the initial evolutionary develo Australia sugg,
ment of modern-day mammalian orders, it seems that most originated followi n= between 100 aJ
the close of the Cretaceous. The classical explanation for this evolutionan Australia, they

lion years. Others such c
andbridge only in the p ~
other continents, includl;:;
ing, therefore, that Eu ro ;:~
region, the Palearctic. It :;;
most distinct of all the bic-
tere are based on the mo.::.-
~cies. The evolutionary !8-
1 fauna of the differe::.
)lant flora, why marsupic.....
td why no southern b eez~
~fly consider the evolutio::.-
is known today.
tammals appeared at aboL
hat the first mammals a::..::
million years ago. Boc.:
tmmals descended from .::
din the Permian about :2 -
y evolved from scales. 0 ::.::
ewlike creature called L:::
~gs. The only surviving eg;-
de the duckbill platy p ~
;us and Zaglossus) foun d ::.
h are by far the most coc-
~th indicate that some ea:->
and still others were orn;::.::-
are likely the ancestors -
1als remained small in siz::
:t 155 million years. Du ~
ance, and range of adap-:2-
md, sea, and air. This woU:..::
nillion years ago.
s one of the greatest c a-~­
!nced (see Chapter 7). T::.::
along with many other te::--
;putable that this event \Y2..o-
ar the present-day Yuca" -
:ation, changes in the ozo.::.::
;terrestrial strike destro\·c.::
vents. Some mammals, ho'"'-
species apparently did g.:
ls, along with their warc-
al.
titial evolutionary develo:;:-
t most originated followiL:,
ttion for this evolutiona:-
FACTORS BEHIND THE MODERN BIOGEOGRAPHICAL REGIONS 309
pattern is that strong adaptive radiation occurred as the mammals evolved to fill
the many vacant niches caused by the catastrophic extinction of the dinosaurs.
However, recent analysis of the DNA structure of modern mammals suggests
that many of the orders may have already begun evolutionary divergence much
earlier in the Cretaceous. It has been suggested that the differences in the genetic
structure of the different mammal orders indicate that mammal divergence may
have begun 100 million years ago. At that time, Laurasia and Gondwanaland
were in the process of breaking into the modern continents. In addition, North
America was split by a shallow sea (called an epeiric sea) that formed when the
sea level rose and flooded the central part of the continental plate. If the DNA-
based reconstruction of the timing of mammal divergence is correct, allopatric
speciation caused by the breakup of Laurasia and Gondwanaland and the devel-
opment of epeiric seas produced the initial divergence that created the different
modern mammal orders. Even if this is the case, the sparsity and small size of
mammal fossils from the Cretaceous indicate that mammals remained small in
number and body size until the burst of adaptive radiation that followed the Late
Cretaceous extinction of the dinosaurs.
One of the most important events in the history of mammal evolution is the
development of the modern marsupial and placental mammals. These two groups
are known as the eutharian mammals. It is generally thought that the marsupials
evolved first, followed by the placentals. The earliest marsupials appear to have
evolved in North America in the Middle to Late Cretaceous, perhaps as early as
100 million years ago. From there they spread southward to South America, then
eastward to Antarctic and Australia, and eastward to Europe (Fig. 10.3). These
early marsupials are known mainly from teeth and jaw fragments. Some appear
to be members of the family Didelphidae, which includes the modern North
American opossum (Didelphis marsupialis). Several other marsupial families are
also represented in these early fossil faunas, indicating that marsupial divergence
must have started prior to the end of the Cretaceous.
Today, marsupials are absent from Europe, represented by only one species
in North America, and they form a relatively small fauna in South America. They
remain dominant in the mammal fauna only in Australia. What happened to this
once far more widespread group of mammals? The restricted distribution of the
marsupials today is tied to continental drift and the rise of the placental mam-
mals. An epeiric sea separated Asia from Europe during the late Cretaceous and
kept the marsupials from colonizing Asia. India, Africa, and New Zealand were
also separated from the other continents by oceans, and marsupials never colo-
nized these regions. The placentals appear to have evolved in northern Asia
shortly after the marsupials. Between 100 million and 65 million years ago, pla-
cental mammals were represented by now extinct groups such as the Creodonta,
which were primitive carnivores, and surviving groups, such as the shrews (Insec-
tavora). By 50 million years ago, placental mammals had spread to Europe, North
America, Africa, India, and South America. Mammal orders such as the Primates
(includes monkeys and apes) , Carnivores (includes dog and cats), and the Lago-
morpha (includes rabbits and hares) were present by this time. Placental mam-
mals were not, however, able to cross over to Australia. At this time, Australia was
separated from the other continents. A few fossil teeth and jaws found in
Australia suggest that a placental species may have been present in Australia
between 100 and 50 million years ago. However, if any placentals were present in
Australia, they went extinct in the Tertiary, leaving the continent to the marsupials.
31 0 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

Continents would have cc

North Ameri<
Continental shelf
exchange of n

~
Early Marsupial sites between the r
Siberi[ e and suggested
migration routes enced by the 1
Manchu~ sia and Gond'

~
Early Placental
~ mammal sites and
suggested
Chi' migration routes Evolution
Although mu'
Central
Asia
mammals, the
mally groupe<
liest angiospe
rarely, flowers
ogy and gene
most closely r
sive efforts fo1
type of plant
angiosperms 1
the flowering
What ty
now agree th
FIGURE 10.3 The Late Cretaceous and Early Tertiary global dispersal of marsupial and they evolved f
placental mammals (after Osborne and Tarling, 1996). the common
maintain that
Finally, analy~
Elsewhere, the arrival and diversification of the placentals likely caused th gests that the
decline of the marsupials due to competition. The gestation of the young withi;: of angiosperrr
the uterus is thought to be more efficient than gestation in the external sac · The timt
found on marsupials, and this may have provided a reproductive advantage to been a topic
placentals. In Europe, competition from placentals led to the complete extinctior. angiosperms,
of the marsupials, while in North and South America the marsupials were greatly were present
reduced in importance and diversity. The early separation of Australia from tb dant during tl
other continents protected the marsupials there from competition with any mam- angiosperms i
mals except bats, and later rats. the modern pl
During the Tertiary, adaptive radiation allowed marsupial mammals to fill Early Tertiary
the many environments offered by Australia. Elsewhere, adaptive radiation an · total flora. Wh
Analysis
allopatric speciation caused by events such as the separation of Europe an ·
North America promoted speciation among the placental mammals. Orders suet est Cretaceou
evolved in the
as the Chiroptera (bats), Rodentia (rodents), Odontoceti (includes toothe ·
marine species such as dolphins and sperm whales), Mysticeti (includes filt e~ did not becom
feeding marine species such as gray whales), Proboscidea (elephants) , Perisso- tologists have
Cretaceous d(
dactyla (includes horses and rhinoceroses), and the Artiodactyla (includes cattle.
continents we
pigs, sheep, antelopes, and giraffes) all evolved between about 50 million and 5
tions of these
million years ago. Convergent evolution (see Chapter 9) by marsupials in Aus-
tralia produced species that were often strikingly similar in form and behavior to ing the Early C
spread from tl
the placental mammals dominant elsewhere.
The fact that the continents were in their modern locations during the angiosperm fl<
Tertiary Period meant that the many terrestrial mammals that evolved in Afri could indicate
by dispersal tc

Continents
Continental shelf
~
Early Marsupial sites
and suggested
migration routes
Early Placental
mammal sites and
suggested
migration routes
) liest angiosperms comes from fossilized leaves, stems, fruits, pollen, and, very
7 rarely, flowers. In addition, there has been much study of modern plant morphol-
) most closely related to the ancient ancestors of the angiosperms. Despite inten-
~ sive efforts for over 200 years, scientists still have not reached consensus on which
I dispersal of marsupial anc
acentals likely caused c:
;tation of the young witt::
ation in the external s<: -
reproductive advantage -
to the complete extinc r; ~ -
he marsupials were grea:...
1tion of Australia from ~=
;ompetition with any m e.= ·
marsupial mammals to =
=re, adaptive radiation <:::.-
>eparation of Europe c.::,_
rrtal mammals. Orders se.::
mtoceti (includes toot.::::-_
, Mysticeti (includes fi.::.~
cidea (elephants), Periss
rtiodactyla (includes car~:.
~n about 50 million anc : ..
r 9) by marsupials in _-\=.....
lar in form and behavio::-
1dern locations during --
nals that evolved in Ai::
FACTORS BEHIND THE MODERN BIOGEOGRAPHICAL REGIONS 311
would have considerable distances and barriers to overcome in order to migrate to
North America and vice versa. The isolation of Australia meant that almost no
exchange of mammals could occur for most of the Teritiary. Thus, the differences
between the mammal species found in the modern faunal realms is strongly influ-
enced by the relatively late speciation of the mammals and the breakup of Laura-
sia and Gondwanaland into separate landmasses by this time.
Evolution of the Flowering Plants
Although much mystery still remains concerning the evolution and spread of the
mammals, the history of the flowering plants (angiosperms-which are also for-
mally grouped as Division Magnoliophyta) is even cloudier. Evidence of the ear-
ogy and genetic structure in order to determine which living species might be
type of plant was the ancestor to the angiosperms, and when and where the
angiosperms first evolved. Indeed, Charles Darwin himself called the origin of
the flowering plants an "abominable mystery."
What type of plant was the ancestor to the angiosperms? Most botanists
now agree that the flowering plants are monophyletic in origin, meaning that
they evolved from a common ancestor. Some paleontologists have suggested that
the common ancestor may have been a type of cycad. Other paleontologists
maintain that the angiosperms may have evolved from seed-bearing ferns.
Finally, analysis of the morphological traits of some primitive living plants sug-
gests that the ancestor may have been related to the modern pines. The question
of angiosperm ancestry remains unresolved.
The time and place of the first appearance of the flowering plants have long
been a topic of great interest. There is good fossil evidence that early
angiosperms, including a number resembling modern magnolias (Magnolia),
were present in the Early Cretaceous. Angiosperms became increasingly abun-
dant during this geologic period. Between 100 million and 65 million years ago,
angiosperms increased from less than 1% of the flora to well over 50% . Many of
the modern plant families appeared in the Middle to Late Cretaceous. During the
Early Tertiary, angiosperms increased to comprise 90% or more of the earth's
total flora. Where did these successful plants first originate and spread from?
Analysis of the fossil leaf structure and geographic distribution of the earli-
est Cretaceous angiosperms has led many biogeographers to conclude that they
evolved in the tropics and then migrated poleward. It is known that angiosperms
did not become dominant in the high latitudes until the Late Cretaceous. Paleon-
tologists have recovered fossil angiosperm leaves, stems, and pollen from Early
Cretaceous deposits in eastern South America and western Africa. These two
continents were joined together as part of Gondwanaland at that time. The loca-
tions of these early angiosperm finds would have been close to the equator dur-
ing the Early Cretaceous and are conformable with a model by which angiosperms
spread from the tropics poleward (Fig. 10.4). In addition, the middle Cretaceous
angiosperm flora of Gondwanaland was more diverse than that of Laurasia. This
could indicate early evolution and speciation in the southern continent, followed
by dispersal to Laurasia. During the Cretaceous, Pangaea continued to break up,
312 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

South Pacific
Ancient Continents
Ancient Continental these modern
shelf Recent]
-- Spread of flowering plants the angiosper
• Area of origin of flowering
preserved fos
plants
deposits frorr
ern Beijing. T
now the olde:
features of th
ering plants n
Althou~
intensely deb
tant. The earl
sida). The inil
have occurrec
evolved from
~
Turkey
One of the me
grasses (Poao
the earth's lm
Early fossils c
to 55 million )
a particularly
increase in gr
drier global c
early grasslan
to have evolv<
ago. The wide:
tionary devel<
ingly, the Me!
Antarctica extensive gra!
GONDWANALAND earth's vegeta
number of ne
and tundra en
FIGURE 10.4 The Late Jurassic to Cretaceous global dispersal of angiosperms (after The exp<
Osborne and Tarling , 1996).
many gymnos
vegetation for
several compe
and the northern continents of Laurasia were separated from Gondwanaland by more efficient
the sea (Fig. 10.4). ovaries and pc
Not all botanists agree with an African-South American center for the evo- tion than are tl
lution and dispersal of the angiosperms. They point out that many of the mos: ferns and mos
primitive forms of flowering plants are found in the South Pacific, including por· with the fiowe1
tions of Fiji, New Caledonia, New Guinea, eastern Australia, and the Malay Arch- persal than exi
ipelago. Recent genetic research has identified the rare tropical shrub Amborella GymnOSJ
as being the living plant most closely related to the ancient ancestor of all the high latitudes i
angiosperms. This small shrub, which has tiny yellow-white flowers and red fruit , is tion of needle
found only on New Caledonia. Comparisons of the DNA of Amborella and ma n ~­ summers are sl
hundreds of species of flowering plants suggest that the first angiosperm arose anc peratures wan
speciation commenced about 135 million years ago. Many botanists conclude tha gymnosperm c
the best explanation for the large numbers of primitive living angiosperms in the distance betwe

Ancient Continents
Ancient Continental
shelf
.- Spread of flowering plants
I Area of origin of flowering
plants
rsal of angiosperms (afte r
~d from Gondwanaland :_
nerican center for the e :-
mt that many of the me:
Juth Pacific, including pc
tralia, and the Malay Ar C:::
; tropical shrub Ambon:
ancient ancestor of all ::::..:
ilite flowers and red frui:.. _
~A of Amborella and m<:::
first angiosperm arose c..:
my botanists conclude ~
e living angiosperms in -~
FACTORS BEHIND THE MODERN BIOGEOGRAPHICAL REGIONS 313
South Pacific region is that this is where the flowering plants first evolved and
these modern species are relics of this early evolution (Fig.10.4).
Recently discovered fossils complicate our understanding of the origin of
the angiosperms even further. Paleontologists from China have found beautifully
preserved fossils of an angiosperm plant, including flowers and seeds, in Jurassic
deposits from China. The site, which is about 130 million years old, is near mod-
ern Beijing. The new fossil plant (Archaefructus liiaoningensis) found at the site is
now the oldest known angiosperm. The age of the fossils and the very primitive
features of the flowers have led the discoverers to suggest that the earliest flow-
ering plants may have evolved in northern Asia (Fig. 10.4).
Although the timing and place of initial angiosperm evolution remain
intensely debated, other events in the evolution of angiosperms are very impor-
tant. The earliest angiosperms all appear to be dicotyledons (Class Magnoliop-
sida). The initial evolution of the monocotyledons (Class Liliopsida) appears to
have occurred in the Late Cretaceous. It is generally agreed that the monocots
evolved from a dicot ancestor. Some of the earliest fossil monocots are palms.
One of the most widespread and important families of all flowering plants are the
grasses (Poacea), which are monocots. Today grasslands comprise about 25% of
the earth's land surface. The grasses appear to have evolved relatively recently.
Early fossils of grass are found in deposits in Tennessee that date from about 65
to 55 million years ago. Commencing around 45 to 30 million years ago, there was
a particularly marked increase in the area of grasslands and savanna. This
increase in grasslands may be associated with the development of cooler and
drier global climatic conditions during the Oligocene and Miocene epochs. The
early grasslands were dominated by c3 species of grasses. The c4 species appear
to have evolved and spread during the Late Miocene, some 10 to 7 million years
ago. The widespread development of grasslands is also associated with the evolu-
tionary development of grazing guilds among the mammalian fauna. Interest-
ingly, the Mesozoic landscapes occupied by the dinosaurs did not contain the
extensive grassy meadows that we take to be such a common feature of the
earth's vegetation. As climate became cooler and drier during the late Tertiary, a
number of new plant species also evolved to occupy the newly forming desert
and tundra environments.
The expansion of the angiosperms led to the displacement and extinction of
many gymnosperms, ferns, and other plants that had dominated the terrestrial
vegetation for hundreds of million of years. It is thought that angiosperms possess
several competitive advantages over other plants. For example, angiosperms have
more efficient vascular systems and are better able to withstand drought. The
ovaries and pollen grains of angiosperms are often better protected from desicca-
tion than are the cones and pollen of gymnosperms or the sporangia and spores of
ferns and mosses. Finally, the coevolution of pollen and fruit-dispersing animals
with the flowering plants produced additional mechanisms for pollen and seed dis-
persal than exists for gymnosperms or ferns, which generally rely on the wind.
Gymnosperms such as pine and spruce, particularly in high elevation and
high latitudes in the northern hemisphere, continued dominant because the reten-
tion of needles during the winter is an advantage for conifers in regions where
summers are short. Evergreens are able to begin photosynthesis quickly once tem-
peratures warm up. In addition, the loss of viable seeds due to the desiccation of
gymnosperm cones is not such a problem in these cooler regions. Finally, the large
distance between the high latitudes of the northern and southern hemisphere and
314 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

the ocean barrier between the continents of Laurasia and those of Gondwana- existence. Pir
land may have restricted the northward dispersal of cold-adapted angiosperms North Ameri
such as southern beech and lessened the competitive pressure on the northerr ern beech is r
gymnosperms. The modern-day restriction of southern beech to the high latitud e~ cool sites tha
and high elevations of the southern hemisphere, and the restriction of pines tc AlthOUf
the northern hemisphere (Fig. 10.5), is directly related to the Late Cretaceotc clear that div'
evolution and distributions of these two genera. Nothofagus likely evolved ir: mammals. Tl
South America around 80 to 90 million years ago. The southern beeches were angiosperms '
then able to disperse to Australia, New Zealand, New Guinea, and Antarcti of the mamm.
Pines evolved about 130 million years ago in Europe. Subsequently, pines contin- not as great a:
ued to evolve and disperse in Europe, northern Asia, and North America. By the
close of the Cretaceous, pines were present in the mid- to higher latitudes of t h~
northern hemisphere, and southern beech was dominant in the high latitudes o:' THE MODERN Bl1
the southern hemisphere.
The Teritary and Quaternary history of southern beech and pine provid = In the follo"
an interesting contrast to the general success of the angiosperms. Over the past 6 ~ focus on som
million years, the range of pine in the northern hemisphere has become relativeh discuss some
large, and many species have evolved. Today there are over 100 pine species ii. timing of whe
of plate tecto
mal and plar
Oceanic faun
Antarctic pas
Modern pine/Modern
nothofagus and fauna of
Fossil pine/Fossil ments that di:
nothofagus

Nearct ic a
The close sim
have led som
realm called t
and fauna , an
discuss the t"
America, Gre
ern Africa, an
is smaller tha
restrial mamn
the Nearctic
species of nat
whelmingly d
sum. In contn
Nearctic. Well
American bu
lope (Antilocc
elk (Cervus ct
bilis), and co:
mammal spec
extinction evt
mammal dive
FIGU RE 10.5 The Late Cretaceous-Early Teritary distribution of the southern beeches Many gt
(Nothofagus) and pines (Pinus). The modern distributions of the two genera. Nearctic. Sorr

1 and those of Gondwanc.-
cold-adapted angiosperll:l5
~ pressure on the norther::
L beech to the high latitu d ~
the restriction of pines t.:
ed to the Late Cretaceo ~
•thofagus likely evolved i=
'he southern beeches we~ ::
w Guinea, and Antarctie<=-
Subsequently, pines contiL-
md North America. By th:e
1- to higher latitudes of tt:e
ant in the high latitudes c.:
n beech and pine provid"'"'
?,iosperms. Over the past c:
·here has become relatiw :
·e over 100 pine species :=
Modern pine/Moder
nothofagus
Fossil pine/Fossil
nothofagus
)
;;~··
I }~
Uiff.t~. ·~
?::J v )
•n of the southern beeches
1e tw o genera .
THE MODERN BIOGEOGRAPHIC REGIONS 3 15
existence. Pines can be found growing from the arctic treeline of Eurasia and
North America to the tropics of southeastern Asia and Central America. South-
ern beech is now represented by a handful of species, and its range is restricted to
cool sites that occupy only small portions of the southern hemisphere.
Although many details of angiosperm evolution remain to be resolved, it is
clear that diversification and spread occurred earlier for flowering plants than for
mammals. The relatively early evolution, diversification, and spread of the
angiosperms during the Cretaceous, compared with the latter evolution and spread
of the mammals, help to explain why the differences between the floral regions are
not as great as the differences between the mammal regions.
THE MODERN BIOGEOGRAPHIC REGION S
In the following consideration of the modern biogeographic regions, we will
focus on some of the better known endemic fauna and flora of each region and
discuss some of the highlights of their biogeographic history. As we will see, the
timing of when different continental plates either diverged or converged because
of plate tectonics is critical in explaining the modern distributions of many mam-
mal and plant families. Although Pielou recognizes distinctive Antarctic and
Oceanic faunal-floral regions, we will not include these in our discussion, for the
Antarctic possesses no terrrestrial mammals, and at the level of family, the flora
and fauna of the Oceanic region can best be considered an amalgamation of ele-
ments that dispersed from continental regions.
Nearctic and Palearctic Regions-The Holarctic
The close similarities in the fauna and flora of the Nearctic and Palearctic regions
have led some biogeographers to treat these two areas as one super region or
realm called the Holarctic. In view of the close relationships between their flora
and fauna , and the historical linkages during the Tertiary and Quaternary, we will
discuss the two regions together here. The Nearctic region includes all of North
America, Greenland, and most of Mexico. The Palearctic includes Europe, north-
ern Africa, and northern Asia. The mammal fauna of the Nearctic and Palearctic
is smaller than that of tropical regions. The Nearctic supports 13 families of ter-
restrial mammals, while the Palearctic supports 18 families. The mammal fauna of
the Nearctic shares many families and genera with the Palearctic. Twenty-one
species of native mammals are found in both realms. The Nearctic fauna is over-
whelmingly dominated by placentals. The one living marsupial genus is the opos-
sum. In contrast, there are some 111 different genera of placental mammals in the
Nearctic. Well-known species that are now endemic to the Nearctic include North
American buffalo (Bison bison), muskox (Ovibos moschatus), pronghorn ante-
lope (Antilocapra americana), bighorn sheep (Ovis canadensis), North American
elk (Cervus candensis), the prairie dog (Cynomys spp.), grizzly bear (Ursus horri-
bilis), and coyote (Canis latrans). During the Pleistocene, the number of large
mammal species in the Nearctic was much larger than it is today. A catastrophic
extinction event at the close of the Pleistocene led to a major decline in large
mammal diversity.
Many genera of mammals that are found in the Palearctic also occur in the
Nearctic. Some better known examples include buffalo (Bison bonansus, called
316 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

bison in Eurasia), moose (Alces alces, called elk in Eurasia), bear (Ursus spp. mammals. The
caribou (Rangifer tarandus, similar to reindeer in Eurasia), and lynx (Lynx spp. the grazing ho
In addition, the Palearctic contains genera that do not occur in the modern nati\·:o about 13 millie
fauna of the Nearctic. These include well-known mammals such as monkeys, pa::- ago allowed tl
das (Ailuropoda), tigers (Panthera), horses (Equus), and camels (Came/us). enter Eurasia.
The angiosperm flora of the Nearctic shares many of its families and ge - of the Antarcti
era with the Palearctic. The Nearctic is richer, possessing 94 different nat i\·:o and allowed di
angiosperm families compared to 69 in the Palearctic. In both regions, speci-= extinction of
of maple, oak, ash, elm, hazel, birch, basswood, poplar, and beech dominate tt:o Antarctica. Wi
deciduous forests. The two regions also share many conifer genera. The Near-- to Europe anc
tic and Palearctic have species of pine, spruce, fir, and larch dominating borec_ aided by the fil
vegetation. The arctic shrubs and herbs of the Nearctic and the Palearctic <L:' The curr
quite similar, even at the level of species. Species such as arctic willow (Sab: Nearctic and F
arctica) and avens (Dryas integrifolia) are common to both regions. The dese:-: Bears likely o
flora of the two regions is, however, quite different in composition. Cacti fc:- America in a r
example are found only in the Nearctic and Paleotropical regions. A number o: Carnivora, whi
desert genera are shared between the Nearctic and Neotropical regions. SoiL= North Americ:
well-known endemic trees of the Nearctic region include tulip tree (Liroder.- eastward and '
dron tulipifera), coastal redwood (Sequoia sempervirons), and Douglas Quaternary. Bl
(Pseudotsuga menziesii). The Palearctic also has many notable endemic tre : origins in Eurc
such as the olive (Olea europaea), pistachio (Pistacia vera), and old worl" the Late Tertia
cedars (Cedrus spp.) of the European and the Mediterranean region, the daw::: North Americ<
redwood (Metasequia glyptostroboides) of China, and the Japanese red cede.:: in the Quatern
(Cryptomeria japonica). Climatic'
The close relationship between the mammals and plants of the Nearctic a - nental glaciati<
Palearctic can be explained by the similar longitudinal positions of the tw - Nearctic and F
regions, and thus their similar range of climate and environments. In additioL and the polar 1
faunal and flora exchanges have been facilitated by the close connection betwee:. dispersed aero
Europe, Asia, and North America during much of the past 65 million years. cooler conditic
Finally, the environments of both regions were strongly impacted by the clima ·::. nonglacial con
and geographic changes associated with late Cenozoic cooling and resulting Que.· plants during 1
ternary glaciations. In the past, the two regions have been closely connected i:. North Am eric<
both the North Atlantic and the North Pacific regions. At approximately 60 to 5= European flon
million years ago, during the Early Eocene, 34 of the 60 mammal genera knm..-:. cantly compare
from Europe also occurred in North America. Early primates, ancestral to OJ- of Europe has
World and New World monkeys and other primates, likely evolved in No ' ruption of envi
America at this time and dispersed to Europe and Africa. A similar close corr -
spondence is evident in European and North American land plants during this Neotropica
period. Exchange of fauna and flora from Europe and North America was
enabled by a land linkage in the area of northeastern Canada, Greenland, anc The Neotropic
northern Europe. islands. It supp
As the North Atlantic grew, the barrier between North America an · diversity of m<
Europe increased over time. At the same time, the European and Asian plate: known endem
fused, allowing dispersal of land plants and animals across Eurasia. Well befor- (Lama), guine<
14 million years ago, Eurasia was one large landmass extending from the Atlanti also shares mar
to the Pacific. By this time, northeastern Asia and northwestern North Americz squirrels, cats, f
were almost joined in the area of Beringia. Sea-level changes due to glaciatim: mammal fauna
during the Late Tertiary and Quaternary produced intermittent land connections other regions. l
between the two continents. The drops in sea level appear to have allowe · related to the I
important exchanges to take place between Late Tertiary Palearctic and Nearcti lies of native 1

:urasia), bear ( Ursus spp.
1sia), and lynx (Lynx spp.
occur in the modern natiH'
nals such as monkeys, par:-
nd camels (Came/us).
my of its families and ger:-
essing 94 different nati\·.::
c. In both regions, spec i~
r, and beech dominate th.::
:onifer genera. The Neare-
d larch dominating borec_
;tic and the Palearctic ar:'
1ch as arctic willow (Sa b
o both regions. The dese:-:
in composition. Cacti fc:-
Jical regions. A number c:
'l"eotropical regions. Sori.::
elude tulip tree ( Liroder. -
rvirons), and Douglas :=
ny notable endemic tr e ~
!Cia vera), and old wo r ~:.
~rranean region, the da w=
td the Japanese red cede_:
I plants of the Nearctic ac
ina! positions of the t''.::
~nvironments. In additio=--.
: close connection betwee=
the past 65 million yea;-s_
y impacted by the clim a~:
cooling and resulting Quz-
been closely connected :..=:
At approximately 60 to~ -
50 mammal genera know=
primates, ancestral to 0 ~::
;, likely evolved in Nor:-_
rica. A similar close corr.::-
an land plants during tb.:s
and North America w e..;:
1 Canada, Greenland, ac:.
reen North America ar.:.
uropean and Asian pla t~
~ross Eurasia. Well befo:=
\tending from the Atlanr::
thwestern North Americ.=.
changes due to glaciatio=
:rmittent land connecti o~
appear to have allowe:.
.ry Palearctic and Nearer::
THE MODERN BIOGEOGRAPHIC REGIONS 317
mammals. The dispersal of horses is a particularly good example. The ancestor of
the grazing horses was the genus Hipparion. It first evolved in North America
about 13 million years ago. A large decline in sea level at about 11.5 million years
ago allowed the Hipparion horses to migrate across the Bering landbridge and
enter Eurasia. The drop in sea level was caused by the establishment and growth
of the Antarctic ice cap. The development of that ice cap caused sea level to drop
and allowed dispersal between Asia and North America but, of course, led to the
extinction of southern beech and almost all other terrestrial organisms in
Antarctica. Within a few hundred thousand years, horses had spread across Asia
to Europe and Africa. The spread of horses, and other grazing mammals, was
aided by the first development of grasslands in the Tertiary.
The current distributions of many of the mammal genera found in the
Nearctic and Palearctic can be traced to later Tertiary dispersal across Beringia.
Bears likely originated in Eurasia in the Late Teritiary and spread to North
America in a number of different dispersal events. Other members of the order
Carnivora, which includes dogs and cats, first evolved in Eurasia and dispersed to
North America. Wolves and cats experienced several waves of migration both
eastward and westward across the landbridge during the Late Tertiary and the
Quaternary. Buffalo, moose, caribou, and the pronghorn antelope also have their
origins in Eurasia and arrived in North America via the Beringian route during
the Late Tertiary and Early Quaternary. Camels, interestingly enough, evolved in
North America some 50 million years ago and dispersed to Eurasia via Beringia
in the Quaternary.
Climatic cooling during the Quaternary, together with the associated conti-
nental glaciations, has had a dramatic impact on the mammalian fauna of the
Nearctic and Palearctic. Cold-adapted mammals such as the muskox, reindeer,
and the polar bear evolved in the Palearctic during the last 1 million years and
dispersed across the northernmost portions of the two regions. The onset of
cooler conditions and the disruptions caused by the shifts between glacial and
nonglacial conditions also produced significant extinctions of mammals and
plants during the Quaternary. The Late Tertiary floras of Europe and eastern
North America were relatively similar in diversity. During the Quaternary, the
European flora, particularly in terms of tree species, has been reduced signifi-
cantly compared to that of North America. The relatively low mammal diversity
of Europe has also been attributed to the impact of climatic cooling and the dis-
ruption of environments by glacial cycles during the Quaternary.
Neotropical Region
The Neotropical region includes South and Central America and the adjacent
islands. It supports an interesting assortment of mammal types and has a higher
diversity of mammals than the Nearctic, consisting of some 23 families. Well-
known endemic mammal genera of the Neotropical Region include llamas
(Lama), guinea pigs (Cavia), and chinchillas (Chincilla). The Neotropical region
also shares many families and genera with the Nearctic, including rabbits, shrews,
squirrels, cats, foxes, raccoons, mountain lions (pumas), and deer. In addition, the
mammal fauna of the Neotropical region has some interesting similarities with
other regions. It includes ilamas, which belong to the family Camelidae and are
related to the Palearctic camels. The Neotropical regions also support two fami-
lies of native monkeys that are more primitive but distantly related to those
318 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

found in the Palearctic, Oriental, and Ethiopian regions. One interesting featur establishment
of the Neotropical region is the diversity of endemic rodents found there. ~ migration of t
includes 12 different endemic families and close to 200 endemic species. Fina ll ~-. deer and cats.'
the Neotropical region contains the highest diversity of marsupial mammals nex: only to the in1
to Australia. The marsupials contribute three families and 84 different species tc> the widesprew
the mammal fauna of the Neotropical region.
The flora of the Neotropical region is notable for its variety. The regio Ethiopian I
possesses 137 different families of flowering plants, making it the most diverse o:
all floral regions. Over 50 families of angiosperms are endemic to the Neotropi- The Ethiopian
cal region. These range from plants of the alpine tundra to the rainforest and th"' Arabian Peni1
desert. A number of other angiosperm families are shared with the adjacen: than Africa an
Nearctic region. Some, such as the cacti, have relatively continuous distributions region possess
across the Isthmus of Panama. A number of shared plant genera and species; families) and l
show an interesting pattern of disjunction in their distributions between th- many well-kn<
Neotropical and the Nearctic. A surprisingly large number of desert specie- zebras (Equu:
including the creosote bush (Larrea tridentata), which is very common in Cali- subgutturosa),
fornia, are found both in the North American desert and in the desert regions a: odonta africar
the southern tip of South America. A large number of Neotropical plant familie~ (Gorilla gorill
and genera are also found in the Australasian or Ethiopian regions. Notable families with
examples besides southern beeches include southern hemisphere coniferous mammal famil
tree genera, Podocarpus and Araucaria, and many genera and species of the contrast, the E
family Proteaceae. The genera Podocarpus and Weinmannia are found in Aus- the N eotropic<
tralian, Ethiopian, and Neotropical regions. In addition, several genera of cus - The asso
ion plants found in the mountain tundra of New Zealand and Tasmania alsc Palearctic can
occur in Chile. These include plants of the genera Donatia, Phyllanchne. mammal fossil
Drapetes, and Gaimardia. years ago. Hov
The high diversity of the Neotropical fauna and flora, and their linkages; placental mam
with other biogeographical regions, reflect both the environmental variety an ration of Afric
the geologic history of the region. The Neotropical region includes environmen " persal of the :
ranging from alpine tundra to temperate forest, tropical rainforest, dry tropica; Miocene, abou
forest, savanna, grasslands, and desert. The range of potential niches is therefor far enough to ·
greater than in the Nearctic or Palearctic regions. The relative importance of th duced a landb1
marsupials in the mammal fauna is explained by the former physical connectioL facilitated the
between South America and the other continents of Gondwanaland. During th mates, camels .
early mammal and angiosperm radiations of the Cretaceous, South America w~ such as the anc
joined to Australia by Antarctica. Africa had already began to move northwest- The connectio
ward and separate from Antarctica by the early Late Cretaceous, about 95 mil - tion of elephar
lion years ago. By the Early Tertiary, about 55 million years ago, South Amerie<:. regions. The c
was well separated from Antarctica but had not yet moved far enough northwar Africa in the L
to be linked to North America. Its mammal fauna, dominated by marsupials, an tually colonize
its flora , containing many Gondwana taxa, evolved in isolation for almost 40 mil- Africa at abm
lion years. This history explains the linkages between the Neotropical, Aus- includes mode
tralasian, and Ethiopian fauna and flora. onized the Pal1
The presence of fauna and flora typical of the Nearctic region is explainec The asso
by the development of a landbridge between the two regions which occurrec Neotropical aJ
with the formation of the Isthmus of Panama about 3 million years ago (see the angiospen
Chapter 8). Some migration across the islands that lay in the area of the presen southern conti
isthmus likely began earlier than this. It is thought that the primate ancestors of mon plant gen
the Neotropical monkeys and the ancestors of the diverse Neotropical rodents However, the
arrived via these biogeographic stepping stones during the Middle Tertiary. The of others such

ms. One interesting feat ur.e
~ rodents found there. 1llli
DO endemic species. Finall:
)f marsupial mammals nex:
. and 84 different species t-
for its variety. The regio-
aking it the most diverse c:
~endemic to the Neotrop:-
.ra to the rainforest and tt::
' shared with the adjace;:::
ly continuous distributio:::..:
I plant genera and spec i~
distributions between t ~=
number of desert species.
;h is very common in c a:.:;.-
:md in the desert regions c_
'Neotropical plant fam ilie:.
~thiopian regions. Not a b~ ::
rn hemisphere conife ro~
genera and species of tt::
tmannia are found in A \:5-
on, several genera of cus::.-
ealand and Tasmania als.::
:ra Donatia, Phyllanchr.-
td flora, and their link ag::-..
environmental variety a;:.:
~ion includes environmer.:..
ical rainforest, dry trop ic-~
otential niches is therefo:-::
relative importance of tt::
'o rmer physical connectic-
}ondwanaland. During t:..::
1ceous, South America w .c__
began to move northwes:-
Cretaceous, about 95
years ago, South Ameri.:::
>Ved far enough northwa:-.:.
ninated by marsupials, a;:;.:
isolation for almost 40 JL1_-
:en the Neotropical, Ar -
earctic region is explain.:-.:
0 regions which occurr.:-,::
t 3 million years ago (~;;-::
r in the area of the pre s ~ ­
lt the primate ancestors
verse Neotropical roder: :..
g the Middle Tertiary. T::.::
THE MODERN BIOGEOGRAPHIC REGIONS 319
establishment of a landbridge around 3 million years ago allowed the northward
migration of the marsupial opossum and the southward migration of placental
deer and cats. This so-called Great American Interchange (see Chapter 8) led not
only to the introduction of species to the Neotropical region, but it also caused
the widespread extinction of many marsupial species in the Neotropical region.
Ethiopian (African) Region
The Ethiopian region consists of sub-Saharan Africa and adjacent portions of the
Arabian Peninsula. The island of Madagascar has a different geologic history
than Africa and is generally excluded from the Ethiopian region. The Ethiopian
region possesses the most diverse mammal fauna of all biogeographic regions (30
families) and has an extremely diverse angiosperm flora (117 families). There are
many well-known endemic mammal genera in the Ethiopian region, including
zebras (Equus burchelli), giraffes (Giraffa camelopardalis), gazelles (Gazella
subgutturosa), wildebeests (Connochaetes taurinus), African elephants (Lox-
odonta africana), baboons (Papio spp.), chimps (Pan troglodytes), and gorillas
(Gorilla gorilla). Although the Ethiopian region shares a number of mammal
families with the adjacent Palearctic and Oriental regions, it shares few land
mammal families with the Neotropical and none with the Australian region. In
contrast, the Ethiopian region shares a number of angiosperm plant families with
the Neotropical and Australian regions.
The association between the fauna of the Ethiopian region and that of the
Palearctic can be attributed to the geologic history of the two regions. The first
mammal fossils found in Africa date from about the Eocene, some 55 to 40 million
years ago. However, despite the lack of direct evidence, it is likely that some early
placental mammals were present in Africa in the Late Cretaceous. The early sepa-
ration of Africa from Australia, South America, and Antarctica prevented the dis-
persal of the southern marsupial fauna to the Ethiopian region. By the Early
Miocene, about 20 million years ago, the African continent had moved northward
far enough to collide with Eurasia in the vicinity of Arabia and Turkey. This pro-
duced a landbridge between the Ethiopian and Palearctic regions. The landbridge
facilitated the southward migration of animals such as the ancestors of the pri-
mates, camels and zebras, which originated in the Nearctic region, and carnivores
such as the ancestors of lions and jackals, which originated in the Palearctic region.
The connection between Africa and Eurasia also allowed the northward migra-
tion of elephants and their relatives, the mammoths, to the Palearctic and Oriental
regions. The order Proboscidea, to which elephants belong, likely evolved in
Africa in the Late Eocene, some 40 million years ago. The mammoths, which even-
tually colonized most of the Palearctic and Nearctic, migrated northward from
Africa at about 2 million years ago. At the same time, the genus Equus, which
includes modern horses and zebras, migrated westward from the Nearctic and col-
onized the Palearctic and Ethiopian regions.
The association between the Ethiopian angiosperm flora and that of the
Neotropical and Australian regions reflects the fact that during the radiation of
the angiosperms in the Late Cretaceous, Africa was still close enough to these
southern continents to exchange flora. This explains the presence of many com-
mon plant genera in the Ethiopian, Australasian, and Neotropical regions today.
However, the relatively early separation of Africa likely precluded the dispersal
of others such as southern beech to the Ethiopian region.
320 CHAPTER 10 REALMS, REGIONS, AND PROVINCES

Oriental Region million years, 1

temperate, bo
The Oriental region includes the Indian subcontinent and adjacent portions of ever, much in
southern Asia. Although it is small in area compared to the regions we have dis- Asian portion:
cussed thus far, it has a very high diversity of both mammals (20 families) an keys, gibbons,
angiosperms (108 families) . Some well-known endemics include the gibbon fam- Fossils of the l
ily (Hylobatidae), Orangutans (Pongo pygmaneus), and the flying lemur family ental region, h
(Cynocephalidae ). A number of well-known species from the Oriental region
have relatives in the Ethiopian region. These include the Indian elephant (Ele-
phas maximus) and the Indian rhinoceros (Rhinoceros unicornis), which hav Australian
relatives in Africa. The Bengal tiger of India (Panthera tigris tigris) has a close rel- The Australim
ative in the Siberian tiger (Panthera tigris altaica), which is found in the south- regional subd
eastern portion of the Palearctic. The tropical angiosperms of the Oriental region most distinct
have many relatives in both the Ethiopian and Australian regions, while the mon- almost complt
tane vegetation contains Palearctic-Nearctic elements, including pines. A well - placental man
known endemic tree of the Oriental region is teak (Tectona grandis). In addition. mals, such as ·
the tropical forests of the region are distinctive in being characterized by a large tralia by hu
number of trees of the family Diptocarpaceae. kangaroos (M
The mammalian fauna of the Oriental region has close ties to the Ethiopia cinereus)). In:
and Palearctic regions and none with the adjacent Australian region. In contrast. laying monotr
the flora has closer ties with the Australian, Ethiopian, and Neotropical region and the echidr
The explanation for these relationships lies in the rather complex geologic his- nia have a rici
tory of the region. During the Early Cretaceous, India and Madagascar were pan early separati
of Gondwanaland and were connected to the southern continents. By the Lat arrival of hum
Cretaceous, some 90 million years ago, the Indian landmass had separated fro m small and con
the rest of the southern continents and had begun a northward movement. By than that of ar
this time, the Indian flora contained many elements of the Australian and African the Oriental, l
angiosperm floras. When India eventually collided with the Asian continen t. In contrc
these plants were able to expand to other portions of the Oriental region. are endemic, ,
The transport of flora to Asia by the Indian subcontinent is evident in th Two very dorr
similarities between the Oriental, Neotropical, and Australasian floras. However.
lyptus is the q
there is no evidence that a mammal fauna of either southern marsupials or pla-
Tasmania, anc
centals existed in India prior to its contact with Asia. India came into contact with
grow from al]
Asia by the Middle Miocene, some 14 million years ago. The transport of south-
accounts for a
ern species to Asia by the northward-moving Indian subcontinent represents a
does not occt
special type of dispersal mechanism referred to as an ark. The name is a poeti
plants to whic
homage to the story of Noah's ark from the Judea-Christian bible. Some of th
region during
earliest mammal fossils from India date from the Miocene and include faun a
the sundering
from both the Palearctic and Ethiopian regions.
the Tertiary. Tl
By the Middle Miocene, Australia and New Guinea had moved northward
adaptive radi<
and were in close proximity to the Malaysian Archipelago. However, Australia
have not bee1
was never directly connected to the Oriental region via a landbridge, and this has
prevented a natural exchange of mammal fauna. It did not preclude the migra- region via the
tion of many other plants and animals, however. Species of the plant family Pro- and animals h
teacea dispersed and colonized southeastern Asia from Australasia. Angiosperm ble northward
families that first evolved in Asia, such as the Myrtaceae, dispersed southward to spread bird gr
Australasia. At this time, bats and birds also dispersed across the stepping stone evolved in Au
route from the Oriental region to Australasia. across the Ma
Many of the Oriental mammals and angiosperms evolved in Southeast origins in Aus
Asia. The Southeast Asian region has been in the lower latitudes for over 100 Nearactic.

t and adjacent portions o::
o the regions we have dis-
tammals (20 families) an.:
:::s include the gibbon fao:;-
ld the flying lemur fai11L:
from the Oriental regi o=
the Indian elephant ( Ele-
os unicornis), which ha' ::
tigris tigris) has a close re:-
tich is found in the souti:.-
rms of the Oriental regio.::.
an regions, while the mor:-
>, including pines. A wel.-
tona grandis). In additio;:;.
g characterized by a larg::
close ties to the Ethiopic=.
tralian region. In contras::.
, and Neotropical regioc.s.
her complex geologic 18-
md Madagascar were pa;-
n continents. By the La:::
!mass had separated fro=
northward movement. B
he Australian and Africa::
v'ith the Asian continer::
1e Oriental region.
:ontinent is evident in tl::::
:tralasian floras. Howe' e:-
uthern marsupials or plc.-
jia came into contact wi~
o. The transport of sourt.-
;ubcontinent represen ts .:.
ark. The name is a poet::
ristian bible. Some of tt::
ocene and include fa ur:_
ea had moved northw<L.:
~!ago. However, AustraL
a land bridge, and this he.;;
I not preclude the migrc.-
:s of the plant family Pre-
Australasia. Angiosper.::.
e, dispersed southward tc
across the stepping stoc::
ns evolved in Southeas·
er latitudes for over l C
THE MODERN BIOGEOGRAPHIC REGIONS 321
million years, resulting in the evolution of a flora and fauna distinctive from the
temperate, boreal, and arctic fauna and flora of the adjacent Palearctic. How-
ever, much interchange did take place between the Palearctic and Southeast
Asian portions of the Oriental regions. The primate ancestors of the Asian mon-
keys, gibbons, and orangutans migrated from the Palearctic during the Tertiary.
Fossils of the Diptocarpaceae, a family that may have first originated in the Ori-
ental region, have been found in Early Tertiary deposits from Alaska.
Australian Region
The Australian region includes Australia, Tasmania, New Guinea, and in some
regional subdivisions New Zealand. Despite its relatively small size, it is the
most distinct of all the biogeographic regions. Its native mammal fauna is
almost completely dominated by marsupials. It is possible that the only native
placental mammals not introduced by humans are bats. Other placental mam-
mals, such as the dingo dog, and perhaps the rat, were likely brought to Aus-
tralia by humans. Unique and well-known endemic mammals include
kangaroos (Macropus), wallabees (Wallabia), and koala bears (Phascolarctos
cinereus)). In addition to marsupials, the region contains the only surviving egg-
laying monotremes, including the duckbill platypus (Ornithorhynchus anatinus)
and the echidnas (Tachyglossus and Zaglossus). Although Australia and Tasma-
nia have a rich marsupial fauna, the islands of New Zealand, because of their
early separation from Gondwanaland, had no non-flying mammals until the
arrival of humans. The mammal diversity of the Australasian region is relatively
small and contains only 10 families. The angiosperm diversity is slightly lower
than that of any other continental region (90 families) and shows affinities with
the Oriental, Ethiopian, and Neotropical regions.
In contrast to the mammals, in which all the marsupials and monotremes
are endemic, only 18 of the angiosperm families are endemic to Australasia.
Two very dominant endemic genera are Eucalyptus and Melaleuca. The Euca-
lyptus is the quintessential Australian tree. It occurs in Australia, New Guinea,
Tasmania, and smaller adjacent islands. There are many different species that
grow from alpine tundra to dry savanna to tropical rainforests. Eucalyptus
accounts for about 95% of the tree biomass in Australia. The genus, however,
does not occur in New Zealand. Fossil evidence suggests that the family of
plants to which the eucalyptus belongs, the Myrtaceae, evolved in the Oriental
region during the Late Cretaceous and likely migrated to Australia following
the sundering of Australia's connection to Antarctic and New Zealand during
the Tertiary. The many modern species of Australian eucalyptus are the result of
adaptive radiation by this Asian angiosperm family. Although the eucalyptus
have not been able to disperse from the Australasian region to the Oriental
region via the stepping stone route of the Malaysian Archipelago, other plants
and animals have been able to disperse. We have already mentioned the possi-
ble northward movement of the Proteaceae family. The ancestors of the wide-
spread bird group, the Corvini, which includes crows, ravens, and magpies, first
evolved in Australasia and then spread throughout the world after migrating
across the Malaysian Archipelago in the Miocene. These birds, which have their
origins in Australasia, can now be found as far away as the Arctic treeline of the
Nearactic.
322 CHAPTER 10 REALMS, REGIONS, AND PROVINCES
KEY WORDS AND TERMS
Biogeographic province
Ark
Biogeographic realm
Biogeographical provincialism
Biogeographic region
Biogeographic line
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Darlington, P. J., Jr. (1965). Biogeography of the
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Kurten, B. (1969). Continental drift and evolu-
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MacDonald, G. M. , Cwynar, L. D., and Whitlock .
C. (1998). The late Quaternary dynamics of
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Morain, S. A. (1984). Systematic and Regional Bio·
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Muller, P. (1974). Aspects of Zoogeography. Dr.
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O 'Brien, S. J. , Menotti-Raymond, M., Murphy, W.
J., Nash, W. G., Wienberg, J. , Stanyon, R.,
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Osborne, R. , and Tarling, D. (1996). The Histori-
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Pearson, L. C. (1995). The Diversity and Evolu-
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Pielou, E. C. (1979). Biogeography. Wiley, New
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Richardson, D. M. , and Runde!, P. W. (1998).
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Cambridge University Press, Cambridge.
Romer, A. S. (1966). Vertebrate Paleontology, 3rd
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Rougier, G.W. , Wible, J.R. , and Novacek, M.J.
(1998). Implications of Deltatheridium speci-
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Savage, R. J. G. (1986). Mammal Evolution: An
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Francisco.
:::wynar, L. D. , and Whitlock_
e Quaternary dynamics of
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Systematic and Regional Bio-

ostrand Reinhold, New York_
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·Jing, D. (1996) . The Histori·

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i). Mammal Evolution: An

Facts on File, New York .
 monkeys of S
CHAPTER 11 Catarrhinii an
(gibbons), an'
together in a ~
BIOGEOGRAPHY AND these other p1
actual basis fc
The get
HUMAN EVOLUTION apparent thrc
our closest re
atives are the
the blood an
Where does the human race come from? How has the environment shaped -. share many 1
How have we shaped the environment? These are basic questions that we all po::::.- tadactyl, hav
der. People seek answers for such universal questions through many differ :::: cases the fee
avenues. After the publication of his book on the origin of species in 1859, Chari"' are perhaps 1
Darwin directly tackled the problem of human origins in his two-volume treatis ~ pencil with o
On the Descent of Man, and Selection in Relation to Sex. In this work, published =. a great range
1871, Darwin made the then startling proposition that humans were related by ~ mates can br
common ancestor to the great apes, such as chimpanzees and gorillas. Darw=. branch with
argued that, like other species, we have evolved to our present physical form aL.: do when the:
degree of intelligence through natural selection. The work and its tenets were, aL.: mals cannot
remain, roundly condemned by some. Others consider Darwin's work a seminC..:. many other
stroke of genius and courage. Since the time of Darwin, scientific research into t ~ and apes, is s
evolution of humans has been a major focus of work by anthropologists, biologis ' because the'
geologists, and paleontologists. This is also the case for biogeographers, who haF the same ot
long been interested in how geography and environmental change have infie- unique cent:
enced human evolution and the early spread of humans around the globe. Indee - focusing on c
the history of our own species remains one of the most fascinating and importaL: Of the prim
of all biogeographic stories. fovea centra
In this chapter, we will review the current state of knowledge regarding t h~ vertebral co
history and geography of human evolution. We will start by looking at our relatior:- for other m
ship to other closely related species of mammals, the primates. We will then trace head. The al
the evolutionary and geographic history of the primates and human ancestors. \\ -- when we co
will also consider the role that environmental change may have played in t h~ jaws contaii
human evolutionary process. We will then examine the rise in the modern huma;: scrapping (i
population size and the geographic dispersal of humans throughout the world. generalist d
although ou
the brains c
THE PRIMATE LINKAGE cally compl<
The sl
Human (Homo sapiens sapiens) are considered by biologists to belong to the arms traits,
order of placental mammals called the Primates. The Primate Order includes ani- in trees. Me
mals such as lemurs, tarsiers, monkeys, apes, and humans. We are the only living trees. Other
members of the Primate family Hominidae and the only living members of the escape prec
genus Homo. The Primate Order is divided by primatologists into two suborders climbers co
and then three infraorders. Humans belong to the suborder Haplorhini, which between br
includes tarsiers, monkeys, and apes. The other suborder, Strepsirhini, includes help identif
lemurs, lorises, and indiriids, which are considered a somewhat primitive group of using prehe
primates. Within the Haplorhini, humans belong to an infraorder called the tageous to
Catarrhinii, which includes the Old World monkeys and the apes. The New World offspring. L

324

e environment shaped ··-
: questions that we all
ts through many diffe r;:-
of species in 1859, Ch a;-~~
in his two-volume tre a t:~~
. In this work, publishec
humans were related b· _
tzees and gorillas. D arv•
present physical form c...:,
>rk and its tenets were. cc
· Darwin's work a semi:.
scientific research into:..::.:
anthropologists, biolo~ ~
biogeographers, who he. _
nental change have i.r:..:':._
around the globe. Inde.:-.:
fascinating and impor:=-
f knowledge regarding :.:
by looking at our relati c:.
·imates. We will then tR
, and human ancesto rs. ':
: may have played in
rise in the modern hu
throughout the world.
ologists to belong to :..: -
imate Order includes --
ns. We are the only U \'-=._
ly living members of :_'--
ogists into two suborc::c
Jorder Haplorhini, \\·t;, ~ ­
er, Strepsirhini, incl u.:_
1ewhat primitive grou;
an infraorder called --
the apes. The New V.'o:-. _
THE PRIMATE LINKAGE 3 25
monkeys of South and Central America appear to be more primitive than the
Catarrhinii and are grouped in the infraorder Platyrrhinii. Humans, the small apes
(gibbons), and the great apes (chimps, orangutans, gorillas) are often grouped
together in a superfamily called the Hominoidea in recognition of the similarity of
these other primates to humans. That is the scientific classification, but what is the
actual basis for claiming that humans are related to other primates?
The genetic relationship between humans and other primates is readily
apparent through DNA analysis. Humans share about 98% of their genome with
our closest relatives, the chimps, and a bit less with gorillas. Our next closest rel-
atives are the orangutans. A close relationship is also supported by similarities in
the blood and protein chemistry of humans and these three primates. Humans
share many physical characteristics with other primates. The primates are pen-
tadactyl, having five fingers and toes. The hands of the primates, and in most
cases the feet, are prehensile, which means they can grasp objects. Human feet
are perhaps the least prehensile of the primates, but most of us can still grasp a
pencil with our toes. The primate clavicle and scapula (shoulder blade) allow for
a great range of movement and a high degree of strength in the arms. Many pri-
mates can brachiate, which means they can use the arms to move from branch to
branch with the body suspended beneath. Brachiating is what human children
do when they swing hand over hand along monkey bars. Most nonprimate mam-
mals cannot brachiate. The primates have a reduced sense of smell compared to
many other mammals. Thus, the snout on most primates, particularly monkeys
and apes, is small and relatively flat. Most primates also have stereoscopic vision
because the eyes are placed on the face in such a way that both eyes can focus on
the same object at once. Most primates also have color vision and possess a
unique central pit in the retina called the fovea centralis, which allows sharp
focusing on one object while retaining visual perception of surrounding features.
Of the primates, only lemurs do not possess color vision, stereoscopy, and a
fovea centralis. The primate cranium is generally high and vaulted. The primate
vertebral column (the spine) attaches lower on the primate skull than is the case
for other mammals. This attachment allows for great mobility of the primate
head. The above features of the primate skull help produce the similarity we see
when we compare the heads and faces of humans, apes, and monkeys. Primate
jaws contain teeth for grinding food (molars), puncturing (canine teeth) , and
scrapping (incisors). This wide range of tooth types suggests an adaptation to a
generalist diet. Human dentition is quite similar to chimp and ape dentation,
although our canines are greatly reduced. Relative to other terrestrial mammals,
the brains of primates are large compared to their body mass and physiologi-
cally complex.
The shared primate features of prehensile hands and feet, shoulder and
arms traits, and color streoscopic vision are best explained as adaptations to life
in trees. Most primates are almost completely arboreal, meaning they live in
trees. Others, such as chimps, spend some time on the ground but climb trees to
escape predators or secure food. Even modern humans are reasonably good
climbers compared to many other mammals. Being able to judge the distance
between branches using stereoscopic vision, seeing color differences that might
help identify food or predators within dense forest canopies, being able to climb
using prehensile hands and feed, and moving swiftly by brachiating are all advan-
tageous to arboreal animals. Primates give birth to relatively small numbers of
offspring. Lemurs produce two or three offspring at each birth, but monkeys and
326 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

apes generally give birth to only one offspring at a time. Again, this is likely ar.
adaptation to arboreal existence. Primate mothers must care for their offspring origins, inc!
for several months in the case of lemurs and up to four years in the case o: tralians. Th1
ancestor fo
chimps. In addition to feeding and protecting them, the mothers must carry off-
years ago. II
spring with them along branches. Broods of more than one offspring would be cise determ
difficult to accommodate. mon ancest
That humans are relatively closely related to primates is indisputable give;:. million yem
the overwhelming genetic, physiological, morphological, and behavioral similari- tion of this
ties. What remains of intense interest and debate is when and how our ancesto _ The anth
diverged from the other primates and we developed into the modern humar. to studying
species. A special type of analysis of genetic variability allows us to construct c. Robert Wa:
molecular clock based on genetic differences between humans and betwee;:. and preserv
humans and other primates. Molecular clock analysis is generally based on t h ~ one study, t
study of mitochondrial DNA, which is a type of DNA that is only passed fron: compared t
dogs were i
mother to offspring. The degree of difference between the mitochodrial DNA :
dog breeds
different species tells us the time that has elapsed since species divergence and interbr·
occurred (Technical Box 11.1). Molecular clock analysis suggests that the humar; concluded 1
domesticatt
tication oft
domesticati
TECHNICAL BOX 11.1 have been
there is no !
DNAha'
Molecular Clocks that died ir
Molecular clocks are developed through the analysis of genetically controlled dif- years old, tl
ferences in individuals and species. It is assumed that two individuals that share the Los Angele
same recent ancestor will possess similar genes and genetic traits. Over time, if the tos) that is
lineages from these individuals remain separated, natural selection and genetic drift ered from 1
will cause a divergence in the genes of the two lineages. For genes that are mainly are tens of 1
influenced by genetic drift, and not strongly influenced by natural selection, the rate
of divergence should be controlled by the natural rate of spontaneous mutation for
the organism. If estimates can be made of the average rate of mutation, the degree
of genetic difference between two populations of species can be compared and the lineage diver
time it took to develop the mutationally controlled genetic differences between the ago. The sear
populations or species can be calculated. primates and
In the 1960s, Allan Wilson and Vincent Sarich of the University of California at science. Traci
Berkeley analyzed the genetically controlled differences in blood proteins from see how geog
humans and other primates and concluded that divergence between ape and human
opment ofm
ancestors occurred 5 to 6 million years ago. Molecular clocks have been devised and
applied to a number of other organisms.
In more recent years, molecular clock research has focused on mitochondrial
DNA (mtDNA). Mitochondrial DNA is found in small energy-producing structures EARLY PRIMATE
in the cytoplasm of cells. Unlike nuclear DNA which controls inherited traits such as
eye color, mtDNA is inherited only from the maternal parent. Thus, the mtDNA of Our knowle<
plants and animals should be an exact copy of the female parent. However, it is most abunda
known that mtDNA has a high rate of spontaneous mutation. This rate of mutation compared to
is about ten times faster than nuclear DNA. Thus, over time the mtDNA of a lineage sparser than
will diverge through mutation from the original maternal pattern. Breeding patterns tion is rare in
or natural selection does not influence the divergence rates and patterns of mtDNA. swamps have
These properties make mtDNA an ideal molecular clock. The longer two lineages
fossils after
have been separated, the greater the divergence in their mtDNA. Wilson and his
colleagues analyzed the mtDNA differences in 147 women of different races and
arose during
a common a1

me. Again, this is likely a;:
ust care for their offsprin~
four years in the case o:
1e mothers must carry ot-
m one offspring would ~
nates is indisputable giYe=
al, and behavioral similar:-
len and how our ancesto:o
I into the modern hum c..:.
ty allows us to construct .: .
:en humans and betwee=
is generally based on tL:
\. that is only passed fr o=
the mitochodrial DNA c:
since species divergenc:-
is suggests that the hum· -
genetically controlled dif-
individuals that share the
tic traits. Over time, if the
selection and genetic drift
For genes that are mainly
natural selection, the rate
>pontaneous mutation fo r
.e of mutation, the degree
can be compared and the
c differences between the
fniversity of California at
; in blood proteins from
: between ape and human
ks have been devised and
)CUSed on mitochondrial
~ rgy-producing structures
)Is inherited traits such as
·ent. Thus, the mtDNA of
le parent. However, it is
ion. This rate of mutation
~the mtDNA of a lineage
attern. Breeding pattern
and patterns of mtDNA
The longer two lineages
mtDNA Wilson and his
~n of different races and
EARLY PRIMATES 327
origins, including Africans, Asians, Caucasians, New Guineans, and Aboriginal Aus-
tralians. They concluded that this mtDNA evidence pointed to a common African
ancestor for all living humans and that this ancestral woman lived about 200,000
years ago. More recent work on molecular clocks, however, suggests that such pre-
cise determinations are beyond the abilities of Wilson's mtDNA analysis. The com-
mon ancestor of all modem humans might have lived from 100,000 years ago to 1
million years ago. In addition, it may be impossible to ascertain the geographic loca-
tion of this common ancestor using Wilson's data.
The anthropological use of molecular clock estimates of divergence is not limited
to studying the evolution of humans or using only samples from living organisms.
Robert Wayne and his colleagues have examined the mtDNA of the bone, teeth,
and preserved tissues of a number of Pleistocene and Holocene mammal species. In
one study, they compared the mtDNA of 67 different modern breeds of dogs and
compared the results to the mtDNA of wolves, coyotes, and jackals. They found that
dogs were indeed closely related to wolves and that much of the modern diversity in
dog breeds can be related to the high degree of genetic diversity in ancestral wolves
and interbreeding between dogs and wolves following dog domestication. They also
concluded that there was likely a number of different locations where wolves were
domesticated by hominids. The most startling conclusion was that the initial domes-
tication of the dog may have begun over 100,000 years ago. His best estimate for dog
domestication is 135,000 years ago, and the most likely hominid involved would
have been the Neanderthals. Archaeologists have responded by pointing out that
there is no fossil evidence linking domesticated dogs with Neanderthals.
DNA has been successfully recovered from the remains of a number of organisms
that died in the past. These include Egyptian mummies that are several thousand
years old, the sabertooth cat (Smilodan fatalis) from the Rancho La Brea tar pits of
Los Angeles that is over 10,000 years old, and brown bear from Alaska (Ursus arc-
los) that is over 40,000 years old. Although there are reports of DNA being recov-
ered from million-year-old magnolia leaves and insect and dinosaur remains that
are tens of millions of years old, many scientists remain skeptical of such claims.
lineage diverged from the chimps and gorillas between 5 and 10 million years
ago. The search for fossils and other physical evidence of the ancestors of modern
primates and modern humans is one of the most intriguing adventures in modern
science. Tracing the fossil evidence of primate and human evolution also helps us
see how geography and environment have been such crucial factors in the devel-
opment of modern species.
EARLY PRIMATES
Our knowledge of early primate evolution comes from the fossil record. The
most abundant fossils are often jaws and teeth; these remains are fairly robust
compared to other parts of the skeleton. The fossil record of the early primates is
sparser than we might hope because these animals were arboreal and fossiliza-
tion is rare in forested settings. Organisms living in environments such as lakes or
swamps have a much better chance of being buried by sediment and preserved as
fossils after death. From the evidence at hand, it seems probable that primates
arose during the Late Cretaceous, about 70 million years ago, and are related by
a common ancestor to insectivores and bats. The first known primates belong to
328 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

an extinct suborder called the Plesiadapiformes. They appear in the North Amer- Eocene, Africa
ican fossil record in the very early Tertiary shortly after 65 million years ago. forested , so a '
These were small animals, the size of squirrels and house cats. Plesiadapiformes prising. The N'
were tree-dwelling and had skeletal features that are similar to those of some been sundered
modern primates, but their dentition was more rodentlike, except for their molars during the Lat'
which were like those of the primates. The Plesiadapiformes hunted small insects mates from mil
and ate seeds. Even at this stage, the primates appeared to have had somewhat What wa:
larger brains relative to their body mass than other mammals with similar niches, with a long tail
such as tree shrews. During the Pliocene, the Plesiadapiformes became very big as a moden
abundant in North America and Europe. These two continents were still joined in sockets that ar
the North Atlantic sector at this time. The climate of the earth was generally visual cortex r1
warm and moist, and this resulted in an extensive and a continuous forest habitat. the brain relat
Despite their abundance in the early Paleocene, by 55 million years ago, the Ple- Aegyptopithecz
siadapiformes had gone extinct and we can trace no living primate group to them. apes than it wa
By the Eocene, about 55 to 38 million years ago, two groups of primates had The Late
become abundant in North America and Europe. One group resembled the mod- million years c:
ern tarsiers, and the other group resembled the modern lorises and lemurs. These Extensive fore!
two fossil groups are called the Omomyids and the Adapids, respectively, and are and it is thoug
likely ancestral to modern tarsiers of the Haplorhini primate suborder and lemurs walk) tree dwe
of the Strepsirhini primate suborder. During this period, primates were present in to southeasterr
Africa, Eurasia, and North America. As a result of plate tectonics, the African con- Some, such as
tinent had moved for enough northward so that primates could disperse there similar to moe
from Eurasia. A continuation of generally warm and moist conditions promoted India, were cor
extensive forest cover that suited the early primates. Toward the end of the is difficult to t1
Eocene, there was a great decrease in Stepsihini primates. The early lemurs went potential candi
extinct in North America, and no later primates are known from that continent extremely wid1
until the arrival of humans. Primates continued to live in Central and South Amer- largely from fc
ica. Platyrrhinii monkeys exist in South and Central America today, and fossil humans. Howe
deposits show that they have been there since the Oligocene, some 30 million orangutans. Wt
years ago. The very early history of the South and Central American Platyrrhinii simply relative:
primates is unclear. It is possible that they dispersed from Africa to South Amer- return to consi1
ica during the Early Eocene. The global decline in Stepsihini primates was likely The abun
due to increasing competition from newly evolving rodents and primates of the higher than to
Catarrhinii infraorder, which are ancestors of modern monkeys, apes, and humans. their large nun
The best fossil evidence of the early Catarrhinii primates comes from the mates suffered
Fayum Depression in Egypt. These Oligocene deposits date from about 31 mil- reasons were ir
lion years and are extremely rich in primate fossils. Included in the fossil fauna is pressure from r
a small, monkeylike primate of the genus Parapithecus that appears ancestral to ronment. In ge1
the Old World monkeys. The name Parapithecus can be translated from its Greek Oligocene. In t
roots as meaning "close to ape." In addition, the deposits contain fossils of a more centrated, it ar
apelike primate of the genus Aegyptopithecus ("Egyptian ape"). Many believe savanna were c
that Aegyptopithecus is a very likely ancestor, or at least a relative, to the modern evolutionary ac
apes and humans. Paleontologists have also found some earlier primates that are to the Ramapi
potential ancestors of Aegyptopithecus, modern apes, and humans. These include open ground.
the small primates of the genus Catopithecus, which are found in the early por- association wit
tion of the Fayum deposits. In addition, fossils have been recovered of an Early and savanna e
Eocene primate that lived in northwestern Africa some 50 million years ago and ithecines was l
possessed teeth that are very much like those of modern apes. Finally, jaw frag- them to be bip
ments of a potential ancestor to Aegyptopithecus have been found in deposits in a gap exists in
Myanmar (Burma). This Asian primate is called Amphipithecus. During the ago. This gap m

appear in the North Amer-
lfter 65 million years ago.
mse cats. Plesiadapiformes
:! similar to those of some
ike, except for their molars
ormes hunted small insect>
:ed to have had somewha:
tmmals with similar niche
adapiformes became very
atinents were still joined ir.
>f the earth was genera U~
t continuous forest habita::..
million years ago, the Pie-
ing primate group to the
wo groups of primates ha:":
group resembled the moc-
1 !arises and lemurs. These
apids, respectively, and are
mate suborder and !em u~
I, primates were present ir.
tectonics, the African cor.-
lates could disperse there
1oist conditions promot e~
;. Toward the end of tl:e
tes. The early lemurs wee:
nown from that continec
t Central and South Am e::--
America today, and fos~
ligocene, some 30 millio=
tral American Platyrrhim
>m Africa to South Ame::--
>sihini primates was like!_
ients and primates of the
onkeys, apes, and humans.
primates comes from tte
; date from about 31 rni:-
uded in the fossil faun a ~
that appears ancestral t.:
translated from its Greei:
s contain fossils of a m o~ e
tian ape"). Many belie\·::
: a relative, to the mode. _
: earlier primates that a:-::
1d humans. These induce
e found in the early po::--
en recovered of an E arl·
50 million years ago an.:
rn apes. Finally, jaw frag-
Jeen found in deposits i.::
rphipithecus. During the
EARLY PRIMATES 329
Eocene, Africa, Europe, and Asia had become closely connected and were largely
forested, so a widespread distribution of primates in Africa and Asia is not sur-
prising. The North Atlantic corridor between Europe and North America had
been sundered at this point. Cooling temperatures, particularly at high latitudes
during the Late Oligocene and Eocene, may have kept these newly evolved pri-
mates from migrating to the New World via Beringia.
What was Aegyptopithecus like? These small primates were tree dwellers,
with a long tail and prehensile hands and feet. In terms of size, they were about as
big as a modern house cat. The skulls of Aegyptopithecus had protected bony eye
sockets that are typical of modern apes and humans. Its snout was reduced, the
visual cortex region of the brain was enlarged, and the olfactory lobes (area of
the brain related to the sense of smell) were reduced. Finally, the dentition of
Aegyptopithecus was similar to that of apes. It was clearly much closer to modern
apes than it was to !arises and tarsiers.
The Late Oligocene and the first half of the Miocene, from about 23 to 14
million years ago, saw a great increase in ape diversity and geographic range.
Extensive forest cover still existed across Africa and most of Eurasia at this time,
and it is thought that these apes were all quadrupedal (using all four limbs to
walk) tree dwellers. Caterrhinii apes were present from Africa to western Europe
to southeastern Asia. Over 20 different genera are known from the fossil record.
Some, such as Pliopithecus, found in western Europe, were relatively small and
similar to modern gibbons. Others, such as Gigantopithecus from China and
India, were considerably larger than modern gorillas. With such an abundance, it
is difficult to trace which Miocene primate might be ancestral to humans. One
potential candidate is a group called the Ramapithecines. These primates were
extremely widespread in the Miocene. The Ramapithecines were first known
largely from fossil teeth and jaws, which are very similar to those of apes and
humans. However, Ramapithecine skulls suggest a closer affinity to modern
orangutans. Which of the Miocene primates is ancestral to humans and which are
simply relatives of our primate ancestors remain controversial questions. We will
return to consider the Ramapithecines in this context in a moment.
The abundance and diversity of the Miocene apes seem to have been much
higher than today. What happened to this array of different species? Despite
their large numbers, large geographic range, and great diversity, the apelike pri-
mates suffered large declines and extinctions in the Late Miocene. Among the
reasons were increasing competition from monkeys in forested areas, increasing
pressure from new carnivore predators, particularly cats, and changes in the envi-
ronment. In general, temperatures and precipitation had been in decline since the
Oligocene. In the temperate and tropical regions where the primates were con-
centrated, it appears that forest cover had begun to decline and large areas of
savanna were developing. Some primate species survived these changes through
evolutionary adaptations to more open ground environments. This brings us back
to the Ramapithecines. They appear to have lived at the edges of forests near
open ground. Fossils of Late Miocene Ramapithecines are generally found in
association with the fossils of animals typical of grassland, brushland, woodland,
and savanna environments. Interestingly, the center of gravity for the Ramap-
ithecines was lower than for modern apes, and this would have made it easier for
them to be bipedal, that is, able to walk on two legs as humans do. Unfortunately,
a gap exists in the fossil record, particularly in Africa, after about 14 million years
ago. This gap makes it difficult to trace the ties, if any do indeed exist, between the
330 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

Ramapithecines and later primates that are more clearly related to mode ~
humans. To find the next candidate for ancestor to modern humans, we have - Miocene
move forward in time about 9 million years.

THE HOMINIDS: AUSTRALOPITHECUS

The term hominid refers to humans and extinct bipedal primates that are direct~_
ancestral or closely related to humans. Some scientists include Ramapithecinpo;:
among the hominids, while other scientists do not. The first widely accept ~
hominids are of the genus Australopithecus, which appeared in Africa about :
million years ago. Before discussing the Australopithecines in detail, let's cor.-
sider the events that may have led up to the development and divergence 0:
bipedal hominids from other primates. Because of the sparse nature of the pr:- FIGURE 11.1
mate fossil record between 14 million and 5 million years ago, our consideratio- possible distribut
must be somewhat speculative.
Of all the modern primates, humans are the only species that is trul:
bipedal. Monkeys and apes can walk on two legs and do so for short distances. - The fragn
general, however, even closely related species such as chimps prefer to m oY ~ vegetation covE
along the ground using both their arms and legs for propulsion. In gorillas an"' arboreal primal
chimps, this takes the form of knuckle walking. Humans do not knuckle walk Ramapithecine
Our bodies are clearly adapted for movement on two feet alone. Alone among a: Initially, these ~
other primates, and indeed all other mammals, humans are bipedal. Although v•e take advantage
take human bipedalism for granted as our natural condition, it is not so easy tc tion in the tree~
explain how this characteristic developed. Primates clearly have their origins · sil fauna and fi<
the trees. As discussed, many facets of human morphology and physiology can b~ not completely
directly attributed to adaptations to arboreal life. In trees, quadrapedal move- Later hominids
ment using prehensile feet and hands is clearly more efficient (not to mentio:;: During the Pli•
safer!) than bipedalism. Like humans, some other modern primates, such <E drier and more
chimps, gorillas, and baboons, are essentially ground-dwelling animals, yet th e ~ climate and veg
have remained quadrapedal. So, the questions we must answer are, what made What wm
our primate ancestors, which were above all creatures of the forest, climb dmn.;. vigorously deb;
from the trees and take up life on the ground, and why did they become bipedal'? ary adaptation
Let's consider the development and implications of bipedalism. forest's edge, b
The movement of some ape species away from a strictly arboreal existence distances than
may have begun by the Miocene, as is evident in certain Ramapithecines. and this would
important factor that may have contributed to this development was environmen- A bipedal stan'
tal change. A trend toward cooler and drier conditions continued through the Lat difficult to tran
Tertiary. This led to a fragmentation of the once continuous African and southern walk. Being abl
Eurasian forest cover. This trend of climatic and vegetation change is coinciden would be a dist
with the evolution of grasses, and associated browsing fauna and predators. Late tocene Africa.
Tertiary deforestation was particularly extensive in Africa where the formerly climbers, and "
extensive forests in the north were replaced by deserts, thorn scrub, and savanna food. Bipedal n
during the Late Miocene, Pliocene, and Pleistocene (Fig. 11.1). Not only was the Chimp infants
world of the Old World primates becoming more arid, it was also becoming more a significant fa
variable in climate and vegetation distribution. The Milankovitch cycles of insola- arms of the ind
tion variability (see Chapter 7) were becoming increasingly important forces in visibility when
the climate system during the Late Tertiary. Increasing concentrations of fine ter- other significan
restrial dust in deep-sea cores off the coast of Africa provide direct evidence of the higher bushes
drying of the continent, loss of forest cover, and increasing variability of climate. cation of safe p

clearly related to moder::
wdern humans, we have 1c
al primates that are direcr;:
;ts include Ramapithecine-s
The first widely accep te.:
tppeared in Africa abom :
hecines in detail, let's co:::-
opment and divergence
te sparse nature of the pr:-
ears ago, our consider ar i~-
only species that is trU:_
jo so for short distances. ·-
as chimps prefer to mo· ::
propulsion. In gorillas ar;:
nans do not knuckle w ··
feet alone. Alone among c.
s are bipedal. Although \ 1 ::
ndition, it is not so eas~ :
!early have their origi ns I::
logy and physiology can c'=
trees, quadrapedal mo\·;;:.
~ efficient (not to mentic-
modern primates, such ~
dwelling animals, yet tb.:
1st answer are, what mac::
; of the forest , climb dov.-
, did they become bipeda.
pedalism.
1 strictly arboreal existec.:::
~rtain Ramapithecines. -~
=lopment was environme:::-
:ontinued through the l z:::
uous African and soutbe;-
tation change is coincide::.
fauna and predators. LG.~::
\frica where the forme:-.
;, thorn scrub, and savan.::.
Ig. 11.1). Not only was t:.::
it was also becoming mo:-::
lankovitch cycles of insol.:
tsingly important forces
concentrations of fin e t.c.:--
,vide direct evidence of :2:.::
.ing variability of climate
THE HOMINIDS: AUSTRALOPITHECUS 331
Miocene Glacial Interglacial - Present
Savanna
and
woodland
FIGURE 11.1 Modern distribution of African rainforest, possible glacial distribution, and
possible distribution during the Miocene (after Foley, 1987).
The fragmentation in forests, the decrease in total forest cover, and shifting
vegetation cover caused by Milankovitch cycles posed a severe challenge to the
arboreal primates and led to many extinctions. Some groups, however, such as the
Ramapithecines were able to adapt to the new environment through evolution.
Initially, these species would have favored forest margin sites where they could
take advantage of savanna and grassland resources, yet also feed and seek protec-
tion in the trees. Some of the earliest Australopithecine fossils are found with fos-
sil fauna and flora indicative of forest and woodland. This suggests a life that was
not completely based on the savanna but perhaps on the forest-savanna ecotone.
Later hominids are more often found with fossils indicating savanna conditions.
During the Pliocene and early Pleistocene, climate was becoming increasingly
drier and more variable, and savanna was expanding at the expense of forest. So,
climate and vegetation may have prompted our ancestors to leave the trees.
What would prompt the development of bipedalism? This topic has been
vigorously debated. There were probably many forces promoting the evolution-
ary adaptation of bipedal movement. On the open ground of the savanna or the
forest's edge, bipedalism is a more energy-efficient form of movement over long
distances than is quadrupedalism. More ground can be crossed with less energy,
and this would be important in moving between widely scattered patches of trees.
A bipedal stance allows the individual to pick up and carry food efficiently. It is
difficult to transport much material if you rely on both your legs and forelimbs to
walk. Being able to transport food off of the savanna to the safety of trees to eat
would be a distinct advantage on the predator-filled plains of Pliocene and Pleis-
tocene Africa. The fossil evidence tells us that Australopithecines were good
climbers, and we can assume they used trees for protection and for foraging for
food. Bipedal movement also makes it easier to transport infants from site to site.
Chimp infants must cling to their mothers when being transported, and falling is
a significant factor in chimp infant mortality. Bipedalism places the head and
arms of the individual relatively high above the landscape. This allows for better
visibility when searching for food and water on the savanna. Being tall presents
other significant advantages. It facilitates the foraging of leaves and fruits from
higher bushes and small trees. It assists in the spotting of predators and identifi-
cation of safe places to run to, such as climbable trees.
332 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

Finally, the Ramapithecines and Australopithecines were generalists _ In 1959 ar

terms of diet and had teeth, like ours, that allowed the biting, scraping, and che - the English pal1
ing of a wide variety of foodstuffs. Unlike baboons, which retain very large a;:~ Gorge region o
dangerous canine teeth, the Ramapithecine and Australopithecine canines we-:: line, and terrest
like ours, very much reduced. Therefore, unlike the modern baboons, they had - - The area is extr
natural defensive weapons on the savanna. What a bipedal individual can do :: many good geo
defend itself, however, is use its free hands for throwing objects and swing~ of the Olduvai
sticks. From the use of objects, rather than teeth, for defense, we can see how ti::: modern geochr
use of tools for other tasks would be a logical development. Once hominids car:::: for early homir
to depend on and develop tools, evolution would favor dexterity of the hands a;:~ son Richard, w<
increasing mental capability required to devise, construct, and apply tools. Con_:_ covering not 01
quently, the development of a highly generalist feeding strategy and dentitio:_ that time, other
which was made necessary by climatic change and the reduction in forests, als - and Kenya, and
contributed to the evolution of bipedalism. Bipedalism was essential for the fu-:- ton of an Aust1
ther evolutionary development that resulted in modern humans. Even if for ~ pologist Donal<
fragmentation and savanna development did not initiate the movement .:
bipedalism, movement on two feet allowed human ancestors to move onto an.:
take advantage of the savanna. As a parting thought on the movement of hum -
ancestors from the trees and onto two feet, we might contemplate the words c·
the eminent physical anthropologist William Howells (1993 p.73); "Had o --
ancestors gone on in the forests, we may suppose that nothing would have ha ~
pened: they would not have turned into bipeds; we would not be here."
Now that we have briefly examined the potential causes and implications o:
bipedalism, let's return to our consideration of the first widely acknowledge.:.
hominid, Australopithecus. The first person to discover and recognize the impo-:-
tance of Australopithecine fossils was a young medical doctor named Rayma n.:
Dart. Dart was originally from Australia and had studied medicine in Britain. t:
1923 he went to South Africa to take up a professorship in anatomy at Witwate-:-
srand University. At that time, the mining of limestone from ancient ca\·::
deposits was producing a number of interesting mammal and reptile fossils. t:
1924, he was given a piece of stone with a skull embedded inside. After muc::.
laborious picking and cleaning, using among other things a pair of knitting ne -
dles, Dart recovered the majority of an immature primate's skull, including ana -
ural limestone cast of the brain. The skull was apelike but possessed molars th2.:
were similar to those of humans. The brain, though small, had a number of fea-
tures that were more human than ape. Intriguingly, the manner in which the spine
attached to the skull suggested a bipedal stance. Dart concluded that the fossi.
belonged to a very intermediary species between apes and humans. In 1925 he
published his findings, naming the newly discovered primate Australopithecus
africanus (southern African ape).
Dart's discovery was not received with much enthusiasm or interest. Indee
it took Dart and his colleague, a Scottish medical doctor named Robert Broom.
many years of hard work excavating South African limestone caves to make their
case that Australopithecus africanus was bipedal and potentially ancestral to mod-
ern humans. By 1950, Dart and Broom had found enough skulls indicative of a
upright position and a bipedal pelvis that anthropologists accepted their conclu-
sions. Australopithecus africanus was firmly placed with the hominids. The work FIGURE 1 1.2
from South Africa indicated that Australopithecus africanus was bipedal and was Afar region of Eth
intermediate in skull and brain features between modern apes and humans. How- found. She walke
ever, exactly how old they were was uncertain. 20 years old at th1
 THE HOMINIDS: AUSTRALOPITHECUS 333

~
ecines were generalists
biting, scraping, and che\1.-
= In 1959 another chapter in the Australopithecine saga opened. In that year,
the English paleontologists Louis and Mary Leakey began digging in the Olduvai
.vhich retain very large arc: Gorge region of east Africa. The gorge contains the fine deposits of lake, shore-
ralopithecine canines we r:: line, and terrestrial environments that were laid down over the past 2 million years.
odern baboons, they had r: .:: The area is extremely arid today and bisected by gullies and valleys, providing for
'i pedal individual can do :.:: many good geologic exposures. In addition, volcanic deposits occur in many parts
•wing objects and swingi;:~ of the Olduvai rock record, and this allows for the dating of the fossil finds using
lefense, we can see how t.:::: modern geochronological techniques. It is one of the richest areas in the world
ment. Once hominids car::.:: for early hominid fossils. For many years, the Leakeys, and more recently their
r dexterity of the hands a;:: son Richard, worked the deposits at Olduvai and other regions of east Africa dis-
uct, and apply tools. Cons::- covering not only further Australopithecines but other hominids as well. Since
ing strategy and dentitio:.. that time, others have found Australopithecines in a number of sites in Tanzania
1e reduction in forests, als.: and Kenya, and as far north as the Afar region of Ethiopia. A 40% complete skele-
m was essential for the h::- ton of an Australopithecus female was found by the American physical anthro-
ern humans. Even if fore:; pologist Donald Johanson and his team in Ethiopia (Fig. 11.2). This remarkable
initiate the movemen t : ·
ncestors to move onto a;::
m the movement of hu m~
t contemplate the words ~
:lls (1993 p.73); "Had o._
tt nothing would have ha;-
mld not be here."
l causes and implications -
first widely acknowled§:-:
..
_,_
- ---..,. .
r and recognize the impc:--
:~1 doctor named Raymo:::.:
lied medicine in Brita in .~ ­
ip in anatomy at Witwac:--
=stone from ancient c~' =
---
1 .
.,._.
__,. a.
,..,.......__ .• _.,
nmal and reptile fossils. : .:.:
Jedded inside. After rn"- -
~;~\
ings a pair of knitting ne-::-
tate's skull, including a 02 _-
• but possessed molars t::._ ,I I
nall, had a number of f.:_ J1
manner in which the sp:.::.-
t concluded that the fo--
~s and humans. In 1925 .:;_
primate Australopith

msiasm or interest. Indee-..:.

tor named Robert Broo::..
estone caves to make til::
)tentially ancestral to me:
>ugh skulls indicative of .=-
;ists accepted their concL
•1.
th the hominids. The wc:-
FIGURE 11.2 Lucy-the fossi I skeleton of a female Australopithecus afarensis from the
:anus was bipedal and
Afar region of Ethiopia. Lucy is one of the most complete early hominid skeletons ever
rn apes and humans. He found. She walked upright and stood about 110 to 120 em (about 4ft) tall. She was perhaps
20 years old at the time of death .
334 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

fossil has been named "Lucy," after the Beatles song "Lucy in the Sky with Dia- by Australopith
monds," which the anthropologists were listening to during the excavation. In about 2.5 millie
1975 Johanson and his colleagues found the remains of a group of 13 Australop- much heavier b
ithecines that apparently died together in some sort of natural disaster such as a etable matter, r
flood or slope failure. This group was called "The First Family_" Perhaps the mos outside of Afric
amazing find has been a set of bipedal footprints that were most likely made by distribution to t
two Australopithecus walking side by side. The tracks were discovered by Mary What wer
Leakey's team in 1976 at Laetoli in Tanzania. The prints were made by hominids small creatures,
and other animals, crossing a landscape that was covered by very freshly deposited kg. Males were ~
volcanic ash. feet, looked mu
Based on the anatomy and dating of the African finds, we know that Aus- skulls were mu•
tralopithecines first appeared between 4 million and 3.8 million years ago and ranging from at
disappeared by 1 million years ago (Fig. 11.3). How many species of Australop- ern human brair
ithecines there were remains a matter of debate. It is possible that between 2.: mass was also rr
and 3 million years ago at least three distinctive species of Australopithecines brains indicate t
existed in Africa. Many authorities conclude that one of the earliest species, Aus- ing and languag
tralopithecus afarensis, persisted until about 3 million years ago when at least two tralopithecus aj
distinctive clades arose. One is represented by Australopithecus africanus intelligence. Bn
(referred to as the gracile line), which were somewhat lightly built and had denti- incipient devel<
tion consistent with an omnivore diet, or a diet with much soft vegetable matter. thought and spe
This species persisted until about 2 million years ago. The other lineage is typified closer to that of
of individuals fo
in social groups.
Ocean
1)1BQ Soii813 C

4
Australopithecus
afarensis
and others
Glacial Interglacial Grassland Woodland

FIGURE 11.3 A simplified history of the hominids. Many authorities recognize additiona l
species of both Australopithecus and early Homo. The progressive cooling of the planet and
the development of glacial interglacial cycles, as evident in cores from the world's oceans,
and progressive decrease of woodland in Africa, as evident from the analysis of fossil soils,
are also indicated. There appears to be a linkage between global cooling and drying, FIGURE 11.4 T
increasing climatic variability, expansion of the African savanna, and hominid evolution Homo sapiens nea
(partly after Haviland, 1989; deMenocal, 1995). beyond Africa .

"Lucy in the Sky with Dit:-
> during the excavation. I=
of a group of 13 Austral o;:---
)f natural disaster such as ::.
t Family." Perhaps the mo,-
t were most likely made c:
s were discovered by Ma.:-::
nts were made by homini"'
ed by very freshly deposite.:
m finds, we know that Ae.>-
l 3.8 million years ago a:_.:
many species of Australo;--
s possible that between : .:
ecies of Australopitheci£:::-.
of the earliest species, A : --
years ago when at least
4ustralopithecus african-
lightly built and had der. _-
nuch soft vegetable m ar: ::~
llle other lineage is typit.:-:
~ Australopithecus
~ afarensis
and others
Jthorities recognize addi t ic - ;.
ssive cooling ofthe planet c.-:
res from the world's ocea ns
)m the analysis of fossil so ~
)al cooling and drying,
1a, and hominid evolutio n
THE HOMINIDS: AUSTRALOPITHECUS 335
by Australopithecus robustus (referred to as the robust line), which lived from
about 2.5 million years ago until approximately 1 million years ago. They were
much heavier built and had jaw muscles and teeth suggesting a diet of coarse veg-
etable matter, nuts, and tubers. No Australopithecine remains have been found
outside of Africa (Fig.11.4), and it is likely that they were restricted in geographic
distribution to that continent.
What were the Australopithecines like in form and behavior? They were
small creatures, with an upright height of about 1 to 1.5 m and a weight of 30 to 60
kg. Males were significantly larger than females. Their lower bodies, including their
feet, looked much more like those of modern humans than apes. However, their
skulls were much more apelike. Australopithecines had relatively small brains,
ranging from about 380 to 485 cu em, which is about one-third the size of a mod-
ern human brain, which averages about 1300 cu em. The ratio of brain size to body
mass was also much less than that of modern humans. The general shapes of their
brains indicate that they would not have had the same capacity for abstract think-
ing and language as modern humans. It is thought by anthropologists that Aus-
tralopithecus africanus was the most advanced form in terms of brains and
intelligence. Brain casts and details from the interior of the cranium show the
incipient development of anatomical features associated with both abstract
thought and speech. The mental capability of the Australopithecines was probably
closer to that of modern chimps and gorillas than modern humans. The collection
of individuals found at the First Family site suggests that Australopithecines lived
in social groups. Such groups are typical of chimps and baboons, as well as humans.
~
------ ....
oR a ~~~ r
u ,'
__ , ... ""' ()
··'
"' ()
.6. Australopithecus
!) H. habilis and/or
H. erectus
'
':> H. erecus
- - H. sapiens
neanderthalensis
'V
FIGURE 11.4 The distribution of Australopithecine, Homo habilis, Homo erectus and
Homo sapiens neanderthalensis finds. It does not appear that Australopithecines ranged
beyond Africa.
336 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

Interestingly, there is no evidence of tool making or the use of fire by Australop- 2.5 to 1.5 millio
ithecus. It is suspected that they may have made use of sticks and rocks as weapon;; pithecus robusu
or to secure food, but they do not appear to have worked rocks into blades o~ Aside fro
scrapers. Paleoenvironmental evidence suggests that the Australopithecine5 habilis represen
lived in the vicinity of the forest savanna border and near groves of trees withic ligence. Associa
the savanna. Recent biochemical analysis of 3-million-year-old Australop- evidence for th(
ithicene bones from South Africa indicates that the Australopithecus africanw in Ethiopia are
diet included fruits and nuts typical of the forest, but also grasses typical of the 2 million years ,
savanna. Australopithecine skeletons suggest that they were better climber5 Homo habilis a
than modern humans and likely used forest trees for food sources and to escape been preserved.
predators. the 1-million-ye
Are the Australopithecines ancestral to modern humans? This topic h ~ tive tools is call(
been hotly debated for over 50 years. Some scientists, such as Louis Leakey, argu- appearance of t
emphatically that they are not. These scientists believe that Australopithecus ~ Lower Paleolitl
related to our genus but had too many apelike features at a relatively late date tc round cobbles t
serve as an ancestor. In this case, the Australopithecines are just one of man ~ that were used
mammal genera that have gone extinct in the past and left no living lineage o:: Oldowan tools J
descendants. Many other scientists believe that the gracile line of Australopithe- wild pigs and e'
cus africanus is a very likely ancestor to the genus Homo. If we assume for c. that tools were 1
moment that the generalist gracile Australopithecines evolved into the genus There is no dou
Homo and thus persisted in this manner, what happened to the robust line? AI: There is even so
agree that the robust line of Australopithecus robustus is unlikely to have beec What is unclear
ancestral to any modern primates or humans. The robust line was a highly spe- scavenging of ki
cialized primate in terms of diet. Their dependence on a low-protein diet specifi - known to hunt <
cally consisting of coarse vegetable matter, nuts, and tubers may have made then:. both hunted an(
unable to persist through increasingly pronounced environmental shifts that typ- terms of teeth o
ified the later portion of the Pleistocene. In addition, the robust Australop- is no evidence tl
ithecines may well have been driven to extinction by competition and predatiou. eaten raw.
from its more generalist hominid cousins of the genus Homo. As simple
resources for ht
high-protein foe
THE HOMINIDS: EARLY HOMO tain the individ1
interaction, and
The oldest fossils of our own genus, Homo, are over 2.5 million years old and communication.
come from east Africa. The first of these remains were found by the Leakeys at discarded tools,
Olduvai Gorge in 1960 and were formally named Homo habilis (handy man) in than individuals
1964. Since that time, fossils of Homo habilis have been found in other portions ing meat caches
of east Africa and southern Africa (Fig. 11.4). The youngest Homo habilis fossils tion and comm
are about 1.5 million years old. appears unlikel~
Homo habilis was a small hominid with a bipedal skeleton very much like modern speech.
the gracile Australopithecines. The big difference between the Australopithecines Suppleme1
and Homo habilis is the skull. In the new genus, the brain increased to 600 to 80 habilis to spen1
cu em. To accommodate this larger brain, the skull was higher and the face more hominids and ar
vertical in appearance. The cheek bones, however, were still relatively broad com- the spread of h<
pared to those of modern humans. In many ways, the anatomical features of another species
Homo habilis appear to make it a good candidate for an evolutionary intermedi· For the ne
ary between the gracile Australopithecines and later species of Homo. There travel to Trinil, J
appears to be much variability within the Homo habilis fossil skulls, and some sci- ical officer in tl
entists recognize these as representing more than one species. Although, it anatomy in Holl
remains debated whether more than one species of Homo lived during the period tion of humans.]

the use of fire by Australo;:-
, sticks and rocks as weapo::.::
10rked rocks into blades e--
hat the Australopithecine-::
near groves of trees with:.::
nillion-year-old Australo;-
Australopithecus africm::.
.t also grasses typical of L:::.::
they were better climbe_
food sources and to esca;o::
rn humans? This topic b..
such as Louis Leakey, arg-_::
~ve that Australopithecus
=s at a relatively late date:
cines are just one of me..::;
md left no living lineage
racile line of Australop i:~
Homo. If we assume fo:-
les evolved into the ge:::
med to the robust line')_:..
us is unlikely to have be_
1bust line was a highly s;-::
n a low-protein diet spe.:-..:.
ubers may have made th.:-.:-
vironmental shifts that r-::
ion, the robust Austra~ c-­
competition and preda:.
:Homo.
r 2.5 million years old :.:;._
re found by the Leak e~-~
mo habilis (handy rna
:en found in other p on~ c­
mgest Homo habilis fos
al skeleton very much ··
'een the Australopithec-...::;
·ain increased to 600 to ~·
s higher and the face c::: .::'-
e still relatively broad
he anatomical feature>
an evolutionary interm.:-.:.
r species of Homo. T.:::---
s fossil skulls, and some ;;.:::
one species. Althoug.t
•mo lived during the pe:-
THE HOMINIDS: EARLY HOMO 337
2.5 to 1.5 million years ago, it is known that Australopithecines, such as Australo-
pithecus robustus, were also living in the same portions of Africa at this time.
Aside from the larger brain, there is good physical evidence that Homo
habilis represents a major step toward the development of modern human intel-
ligence. Associated with some of the Homo habilis sites is the first indisputable
evidence for the manufacture and use of specialized tools. Tools found at Hadar
in Ethiopia are 2.5 million years old, and many sites have revealed tools that are
2 million years old. The tools that we know of are made of stone. It is likely that
Homo habilis also made tools from wood, bones, and sinew, but these have not
been preserved. The stone tools that have been found do not change much during
the 1-million-year period of Homo habilis's existence. This collection of distinc-
tive tools is called the Oldowan Tradition in recognition of the Olduvai finds. The
appearance of these tools marks the start of the archaeological period called the
Lower Paleolithic (old stone age). The Oldowan tool assemblage includes large
round cobbles that were likely used as hammers, cobbles with sharpened edges
that were used as hand axes, and sharp stone flakes that were used as blades.
Oldowan tools have been found in association with the bones of animals such as
wild pigs and even elephants. The bones of these mammals show clear evidence
that tools were used to break them open for marrow or to scrape meat off bones.
There is no doubt that Homo habilis used meat as an important part of its diet.
There is even some evidence that they may have preyed upon Australopithecines.
What is unclear is the degree to which meat was obtained by hunting versus the
scavenging of kills made by predators such as lions. Chimps in the wild have been
known to hunt other mammals, such as monkeys, so it is likely that Homo habilis
both hunted and scavenged meat. Without tools, humans are poorly equipped in
terms of teeth or nails to hunt, skin, and remove meat form other animals. There
is no evidence that Homo habilis used fire, and thus the meat they consumed was
eaten raw.
As simple as they were, the Oldowan tools greatly increased the food
resources for humans, particularly in terms of high-protein meat. The increase in
high-protein food decreased the foraging time needed to find enough food to sus-
tain the individual or group. This allowed more time for tool making and social
interaction, and may have contributed to the further evolution of intelligence and
communication. Some sites contain large assemblages of butchered animals and
discarded tools, suggesting the butchering activities by groups of hominids rather
than individuals. Group activity in hunting, scavenging, butchering, and protect-
ing meat caches may have promoted the further development of social interac-
tion and communication. Although they clearly lived and worked together, it
appears unlikely that Homo habilis had the physiological or mental capacity for
modern speech.
Supplementing vegetable matter with meat would have also allowed Homo
habilis to spend more time on the savanna and travel further than earlier
hominids and apes. However, the true embrace of a savanna-based existence and
the spread of hominids beyond Africa would have to await the evolution of yet
another species of Homo.
For the next step in our study of human evolution, we will leave Africa and
travel to Trinil, Java, in the year 1891. In that year, Eugence Dubois, a young med-
ical officer in the Dutch Army, made a startling discovery. Dubois had studied
anatomy in Holland and had become intrigued with Darwin's ideas on the evolu-
tion of humans. He was also familiar with the works of Alfred Russel Wallace, who
338 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

as early as 1855 had suggested that humans and apes might share a common ances· Leakey) and A
tor, and the fossils of that ancestor might be found in Southeast Asia. Dubois con· the boy had die
eluded that Southeast Asia was indeed a likely spot to find fossil evidence of early iment; this pre~
human ancestry. He was so convinced of his conclusions that he joined the Dutch reached adulthc
Army in order to get stationed in the colony of the Dutch East Indies, which today em tall and wei
forms modern Indonesia. At first, Dubois had no official support for his paleonto· erectus are very
logical endeavors, but after publishing several speculative articles, he was able to long and relativ
secure full authorization, along with two engineers and a team of convict laborers. ence in the size
For two years, Dubois searched the island of Sumatra in vain for early hominids. Their
hominid fossils. He then moved his expedition to Java, where he discovered a had a very prorr
skull cap and a femur that appeared to share features of both humans and ape foreheads and r
In 1894, he published his account and suggested that this was indeed an early what modern n
hominid. Dubois called the fossil species Pithecanthropus erectus (upright ape- preted as an ad
man). The fossils that Dubois recovered were popularly referred to as "Java stature and nas<:
Man." Both scientists and the public greeted Dubois' conclusions with consider· tionary adaptati
able skepticism. We should remember that his findings were announced about 3C Homo erectus n
years before Dart's early hominid discoveries in South Africa and almost 7 of earlier homo
years before the Leakeys' work in east Africa. Dubois was shattered by the poor and throat of H .
reception of his discovery, and for a long period of time he simply kept the fossil s highly controver
hidden beneath the floorboards of his home in Holland. We now know tha The geogn
Dubois was correct; his fossils were indeed from an early hominid that we classify and Asia is an t
today as Homo erectus. We also know that Homo erectus is younger than Homo from the Atlant
habilis and older than Homo sapiens. and to the Pacifi
As the twentieth century progressed, more Homo erectus fossils were foun d few other terres1
in Java and in China. The most famous of these was from Zhoukoudian near Bei· It is likely that tl
jing. These fossils, which came to be known as "Peking Man" (Peking is an older generated in par
spelling for the capital city of China, Beijing), were unfortunately lost during tions in Africa aJ
World War II. During the last 50 years, fossils of Homo erectus have also been while the variati•
found in several parts of western Europe, the Republic of Georgia, and in south pronounced. Inc
and east Africa (Fig. 11.4). There is much temporal and geographic variability in mated the adapt
details of the skulls and other bones of Homo erectus. As with other early addition, the mo
hominids, there is debate on the taxonomy of these finds, and some paleontolo· may have been
gists recognize different species or subspecies of Homo during this period. Some changes in clima
authorities classify the earliest Homo erectus-type fossils as a separate species. populations shif1
Homo ergaster. We will stick with a conservative interpretation that all of these resources shifted
finds can simply be referred to as Homo erectus. The age of Homo erectus fossil s The geogra
spans the period of about 1.7 million to approximately 300,000 years ago. The itated by increas
appearance of Homo erectus coincides with increasingly cold, dry, and variable Homo erectus an
climatic conditions associated with the Pleistocene and its glacial-interglacial and varied chop
cycles. This had led some paleontologists to suggest a link between climatic cleavers. Similar
change and the evolution of Homo erectus. There is some evidence that Homo are found from
erectus may have persisted in parts of Asia until less than 100,000 years ago. The choice of stones
dates from the different localities show that the oldest Homo erectus fossils come hearths have bee
from Africa and Georgia. Most of those from Europe and China date from about was being used b~
700,000 to 500,000 years ago, although some fragmentary evidence suggests ago. In addition,
Homo erectus may have spread to Eurasia before 900,000 years ago. In any case. vived in relative!'
with Homo erectus we have the first evidence of hominids leaving Africa and fire. Fire not only
spreading extensively through Asia and Europe. greatly enhanced
Homo erectus were tall and strong individuals. The most complete skeleton tiona! nutrients a
is that of an adolescent boy found by Richard Leakey (the son of Louis and Mary contamination in

ight share a common ances-
outheast Asia. Dubois cor..-
find fossil evidence of ear~'
1s that he joined the Dutc-::
ch East Indies, which to de:.
al support for his paleontC'-
tive articles, he was able :-
a team of convict laborers.
Sumatra in vain for ear:
va, where he discovered ::.
. of both humans and a
t this was indeed an ear:
opus erectus (upright ap:e-
tlarly referred to as "Ja\ _
conclusions with conside::--
; were announced abou t :
·uth Africa and almost -
was shattered by the pac-
e he simply kept the foss:....
!land. We now know th:-
·ly hominid that we class::
:tus is younger than H 01~
J erectus fossils were follil::.
,m Zhoukoudian near B.=--
~ Man" (Peking is an o!C:e-
unfortunately lost duri::=
no erectus have also be:::-
c of Georgia, and in soc·-
d geographic variabili ty
ctus. As with other ea::-.
nds, and some paleonto::-
o during this period. So;::,-
ssils as a separate spec::::
·pretation that all of thcs.=
ge of Homo erectus foss::.
!ly 300,000 years ago. E=
gly cold, dry, and vari a,., -
md its glacial-intergla
t a link between climc-
;ome evidence that Ho,...-
lan 100,000 years ago. T.-:.:
Homo erectus fossils co::::
md China date from a
tentary evidence sugges:...
000 years ago. In any c<:s::
ninids leaving Africa a:;_
1e most complete skele: , -
the son of Louis and l\12..
THE HOMINIDS: EARLY HOMO 339
Leakey) and Alan Walker at Nariokotome in Kenya, east Africa. It seems that
the boy had died in shallow water and his body had been quickly covered by sed-
iment; this preserved his skeleton from scavengers. It is estimated that had he
reached adulthood the so-called Nariokotome Boy would have stood close to 180
em tall and weighed about 70 kg. Aside from the skulls, the skeletons of Homo
erectus are very similar to those of modern humans and appear well adapted to
long and relatively fast movement across open country. There is much less differ-
ence in the size of male and female Homo erectus than is the case with earlier
hominids. Their skulls were slightly wider than those of modern humans, and they
had a very prominent bony ridge above their eyes. They also had strongly sloping
foreheads and receding chins. A relatively wide nasal opening suggests a some-
what modern nose shape and nasal passages. These features have been inter-
preted as an adaptation to conserve body moisture during breathing. Both the
stature and nasal configuration of Homo erectus have been interpreted as evolu-
tionary adaptations to life on the hot, dry, open savanna. The brain capacity of
Homo erectus ranged from 900 to 1200 cu em and was therefore larger than that
of earlier homonids but slightly smaller than modern humans. The brain shape
and throat of Homo erectus may have allowed modern speech, but this remains
highly controversial among physical anthropologists.
The geographic expansion of Homo erectus out of Africa and into Europe
and Asia is an extraordinary chapter in human history. Hominids now ranged
from the Atlantic Coast in Europe to the shores of the Indian Ocean in Africa
and to the Pacific Coast in southeastern Asia. No other Quaternary primate and
few other terrestrial mammals aside from hominids have had such a huge range.
It is likely that the evolution of Homo erectus and its geographic expansion were
generated in part by environmental change. As the Pleistocene proceeded, condi-
tions in Africa and adjacent portions of Europe and Asia became generally drier,
while the variations in climate associated with Milankovitch cycles became more
pronounced. Increasing areas of savanna and open landscapes may have pro-
moted the adaptations to dry conditions and mobility seen in Homo erectus. In
addition, the movement of Homo erectus and eventual expansion beyond Africa
may have been promoted by the adjustments in foraging areas in response to
changes in climate and vegetation. It is reasonable to think that Homo erectus
populations shifted their ranges as the distributions of vegetation and animal
resources shifted in response to climatic change.
The geographic expansion of Homo erectus also appears to have been facil -
itated by increasing cultural sophistication. The Lower Paleolithic stone tools of
Homo erectus are called the Acheulean Tradition and include much more refined
and varied choppers and scrapers, as well as finely developed hand axes and
cleavers. Similar levels of workmanship in stone, but not exactly similar tool kits,
are found from Africa to China. Homo erectus were also more particular in
choice of stones used to make tools and particularly favored flint. Evidence of
hearths have been discovered at several Homo erectus sites. It appears that fire
was being used by at least 250,000 years ago and perhaps as early as 500,000 years
ago. In addition, it is unlikely that populations of Homo erectus could have sur-
vived in relatively cool regions such as China and Europe if they did not possess
fire. Fire not only provided warmth and protection against predators, but it also
greatly enhanced the availability of food resources. Cooking often releases addi-
tional nutrients and proteins in meats and vegetables. It can also kill bacterial
contamination in meats and detoxify certain plants that would be poisonous if
340 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

eaten uncooked. In addition to stone tools and fire, it is likely that Homo erectus Java only 50,0
fashioned tools from wood, bamboo, and bone, and in Southeast Asia they may Africa Model.
have been able to cross water on rafts. Populations in cool areas likely used ani- modern huma1
mal skins for clothing and to construct shelters. Paleontological and pathological Africa some 21
evidence suggests that Homo erectus hunted and/or scavenged in groups and been widely qu
took care of group members who were ill or infirm. As technologically advanced Alternati
and geographically expansive as they were compared with earlier hominids, we skull features i1
find no evidence for some important attributes of modern humans in Homo erec- tures found in I
tus. There is no evidence, for example, that they developed ritualistic behavior. large bony bro
such as formal burial or cremation of the dead, or that they produced nonutilitar- Aboriginal fos:
ian items for ritual ceremonies or art. Such features, when they do exist, provide posed that Hor.
firm evidence of the ability for abstract thought, and sophisticated communica- and then sprea1
tion and culture that we associate with modern humans. For this, we must await Out of Asia Me
the appearance of our own species, Homo sapiens. It has als
Homo sapiens '
Europe, and/or
THE HOMINIDS: HOMO SAPIENS Africa, Europe
and genetics th
All humans who are alive today are members of the taxon Homo sapiens sapiens. ens. Some rese<
The addition of the second subspecies name, sapiens, is to differentiate modern origins for the '
humans from the oldest Homo sapiens fossils, which possess a more prominent Asians may ha
brow line and thicker skull than is found in present-day humans. These early southern Asian
forms are generally referred to as archaic Homo sapiens. Homo sapiens sapiens erectus living ir
are referred to as modern humans. In addition, the term Homo sapiens sapiens evolved from 1
also recognizes the difference between modern humans and an extinct hominid called the Molt
of the recent past, Homo sapiens neanderthalensis (Neanderthal man). It is likely by hominids we
that Homo sapiens evolved from Homo erectus. Archaic Homo sapiens, with in geographical
brain capacities similar to modern humans but having some traits similar to holds that mod1
Homo erectus, appear in Asia, Africa, and Europe by about 400,000 years ago. ing between ea1
Although their fossils and tools are rare, with the Archaics there is an increased populations fro
diversity and refinement of stone tools and we enter the Middle Paleolithic (mid- tralia and Euro
dle stone age). In 1967 Richard Leakey found the skull of an early Homo sapiens ing the origin
sapiens near the Omo River in Kenya. This skull dates to about 130,000 years ago anthropologists
and provides the earliest evidence of essentially modern humans. Fossils indicate lively debate an
that Homo sapiens were present in Europe and Asia perhaps 100,000 to 90,000 The histm
years ago. Neanderthal fo:
Before we continue our consideration of our own Homo sapiens sapiens found in 1856 ir
ancestors, we need to consider two important facets of the Homo sapiens evolu- for limestone u
tionary story. The first is where did Homo sapiens first evolve? The second ques- quite thick, had
tion is how were the Neanderthals related to us and what became of them? prominent brov
As far as we know, all earlier hominid ancestors evolved first in Africa. this odd skull c<
Many scientists believe that this is also the case for Homo sapiens. Fossils with a lost Cossack ~
intermediate traits between Homo erectus and archaic Homo sapiens have been posed that the s
found in Ethiopian deposits that are about 1 million years old. In this scenario. anthropologist,
Homo sapiens evolved first in Africa and then spread through Europe and Asia human and clas:
between about 700,000 and 400,000 years ago. They either displaced and cause next 50 years, I'
the extinction of Homo erectus, or absorbed them through interbreeding. Homo tions or were re
sapiens with their larger cranial capacity and demonstrably superior technica: century, it was
capabilities was likely a superior competitor than Homo erectus. There is some hominid closely
controversial evidence that the last Homo erectus may have disappeared fr or::. fossils date frorr

t is likely that Homo erec
in Southeast Asia they Jlli_
n cool areas likely used a::_-
)ntological and patholo gi ~·
•r scavenged in groups a::.~
\s technologically adva nc~.:.
:d with earlier hominids. " :
dern humans in Homo er:?_-
reloped ritualistic beha\ic
t they produced nonutili tE..- -
when they do exist, pr o\~ :_
i sophisticated communi.::::-
ans. For this, we must aw=
axon Homo sapiens sapi.:
; is to differentiate moce::-
1 possess a more promin7-
lt-day humans. These er
•iens. Homo sapiens sap1c
term Homo sapiens sap:
ans and an extinct homi::.....::
eanderthal man). It is lik::
rchaic Homo sapiens, w:
ving some traits similar
'Y about 400,000 years "-=
chaics there is an increa_--= _
he Middle Paleolithic (c::...,~
Jl of an early Homo sap z~
; to about 130,000 years ~=
~rn humans. Fossils ind i~
1 perhaps 100,000 to 90. =
Jwn Homo sapiens sap;:. _
)f the Homo sapiens e\T'.~
:t evolve? The second q•.: :::-
vhat became of them ?
Homo sapiens. Fossils
[c Homo sapiens have t);:-_
years old. In this scene.:-·
i through Europe and ..;
~ ither displaced and caG:::-_
~ough interbreeding. H e
nstrably superior tech£:.:.
Jmo erectus. There is sc:-
tay have disappeared t-
THE HOMINIDS: HOMO SAPIENS 341
Java only 50,000 years ago. The scenario outlined above is called the Out of
Africa Model. Molecular clock analysis (Technical Box 11.1) suggests that all
modern humans are descended from one Homo sapiens female who lived in
Africa some 200,000 years ago. This conclusion, called the Eve Hypothesis, has
been widely questioned, however, by molecular geneticists and anthropologists.
Alternatively, some physical anthropologists point to certain distinctive
skull features in early Asian and Australian Homo sapiens that are similar to fea-
tures found in late Homo erectus fossils from Java. Some of these features, such as
large bony brows, have even been found in several relatively recent Australian
Aboriginal fossils dating from the past 40,000 to 10,000 years. It has been pro-
posed that Homo sapiens may have evolved from Homo erectus in southern Asia
and then spread to Australia, northern Asia, Europe, and Africa. This is called the
Out of Asia Model.
It has also been suggested, on the basis of morphological evidence, that
Homo sapiens evolved from two or more separate events and lineages in Africa,
Europe, and/or Asia. In this latter case, parallel evolution occurring separately in
Africa, Europe, and Asia produced lineages that were so similar in both anatomy
and genetics that they could interbreed and produce modern Homo sapiens sapi-
ens. Some researchers suggest that there may have even been two separate Asian
origins for the evolution of Homo sapiens from Homo erectus. Modern northern
Asians may have originated from populations of Homo erectus in China, and
southern Asians and Australians may have originated from populations of Homo
erectus living in Java and adjacent islands. The scenario in which Homo sapiens
evolved from Homo erectus separately at two or more geographic locations is
called the Multiregional Model. In this case, it is possible that parallel evolution
by hominids would have produced similar morphological and physiological traits
in geographically separated populations. One version of the Multiregional Model
holds that modern humans in Europe and Asia may have arisen from interbreed-
ing between earlier European and Asian hominids and expanding Homo sapiens
populations from Africa. Some genetic and skull morphology evidence from Aus-
tralia and Europe appears to support this possibility. Which hypothesis concern-
ing the origins of modern humans is favored at present? Today, many
anthropologists support the Out of Africa Model, but this remains a topic of
lively debate and intense research.
The history of the Neanderthals is an extremely interesting story. The first
Neanderthal fossils that were recognized as representing extinct hominids were
found in 1856 in the Neander Valley of Germany. The quarrying of cave deposits
for limestone unearthed a rather unusual hominid skull cap. The skull cap was
quite thick, had a pronounced sloping of the forehead , and displayed a very thick,
prominent brow. The nineteenth-century scientists did not know what to make of
this odd skull cap. It was suggested at first that it might have been the remains of
a lost Cossack soldier from the Napoleonic Wars. Alternatively, it was also pro-
posed that the skull cap might have belonged to an infirm Dutchman. An English
anthropologist, William King, suggested that the skull might be from an extinct
human and classified the fossil as a new species, Homo neanderthalensis. Over the
next 50 years, Neanderthal remains continued to surface in European excava-
tions or were recognized in existing museum collections. By the early twentieth
century, it was widely accepted that Neanderthals were indeed an extinct
hominid closely related to modern humans. We now know that most Neanderthal
fossils date from approximately 130,000 years ago to 35,000 years ago.
342 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

The numerous Neanderthal fossil sites provide a fairly clear picture of- - Our skull~
geographic distribution of these hominids (Fig. 11.4). They appear to have Jj , =-- tions to a warn
exclusively in Europe and the Near East. It is possible that there were some our similarity ir
ulations along the Mediterranean Coast of Africa, but this remains uncertain. conjecture. Em
now seems quite clear that Neanderthals did not live in east or southeastern ~ stooped, brutisl-
nor did they live in sub-Saharan Africa. It is very likely that they evolved ErL tions led to the
Archaic Homo sapiens in either Europe or the Near East. Thus, they are the o=.. on a crowded s
hominids that unquestionably did not first evolve in sub-Saharan Africa. other passenger
Because of their sloping skull, prominent brow, recessed chin, and hec. grapher, Jared _
build, it was supposed that Neanderthals represented a primitive ancestor · Neanderthals w
modern humans. However, both the conclusions that Neanderthals were e\-o__ bulbous nose, d,
tionarily primitive or were ancestral to modern humans are wrong. would appear al
Like modern humans, Neanderthals evolved from archaic Homo sapic how crowded.
Evidence from Atapuerca, Spain, suggests an early evolution of the Neandertt :_ What bee
commencing by 300,000 years ago. The stocky build, thick bones, large nose. - short answer is
heavy jaws and dentition of the Neanderthals were genotypic features and can-- Europe and th1
seen in the skeletons of Neanderthal children. It is likely that the Neandertt:::... Homo sapiens 1
evolved these features as adaptations to life as hunters in the cold environm :.- coldest conditic
of glacial age Europe. The stocky build would have maintained body warmth c:...: for foraging or
ing the cold winters. The large muscles and thick bones would have been a b ;::: In addition, it i
fit to hunters who went after game such as deer, mammoth, and woolly rhinoce::- food sources tl
The maximum population size and widest geographic extent of Neanderth::.._ shown by Hom
appear to have occurred during the period 100,000 to 40,000 years ago, will - - cates a more raJ
coincides with generally cold conditions (see Chapter 8). cope with chanf
The Neanderthal tools from this period are classified as Middle Paleoli Did moder
and are typified by the Mousterian Tradition, which appears to have eva ]\·:-_ from the Near E
directly from the older Acheulean Tradition. The Mousterian tools were mo:-: regions during tl
finely made and more diverse, including borers, simple saws, and points that \\- ::-: either group fou
attached to wooden poles to form spears. Neanderthal spears have been fo - .: hominids huntir
embedded in the fossil remains of large mammals, and provide evidence of ---= derthals and me
hunting skills of the Neanderthals. Neanderthals used fire and certainly fashi o n~.: think that the t\
clothing and shelter from animal hides. They lived in groups and cared for infu:::: interact directly,
members. They also had ritualistic practices that are evident in formal burials th2. foragers and res
have been found in some cave sites. This includes the placing of flowers in t2: possible that the
grave or the arrangement of animal skulls around the grave. Carved symbo!:..
strategies to cop
plaques called churingas have been found at some Neanderthal sites and pro\·i ·=
breeding may ha
further evidence of symbolic abstract thought. Most, though not all, anthropolc-
has been suggest
gists believe that Neanderthals had the capacity for speech, but perhaps not t.:::
certain facial fee
full range of sounds used by modern humans. Impressively, the world 's firs:"
from Europe a1
known musical instrument is a small flute made from bone and found at an 80,00
derthalensis and
to 40,000 year old Neanderthal site in Slovenia. Rather than being evolutionari:_
from Neanderth;
primitive, the Neanderthals were intelligent and possessed specialized adaptz-
tions to live by hunting and gathering in a cool climate region. much difference
What was the relationship of the Neanderthals to us? Modern-lookin; breeding betwet
Homo sapiens sapiens had likely evolved in Africa from Archaic Homo sapier_ archaeological e·
and were likely present in Asia during the same period of time that Neandertha:s technologies be
were present in Europe and the Near East. Thus, modern humans evolved sepa- between the N e<
rately from Neanderthals and not as their descendants. Molecular clock analysi5 40,000 years ago
suggests that the divergence between Homo sapiens neananderthalensis anc clear. By 35,000
Homo sapiens sapiens occurred several hundred thousand years ago. left on the planet

a fairly clear picture of ti:::
. They appear to have li\·e.::
~ that there were some p o~­
It this remains uncertain. =
n east or southeastern As~
:ely that they evolved frc-
~ast. Thus, they are the o
ub-Saharan Africa.
v, recessed chin, and hec:-
ed a primitive ancestor :,
t Neanderthals were e\·o__
ns are wrong.
·om archaic Homo sapic
olution of the Neanden b ... ,
thick bones, large nose. c.::._
notypic features and car. =-,~
kely that the Neand en t~
rs in the cold environmc-=
lintained body warmth c-:.=
:s would have been a be:
1oth, and woolly rhinocc-::-
Iic extent of Neande-"-
to 40,000 years ago, w~-
8).
sified as Middle Paleoli:...::..
1 appears to have eY oi~ :-.:.
ousterian tools were r::
; saws, and points that
tal spears have been fo
nd provide evidence o:: ~­
fire and certainly fashio::._
~roups and cared for iK::
1ident in formal burials · -
1e placing of flowers in -·
he grave. Carved sym:C,
anderthal sites and pro'
though not all, anthro;:,-
peech, but perhaps no: :
lressively, the world 's ::..
Jone and found at an SC
:r than being evoluti o~:E...-:
;sessed specialized ade:;:
e region.
1ls to us? Modern-J oe..
om Archaic Homo sa;
l of time that Neande:--
lern humans evolved s_.
:s. Molecular clock anc...
rts neananderthalensi_
>and years ago.
THE HOMINIDS: HOMO SAPIENS 343
Our skulls, dentition, and physique suggest a more generalist set of adapta-
tions to a warmer climatic environment. Although we share common ancestry,
our similarity in physical appearance to Neanderthals has long been a matter of
conjecture. Early anthropological reconstructions portrayed Neanderthals as
stooped, brutish creatures with hanging arms and apelike faces. Later reconstruc-
tions led to the often repeated suggestion that if we were to meet a Neanderthal
on a crowded subway we would not notice much difference between him and
other passengers. More recently, the eminent physiologist, ecologist, and biogeo-
grapher, Jared Diamond, has dispelled this myth. He points out that although
Neanderthals were no slouching brutes, the huge barrel chest, massive arms, giant
bulbous nose, deeply receding chin, prominent brow ridge, and sloping forehead
would appear absolutely jarring to modern humans riding on a subway, no matter
how crowded.
What became of the Neanderthals has long been a topic of debate. A
short answer is that they went extinct and were replaced in their range in
Europe and the Near East by Homo sapiens sapiens. The generalist species
Homo sapiens sapiens had a much more energy-efficient build for all but the
coldest conditions. Modern humans were better able to travel long distances
for foraging or to escape inhospitable climatic conditions than Neanderthals.
In addition, it is thought that Homo sapiens sapiens used a wider variety of
food sources than Neanderthals. Finally, the speed of technical innovation
shown by Homo sapiens sapiens in tool development and tool diversity indi-
cates a more rapid ability to shift and develop tools and survival techniques to
cope with changing environments.
Did modern humans actually drive Neanderthals extinct? There is evidence
from the Near East that Neanderthals and humans lived in the same geographic
regions during the end of the Neanderthal period. There is no clear evidence that
either group fought with or hunted the other. At first, we might think the idea of
hominids hunting other hominids preposterous, but we know that both Nean-
derthals and modern humans practiced cannibalism, so it is not outrageous to
think that the two species might have hunted one another also. If they did not
interact directly, perhaps Homo sapiens sapiens were simply better hunters and
foragers and resource competition led to the decline of the Neanderthals. It is
possible that they could not adopt a more generalist diet or evolve new tools and
strategies to cope with dwindling game. Finally, it has been proposed that inter-
breeding may have occurred between Neanderthals and Homo sapiens sapiens. It
has been suggested that some European and Near Eastern populations preserve
certain facial features typical of Neanderthals. In addition, some human fossils
from Europe appear to represent hybrids between Homo sapiens neanan-
derthalensis and Homo sapiens sapiens. However, comparison of DNA extracted
from Neanderthal bones with DNA of modern humans suggests that there is too
much difference between the two subspecies for there to have been any inter-
breeding between Neanderthals and us. In addition, there is no compelling
archaeological evidence for interaction, such as trade of tools or sharing of tool
technologies between Neanderthals and modern humans. What occurred
between the Neanderthals and Homo sapiens sapiens in that distant time some
40,000 years ago will long remain a topic of keen interest, but the results are
clear. By 35,000 years ago, the Neanderthals were extinct and one hominid was
left on the planet- modern humans.
344 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

THE GEOGRAPHIC EXPANSION

OF MODERN HUMANS

The geographic expansion and population growth of Homo sapiens sapiens ~

one of the most dramatic and significant events in the recent biogeographic his-
tory of the earth. At 100,000 years ago, it is possible that Homo sapiens sapiens
were present only in Africa. By 90,000 years ago, they were also present in the
Near East but were subsequently replaced by Neanderthals as climate coolec
during the last major episode of Quaternary glaciation. The early Homo sapien~
sapiens were adapted to life in relatively warm conditions on the open country-
side and utilized a variety of Middle Paleolithic tools that had their roots in the
Acheulean Tradition. Around 50,000 years ago, there seems to have been a rapic
expansion of Homo sapiens sapiens in and out of Africa. They created art, prac-
ticed ritual burials, and displayed other indications of sophisticated culture. Th ~
period has been called the Human Revolution. By no later than 40,000 years ago.
Homo sapiens sapiens was present throughout Africa and Eurasia (Fig. 11.5)
There is widespread evidence that people colonized northern Siberia by 35,0()(
to 25,000 years ago. The late spread of humans from southern Asia to northerr 8
Asia may indicate that a relatively high degree of technical and cultural develop- 7
ment was required before people could settle this cold region. However, there ~ ~6
0
some evidence from northern Siberia that small groups of hominids may have ~5
entered the region significantly earlier. In Europe, the earliest Homo sapiens 04
sapiens were widespread by about 40,000 years ago. They are called Cro- (/)
c
Magnons after a cave near Les Eyzies, France, where some of their remains were ~3
i:ii 2
found. These particular early modern humans were tall and slender people witt
large craniums. The average size of the Cro-Magnons was larger than that o: 1
most subsequent humans until the twentieth century. This is because they appar- 0
2-5 million
ently obtained a highly nutritious diet from hunting and gathering compared tc
later agrarian peoples.
The preserved stone, bone, and wooden artifacts of the early EuropeaL FIGURE 11.5 Tl
and western Asian Homo sapiens sapiens display much greater variety anc Gamble, 1994; Bro
greater finesse than the Mousterian Tradition artifacts of the Neanderthals. We population.
classify these early modern human artifacts and the culture they reflect as Upp e ~
Paleolithic. Some important new tools included chisels and wooden spear throw-
ers that increased the distance a projectile could be launched. We also see th Homo erectus m
clear development of representational art. Wood, bone, and stone carvings o: passages betwee
animals were made, as were Venus figurines, which are carvings and baked clay lace's Line (see 1
figures of women with greatly enhanced hips and breasts. Venus figures are The first undispu
found from western Europe to the Near East to Siberia and indicate the sharin§. years ago, and nE
of artistic and symbolic ideas over very great distances even in this early period. The timing
The walls of at least 200 caves in France and Spain contain beautiful, somewha ica remain contr
abstracted paintings of animals. The oldest of such artworks in Europe date back cial maximum lc
about 15,000 to 32,000 years. Early paintings and drawings also appear on rock s Siberia and Alas
at a number of sites in Australia. Across Europe, Asia, Australia, and Africa we bridge from Sib(
see greater geographic variability in tools and other artifacts than was the case Territory of Can
with Neanderthals and earlier species of hominids. People were devising ne\Y Beringia suppon
tools, art forms, and other innovations to suit the resources and requirements of and would have
their environments. from modern hu
At the same time that Homo sapiens sapiens was expanding throughout ican and South 1
Europe and northern Eurasia, they were also colonizing Australia. Although Siberia. As the C
 THE GEOGRAPHIC EXPANSION OF MODERN HUMANS 345

' Homo sapiens sapiens _

recent biogeographic ~
hat Homo sapiens sap ier_
y were also present in ~ =
lerthals as climate cool:'.:
1. The early Homo sap ier..
ions on the open count~ ­
that had their roots in ::.=
eems to have been a ra,;-:.:
ca. They created art, prc:- "
Tazmania
sophisticated culture. T.:......
ater than 40,000 years a ~ ­
a and Eurasia (Fig. 1 ~ ..:
torthern Siberia by 35.=
;outhern Asia to northe::- 8
lical and cultural develc: 7
l region. However, there ~6
0
tps of hominids may he· : ~5
he earliest Homo sap:= 04
(/)
~o. They are called C:-:- c
~ 3
ome of their remains \\· ===-~
i:ii 2
11 and slender people '>'>:...:
IS was larger than tha: 1
'his is because they app .: OJ
2-5 million
;\' I I I I I I I I I =rY I
10,000 9,000 8,000 7,000 6,000 5,000 4,000 3,000 2,000 1,000 0
nd gathering compare.:
Time (Years before present)

ts of the early Europ.:z-
nuch greater variety 2..::._
s of the Neandertha ls.. ";
lture they reflect as u r~·­
and wooden spear thrc
aunched. We also see --
ne, and stone carvi n ~
e carvings and baked -
>reasts. Venus figur es -=.:
a and indicate the sha: _
; even in this early pe::: _
ntain beautiful, somew--
vorks in Europe date c.::....
rings also appear on r ~-
, Australia, and Afrj ~ ~
1rtifacts than was the .::.:
'eople were devising :::.:: -
lrces and requirem e n ~
ras expanding thro u ~ ".: -
izing Australia. Alth0 _
FIGURE 11.5 The general geographic spread of Homo sapiens sapiens (data from
Gamble, 1994; Brown and Lomolino, 1998; Fagan, 1998) and the growth of hominid
population.
Homo erectus may have been able to cross small areas of water, the larger water
passages between southeastern Asia and New Guinea-Australia that form Wal-
lace's Line (see Chapter 10) kept these earlier hominids from reaching Australia.
The first undisputed evidence of human presence in Australia dates to about 40,000
years ago, and new evidence suggests human presence as early as 60,000 years ago.
The timing and arrival route of the first humans in North and South Amer-
ica remain controversial. The drop in sea level that occurred during the last gla-
cial maximum lowered the Bering Sea enough to produce a landbridge between
Siberia and Alaska. It is indisputable that some humans crossed the Bering Land-
bridge from Siberia and entered the ice-free portions of Alaska and the Yukon
Territory of Canada by at least 15,000 years ago. During this period, the region of
Beringia supported a number of large mammals, including mammoth and bison,
and would have provided good hunting grounds for humans. Genetic evidence
from modern human populations suggests a linkage between many North Amer-
ican and South American native peoples and people living near Lake Baikal in
Siberia. As the Cordilleran and Laurentide ice sheets melted between 14,000 and
346 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

12,000 years ago, a corridor of tundra, grassland, and shrubby habitat opene: Pacific. Genetic
between Alaska and the Great Plains. This corridor would have served as a rou t~ first immigrant1
for the rapid southward migration of humans and animals into the heart of Nor '- Melanesian Isla
America. By 12,000 years ago, people were present throughout North and Sou '- by later arrivah
America. The initial peopling of the New World via Siberia is supported by e\ :- of the most imi
dence from anatomical and genetic analysis that shows an affinity between nati,-=- the Lapita Cull
peoples of North and South America and people in northeastern Asia. reached islands
There is some evidence from sites in eastern North America and Brazil th.: people who in c:
people were present in North and South America by 15,000 to 30,000 years ago. T. duced ceramic 1
this evidence is correct, it means that humans had to have crossed from Sibe r~ area of Fiji until
and entered central North America before the start of the last glacial maximur:;:, and 700 years a.
some 30,000 years ago. The fact that very few sites of this suggested age are knO\C as far north as I
is a marked contrast to the widespread evidence of humans across North an- Polynesians rna:
South America after the end of the last glaciation 12,000 years ago. If people wer= sians were exce
present between 30,000 and 15,000 years ago, it would seem that they existed c to their initial c
very low population levels and made few, if any, implements from stone. marine trade lin
There is also some evidence that prehistoric people arriving in North an: Hawaii.
South America may have used several different migration routes. Stone poin= Thegeogr;
found in North and South America show similarities to similar aged points fro- population size.
northwestern Europe. This has led to the suggestion that this technological trad:- existed, the pop
tion was introduced to North America by people moving across the Nor .__ 10,000 years age
Atlantic. The 8000-year-old skeletal remains of a male human found in Washing- likely a few mil
ton State bears some resemblance to modern western Asians and Polynesiall5. growth was pro
suggesting the migration of some groups by water across the North Pacifi - people. Some an
Finally, there is evidence that people may have moved south from Beringia alon; sapiens sapiens
the Pacific coastline of North America. In this case, human migration into cent re... human populati
North America would not have had to wait until the continental ice sheets ha: gathering practi
retreated. mobility and lar
Prehistoric Asian migration to North America may have continued int sapiens sapiens 1
more recent time. The ancestors of the modern Navajo and Apache of the Soutt- of people and ra
west and the NaDene of northwestern Canada likely migrated from northeaster:::. tition with othe1
Asia in the last 5000 years. Their languages are related to one another, and a:. organization an<
show linkages to northern Chinese-Tibetan languages. The Aleuts and Inuits (fo - ported in smalle
merly called Eskimos) of northern Alaska, northern Canada, and Greenland ar= sure, fueling ne"
very closely related to the Chukchis and other native peoples in northern Sibe ri ~ During thf
The first humans in the High Arctic of the New World, including the ancestors o: more marked an
the modern Inuit, likely migrated from Asia between 4000 and 1000 years ago. Only a small por
There are other portions of the inhabited world that were settled relative!_ habited regions.
recently by humans. These are islands that required relatively long voyages tc tural innovation:
reach them. Jamaica and the Lesser Antilles of the Caribbean region were settle.: pottery and ther
less than 5000 years ago. Madagascar, off the coast of Africa, was settled aboc capacity of the
2000 years ago. Some islands, including several in the Indian Ocean and th"" gested that the d
Commander Islands off the coast of Kamtchatka, Russia, appear to have tm: increase in the 1
been reached by humans only in the last few centuries. area, and the gee
Perhaps the most impressive story of human expansion to remote islan " There is or
involves the peopling of Hawaii and the other islands of the Pacific. Th - graph in Figure :
Pacific Islands appear to have been settled by waves of migrants originating li:. Europe and adj;
southeastern Asia. The first humans arrived on the western Melanesian Islan - 1700s. The most
near New Guinea about 45,000 years ago. These early peoples were able tc between A.D. 13
cross large expanses of water but were not able to spread into the easte r;:: died. Some Eun

1d shrubby habitat opene=
·ould have served as a ro u: ~
mals into the heart of Non:.
1roughout North and Sou" -
Siberia is supported by eY-:-
rs an affinity between nat i'· ~
ortheastern Asia.
rth America and Brazil t h~
l5,000 to 30,000 years ago.=...
> have crossed from Sibe:-
>f the last glacial maximu=
llis suggested age are knov-
: humans across North a::,_
100 years ago. If people \\ e-::
ld seem that they existed ~
ments from stone.
~ople arriving in North a::,_
~ration routes. Stone poi=."
to similar aged points frc;-
hat this technological trc.-
moving across the Ko;-...:;
e human found in Washi.::r
rn Asians and Polynesic::...
r across the North Pace_
:i south from Beringia alc-
Jman migration into c en::.:-~
: continental ice sheets t.
1 may have continued ~ :
o and Apache of the Soe:= -
migrated from northe a s:~­
ted to one another, and
:. The Aleuts and Inuits (:.:--
Canada, and Greenland -:.:
peoples in northern Sibe:-
:i, including the ancesto. _ _
4000 and 1000 years age
_that were settled relati\ :-
relatively long voyages.
rib bean region were se r.~ :: _
)f Africa, was settled a
the Indian Ocean and :-
~ussia, appear to have =-.
s.
xpansion to remote ish:__
slands of the Pacific. -=--
s of migrants originatin_§: -
vestern Melanesian Isle.:..:..
~ arly peoples were able
:o spread into the eas t:--
THE GEOGRAPHIC EXPANSION OF MODERN HUMANS 34 7
Pacific. Genetic and linguistic evidence suggests that the descendants of these
first immigrants live today in the mountainous interiors of some of the western
Melanesian Islands. Their ancestors were probably displaced from coastal areas
by later arrivals and survived only in the mountainous interiors of islands. One
of the most important later groups to expand in Melanesia were the people of
the Lapita Culture who spread into Melanesia at around 3500 years ago and
reached islands as far eastward as Fiji. The Lapita were a relatively sophisticated
people who in addition to being excellent sailors, practiced agriculture and pro-
duced ceramic pottery. Subsequent human migrants did not spread beyond the
area of Fiji until the Polynesians expansion phase occurring between about 2100
and 700 years ago. The Polynesians were excellent sailors who colonized islands
as far north as Hawaii and eventually as far east as Easter Island. To the south,
Polynesians most likely reached New Zealand about 700 years ago. The Polyne-
sians were excellent navigators, relying on the stars to chart courses. In addition
to their initial colonization voyages, Polynesian peoples in Hawaii maintained
marine trade linkages with Tahiti for a number of years following their arrival on
Hawaii.
The geographic spread of modern humans has been followed by a growth in
population size. Over most of the time during which Homo sapiens sapiens have
existed, the population level of humans has remained relatively low. Until about
10,000 years ago the entire human population of the planet at any one time was
likely a few million individuals. During much of this period, human population
growth was promoted in part by the colonization of new parts of the world by
people. Some anthropologists have suggested that from the appearance of Homo
sapiens sapiens onward there has been a positive feedback linkage between
human population size and cultural and technical innovation. The hunting and
gathering practices of early Homo sapiens sapiens appear to have required
mobility and large ranges for movement. There is some evidence that as Homo
sapiens sapiens populations grew in the Mesolithic and Neolithic, the movement
of people and range available for foraging became more restricted due to compe-
tition with other bands of humans. This may have spurred on changes in social
organization and technologies that allowed larger numbers of people to be sup-
ported in smaller areas. These successes in turn created greater population pres-
sure, fueling new innovations in social organization and technology.
During the period 10,000 years ago to about 300 years ago, there was a
more marked and steady increase in the world population of humans (Fig. 11.5).
Only a small portion of this growth can be attributed to the colonization of unin-
habited regions. The increasing size of population was largely promoted by cul-
tural innovations such as the development of agriculture and the development of
pottery and then tools made of metals. These innovations increased the carrying
capacity of the land for humans. The paleontologist, Niles Eldredge, has sug-
gested that the development of agriculture for humans can be likened to a major
increase in the niche of a species. More people could be supported in a given
area, and the geographic range of humans could expand.
There is one notable decline in world population that is apparent from the
graph in Figure 11.5. This decline is the reuslt of a series of plagues that affected
Europe and adjacent areas starting around A.D. 1347 and continued into the
1700s. The most severe of these outbreaks was the Black Death which occurred
between A.D. 1348 and 1351 when perhaps one-half of the population of Europe
died. Some European cities lost two-thirds of their population. The plagues of
348 CHAPTER 11 BIOGEOGRAPHY AND HUMAN EVOLUTION

this period were caused by the bacteria Yersinia pestis, which is carried in --~ _.::_,[enocal, P. B. (1995). Plio-]
blood of rats and transferred to humans by flea bites. Forms of the plague .:limate. Science 270, 53-59
also be transferred to people through exposure to bodily fluids such as sali\ _ ::.::mond, J. (1992). The Third
Early reports of the plague come from China in 224 B.C. Originating in Asia, ·""-= Evolution and Future ofth
plague was possibly first transmitted to western Europeans when an army, besi-;- Ha rper Perennial, New Yo
ing an Italian trading post on the Crimean Peninsula, catapulted plague-infesl- ~ -~ an. B. M. (1998). People of
corpses into the town. The disease spread rapidly due to the proliferation of tra..:.: lnrroduction to World Preh
linkages in Europe and the increasing number of people living in crowded a::.~ Longman, NY.
relatively unsanitary cities. Aside from being a horrible human tragedy, ·- = =_:ey. R. (1987). Another Unit
;nan, London.
plague and its impact provide stark evidence of the potential rapid spread - ~
::: ::...;nble, C. (1994) . Timewalkt
profound damage that can be caused by invading organisms. of Global Colonization. H:
In the last 200 years, the size of the human population has exploded fro= Press, Cambridge, MA.
about half a billion to over six billion people. The recent explosive growth ~ -"2\ iland, W. A. (1989). Anthr,
human population can be attributed to increasing rates of industrial, techni ·ion. Holt, Rinehart and W
and scientific development and the rapid transfer of such new developmeL- ~ :J . d en , C. (1999). Were Span
throughout the world. These developments have served to increase agricultu ~ first Americans? Science 2~
output and decrease human mortality rates in many parts of the world. The ra p: ~ -:': v\\ ells, W. (1993). Getting H,
development of new technologies and the shift of the socioeconomic syste= Washington, DC.
away from an agrarian base toward a techno-industrial base that has occu rr ~: -:':2blin, J. (1999). The Quest fc
ogy 52, 26-32.
since about 1800 is called the Industrial Revolution. In addition to leading :
-~s . C. M. (1993). Tertiary m:
increased human population size, the Industrial Revolution has seen a shift of tt:o
:he context of changing cli1
world's human population to the cities from rural areas. The remarkable g ~­ nd tectonic events. Annua
graphic expansion and population growth of Homo sapiens sapiens have had -'- ogy and Systematics, 467-5<
large impact on the environment and other species with which we share tt~ _::tanson, D., Johanson, L., an
earth. In the next chapter we will examine the role of our species as an agent - Ancestors: In Search of Hu.
both evolution and extinction. lard Books, New York.
- = ~ e n hans, H., F., Daryl, R . P. ,:
(1997). Early humans and r
KEY WORDS AND TERMS Holocene sea levels in the '
Islands-Hecate Strait, Briti
Acheulean tradition Industrial Revolution Oldowan tradition Canada. Science 277,71-74
Bipedal Molecular clock Out of Africa model -2:ohlin, W. S., and Harper, A.
Hominid Mousterian tradition Out of Asia model of the continents: Australia
Human Revolution Multiregional model Primate Biological Aspects of Hum,
Cambridge Studies in Biolc
ogy (ed. C. G. N. Mascie-Ta
Lasker), pp. 14--40. Cambri
Press, Cambridge.
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Macchiarelli, R., Napoleone, G., Papini, M. , bones show. Science 286, 18-19.
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Villa, I. (1998). A one-million-year-old Homo Selection in Relation to Sex. Murray, London.
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to the proliferation of tra-=-=
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)rrible human tragedy. ::::.=
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•f our species as an ager:::
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CHAPTER 12 some organism
pies of how h1
genotypic and J
tionary respon:
HUMANS AS A FORCE chemicals such
few instances o
IN EVOLUTION of insect and m
cides. Over 400
AND EXTINCTION one or more ci
sessed genes p
were able to st
killed by the pe

In the previous chapter, we saw how the history of humans is itself a fascin a ·- _ Animal and
biogeographical story in which evolutionary forces, extinction, dispersal, -.:
geography all played a role. The end result of this saga was the rise of mod :-:: The most impo
humans to a global population size of six billion individuals and a geographic - come from the
tribution that includes every continent, with forays to the depths of the ocean - .: domestication l
the nearer reaches of outer space. What impact have the numerous and wi ·=- ulation increas(
spread members of the human species had on the rest of the biosphere? We co .: animals are the
easily focus an entire textbook on this important question. In this chapter we '' - cally innovativE
have to restrict our scope to looking at the role of humans in two key areas. Fl.f5"_ have come intc
we will examine how humans have influenced the evolutionary development for a prolonged
plant and animals species, particularly through domestication. The domesticatic- develop stratifi
of plants and animals and the development of agriculture are central to -..: such as farmer
growth of complex human societies, technological advancements, and increases - Because of the
human population size that have occurred over the past 10,000 years. Second. : geographers, in
will consider the sobering evidence of how humans have become a major force ~ made the patter
the extinction of plant and animal species around the world either directly landscape chan
hunting or by causing landscape alterations and environmental change. domesticated p
ancestral speciE
domestication '
HUMANS AS AN EVOLUTIONARY FORCE resources for br
careful use of n;
Humans can influence the evolutionary development of species in a number • cultural yields
different ways. Changes in the landscape caused by agricultural land clearan ::c important area
can lead to strong selective pressure for traits that allow wild species to live r::. ment of agricult
open fields or utilize crop and pasture plants for food. The removal of predato:-:: Domesticl
by humans can cause the remaining species to evolve adaptations that decrea::~ come to depen'
defenses against predation but increase foraging efficiency or reproductive rat .:. practical or oth,
Hunting pressure by humans can cause compensatory evolutionary develo;- provide food. 0
ments in prey species. For example, there is evidence that hunting by Near:- purposes such
derthals and early modern humans led to a decrease in the body size of the sp - selectively bree1
tortoise (Testuda gracea) in the Mediterranean region. Small body size may ha' :- traits can includ
been a beneficial adaptation because smaller individuals are less visible in tt:- tion in animals, '
landscape and less attractive to hunters as they provide lower nutritional vale:: resulting domes
than large prey. ior from their '-'
In more recent times, the introduction of pesticides and pollutants into th:: corn is probabl)
environment has exerted selective pressure and led to evolutionary change li: authorities and

350

mmans is itself a fascin a ~
s, extinction, dispersal. c.:;._
;aga was the rise of moc:--
viduals and a geographic -"
1 the depths of the ocean ::..=_:
ve the numerous and w~:..::
t of the biosphere? We co
:stion. In this chapter we
mans in two key areas. F:..
volutionary developmer.:
:stication. The domesti
riculture are central to --
vancements, and incre as~ -
>ast 10,000 years. Secon ·
ave become a major for.:-::
the world either directh
·onmental change.
1t of species in a numb;;-
agricultural land clear.:..:.
:1llow wild species to b =
d. The removal of pre ~:
·e adaptations that deer::
;iency or reproductive ~~ _
tory evolutionary de\·;e _
nee that hunting by "\\::
in the body size of the s:-
n. Small body size ma\· ·-
duals are less visible ;.;:,
vide lower nutritional " -
:ides and pollutants in · -
I to evolutionary char:i=
HUMANS AS AN EVOLUTIONARY FORCE 351
some organisms. One of the most widespread and economically significant exam-
ples of how human-introduced selective pressure has recently influenced the
genotypic and phenotypic characteristics of other species comes from the evolu-
tionary response of insects and mites to pesticides. When the use of insecticide
chemicals such as pyrethroids became widespread in the 1950s there were very
few instances of natural resistance by insects. By the late 1980s many populations
of insect and mite species were almost completely resistant to the common pesti-
cides. Over 400 species of insects were reported to have developed resistance to
one or more classes of pesticides in this 30 year period. Individuals which pos-
sessed genes producing resistance to pesticides were selected for because they
were able to survive and reproduce, while individuals without such genes were
killed by the pesticides.
Animal and Plant Domestication
The most important examples of human impact on the evolution of other species
come from the domestication of plants and animals. In addition, the process of
domestication has been of critical importance to the geographic spread and pop-
ulation increase of humans over the past 10,000 years. Domesticated plants and
animals are the basis of agriculture, and without agriculture the complex, techni-
cally innovative societies and large human populations that exist today could not
have come into being. Agriculture allowed people to become sedentary (living
for a prolonged period in one place) , establish permanent villages and towns, and
develop stratified societies that included specialized and dedicated segments
such as farmers, artisans, soldiers, religious leaders, educators, and governors.
Because of the crucial importance of domestication and agriculture, some bio-
geographers, including the influential figure, Carl 0. Sauer of Berkeley, have
made the patterns and processes of plant and animal domestication, and resulting
landscape change, a central focus of their work. Understanding the origins of
domesticated plants and animals is of more than strictly academic interest. The
ancestral species of crop plants may often still be found in areas where early
domestication occurred and may provide important crop strains and genetic
resources for breeding and agriculture today. It has been shown, for example, that
careful use of native varieties of potatoes can aid in maintaining sustainable agri-
cultural yields in Andean farming regions. Let's explore the fascinating and
important area of biogeographical research into domestication and the develop-
ment of agriculture.
Domestication refers to the process by which plant and animal species
come to depend on humans for survival while in turn providing humans with
practical or other benefits. Many domesticated species, such as corn (Zea mays)
provide food. Others, such as garden roses (Rosa spp.) are used for nonpractical
purposes such as providing color in gardens. During domestication, humans
selectively breed plants or animals to propagate and enhance desired traits. These
traits can include docility and trainability, increased meat, milk, or wool produc-
tion in animals, or increased fruit size and production in plants. In many cases, the
resulting domesticated species are markedly different in morphology and behav-
ior from their wild progenitors. For example, the wild ancestor of domesticated
corn is probably a small grass called teosinte (Zea mexicana, according to some
authorities and Zea mays mexicana according to others). Some of the first
352 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

archaeological evidence of corn domestication comes from ears and kernels p:-~ and unconsci01
served in caves in the Tehuacan Valley of Mexico. Through a process of natu: _ was done inad1
mutation and then domestication that extended from about 7000 to 500 ye-- tion practiced 1
ago, people in the highlands of southern Mexico produced increasingly la r~ ~ selection when
eared and large-seeded domestic corn from this small-eared and small-seed=- biological envi1
ancestor (Fig. 12.1). Domesticated corn provides several advantages 0'- Homo sa]
teosinte for humans. Domesticated corn has larger and more nutritious kernc domestication <
and more kernels per ear. It has been estimated that a single kernel of mod - were hunters a
corn contains as much nutrition as an entire ear of the earliest domesticated va:- nuts, carrion, ai
ety. In addition, the kernels of domesticated corn do not fall off the ear eas:_ establish perm
This makes it simple and efficient to harvest corn. However, the adherence of ---= response to th<
kernels to the ears means that domesticated corn cannot disperse its own sec hunting and g<
Corn seed dispersal must be done by humans. As a result, domesticated com _ areas, such as p
entirely dependent on humans for its survival as a species. If humans did ture over the 1
plant and tend domesticated corn, it would go extinct almost immediately. Ma:: America, Afric.
domesticated plants and animals would not survive without human interventi =. The earlit
Although domestication and agriculture are tightly intertwined, the word "ag::--- The domestic c
culture" has a different specific meaning than domestication. Agriculture is .-...: The modern de
cultivation of domesticated plants and animals for human use. created by dorr
Charles Darwin was much interested in the selective pressures that hum ~ ­ dog might have
apply to species during domestication. He saw that the domestication proc?-< hunt and live ir
was much like natural selection in nature, except that during domesticati =. game and perh
human actions were the most important selection force. Darwin recognized tl:..:. easy to imagim
humans applied both conscious selection when they purposefully bred Wolf cubs that
removed certain individuals to achieve desired genotypically controlled trac:. human captors.
would have wm
as loyalty and r
captive wolf pc
what many con
sapiens sapiens
evolution of tht
15,000 to 11,00!
genetics of moe
of the wolf m
100,000 years <
date of dog do1
huahuas to hug
resulting evolut
Some par
mal domesticat
Asia. The ance~
Mountains of
humans startin1

]I
em
size and have c
wild sheep con
remains of she
10,000 years ag
hunters were ki
5,000 BC 2,000 AD
about 10,000 yc
FIGURE 12.1 The gradual development of modern corn from teosinte. male goat and
)N
from ears and kernels pre-
trough a process of nature_
n about 7000 to 500 ye ~
·oduced increasingly large-
'lll-eared and small-seede.:
several advantages o,·e:-
nd more nutritious kern e ~
a single kernel of mode:::
earliest domesticated var:-
, not fall off the ear easi:. ·
Never, the adherence of tt::
mot disperse its own seeCs.
esult, domesticated corn ::
species. If humans did nC>-
almost immediately. MaL_
ithout human interventio:::.
1tertwined, the word "ag:::-
tication. Agriculture is tt::
nan use.
:tive pressures that hum a:::.
the domestication procn
:hat during domesticat ic:::.
ce. Darwin recognized t'--
lley purposefully bred --
atypically controlled trc.:_
--,
2,000 AD
>m teosinte.
HUMANS AS AN EVOLUTIONARY FORCE 353
and unconscious selection when maintenance or removal of certain genotypes
was done inadvertently. We can consider both conscious and unconscious selec-
tion practiced by humans to be forms of artificial selection in contrast to natural
selection where the selective pressures are generated by the natural physical and
biological environment.
Homo sapiens sapiens are the only hominids conclusively shown to practice
domestication and agriculture. Earlier hominids, and early Homo sapiens sapiens,
were hunters and gatherers who relied on naturally occurring vegetation, fruits,
nuts, carrion, and game for subsistence. Hunters and gatherers generally do not
establish permanent settlements such as villages, They move their camps in
response to the movement of game and changes in the season. Subsistence by
hunting and gathering is still practiced by people in some extremely remote
areas, such as portions of the Amazon Basin. Prior to the spread of European cul-
ture over the past 500 years, many people in areas of North America, South
America, Africa, and Australasia lived by hunting and gathering.
The earliest domesticated species was probably the dog (Canis familiaris).
The domestic dog appears to be a direct descendant of the wolf (Canis lupus).
The modern dog, encompassing all of the different breeds we have today, was
created by domestication by humans. We can speculate how domestication of the
dog might have occurred. Wolves are relatively intelligent and social animals that
hunt and live in packs. It is likely that humans and wolves competed for the same
game and perhaps for protection in caves. Because of this close proximity, it is
easy to imagine how wolf cubs could have been acquired and raised by humans.
Wolf cubs that were particularly vicious would perhaps have been killed by their
human captors. Other cubs that were not inclined to bond with the human group
would have wandered away from the humans. Eventually, selection for traits such
as loyalty and nonaggressive behavior toward humans would be dominant in the
captive wolf population. After thousands of years, domestication has produced
what many consider a new species, the domestic dog. Fossil evidence from Homo
sapiens sapiens sites in Europe suggests that the domestication of the wolf and
evolution of the modern dog were taking place in Eurasia and North America by
15,000 to 11,000 years ago. Interestingly, a molecular clock estimate based on the
genetics of modern dog breeds and wolves suggests that the initial domestication
of the wolf may have been started by Homo sapiens neanderthalensis over
100,000 years ago (see Technical Box in Chapter 11). Regardless of the initial
date of dog domestication, all of the amazing breeds of dogs, from the tiny chi-
huahuas to huge Saint Bernards were developed through artificial selection and
resulting evolutionary change caused by humans.
Some particularly good fossil evidence of the rate and timing of early ani-
mal domestication comes from the history of goats and sheep in southwestern
Asia. The ancestors of the modern domestic goats and sheep lived in the Zagros
Mountains of Iran and Iraq and were hunted by Neanderthals, and modern
humans starting at least 40,000 years ago. Modern goats and sheep are smaller in
size and have different horns and coats than their wild ancestors. The coats of
wild sheep contain far less wool than is the case for domesticated sheep. The
remains of sheep and goats found at hominid sites that are older than about
10,000 years ago show no evidence of domestication. In addition, it is clear that
hunters were killing and eating female and male goats and sheep of all ages. After
about 10,000 years ago, there is a shift toward a greater abundance of pre-adult
male goat and sheep skeletons at hominid sites. It is possible that at this time
354 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

humans realized that by taking pre-adult rams and sparing females and sexuall~
mature rams for reproduction, the populations of sheep and goats would remai::
abundant for hunting. Many parts of the world have similar hunting laws toda:
allowing, for example, the hunting of male deer but not female ones. Shortly fo:-
lowing the shift to selective hunting of young males, the development of domes -
cated sheep and goat species is evidenced by decreases in the size of goat an.:
sheep skeletons and changes in horn morphology. It seems clear that shortly aft -
10,000 years ago people were keeping herds of these animals. By 9000 years age Ocampo
goat and sheep were being introduced and raised by people at sites that were o :- Cave
(7000 BC)
side the natural ranges of wild goats and sheep. Squash?
So, it is thought that human interaction with goats, sheep, and other dom e~ Gourd?
ticated herding animals first took the form of hunting, then selective hunting, a c.: Scarlet
Runner Tehuc
finally herding. In some locations, such as the Zagros Mountains, goat and shee; Bean (6000
hunters may have begun the process of domestication by driving small wild her ·- (4000 BC) Maize
Common Squa<
into areas closer to human habitation or where the herds could be supported b ~ Bean Gourc
water and vegetation but also be hunted easily. True domestication, caused : (5000
Maize
artificial selection and resulting genetic changes, could ensue only if natural wil.:
Aya
populations were not allowed to breed with the herded ones. Natural populatio - (60(
may have been kept away from human herds either by fencing or eradication.:: Lim
Con
is likely that at first inadvertent actions by people kept the wild populations awa:
from herded animals. Humans may have observed how some of the herded in ·_
viduals had desirable traits, such as long coats, high milk production, or larg;:
amounts of meat, and selectively bred animals in the herds that had those trai " FIGURE 12.2
Evidence from archaeological sites in the Zagros Mountains and observations o: around the worl<
recent and modern primitive herding peoples, such as reindeer herders in nont- (after Simmons,
ern Eurasia, appear to support this model of early domestication.
Over the past 10,000 years, many animals have been domesticated in differ-
ent parts of the world, and we often have reasonable archaeological informatio:.
on the early period of domestication (Fig. 12.2). For example, pigs (Sus scrofa north to easte
and cattle were domesticated in southwestern Asia at about the same time as Israel (Fig. 12.
sheep and goats. The horse (Equus equus) and bactrian camel (Camelus bactri- Crescent. Tod;
anus) were domesticated in central Asia between 5000 and 3500 years ago. Wate:- wheat, barley,
buffalo (Bubalus bubalis) were domesticated in India and adjacent southeasten: Crescent. Man
Asia about 4500 years ago. Llama (Lama guanicoe) were domesticated in Soutt. and agricultun
America about 3500 years ago. In areas such as central Eurasia, portions of the the remains of
Near East, northern Eurasia, and parts of Africa, human cultures developed tha: the melted anc
were organized around herding domesticated animals such as sheep, horses, cat- Many tells ha'
tle, camels, and reindeer. These people supplemented the meat and milk obtainec artifacts dating
from their herds with hunting and gathering. The human herders migrated an ings and artifa
moved along with their herds during seasonal changes in grazing areas. This for and animal rer
of subsistence is called pastoral nomadicism and is still practiced in some areas o. development c
the world today. nent villages. 1
Some of the earliest evidence for the domestication of plants comes from that dates bad
southwestern Asia and dates from about the same period as the domestication of the early dome
goats and sheep. Indeed, a variety of important food plants, including whea t plant cultivatic
(Triticum), rye (Secale cereale), barley (Hordeum vulgare), lentils (Lens culi- called Tell Abt
naris), chickpeas (Cicer arietinum), and peas (Pisum sativum) were all domesti- early as 10,900
cated in the Near East about 11,000 to 9000 years ago. In southwestern Asia, the Htiytik located
archaeological sites with evidence of early plant domestication occur along an ment supporte
inverted crescent that extends from the mouth of the Tigris and Euphrates rivers. bined with hun
>N HUMANS AS AN EVOLUTIONARY FORCE 355

>aring females and sexuall_

Y""
r
Cayonu . Palegawra
ep and goats would remai.::: (7000 BC) Shamdar Cave ~
Einkom (9000 BC) (12000 BC) ~o
similar hunting laws tod a' Pea Sheep? Dog
ot female ones. Shortly fol- 1
Lentil o Central Asia
te development of dome st~ ­ Sheep /(3000 BC)
Pig Horse
.ses in the size of goat an.: Goat? .· ···""""' ....,_ (1500 BC)
:ems clear that shortly afte:- D~_
l[:j
· -..... Bactrian "'
:mimals. By 9000 years age
~ople at sites that were ou:-
Ocampo
Cave
(7000 BC)
' camel

;;;-·
Squash? q Pan Po
ts, sheep, and other dome5- Gourd? q. (3750 BC)
then selective hunting, an: Scarlet / )Foxtail
(}, Millet
Mountains, goat and shee_: Runner Tehuacan ; · )_Cattle
by driving small wild herc'-
~rds could be supported I _
: domestication, caused _
Bean
(4000 BC)
Common
Bean
(6000 Bq
Maize?
Squash
Gourds
(5000 BC)
'""'- -~

· South
1 America
Egypt
. .Highland (3000 BC)
~ Donkey
(2000 BC) Jericho
(2000 BC)
Dromedary
Jarmo ~Ali Kosh
(:J
~_! Cabbage

(6700 BC) (7500 BC)

i ensue only if natural \\i.::
i ones. Natural populatior..::
y fencing or eradication.=
• the wild populations awe.
w some of the herded inC.-
milk production, or l ar~::
herds that had those trai:s.
mtains and observations
reindeer herders in norr---
nestication.
een domesticated in diffe:--
lrchaeological informat ic:
example, pigs (Sus sera_-'.;
1t about the same tim e ~
an camel (Camelus bac-:r-
' and 3500 years ago. Wa:::--
and adjacent southeaste:-::.
ere domesticated in Soe:..:
a! Eurasia, portions of t:.=
an cultures developed tt2.
such as sheep, horses. c-2. ·-
he meat and milk obtair..:-_
nan herders migrated a:-..:
in grazing areas. This fo:-
practiced in some areas -
tion of plants comes frc-
od as the domesticatio
,d plants, including whe_
tlgare), lentils (Lens cu.
:ativum) were all dom e5:
In southwestern Asia. L::
testication occur along =-
igris and Euphrates ri\
Maize Guinea Pig (7000 BC) Einkom Sheep
Ayacucho (1500 BC) Einkom Emmer Goat?
(6000 BC) ~ Llama
Emmer \J
Barley Barley • (7000 BC)
Lima Bean Alpaca
Pea Lentil Einkom
Common Bean Pea Emmer?
Lentil
Centers of domestication • Fertile crescent
FIGURE 12.2 Areas and minimum ages for domestication of selected animals and plants
around the world . Important centers of domestication and agricultural origins are indicated
(after Simmons, 1989).
north to eastern Turkey, and then south to the coastal regions of Lebanon and
Israel (Fig. 12.2). This region of early plant domestication is called the Fertile
Crescent. Today, many of the wild ancestors of domesticated plants such as
wheat, barley, lentils, peas, and chickpeas are found in the vicinity of the Fertile
Crescent. Many of the archaeological sites with evidence of early domestication
and agriculture occur as low mounds, locally called tells. The tells are formed by
the remains of human habitation. In most cases, these remains are dominated by
the melted and broken mud bricks of structures such as houses and village walls.
Many tells have been occupied and abandoned many times and have layers of
artifacts dating from many different periods. In some instances, the earliest build-
ings and artifacts date from the very dawn of domestication. Studying the plant
and animal remains from the tells shows how the process of domestication and
development of agriculture went hand in hand with the development of perma-
nent villages. The biblical city of Jericho (Fig. 12.2) is represented by such a tell
that dates back to at least 10,000 years ago and provides important evidence for
the early domestication of wheat. Some of the earliest evidence of domesticated
plant cultivation and use comes from a site along the Euphrates River in Iraq
called Tell Abu Hureyera. Finds from this site suggest that rye was used there as
early as 10,900 years ago. Perhaps the best known settlement of this age is Catal
Hiiyiik located in eastern Turkey (Fig. 12.2). By about 9500 years ago, this settle-
ment supported some 10,000 inhabitants based on rudimentary agriculture com-
bined with hunting and gathering.
356 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

We can speculate how hunting and gathering peoples living in the Ferti..:: and wild emr
Crescent and other parts of the world initially domesticated plants. These pea ~= 10,000 years
likely used the wild ancestors of domesticated plants as food sources for a ve=-=- these species,
long period prior to domestication. Flint sickle blades from the Fertile Cresce::.- native range
date back at least 12,000 years ago and suggest the harvesting of the wild anc = wheat (T. ae
tors of wheat. Many of the wild progenitors of domesticated plants favor dis- between emn
turbed environments such as burned areas or areas with exposed mineral soLs. a hybrid, mod
These plants may have seeded into land near camp sites that were disturbed . 42 compared
human activity. Humans would have been able to observe the ecology of the:= of einkorn th:
plants at close quarters. They could have observed on what types of soils the foo: seeds than nc
plants grew best and which varieties of food plants provided the most seeds a- also has seed
fruit. The domestication of plants likely began as selective cultivation of \\ · - The naked se
food plants, perhaps by transporting seeds to new camp sites or changing lo logical evider
environmental conditions to suit favored plants. Hunting and gathering peop::c cent by about
who lived in more recent times have been shown to practice such cultivation. F in the wheats
example, Australian aborigines were known to have replanted small wild yar::..! itous acciden
after harvesting the larger ones. They also used burning and flooding to alre- hybrids and
environmental conditions to benefit the growth of certain edible plants. Eart genetic event
human cultivators would also have noticed that some individuals of the sarr::c wheat would
species produced larger seeds and fruits or more abundant seeds and fruits. Th e~ survive.
may have selectively planted seeds from those. AlthOU!
As in the case of animals, restricting the flow of wild genes into the pia;:: people aroun
population that is being domesticated is very important. However, it is much m o~ seen, corn d•
difficult to restrict the flow of pollen than it is to control contact between cui ·_ America. Lirr
vated and wild animals. Pollen is transported by the wind or insects and can ea s il.~ and South A1
move from wild to domesticated plants growing near each other. Interestino :· come from A
many of the ancestors of important early domesticated plants, such as wheat, typ:- haps adjacen
cally self-pollinate more frequently than they are fertilized by other individuals. bage (Brassic
Therefore, it is less likely that pollen from a nonfavored wild stock of plants waul.::
introduce genes into plant populations that humans were cultivating.
Questions
Seeds from archaeological sites in the Fertile Crescent show that betwee::
11,000 and about 8000 years ago, cultivation of wild plants led to domesticate"' Two question
varieties of grains such as wheat and barley that had larger seeds, more nutritiot:S archaeologist
seeds, larger seed production, and in many cases, seeds that more strongl:· and second, '
adhered to the plant. An important example of such evolutionary change due tc Surprisingly, 1
domestication is provided by the early history of wheat. Because of its nutritious and take up a
seeds, high yield, and wide environmental tolerance, wheat is the most importa : ural environr
domesticated grain in the world today. About 20% of the calories consumed by kilocalories o
the entire human population of the world are derived from wheat. Archaeologi- In contrast, t
cal evidence and data from the modern genetics of domesticated and wild whea~ protecting en
provide information on the history of this important crop plant. A number of wil - ries of energ)
species of the wheat genus are found in southwestern Asia today. One of the ear- energy perspt
liest cultivated species was probably wild einkorn wheat (Triticum boeoticum). subsistence a
which through artificial selection gave rise to domesticated einkorn (T. monococ- ing leaves les
cum). Domesticated einkorn has larger seeds that are more strongly attached to gathering in r
the rachis (central stem) of the plant compared to wild einkorn. A similar history available to l
is apparent for domesticated emmer wheat (T. dicoccum), which arose from the flavorful , or e
wild species T. dicoccoides. Some taxonomists now combine both einkorn species ering is a relc
together as T. monococcum and put both emmer species together as varieties of a Two factors n
species called T. turgidum. Archaeological evidence indicates that both wild einkorn culture. The 1
 >N
·eoples living in the FertiJ-
icated plants. These people
as food sources for a vef\
s from the Fertile Crescen:
trvesting of the wild ance5-
testicated plants favor di5-
.vith exposed mineral soils.
ites that were disturbed ....
'serve the ecology of thes::
what types of soils the focx:
1rovided the most seeds o~
~lective cultivation of wil-
mp sites or changing locc.
tting and gathering peop[::
·actice such cultivation. Fe~
replanted small wild yar.s
ning and flooding to altc::-
ertain edible plants. Ear:_
1e individuals of the saiL::
dant seeds and fruits. The-
f wild genes into the p i a;:~
t. However, it is much m o::-~
trol contact between cuJ:.::-
nd or insects and can easi:
r each other. Interestin~ :
plants, such as wheat, t ~·~.-
ilized by other individu<U..
I wild stock of plants woU:.:
:re cultivating.
rescent show that betwee::.
plants led to domesticate.:
rger seeds, more nutritim
seeds that more strong._
volutionary change due : -
tt. Because of its nutritio:......
heat is the most impon a;:
· the calories consumed t
from wheat. Archaeolos-
mesticated and wild whe~
op plant. A number of \\i.:.:
1\sia today. One of the ec:-
teat (Triticum boeoticw r.
ated einkorn (T. monoco.-
more strongly attached : -
einkorn. A similar histo:-
tm), which arose from rl:.=
1bine both einkorn spec:::--
~s together as varieties o:: _
ates that both wild einko:-
HUMANS AS AN EVOLUTIONARY FORCE 357
and wild emmer wheat were being cultivated in the Fertile Crescent by almost
10,000 years ago. In addition to cultivation within the natural distributions of
these species, wheat had also been introduced and cultivated in areas outside its
native range on the higher floodplains of the Euphrates River. Modern bread
wheat (T. aestivum) appears to be a hybrid produced by cross-fertilization
between emmer wheat and a wild wheat species (T. tauschii). In addition to being
a hybrid, modern bread wheat is also a polyploid, having a chromosomal count of
42 compared to 28 for emmer wheat, which is itself probably a polypolid progeny
of einkorn that has a chromosome count of 14. Polyploid plants often have larger
seeds than normal diploid varieties. Aside from having larger seeds, bread wheat
also has seeds that lack the plumed bracts that cover the seeds of other wheat.
The naked seeds of bread wheat are much easier to process for food. Archaeo-
logical evidence shows that bread wheat was being cultivated in the Fertile Cres-
cent by about 8000 years ago. Events such as the mutation that caused polyploidy
in the wheats and cross-fertilization between emmer and T. tauschii were fortu-
itous accidents that improved the food value of wheat. Through selecting these
hybrids and polyploids for cultivation, humans capitalized on these chance
genetic events. As is the case with corn, should humans cease to cultivate it, bread
wheat would not be able to disperse adequate numbers of seeds for the species to
survive.
Although many important domestic species come from southwestern Asia,
people around the world domesticated plants (Fig. 12.2). As we have already
seen, corn domestication occurred in Central America and spread to South
America. Lima beans, common beans, potato, and squash also come from Central
and South American areas. Sorghum, finger millet, and some yams (Dioscorea)
come from Africa. Domesticated Asian rice (Oryza sativa) is from India and per-
haps adjacent portions of eastern Asia. Foxtail millett (Setaria italica) and cab-
bage (Brassica oleracea) come from China.
Questions of the Orig in and Spread of Agriculture
Two questions regarding domestication have long fascinated biogeographers and
archaeologists. First, why did hunting and gathering people turn to agriculture,
and second, what was the geographic pattern of agricultural origin and spread?
Surprisingly, the impetus for people to abandon a hunting and gathering lifestyle
and take up agriculture is not that clear. Experiments have shown that in the nat-
ural environment of southwestern Asia a hunter-gatherer can obtain up to 50
kilocalories of food energy for every 1 kilocalorie of energy expended on work.
In contrast, because of energy needed for field preparation, sowing seeds, and
protecting crops, low-technology subsistence farming nets only about 17 kilocalo-
ries of energy for every 1 kilocalorie expended by the farmer. From a per capita
energy perspective, there seems no reason to shift from hunting and gathering to
subsistence agriculture. In addition, it has also been shown that subsistence farm-
ing leaves less free time for recreation and other activities than does hunting and
gathering in relatively resource-rich environments. Finally, of the fruits and seeds
available to hunter-gatherers, grains such as wheat are not the most nutritious,
flavorful, or easiest to process. From the individual's viewpoint, hunting and gath-
ering is a relatively attractive mode of life compared to subsistence agriculture.
Two factors may help explain how and why hunters and gatherers turned to agri-
culture. The first is the external pressure caused by environmental change, and
358 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

the second is the intrinsic dynamics of human populations, particularly in the fc_ environmental
of pressure from growth. example, the de
The premise that changes in human culture are driven mainly by changes _ 8000 years ago,
environment is often referred to as environmental determinism. In addit i _ dent with this J:
environmental determinists believe that differences in present cultures an instantane01
largely be explained by geographic differences in environment. Such theor~=-­ Fertile Crescen
were particularly popular with geographers, anthropologists, and biologists in ,:. ~ shift from depe
nineteenth and early twentieth centuries. In the mid-twentieth century, some mals such as g
the most divisive debates within academic geography erupted over the validity grains. During t
environmental determinism. In the early twentieth century, the archaeologist . ticated in the n
G. Childe advanced the Oasis Theory to explain how environmental change :c: from the early
to the domestication of plants and animals and the development of agricultur tributed as muc
southwestern Asia and adjacent Africa. Childe supposed that the climate that once dome
southwestern Asia and northern Africa had been cool and wet during Pleistoc - = is a cultural dyr
glaciations. He proposed that during the last glacial maximum, about 20,000 yec._ The isola1
ago, the climate of the region was moist, vegetation was lush, and game plentili and game has
As the global climate warmed at the close of the Pleistocene, southwestern As lifestyle. Agricu
and northern Africa became warmer and drier. Under these conditions, pro d · ~­ in the context <
tive vegetation, game animals, and humans all became restricted to sites ne:o - other humans. l
water, such as river valleys, lake shores, and oases. By being forced into such cl o:~ farming is grea1
contact, humans were able to observe the ecology of plants and animals that th~_ land than does
relied upon. Eventually, this led to the selection of species and individuals for ct:..- ported on a srr
tivation. As attractive as Childe's original hypothesis sounds, it has a fatal fia storage of grail
Since the 1960s, fossil pollen records have been recovered from the sediments c: environmental
a number of lakes in southwestern Asia. Similar records have been obtained fro- yields of crops.
dried up lakes in Egypt. These records show that the climate of the last glacic... where, or in ex1
maximum was actually quite dry. Vegetation in southwestern Asia and adjace;:;.- Third, in most <
Africa was sparser and more desertlike during the late Pleistocene than it w~ tions. They mm
11,000 to 8000 years ago when initial domestication and the development of agr:- changes in plan
culture occurred. This is exactly opposite to what Childe had envisioned when h~ young, the age·
constructed the Oasis Theory. Childe's hypothesis was discredited. such movemen·
The possibility that climatic change may have prompted plant and animC: over hunting a
domestication and the development of agriculture in southwestern Asia is no" remain near an
being reexamined. Evidence from a number of different sources suggests that th~ meant that nat1
general pattern of warming and increasing moisture during the Pleistocene- siderations, it h
Holocene transition was interrupted by a rapid and pronounced episode o:" opment of agr
cooler and drier conditions around 12,000 years ago. This episode correlates witt sapiens sapiens
the Younger Dryas cooling event. The Younger Dryas event was centered on th mals allowed f,
North Atlantic region but affected the climate of much of the world. The Younge~ trips would be
Dryas was likely caused by the rapid drainage of glacial meltwater from North area would be r
America into the North Atlantic Ocean. It has been suggested that decreases in stratified societ
plant and game resources during this period of rapid climatic change may haY ally subject oth
caused extreme stress to hunters and gatherers. The movement toward domesti- that people we
cation and the development of agriculture may have been a response to the rapid feedback was s<
decline in natural resources. cation of agricu
When we consider the world at large, two important facts argue against envi- The geog1
ronmental determinism as the sole or universal factor that led to the domestica- agriculture hav
tion of plants and animals and subsequent development of agriculture First, the first half of th
domestication of plants and animals occurred at different times in different part domestication
of the world. It is often difficult to find any convincing evidence for significant spread from th
\1
)ns, particularly in the fa c~
riven mainly by changes i.L.
!leterminism. In additio
in present cultures ca.;:
vironment. Such theoriQ
)gists, and biologists in th.:
wentieth century, some c:
rupted over the validity c:
1tury, the archaeologist \ -
~nvironmental change !e..:
elopment of agriculture i:.
•osed that the climate c:
nd wet during Pleistoce rr~
<imum, about 20,000 yea;-s
slush, and game plentift:.
[Ocene, southwestern As:.:
these conditions, produ:-
te restricted to sites ne:=:
:ing forced into such clos::
<:~nts and animals that th.:
[es and individuals for Cl-·_
munds, it has a fatal fia
red from the sediments .
; have been obtained fro-
climate of the last gl ac-~
1estern Asia and adj ace-
.e Pleistocene than it \Y.:_
I the development of ag::--
e had envisioned when ::::
liscredited.
ompted plant and ani~
;outhwestern Asia is nc
t sources suggests that t.:.=
during the Pleistocer:.::-
. pronounced episode _
1is episode correlates \\-::-
~vent was centered on ;~=
)f the world. The You n~::­
[al meltwater from No:--
ggested that decreases -
:limatic change may he:·::
wement toward domes:.:-
~n a response to the ra;: •.:.
tt facts argue against en'--
hat led to the domest i;::
tt of agriculture First. ~­
nt times in different pc.:-
g evidence for signifi ~- -
HUMANS AS AN EVOLUTIONARY FORCE 359
environmental changes during initial domestication in these other regions. For
example, the domestication of corn in Central America did not begin until about
8000 years ago, and there is no evidence of major environmental changes coinci-
dent with this process. Second, domestication and the shift to agriculture was not
an instantaneous event. Evidence from Tell Abu Hureyra and other sites in the
Fertile Crescent shows that it took several hundred years for the inhabitants to
shift from dependence on wild plants, including wild einkorn wheat, and wild ani-
mals such as gazelle (Gazella subgutturosa) to goats, sheep, and domesticated
grains. During the next two to three thousand years, additional plants were domes-
ticated in the region. In Mexico and Central America, it took perhaps 5000 years
from the early start of the domestication of corn until agricultural crops con-
tributed as much as hunting or gathering to the food used by humans. It appears
that once domestication and the development of agriculture is set in motion, there
is a cultural dynamic that propels it, regardless of environmental change.
The isolated hunter and gatherer in an environment rich with food plants
and game has little obvious inducement to develop and adopt an agricultural
lifestyle. Agriculture does, however, provide several advantages when considered
in the context of an increasing population of humans that are not isolated from
other humans. First, although the effort required to obtain energy via subsistence
farming is greater, a cultivated landscape provides much more energy per unit of
land than does the harvesting of wild plants. More people can thereby be sup-
ported on a smaller area of land. Second, agricultural systems that permit the
storage of grains or the keeping of herds allow people to use these reserves if
environmental factors, such as drought or a particularly cold year, decrease the
yields of crops. Hunters and gatherers can only move on and try their luck else-
where, or in extreme cases perish, during periods of difficult climatic conditions.
Third, in most environments, hunting and gathering requires mobility of popula-
tions. They must be able to move to follow game or take advantage of seasonal
changes in plant availability. It is difficult to make large movements with the very
young, the aged, and the infirm. In addition, as human populations increased,
such movements would bring different bands of people into contact and conflict
over hunting and gathering grounds. It would be advantageous for a band to
remain near and defend a particularly good fishing or hunting site, even if this
meant that natural plant resources were less than optimal. In view of these con-
siderations, it has been suggested that one crucial factor in prompting the devel-
opment of agriculture was the growth and geographic expansion of Homo
sapiens sapiens populations during the Holocene. Domesticated plants and ani-
mals allowed for a sedentary life, where the danger of conflict during foraging
trips would be reduced and population size for defending resources in a small
area would be maximized. With permanent habitation and larger populations, the
stratified societies that developed would increase in power to defend and eventu-
ally subject other groups. Permanent settlements of large populations also meant
that people were now tied to agriculture in order to sustain society. A dynamic
feedback was set in place that continued to drive domestication and the intensifi-
cation of agricultural production.
The geography of initial plant and animal domestication and the spread of
agriculture have long been a topic of intense interest to biogeographers. In the
first half of the twentieth century, a number of researchers speculated that
domestication and agriculture developed in the Fertile Crescent and then
spread from there throughout the world. Alternatively, the geographer Carl 0.
360 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

Sauer proposed that domestication and agriculture first developed in southeas::- and northern c
ern Asia and then spread from there. To support such claims, some scientis:. people, of the
have pointed to the antiquity of agriculture in the Fertile Crescent relative : complexes sud
other parts of the world. Evidence for a southeastern Asian origin came fro= ument in Ariz•
the fact that the bottle gourd (Langenaria siceraria) was an early domesticate ·- three crops. Ne
both Mexico and Thailand. The problem with such theories that envision or::: presence of cer
and only one, center of the origination of domestication and agriculture is t t~ west and Mesc
there is no convincing archaeological or historical evidence to support · :: diffused along
movement of people, crops, or technologies from the Fertile Crescent or As_ The spre;
over great distances in the early Holocene. The initiation of corn domesticatio- and agricultun
in Mexico or potato domestication in South America began about 8000 yea: . the world. In S<
ago and appears to be local innovations that took advantage of existing plan::. fied societies 1:
Similarly, the domestication of finger millet in Africa or rice in Asia also appea:. societies were
to have been local innovations. There is no reason to suspect that people _ Africa, Europe
many parts of the world were not all capable of independently domestica til:~ eastern North,
plants and animals and developing agriculture. of domesticate
Despite the universal ability of humans to domesticate plants and anim<L:3 developed. No1
and to develop agriculture, both archaeological and genetic evidence from cr : Eurasia and A
plants shows that certain areas of the world were centers of domestication for t :: lead to further
early development of agriculture. Other areas either developed agriculture aft -: western North
contact with people and crops from one of these centers, or they never develope - such as wheat
it at all. Notable centers of initial plant domestication and agricultural develor- Today, for exar
ment include southwestern Asia, a broad band of central Africa, India, and eas:- while China is
ern Asia, Central America, and South America (Fig. 12.2). People in areas such co Domestic
northern Europe, North America, and Australia practiced agriculture only after i: primarily beca1
was introduced from elsewhere. cated varies frc
The geographic diffusion of agriculture from centers of domestication h~ contained man
long been of interest to biogeographers and anthropologists. The geographic di:- barley, rye, len1
fusion of domesticated plants and animals in Europe and pre-Columbian No ~ cation. Both gr
America has been studied with particular intensity. Crops such as wheat and ba:- domesticated a
ley moved from the core of the Fertile Crescent to adjacent areas of southwes:- and variety of
ern Asia within a few hundred years. By 8000 years ago, these crops an · development o
agricultural practices had spread to Greece. Between 7000 and 6000 years ago. and South Arne
grains originating in the Fertile Crescent were grown across the European conrj- as corn and po
nent from northern France to close to the Russian border. By 5000 years ago, th- from the lama
growing of grain had spread to the climatic limits of such crops in Scandina ic. explain the slo
and Britain. In North America, the cultivation of corn spread from central Amer- completely agr
ica to the southwestern United States about 3000 years ago and reached its as North Ame1
northeastern climatic limits of corn cultivation in southern Ontario about 70C ply did not pos
years ago. The diffusion of domesticated plants and animals, as well as agricul- the basis for tl
tural practices, in Europe, North America, and other parts of the world was like!~ North Americ<
facilitated by many factors, including the movement of people and the transfer o: that possess the
materials and ideas from one group of people to another. ing ease, and r
The cultivation of wheat and other grains formed the foundation for th corn such imp
rise of cities and civilizations in the Fertile Crescent. The diffusion of these crops remarkable ab
across Europe was followed by the diffusion of technical practices such as pottery animals, they 1
making and metal smelting. In the Americas, the cultivation of corn, along with raw materials <
beans and squash, satisfied a similar function serving as the foundation for the ticate because
Mesoamerican civilizations such as the Maya and Aztecs, and the Inca in South the advent of I
America. (Mesoamerica is an archaeological term that refers to southern Mexico and the splicin
 IN
~st developed in southeasv
tch claims, some scienti ~ - ­
,·ertile Crescent relative tc
n Asian origin came fro
;as an early domesticate i=
heories that envision one
ion and agriculture is the.:
evidence to support tl:::
~ Fertile Crescent or A s2~
tion of corn domestica tio;::
a began about 8000 ye~
vantage of existing planL.•
Jr rice in Asia also app ea: _
to suspect that people ::.
ependently domesticate;
:sticate plants and animL
enetic evidence from cr _ ~
rs of domestication fo r t.:.::
eveloped agriculture af:::-
·s, or they never develope.:
and agricultural develo;-
ral Africa, India, and e as:~
.2). People in areas su ch ~
:ed agriculture only afte:-
1ters of domestication .t:-=..
ogists. The geographic c~
md pre-Columbian No;--
lps such as wheat and
jacent areas of south \\·~-­
us ago, these crops ~.::.
7000 and 6000 years a5
cross the European co;::.
ier. By 5000 years ago. C::
mch crops in Scandinc.•.
pread from central Arn e~
ears ago and reached --
tthern Ontario abou t -
nimals, as well as agri __
rts of the world was like
people and the transfe L"
~r.
:d the foundation fo r :..::..
te diffusion of these ere_
I practices such as pott::;-
;ation of corn, along w::.:
as the foundation fo r :..:::;:
:cs, and the Inca in S o:~.-­
~efers to southern Mex::::
HUMANS AS AN EVOLUTIONARY FORCE 361
and northern central America.) The pre-Columbian Pueblo builders, or Anasazi
people, of the southwestern United States built considerable towns and village
complexes such as those found at Mesa Verde Colorado, Wupatki National Mon-
ument in Arizona, and Chaco Canyon, New Mexico, on the basis of the same
three crops. Not only were these crops introduced from Mesoamerica, but the
presence of ceremonial ball courts in pre-Columbian sites throughout the South-
west and Mesoamerica shows that other aspects of culture and technology also
diffused along with agriculture.
The spread of both nomadic pastoralism, based on domesticated animals,
and agriculture, based on crops and animals, occurred in many other regions of
the world. In some cases, climate, soils, and available crops led to complex strati-
fied societies based on a variety of crops and animals. By 2000 years ago, such
societies were present in Mesoamerica and South America, much of northern
Africa, Europe, and southern Asia. In other regions, such as southwestern and
eastern North America, simpler agricultural societies, based on a smaller number
of domesticated species and often having only semipermanent villages, had also
developed. Nomadic pastoralism was dominant in grassland and desert regions of
Eurasia and Africa. European expansion, following the fifteenth century, would
lead to further expansion of agriculture to areas such as Australia and central and
western North America. It would also lead to the worldwide diffusion of crops
such as wheat from the Near East, rice from Asia, and corn from the Americas.
Today, for example, California is one of the largest producers of rice in the world,
while China is a major producer of corn .
Domestication originated and took hold in some areas and not in others
primarily because the availability of wild plants and animals that can be domesti-
cated varies from geographic region to geographic region. The Fertile Crescent
contained many wild plant and animal species such as the ancestors of wheat,
barley, rye, lentils, chickpeas, sheep, goats, and pigs, which were ideal for domesti-
cation. Both grains, which are high in carbohydrates, and animals which could be
domesticated and furnish meats high in proteins, were available. This abundance
and variety of plants and animals that could be domesticated explains the rapid
development of an agriculture-based society there. Southern Mexico and Central
and South America also had a number of species suitable for domestication, such
as corn and potatoes, but had fewer high-yield plants and no large animals aside
from the lama and alpaca for domestication. These more limited resources may
explain the slow rate of transition from hunting and gathering to a more or less
completely agricultural society in Central and South America. Finally, areas such
as North America, northwestern Europe, Australia, and many other regions sim-
ply did not possess plant and animal species that could be domesticated and form
the basis for the transition to an agricultural society. In northwestern Europe,
North America, or Australia, for example, there are no native species of plants
that possess the combined properties of nutrition, crop yield, harvesting-process-
ing ease, and range of growth conditions that have made wheat, Asian rice, and
corn such important crops around the world. Although humans have shown a
remarkable ability to shape the evolutionary development of many plants and
animals, they have required the proper raw materials to accomplish this. Those
raw materials are the plant and animal species that humans were able to domes-
ticate because of their morphology, physiology, or behavior. Perhaps, now, with
the advent of biochemical techniques that allow the alteration of genetic codes
and the splicing of new genes into organisms, we will shape species and fashion
362 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

new ones that even more specifically suit our perceived needs. The wisdom humans madE
such manipulations of the evolutionary process remains highly controversial. extinct, huma
that coevolve
evolve adapti1
HUMANS AS A FORCE OF EXTINCTION have seen in C
cially introduc
The growth and geographic spread of Homo sapiens sapiens populations h_ them, the pre~
been a remarkable episode in the biogeographical history of the earth. Howe\- : by the new p
this growth and expansion has not been without significant cost. Perhaps tt= Hominids live
most regrettable byproduct of human success has been the global extinction oi _ tory. Other ar
great many plant and animal species. Unfortunately, the rate at which our speci- = past 60,000 ye
has caused other species to go extinct appears directly tied to human populati - the presence c
size. As human population has exploded over the past few centuries, so has t = causing signifi
number of plants and animals that we have lost forever to extinction. Nil : dence for the
Eldredge has compared the magnitude of these recent extinctions to five ear ti~­ areas such as
mass extinctions represented in the geologic record. These are the Up - where people
Ordovician, Upper Devonian, Permo-Triassic, Upper Triassic, and Cretaceo -- of these exam
Tertiary extinction events (see Chapter 9). He coined the term sixth extinction - Fossil ev
refer to the episode of extinctions over the past 10,000 years or so that have be - 12,000 years a
caused by humans. All of the five earlier mass extinctions were caused by car.:o- that is remarl
strophic geologic events or strikes by extraterrestrial objects such as meteo - which the adt
Although it is arguable that the number of extinctions over the past 10,000 yea;: often referrec
does not yet equal the massive loss of species during these earlier geologic p e~­ tained specie~
ods, neither have these earlier extinctions been caused by the actions of or::o columbi) and
species-that species being Homo sapiens sapiens. horses (Equuj
Human beings have caused the extinction of other species through b o~ shastensis), sa
direct and indirect processes. In a number of cases, direct predation by huma - simus). The yc
has led to extinction. In other cases, the removal or reduction of prey species h - suggest that al
created trophic cascades that have led to the extinction of predators. Huma - other species,
have also introduced alien species into environments, and these have driY :::. the 79 large m
other species to extinction through predation or competition. Land clearance he.> ago, only 22 SJ
caused species of plants and animals to go extinct due to loss of habitat. In man: nent 500 year
cases, extinctions have been due to combinations of factors, such as aggress i\-- mammal spec
hunting coupled with the destruction of habitat due to land clearance. mammals rem
Extinctions caused by modern humans can be divided into two general cat- muskox (Ovit
egories. The first category is a series of prehistoric extinctions caused by the initi<L mals did not ~
geographic expansion of human beings from Africa and Eurasia. The second ca - mammal extir
egory is a series of historic extinctions caused by the combination of Europear: Two setf
colonization and socioeconomic expansion, and the rapid global increase i.;l extinctions th:
human population. Let's look at both types of events. first hypothes
hypothesis, lm
vegetation at 1
Prehistoric Extinctions
This was beca
Prehistoric extinctions caused by the initial geographic expansion of moderL tions, had larg
humans can be regarded as being similar to the introduction of a nonselective (or populations, a
euryphagous) predator into a new environment. Much of the remarkable success same genera, <
of modern humans in terms of geographic range and population size can be interglacial tn
attributed to our relatively wide generalized niche. Humans are omnivores the last glacial
adapted to take advantage of a number of different prey species and plants. The The sec
increasingly sophisticated tools and social organization of prehistoric modern extinction din

:d needs. The wisdom o:'
highly controversial.
sapiens populations ha>
·y of the earth. Howe\· e~
[ficant cost. Perhaps th:-
he global extinction o f~
rate at which our sp ec i~
ed to human populatic:.
few centuries, so has tt:-
:ver to extinction. t\ile<
:xtinctions to five e ar li :-~
. These are the Upp.:-
'riassic, and Cretaceot.:s-
~ term sixth extinction :
~ars or so that have bee=
ns were caused by cat::.-
>bjects such as met e o~
ver the past 10,000 yea:
~se earlier geologic pe:::-
j by the actions of o;:;:-
er species through bo· -
ct predation by burn a::...
ction of prey species h::
n of predators. Humn-
and these have driY.:-:-
tion. Land clearance t~
loss of habitat. In rna::
~tors, such as aggressi· :
nd clearance.
led into two general cc.·-
ions caused by the inic.:_
Eurasia. The second c.-:.·-
mbination of Europe.::.::
apid global increase --
:: expansion of mod --
ion of a nonselective --
. the remarkable succh
population size can ~­
fumans are omni\' o~=
' species and plants. T..:.:
of prehistoric mod.::-.
HUMANS AS A FORCE OF EXTINCTION 363
humans made them a very efficient predator. If a favored prey species goes
extinct, humans can shift reliance to other animal prey and plants. Prey species
that coevolve in the same geographic region as a nonselective predator will
evolve adaptive strategies to persist in the face of this predation. However, as we
have seen in Chapters 4 and 8, when nonselective predators colonize or are artifi-
cially introduced to an environment where prey species have not coevolved with
them, the prey often are not adapted to escape or otherwise withstand predation
by the new predator. The results can often be extinction of the prey species.
Hominids lived and evolved in Africa and southern Eurasia for most of their his-
tory. Other animals in these regions evolved with hominids as predators. In the
past 60,000 years, humans spread to regions where animals had evolved without
the presence of hominid predators. We have little evidence of prehistoric humans
causing significant widespread extinction of animals in Africa. We do have evi-
dence for the role of modern humans in large prehistoric extinction events in
areas such as North America, Australia, and the islands of the South Pacific
where people have spread only in the last 60,000 years or less. Let's look at each
of these examples in turn.
Fossil evidence from many sites shows that until the late Pleistocene, about
12,000 years ago, North America contained a richness of large mammal species
that is remarkable by modern standards. Large mammals are those species in
which the adults are greater than 44 kg in weight. The large mammal fauna is
often referred to as the megafauna. Among other things, the megafauna con-
tained species of mammoth (Mammuthus primigenius, M. jeffersonii, and M.
columbi) and mastadon (Mammut americanum) related to elephants (Fig. 12.3),
horses (Equus spp.), camels (Camelops hesternus), ground sloths (Nothrotheriops
shastensis), sabertooth cats (Smilodon fatalis), and short-faced bear (Arctodus
simus). The youngest radiocarbon dates on the remains of these extinct mammals
suggest that all of the North American species of these large mammals, and many
other species, disappeared around 10,000 to 4000 years ago (Fig. 12.4). In fact, of
the 79large mammal species that existed in North America at about 20,000 years
ago, only 22 species were present when the Europeans first colonized the conti-
nent 500 years ago. This represents the extinction of roughly 70% of all large
mammal species in North America. When the Europeans arrived, the largest
mammals remaining in North America were species such as bison (Bison bison),
muskox (Ovibus moschatus) in the far north, and bears. Interestingly, small mam-
mals did not suffer the same high rate of extinction during the period of large
mammal extinctions 10,000 to 14,000 years ago.
Two sets of hypotheses have been constructed to explain the megafauna!
extinctions that occurred in North America at the close of the Pleistocene. The
first hypothesis attributes the extinctions to climatic change. According to this
hypothesis, large mammals could not cope with the rapid changes in climate and
vegetation at the end of the last glacial maximum and beginning of the Holocene.
This was because they were more sensitive to climatic and environmental condi-
tions, had larger resource requirements, larger individual feeding ranges, smaller
populations, and lower reproductive rates. The flaw in this argument is that these
same genera, and indeed many of the same species, had survived earlier glacial to
interglacial transitions in the Pleistocene. What was so different about the end of
the last glacial episode that made it lethal to the North American large mammals?
The second hypothesis attributes the North American large mammal
extinction directly to humans. This hypothesis has been particularly well devel-
 oped and artie;
sity of Arizonc
humans entere
ago and spreac
ago. Martin th'
hunting and g;
game such as h
interior of Nor
different speci1
been preyed UJ
behavioral ad<
humans would
enced rapid pc
bers of human
species. Local '
would cause h
geographic exi
apparent rapic
FIGURE 12.3 The skeleton of a Jefferson mammoth (Mammuthus jeffersonii) at the America 12,001
American Museum of Natural History in New York. This species of mammoth was
geographically
approximately the same size as living elephants and was present in midwestern and
ulation of hum
eastern North America during the end of the Pleistocene. Mammoths were the largest
mammals in North America at that time. By around 10,000 years, all species of mammot hs
uals of the larg'
in North America had become extinct. Hunting by humans was likely a major factor in th e hunted and po1
extinction of mammoths and other large mammals in North America. The hypothesi2
front. The mov
(/)
are somewhat i
c
0 the Blitzkrieg 1
·-e lightning war.)

~-~al 80~
60
40
J What evi
America? The
very similar th1
~ 20 ~------- ence of large p
0Q5 o0~~~==~~~--~---~--~--T
-=; F 1 1 fasten the poin
..c 3.0 2.5 2.0 1.5 1.0 0.0
§ Millions of years points (Fig. 12
z before present where the poin
by a teenager r
ian Institution.
oindians. The C
are somewhat
sites in the regi
with these tool
cally with moe
introduction of
00~~~-r~~~~~-r~--~~~~-;~.-~ 12,000 years ag
34 32 30 28 26 24 22 20 18 16 where Clovis I
Radiocarbon years before present
(x1000)
moths. The site
the bones of b
FIGURE 12.4 Radiocarbon dated remains of extinct Pleistocene large mammal species ir;
12,000 to 13,00
North America and a comparison of the late Pleistocene extinctions with known North
Clovis points a
American mammal extinctions over the past 3 million years (after Meltzer and Mead, 1982;
extinct large an
Martin, 1984, 1999). A Clovis point is shown on the diagram.

364

Tiuthus jeffersonii) at the
:s of mammoth was
:mt in midwestern and
1moths were the largest
1rs, all species of mamm o r-~
s likely a major factor in thE
me rica.
_)
/0.0
ene large mammal spe c i ~
ions with known Nort h
ter Meltzer and Mead , 19E::.
HUMANS AS A FORCE OF EXTINCTION 365
oped and articulated by the American geoscientist PaulS. Martin of the Univer-
sity of Arizona. As we have reviewed earlier, there is compelling evidence that
humans entered Alaska and the Yukon Territory from Siberia about 15,000 years
ago and spread southward as the Laurentide ice sheet melted about 13,000 years
ago. Martin theorizes that these newly arrived immigrants brought with them a
hunting and gathering culture from Siberia that included acquisition of large
game such as horse, bison, and mammoth. When these hunters first arrived in the
interior of North America about 13,000 years ago, they would have found many
different species of large mammals for hunting. These species would never have
been preyed upon by human hunters and would not have developed physical or
behavioral adaptations to human hunting pressure. Martin believes that the
humans would have had great initial success in hunting and as a result experi-
enced rapid population growth. Ease of hunting, coupled with increasing num-
bers of human hunters, would lead to local extinction of large mammal prey
species. Local extinction of this type by a predator is called overkill. This in turn
would cause humans to move generally southward in search of new prey. The
geographic expansion of humans southward in search of new prey explains the
apparent rapid spread of humans from Canada to the southern tip of South
America 12,000 years ago. Behind the southward-moving front of the large and
geographically expanding population of human hunters would be a sparser pop-
ulation of humans who would likely hunt the remaining large mammals. Individ-
uals of the large mammals that escaped the first arrival of humans would likely be
hunted and potentially driven extinct by the humans behind the advancing front.
The hypothesized expanding front of human hunters is sometimes called the kill
front. The movement and actions of the human hunters under Martin's scenario
are somewhat like a rapidly advancing army, and his hypothesis is referred to as
the Blitzkrieg Theory of Pleistocene Overkill. (Blitzkrieg is the German word for
lightning war.)
What evidence is there to substantiate the Blitzkrieg Theory in North
America? The stone tools used by people in North America 12,000 years ago are
very similar throughout the entire continent. They are characterized by the pres-
ence of large points with fluted bases. The fluting appears to have been used to
fasten the points to spear shafts. These very distinctive artifacts are called Clovis
points (Fig. 12.4) after the town near the archaeological site in New Mexico
where the points were first discovered. The first Clovis points were found in 1929
by a teenager named Ridgely Whiteman who reported his find to the Smithson-
ian Institution. The people who used these points are often referred to as Pale-
oindians. The Clovis fluted points and other features of the Paleoindian tool kit
are somewhat similar to prehistoric tools from 20,000-year-old archaeological
sites in the region around Lake Baikal in Siberia. The Siberian peoples associated
with these tools appear to have hunted mammoths and have been linked geneti-
cally with modern North American native peoples. This suggests the possible
introduction of this hunting culture and technology from Siberia about 13,000 to
12,000 years ago, as Martin suggests. In North America there are at least 13 sites
where Clovis points and tools are found in association with butchered mam-
moths. The site in New Mexico where the points were first discovered included
the bones of butchered mammoths. All of these types of sites date from about
12,000 to 13,000 years ago, and many contain the remains of several mammoths.
Clovis points are also found in association with the remains of other living and
extinct large and small mammals. This evidence shows that at the time of the late
366 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

Pleistocene extinctions, the Paleoindians in North America were gener~ and 700 years l
hunters and were capable of hunting very large mammals. New Zealand.
The evidence that humans played a role in the extinction of many Ia:-:: aside from bat~
mammals in North America appears very strong, but it cannot be said that hu iL o ~ for example, h
hunting alone was responsible. It is likely that the North American biota ranged in size f
under stress during the late Pleistocene as climate warmed and vegeta · that when hum
changed. The foraging conditions and ranges for the large mammals were shif .; ~ ~ ago, they slaugl
geographically and in some cases contracting. The addition of humans, who w - ~: moa species w
highly sophisticated and generalist predators, was more than the already en vir -- extinction of th
mentally stressed mammals could withstand. The removal of herbivores such .:._ focus to other ·
mammoths, mastodons, horses, and camels by hunting likely produced tro p~ _ nested along th
cascades that led to the extinction of large predators such as sabertooth following Poly1
Finally, it has recently been suggested that humans and their accompanying dO"=- other Pacific Is
may have inadvertently introduced diseases that contributed to the extinction covered and 01
the American megafauna. Hawaii shows t
The magnitude of late Pleistocene mammal extinctions is no less sever extinct followir
Australia than in North America. Indeed, the Australian extinctions were broa hunted princip<
in scope. All marsupials weighing more than 100 kg disappeared between abo _ Flightless birds
60,000 and 15,000 years ago. These included giant herbivorous and carnivora_ of flightless bir
kangaroos, representing a loss of 19 species. In addition, almost 60% of all rna ___ extinct followin
pial species weighing between 10 and 100 kg suffered extinction during ·- Aside fro
period. Reptiles and birds also went extinct during this period. These include a -- tion of island
m-long carnivorous monitor lizard (Megalania prisca) and a flightless ostrich-size.: expanded aero
bird (Genvornis newtoni). In all, at least 60 vertebrate animal species disappeare.: animals, includ
from Australia during this time. rats. All these r
What was the role of humans in the Australian late Pleistocene extinctioiS contributed to
and why was the event so much broader in terms of the size and type of extinct a£.::- ber of crop spe
mals? There is good evidence that humans first arrived in Australia about 60,000 : (Colocasia esc,
53,000 years ago. This time period corresponds with the last known ages for sweet potato (
large flightless bird Genyornis newtoni and a number of other marsupial speci .: . species. In ord1
Hunting by humans was likely a factor in these extinctions. Fossil pollen and cb<L- vegetation. On
coal in lake sediments also show evidence of increased fires and losses of forest ar...: ture. Finally, tho
shrubland during the period 60,000 to about 20,000 years ago. Humans may ha,-:: ing materials a
been responsible for setting frequent fires, which transformed the interior of t :: island vegetati<
continent to more desertlike conditions. Thus, humans may have killed off speci : on the Hawaiia
both by hunting and by creating fires that changed the vegetation. Detailed mo~­ inated by plant
phological and chemical analysis of the bones and eggs of extinct Australian ven ~­ estimated that
brates suggests that many were browsers who lived by eating the leaves of shru~ to extinction. 1.
and trees. The replacement of forest and shrubland by desert would have destroy -' to the extincti
their habitat. The loss of habitat explains why so many small as well as large ani- destroying hab
mals went extinct during the late Pleistocene in Australia. Trophic cascades wouJ · The Polyn
have led to the extinction of large predators such as Megalania prisca. Natural cli- the Europeans
matic change likely played some role in the latter prehistoric faunal extinctions ir; 1800, they four
Australia. During the last glacial maximum 20,000 years ago, Australia became ven - extensive areas
dry and experienced increased desertification. soils, wood, anc
In both North America and Australia, it might be argued that natural cli - areas had been
matic change, in addition to human activity, may have contributed to prehistori gus). Charcoal J
extinctions. In the case of the Pacific Islands, we have clear evidence of the powe ing. Some of tt
of prehistoric humans to cause widespread extinction in the absence of any other potato. Howevc
environmental changes. The eastward expansion of Polynesians between 21 00 this crop. Othe
IN
America were gener a l~:
1als.
: extinction of many la r~:­
cannot be said that humc..:-.
.J'orth American biota wc.o
: warmed and vegetatic:.
rge mammals were shiftli:;
ition of humans, who "' e:-:-
~ than the already enviro::.-
oval of herbivores such ~
tg likely produced tropL:
·s such as sabertooth ca:s.
j their accompanying do;
·ibuted to the extinctio- -
nctions is no less sever
n extinctions were broac:-
isappeared between a
rbivorous and carnivorc-
1, almost 60% of all maE:..-
red extinction during ~
; period. These include 2 - _
end a flightless ostrich-siz;:-_
mimal species disappea::-.::.
1te Pleistocene extinctic:......
size and type of extinct c.=.-
.n Australia about 60,00C -
1e last known ages for _._
of other marsupial spec:-
ons. Fossil pollen and c ~
ires and losses of forest c.::._
:J.rs ago. Humans may be. _
;formed the interior of ---
may have killed off spec.::-
vegetation. Detailed r:u cr
of extinct Australian Ye:--
eatirlg the leaves of shr.
~sert would have destro: :::_
small as well as large c.::;..
lia. Trophic cascades wo __
~ga lania prisca. NaturaJ-=:
istoric faunal extinctioc.s -
ago, Australia became \._
Je argued that natural -
contributed to prehistc::-
lear evidence of the po"' ::
n the absence of any O~::'
:>olynesians between := ~
HUMANS AS A FORCE OF EXTINCTION 367
and 700 years brought humans to many uninhabited islands such as Hawaii and
New Zealand. Although these remote islands did not possess mammal faunas
aside from bats, they did have distinctive bird faunas (avifauna). New Zealand,
for example, had 11 species of large flightless birds called moas. These birds
ranged in size from 20 kg to a staggering 250 kg. Archaeological evidence shows
that when humans first arrived on the islands of New Zealand about 700 years
ago, they slaughtered and ate moas in huge numbers. It is estimated that all of the
moa species were driven to extinction within about 100 years. Following the
extinction of the moas, there is archaeological evidence that humans turned their
focus to other prey, including now extinct species of penguin and petrels which
nested along the coast. At least 77 species of birds went extinct in New Zealand
following Polynesian colonization. Similar avifauna! extinctions are evident on
other Pacific Islands that were formerly uninhabited by humans. Hawaii was dis-
covered and occupied by humans about 1500 years ago. Fossil evidence from
Hawaii shows that at least 35 species, and perhaps over 55 species, of birds went
extinct following the arrival of the Polynesians. In some cases, these birds were
hunted principally to supply feathers for ornamentation of clothing and jewelry.
Flightless birds were particularly vulnerable to extirlction. Hawaii had a number
of flightless birds, including eight species of geese and an ibis species that went
extinct following the arrival of the Polynesians.
Aside from direct hunting, several other factors contributed to the extinc-
tion of island biota. The Polynesians brought agriculture with them as they
expanded across the Pacific. They transported and introduced domesticated
animals, including dogs and pigs. In addition, they inadvertently introduced
rats. All these mammals preyed upon bird, reptile, and invertebrate faunas and
contributed to Pacific Island extinctions. The Polynesians also brought a num-
ber of crop species and edible plants, including coconut (Cocos nucifera), taro
(Co locasia esculenta), yams (Discorea), Breadfruit (A rtocarpus communis),
sweet potato (Ipomea batatas) and other plants, together with many weedy
species. In order to grow taro, the Polynesians cleared large areas of lowland
vegetation. On wetter sites, the Polynesians practiced slash and burn agricul-
ture. Finally, they cut down additional forest at low to mid-elevations for build-
ing materials and firewood. Following these clearances, much of the original
island vegetation was replaced by plants introduced by the Polynesians. Today
on the Hawaiian Islands almost all vegetation below 760 min elevation is dom-
inated by plants introduced by the Polynesians and later by the Europeans. It is
estimated that about 130 species, or 10% , of Hawaii's native plants were driven
to extinction. Undoubtedly, this landscape alteration significantly contributed
to the extinction of birds and other animals such as snails and insects by
destroying habitat.
The Polynesians dramatically changed the landscape on other islands. When
the Europeans first arrived and settled in New Zealand between A.D. 1642 and
1800, they found the lowland landscape largely dominated by grasslands and
extensive areas of bracken ferns (Pteridium aquilinum) . Evidence from fossil
soils, wood, and pollen shows that, prior to the arrival of the Polynesians, these
areas had been dominated by forests of trees such as southern beech (Nothofa-
gus). Charcoal found in soils shows that the forest clearance was caused by burn-
ing. Some of the forest was likely cleared to facilitate the cultivation of sweet
potato. However, there was also much clearing beyond the agricultural limits of
this crop. Other areas were likely burned to ease the hunting of moas and to
368 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

encourage the growth of bracken, which was used as food by the Polynesians.-:-- 100
clearance of forest on Easter Island was even more extreme. At its peak, the Pe.
90
nesian population on Easter Island numbered about 7000 people. The peo;
there had a very advanced culture and social structure. Polynesians cleared fo:-:-
for agriculture and firewood, as well as to build and transport the island's fa JLc:-_
80 •c I~

monumental stone heads. Fossil pollen evidence shows that the present tree.i"' (/)
70
<ll
vegetation of the island was strictly the result of land clearance by hum~ ~ ·g 60
Q.
Again, the destruction of habitat must had severe consequences for native pl~ ­ (/)

and animal species. In the case of Easter Island, the destruction of the fores t C:.S ~50
<ll
led to the collapse of Polynesian society on the island and a regression in cui __ ..c
to very primitive conditions. The first Europeans to visit the island in the e i~ ­ ~ 40
z
teenth century found only a small number of islanders living in primitive • 30
huts and caves. The surviving Easter Islanders had no memory of their earlier _
ture and how the great stone heads were carved or transported. 20
Although we have focused on North America, Australia, and the Pa ·..:,
10
Islands, the geographic expansion of Homo sapiens sapiens is associated "''" -
increased extinction rates in many other parts of the world. Such areas inclt:.2i:: 0
northern Eurasia, islands in the Mediterranean Sea, and the island of Mad agas --- (J)
N
(J)
l1"l
<0 <0
off Africa. Humans arrived in Madagascar about 2000 years ago. All animals v.ri - _ ~ ~

6 6(')
body mass larger than 12 kg were driven to extinction shortly thereafter. It wo' 0
<0 <0
be wrong to think that the Native Americans, Australians, or Polynesian islancic:-
are in any way worse than other groups of humans in terms of their intentio- - FIGURE 12 .5
impact on the prehistoric environment. The aim of these and other peoples was ;:: _ 1600 to 1960 to tr
to produce massive extinctions of other species; it was simply to survive.

Hist oric Ext inctions increase in bin

The last 500 years have witnessed a wave of extinctions for plants and anim~ gone extinct in
that is greater in magnitude and geographic scope than any of the prehisto:--h Freshwat1
extinction events (Fig. 12.5). In the case of mammals, at least 88 species have go;:;:: the historic per
extinct since A.D. 1500. About 52% of the mammal species lost were roden' have gone exti
Over 3% belonged to our own order, the primates. A careful analysis of his tori • rather than st1
records shows that at least 51 mammal species have disappeared in the tin::: species have bE
period from 1750 to the present. The others went extinct sometime between 1 -o not escape fron
and 1750. Over 73% of those extinctions have occurred on islands, mostly in tt::: Historic E
Caribbean (Fig.12.5). The Caribbean losses include a species of monkey (Xenothr:- isms. Over the J
mcgregori) that once lived on Jamaica. Over 73% of all continental mamm<U 194 species of
that went extinct during the historic period were from Australia. The histor:: species are kno
mammal extinctions in Australia range from small marsupials to species as larg.: tions have occu
as kangaroos, and they rival the number of species lost during the prehistoric-la ~ tions is direct!'
Pleistocene extinctions. In contrast, North America has experienced one kno""-:: known historic
mammal species extinction since 1500. That extinct species is the rabbit Sylvila- ber of species lc
gus insonus, which was last reported from Mexico in 1991. One famous su known to scie1
species of mammal that has gone extinct in North America is the Californic. plants. We will
grizzly bear, which was a variety of the grizzly bear (Ursus arctos). Hunting b~ What we can b1
Spanish, Mexican, and American settlers was the primary cause of this demise o: gone extinct co
this distinctive variety of grizzly. The last California grizzly died in the San Fran- the past 500 ye;
cisco Zoo in the early twentieth century. The extin'
For birds, the rate of extinction over the past 500 years has been even mar many causes, in
severe than for mammals. It is estimated that humans have caused a 1000-fol duced alien spe
ON
HUMANS AS A FORCE OF EXTINCTION 369

'ood by the Polynesians. T.::.:: 100

:treme. At its peak, the Po:_
90
1t 7000 people. The peo;.:- r Islands
e. Polynesians cleared for.:-:- 80 • Continents
:ransport the island's farr:= _
ws that the present tr ee ~~ C/)
70
Q)
land clearance by humc:.::. ·~ 60
c..
nsequences for native pi::.: C/)

estruction of the forest L 0 50

Q;
land a regression in cult:=_ ..0

visit the island in the ei :-- ~ 40

z
lers living in primitive r. _ 30
memory of their earlier .:.-_
ansported. 20
L, Australia, and the Pa--
. sapiens is associated 10

~ world. Such areas inc ~:

0
n.d the island of Madagas- 0>
(\J
0>
I!)
0>
0>
0> 0>
<t
0>
I'-
0>
0
0>
C')
0>
<0
0>
0>
0>
(\J
0>
I!)
0
<0
<0 <0 <0 ;:::: I'- I'- CXl CXl CXl CXl 0> 0> 0>
years ago. All animals \\:- - ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~

60 6C') 6<0 60 6(\J 6I!) 6CXl 6 6<t 6 60 6C')

L shortly thereafter. It \ \ '0'
<0 <0 <0 I'- I'- I'- I'- co CXl
I'-
CXl 0> 0>
ians, or Polynesian islanL:.
n terms of their intemi-- FIGURE 12.5 Total historical animal extinctions on continents and islands from A.D.
;e and other peoples was - 1600 to 1960 to the 1990s (after World Conservation Monitoring Center, 1992).

simply to survive.

ions for plants and ani.;::.
than any of the pre hi s: .:~
tt least 88 species have ~
l species lost were rocie-
:.:areful analysis of histo:-:.
re disappeared in the ::::.:-
net sometime between ~~·
ed on islands, mostly i2 --
,ecies of monkey (Xen o: --;:
Jf all continental rna- -
1rsupials to species as I:::=
t during the prehistoric-
las experienced one kr: -
pecies is the rabbit S\·:·
in 1991. One famou s ~­
America is the Ca!ifc-
(Ursus arctos). Hunt in~
1ary cause of this demis::
rizzly died in the San F:-
) years has been even ::;::
n.s have caused a 1000-:
increase in bird extinctions. One estimate suggests that about 219 species have
gone extinct in the historic period; of those, about 40% were island species.
Freshwater fish have also suffered greatly increased rates of extinction in
the historic period. Over the past 100 years, at least 172 species of freshwater fish
have gone extinct. Most of the extinct species were those that lived in lakes,
rather than streams and rivers, or spring pools. A large proportion of those
species have been cichlid fish from Lake Victoria in Africa. Lake fish often can-
not escape from overfishing or environmental changes caused by humans.
Historic extinctions have also been high for many other groups of organ-
isms. Over the past 500 years, at least 242 species of mollusks have been lost, and
194 species of insects have disappeared. Of the flowering plants, at least 713
species are known to have gone extinct. Again, a high proportion of these extinc-
tions have occurred on islands. It is distressing to note that the number of extinc-
tions is directly proportional to human population growth. Unfortunately, the
known historic extinctions represent only minimum estimates of the actual num-
ber of species lost. Undoubtedly, hundreds of species went extinct before they were
known to scientists. This is particularly true for insects, mollusks, and smaller
plants. We will likely never know what most of these lost organisms were like.
What we can be certain of is that thousands of species of plants and animals have
gone extinct concomitant with the human population's unparalleled growth over
the past 500 years.
The extinctions of plants and animals observed over the past 500 years have
many causes, including habitat alteration, hunting and fishing, pollution, and intro-
duced alien species of predators, competitors, and pathogens. Habitat alteration is
370 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

perhaps one of the most important forces driving historic extinctions. As h predator began
populations have grown, they have altered terrestrial landscapes for agricuJ. _ - caused the extir
urban development, and resource extraction such as mining and logging. In manyc
alterations of the landscape destroy the habitat of terrestrial organisms. H - illustrated by tl
alteration of the environment also impacts heavily on freshwater and coe:- on Muritius Isl
ecosystems. In the case of extinct fish, about 70% of the species lost over the ;:- prey, and so the
100 years were victims to habitat alterations such as the damming of ri,-_ peans also clem
changes in lake water levels, additional sedimentation due to human land ·- addition, cats, r:
changes in river channels and lake bottoms, and disruption of aquatic vegetatiL_ the dodo nestli
We also have good historical evidence that aggressive hunting by E •...:- period, a simila
peans played a central role in the extinctions of a number of animals such as·- Pezophaps soli1
great auk (Pinguinus impennis) and Steller's seacow (Hydrodarmalis gigas).-:-- Ocean. This ext
great auk was a flightless seabird of the North Atlantic that was very much · • which the three
the penguins of the southern hemisphere. It lived in great numbers on r - torical evidence
coastlines from Scotland to Newfoundland, Canada. During the 1800s, Europ~ of freshwater c
sealers, fishermen, and sailors began to intensively hunt the great auk for f North Americm
and to provide fish bait. Hunters were able to herd the birds straight from -- It is unkn
shore to their deaths in the holds of ships. In addition, egg collectors in E ur ::. have gone extir
paid high prices for great auk eggs; the taking of eggs for collectors may hi: • the number at c
been a major factor in the final demise of the birds. By 1844 the great auk '- feeding and occ
gone extinct. Steller's seacow, which was similar to manatees found in Flof.- were thus partie
was first discovered in the Bering Sea in 1741 by George W. Steller, the natur~ corresponds wit
on a Russian expedition led byVitus Bering. The seacow became a prime prey : _ the twentieth ce
Russian sealers, and by 1768 it was driven into extinction. In addition to hun ·- _ to develop a fisl
animals for food, sport, and collecting, Europeans hunted predatory animals : are predators tt
protect their domesticated livestock. A large predatory marsupial called the T::....-- lids also experic
manian Tiger (Thalacinus cynocephalus) was a direct victim of European era- - bait or were in:
cation efforts. European colonists, perceiving the Tasmanian Tiger to be a rn a~ byproduct of th
threat to sheep herds, launched a vigorous hunting campaign against the anim:= introduced aqu
and by the 1930s it had disappeared. throughout the
Pollution has been an especially important factor in the extinction of rna::: plant cover dest
fish and other aquatic species. Many rivers and lakes have received large amou;:::... tion and decrea:
of pollution either through the direct dumping of sewage and waste or thro ::· culture and inc
inadvertent spills and the runoff of polluted waters from urban and agricult u;:.:... increased input
areas. Pollution has caused or played a major role in the historic extinction tributed to fish
three species of chub (Evarra) from the Mexico City region. It has also b e~ caused by hum<
blamed for the extinction of the fish species such as Chondrostoma scodrensis = aiding local peo
Lake Scadar, Montenegro, and Brycon acuminatus and Leptolebias marmora r~ ecosystem and t
in Brazil. In many c
As European exploration led to frequent travel and trade between con_- been species tha
nents and islands, alien species of plants and animals were introduced either ~ 9 we saw how~
accident or design. In Chapter 4 we discussed how the accidental introduction o: population sizes
mosquitoes carrying avian malaria led to the extinction of many bird species i::: bility of island ~
Hawaii. Organisms that are purposely introduced to new areas can have simile:.: human hunting
disastrous results. It is often not possible to foresee the extinctions that rna: change. Histori1
result from such introductions. The French introduced the African land sn a:.. species with sm:
(Achatina fulica) to the islands of French Polynesia as a source of food. Howevc (Ectopistes migr
the African snails quickly became serious pests to native vegetation and crops. Tc turtle doves. Wh
remedy this situation, the predatory snail Euglandia rosea was introduced rc dant bird in the
the islands to combat the African land snail. Unfortunately, this nonselectiY lion passenger ,

~ic extinctions. As humar:
mdscapes for agriculture.
aining and logging. SueZ;
~stria! organisms. Huma;:;
1 freshwater and coastc.:
species lost over the pas:
the damming of rivers
due to human land us;
on of aquatic vegetatio
·essive hunting by Eur0-
1er of animals such as tL:
'ydrodarmalis gigas). 11.=
that was very much li.k::
great numbers on rocl-:
ring the 1800s, Europe a=
1t the great auk for foe):.
e birds straight from tl::
egg collectors in Euro ~
for collectors may ha\ =
v 1844 the great auk he.:.
matees found in FloriC=..
~ W. Steller, the naturals:·
' became a prime prey ic:
m. In addition to hunti.:..;:
ted predatory animals : ·
marsupial called the T&
ictim of European erac-
mian Tiger to be a majc-
paign against the anima:.::
in the extinction of rna=
'e received large amou:.::.
1ge and waste or thro L~ ­
m urban and agricul ttL~
the historic extinction -
region. It has also beE::
ondrostoma scodrensis _
Leptolebias marmora;-
md trade between cm:=-
rere introduced either t 9 we saw how such species are generally more prone to extinction. The small
tccidental introduction c
1 of many bird species=
~w areas can have simL
the extinctions that IIl2
d the African land sL::....
source of food. Howe\·c:
~vegetation and crops.-::
rosea was introduced :
mately, this nonselecr:· =
HUMANS AS A FORCE OF EXTINCTION 371
predator began to prey upon native island snails. Predation by Euglandia rosea
caused the extinction of many native Pacific Island snail species.
In many cases, historic extinctions result from a combination of factors, as is
illustrated by the dodo bird (Raphus cucullatus). The dodo was first discovered
on Muritius Island by European sailors in 1507. The flightless birds were easy
prey, and so the island became a stop for providing meat for sailing ships. Euro-
peans also cleared the native forests, which likely deprived the dodos of food. In
addition, cats, rats, and pigs introduced by the Europeans probably preyed upon
the dodo nestlings and eggs. By 1681 the dodo was extinct. During the same
period, a similar fate befell the dodo 's nearest relatives, Raphus solitaires and
Pezophaps solitaria, flightless birds that lived on nearby islands in the Indian
Ocean. This extinction event represents the loss of the entire Raphidae family to
which the three species belonged. In more recent times, we have even clearer his-
torical evidence of how a number of interacting factors produced the extinction
of freshwater cichlids in Lake Victoria and the continental extinction of the
North American passenger pigeon. Let's look at both of these examples.
It is unknown precisely how many species of cichlid fish (Haplochromis)
have gone extinct in Lake Victoria in recent decades, but some estimates place
the number at over 100. Many of these species were very specialized in terms of
feeding and occurred in low population numbers in small areas of the lake. They
were thus particularly prone to extinction. The widespread extinction of cichlids
corresponds with a number of human-induced changes in Lake Victoria. During
the twentieth century, the Nile perch (Lates niloticus) was introduced to the lake
to develop a fishery that would support people living around the lake. The perch
are predators that feed on the cichlids. After the introduction of perch, the cich-
lids also experienced increased fishing pressure. Cichlids were caught for use as
bait or were inadvertently caught in perch nets. Cichlids were also killed as a
byproduct of the application of fish poison to catch perch. By the late 1980s, an
introduced aquatic plant, water hyacinth (Eichhornia crassipes), had spread
throughout the lake and covered many shallows with dense floating foliage. The
plant cover destroyed the cichlid habitat by causing decreased sunlight penetra-
tion and decreased water oxygen concentrations. Finally, land clearance for agri-
culture and increased human population density around the lake have led to
increased inputs of pesticides, fertilizer, and sediment, all of which have con-
tributed to fish mortality and the loss of cichlid species. Some of the changes
caused by humans to Lake Victoria were instituted with the best intentions of
aiding local peoples, but the end result has been a catastrophic disruption of the
ecosystem and the extinction of many species.
In many cases, the animals that have gone extinct in historic times have
been species that had small population sizes and were highly endemic. In Chapter
population sizes, highly endemic distributions, and often limited dispersal capa-
bility of island species made them particularly susceptible to extinction due to
human hunting, introduced predator and competitor species, and landscape
change. Historic extinctions, however, have certainly not been restricted to
species with small population sizes or endemic distributions. Passenger pigeons
(Ectopistes migratorius) were beautiful birds that resembled morning doves and
turtle doves. When the Europeans first arrived in North America, the most abun-
dant bird in the continent was the passenger pigeon. It is estimated that five bil-
lion passenger pigeons inhabited the eastern half of North America. That is
372 CHAPTER 12 HUMANS AS A FORCE IN EVOLUTION AND EXTINCTION

almost equal to the total number of all birds that inhabit the continent today! 11:= ::.:::J . M., and Walker, M. J. C.
passenger pigeons were strong fliers, and the geographic range of the bir nary Environmental Chan
extended from the Gulf of Mexico to southern Canada. They traveled in gre ~ Human Perspectives. Wile:
::.. mler, M. A (1992). Indep<
flocks that numbered in the millions, and it is said that their passing would dark =
and recent genetic eviden<
the sky. During the mid-1800s two factors began to seriously impact on the p ~
~a tion. Economic Botany '
senger pigeon. First, land clearance for agriculture intensified and expand<:-.:. E :.:mler, M. A , and Byrne, R.
westward from the Atlantic Coast states to the edges of the Great Plains. ~ ;::a! genetics of domesticati<
destroyed the forest habitat of the pigeons. The pigeons relied on oaks, chestn _ agriculture. Current Anthrc
and beech as principal sources of food. The birds also required trees to nest. Ti :: ::.~vw n , J. H ., and Lomolino, ~
cutting of a favored nesting tree could destroy over 100 nests. Second, the pigeo:::.. oeography, 2nd edition. Si1
were hunted in great numbers. Random shots into the sky would bring down se'- underland, MA.
eral birds when flocks were passing. It is estimated that one day of hunting us· .: :.-.:ddi hy, L. W., and Stone, C.
shotguns could yield up to 25,000 birds. The birds were also caught in nets wb ~­ o. Native Hawaiian Vegeta
resting and got stuck in sticky lime spread out on the ground. Millions were kille-= Hawaii, Honolulu.
:amond, J. M. (1984). Norm;
each year. The expansion of the railroad in the 1850s led to increased hunting "--'
Ia ted populations. In Extin
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:\itecki), pp. 191-246. Uni'
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newly planted crops. To get some perspective on this wide-scale slaughter, it ~ The Evolution and Future
worth noting that in 1869 one Michigan county sent 7.5 million birds to marke _ m al. Harper Perennial, Ne
in the east. During the 1870s and 1880s, a drastic decline was observed in ~= .amond, J. M. (1997). Guns,
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he human "niche" and tht
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CHAPTER

DESCRIPTION AND
INTERPRETATION OF
BIOGEOGRAPHIC
DISTRIBUTIONS

What can the size and shape of the geographic distributions of plants and animals
tell us about their ecology and evolutionary history? For example, does examination
of the size and shape of the geographic distributions of bird species allow us to draw
conclusions regarding underlying general relationships that might exist between all
birds and the environment? Is it possible that by analyzing maps of modern distri-
bution of a plant species we can untangle the long history of evolution and dispersal
that species and its ancestors have experienced? For over 150 years biogeographers
have pondered and debated these questions. The ecological analysis of geographic
distributions remains of keen interest and continues to evolve as a subdiscipline
within biogeography and ecology. In the early 1980s, the Argentine biogeographer
Eduardo Rapoport applied the word areography to the ecological analysis of mod-
ern plant and animals distributions, while the American biogeographer James
Brown employed the term macroecology to describe such work. Many historical
biogeographers have made their life's work out of the careful analysis of modern
geographic distributions and the use of these distributions to infer evolutionary his-
tory. Such work is sometimes referred to as phylogenetic biogeography.
In this chapter we will explore the ecological and evolutionary interpreta-
tions that are often made from the modern geographic distributions of plants and
animals. We will see that many modern distributions are the product of fascinat-
ing biogeographic histories. In our analysis of geographic distributions, we will
use the term range to refer to the area permanently occupied by a plant or animal
taxon. This term can also be applied to the browsing, foraging, or hunting area of
an individual animal, but we will not use the word in that sense in this chapter.
Any meaningful analysis of the size and shape of a species' ranges requires reli-
able mapping of the geographic distributions of plant and animal species. We will
first consider how geographic distributions maps are generated. We will then look
at some general relationships that have been observed between the physiology
and morphology of species and the sizes and shapes of their geographic ranges.
We will follow this by outlining some of the frequently encountered general dis-
tributional patterns that biogeographers have come to recognize. We will look at
how these general patterns have been interpreted in terms of species' ecology
and evolutionary history. We will then critically examine the ways in which the
analysis of modern distributions has been used to infer the evolutionary history
of plants and animals.

377
378 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

MAPPING BIOGEOGRAPHIC DISTRIBUTIONS

In this book we have examined many maps portraying the distributions of difiC":"-
r.~
. 6
...
ent plant and animal species. We have assumed that these maps are valid re ::-=- "·
. '
sentations of the spatial distributions of these organisms, and we have base: ·I!!>
some important conclusions on the maps. Let's now consider how such biog- -
graphic maps are made and the uncertainties that they may contain.
Biogeographic maps depicting the distributions of plants and animals <::=
derived from spatial networks of observations that record the presence or abser.:-
of species. In collecting such data, plant and animal species are identified in -:.. =
field or sampled and taken back to the laboratory for identification. Field obsen .:_-
tions and identifications that do not require collecting or harming organisml' c..:-=
best because they are the least disruptive to the environment. The latitude, lon:-
tude, elevation, and environmental setting of each site is carefully recorded. ·-
some cases, these data are used to produce dot maps that simply show the si:=-
where the species has been found and recorded (Fig. 13.1). The more sites visit .::..
the more accurate the maps will be. Throughout this book we have present.:-.:.
maps that use shading or boundary lines to represent the geographic ranges
. 6
species. These types of maps, in which the area occupied by an entity (such as _
plant or animal species) is delineated by continuous shading or circumscribed ~
'\.. -
r. , ;.;:;,.<)\
g _..·.·
an unbroken line, are called choropleth maps. The data from dot maps are extr -;-
--~
olated to make choropleth maps. The extrapolations are done using knowledge '- _
the environmental requirements of the species and the environmental conditio::..
of the area being mapped. In the case of plants, information from airphotos L
other remote sensing imagery is sometimes used to supplement field observatio=:.s.
Choropleth maps are popular because they rapidly convey a general impression L:
the geographic areas in which the species is found and the areas in which it ...::
absent (Fig. 13.1). However, information from biogeographical choropleth rn a~
must be used with some caution, as will now be explained. (
Choropleth maps often include large areas in which the presence or abs e n~
of the species is extrapolated and not directly observed. In a number of cases, su -
extrapolations have been shown to be incorrect. For example, it has long bee:. FIGURE 13 .1
thought that big cone Douglas fir (Pseudotsuga macrocarpa) grows both in tl:.:: mariana) in Nort
mountains of southern California and in adjacent highlands of the Baja PeninstL.2 the same data. C
of Mexico. Because the general environment of the Mexican mountains is similc.:::- that the density ,
to that in southern California, the range of the big cone Douglas fir was inferred : ~ northern bound<
extend into Mexico. Biogeographic maps have been made that show the speci = and stands.
growing in northern Mexico. Recent detailed botanical surveys found that the:::
are actually no big cone Douglas firs in Mexico. It can be concluded that tt::
species is a very narrow endemic restricted to southern California. Incorre . Classic examp
extrapolations of the presence or absence of the species can produce erroneoe..;: from the deta
maps and misleading interpretations of biogeographic distributions. (Clematis fren
Simple choropleth maps present the illusion that the plant or animal speci ~ momys manti<
is distributed evenly and continuously throughout the shaded or delineated are2.. animals as con
Such even and continuous distributions do not occur in nature. Within the when its distri
mapped boundary of the species there will be areas of high- and low-populatio;: may occur as r
density. Neither are plants and animals distributed continuously within their by large and s
mapped geographic ranges. Instead, they occur as scattered individuals anc species becom
groups (metapopulations) separated by small and large areas where they ar als in the vicin
absent. Such discontinuous distributions are referred to as being punctiform. vey this fragn
>HIC DISTRIBUTIONS MAPPING BIOGEOGRAPHIC DISTRIBUTIONS 379

,...
•
~the distributions of differ-
hese maps are valid repre- ""·
:•. D
nisms, and we have base.:
consider how such biogeo-
y may contain.
•••
-.,• . .....,..'
of plants and animals ar" --i• , •
.. • ,#
)[d the presence or abse n c~ • \It .....!•
pecies are identified in tl:." • •
ientification. Field obsem :.-
••
~ or harming organisms a;"
)nment. The latitude, long:-
te is carefully recorded . I=
that simply show the si te-;
3.1). The more sites visite.:
s book we have presente.:
t the geographic ranges c-:
1ied by an entity (such as .:. ""·
hading or circumscribed c:
1 from dot maps are extra;--
re done using knowledge c-
e environmental conditio::....
rmation from airphotos c-
,plement field observatio::.s.
vey a general impression c
nd the areas in which it ~
'graphical choropleth m2.;::--
1ed.
ich the presence or abs e n.:-~
l. In a number of cases, s· ·--
example, it has long bee- FIGURE 13. 1 A dot map based on recorded collection sites for black spruce (Picea
'Ocarpa) grows both in t~~ mariana) in North America (after Porsild and Cody, 1980) and a choropleth map based on
lands of the Baja PeninsL the same data. Detailed surveys based on field observations and airphoto analysis show
exican mountains is sirnili: that the density of spruce decreases dramatically near the northern treeline. The actual
: Douglas fir was inferred : _ northern boundary is relatively diffuse and composed of scattered disjunct individuals
nade that show the speci:::- and stands.
al surveys found that the:-:
can be concluded that ~=
them California. Incorrc.::- Classic examples of how species occur in such a discontinuous manner comes
:ies can produce erroneo_ from the detailed mappings of the distribution of wild plants such as clematis
distributions. (Clematis fremontii) in Missouri or wild animals such as pocket gophers (Tho-
the plant or animal speci:::-- momys monticola) in California. A simple choropleth map presents plants and
shaded or delineated arc:.:. animals as continuously distributed throughout their ranges (Fig. 13.2). However,
cur in nature. Within r:::: when its distribution is examined in increasing detail, it is seen that the species
f high- and low-populatic.:: may occur as relatively small metapopulations and individuals that are separated
continuously within the..::- by large and small areas of unoccupied land. In many cases, the distributions of
scattered individuals a::.: species become extremely fragmented into isolated metapopulations or individu-
:trge areas where they a=-: als in the vicinity of the species' geographic limits. Choropleth maps do not con-
d to as being punctifor=. vey this fragmentation and the diffuse nature of geographic range limits. In
380 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

0.2

\
0.16
(/)
Q)
'(3
Q)
g-0 .12
0
c
0
t 0.08
0
c.
e
a..
0.04

100 km
0
0

FIGU RE 13.3
10 km (after Brown, ·

range size oJ
thirds the si:
Canada. Mo~
have medim
Regardless c
animal speci
the continen
1 km
backward J <
FIGURE 13.2 The typical geographic distributio n of a plant species. The distribution range sizes f<
appears continuous on a cho ropleth map, but when viewed at a smalle r scale we can see tion, consisti
that the species actually occurs as scattered metapopu latio ns and whe n view ed at an eve n
ranges, is sor
smaller scale the species is seen to occur as scattered individuals.
What a
of species ra
the hollow c
summary, biogeographic maps are useful tools, but are usually very crude approx- their ecologi
imations of the actual spatial distributions of plants and animals. logical factO!
have broad
compete effc
BIOGEOGRAPHY OF RANGE habitats. The
SIZE AND RANGE SHAPE of environm
species withi
We have seen in preceding chapters that some plants and animals, such as mod- The widespr<
ern humans, have extremely large geographic ranges. Other taxa, such as the mental and c
Devil's Hole pupfish (Cyprindon diab olis), which is restricted to one tiny desen geographic a
spring in Nevada, or the Catalina mahogany (Cercocarpus traskiae), which i A meri<
restricted to one small valley on Catalina Island off the coast of California, have large geogn
very small ranges. We might ask, is there a typical or average range size for range, while
species? If we plot the range sizes of North American bird species (Fig. 13.3), we ties. Brown
find that the majority of species have ranges of less than 5 million sq km. A few plays in detc
species have ranges greater than 10 million sq km. For North and Central Amer- eralist speci
ican mammal species we see a similar pattern. Most mammal species have a which allow
HIC DISTRIBUTIONS
..•
m (after Brown, 1995).
species. The distributio n
a smaller scale we can see
3nd when viewed at an eve-
als.
Lisually very crude apprc -
d animals.
md animals, such as mo.: ·
:. Other taxa, such as ::..:
stricted to one tiny des;:-
carpus traskiae), whic
e coast of California, he.· :
)r average range size ::::
bird species (Fig. 13.3). "'":
an 5 million sq km. A :~
North and Central An:~-
mammal species ha,-e _
BIOGEOGRAPHY OF RANGE SIZE AND RANGE SHAPE 381
0.2
0.16
en
Q)
'(3
Q)
~0.12
0 Hollow curve distribution
c '\
0
:e0 0.08 ""
e
a.
0..
·"
"" '
0.04
' ' --..
0 --_,__
0 5 20 25
Area of geographic range (106 km2)
FIGURE 13.3 Size of the geographic ranges on North and Central American land birds
range size of less than 2 million sq km. A range of 1 million sq km is about two-
thirds the size of Alaska, and a range of 10 million sq km is about the size of
Canada. Most groups of organisms show a similar pattern: the majority of species
have medium to small ranges, and only a few species have very large ranges.
Regardless of which continent is studied, it has been found that most terrestrial
animal species have ranges that are less than one-fifth to one-fourth the size of
the continent they live on. In general, a lognormal curve having the shape of a
backward J describes the relationship between species numbers and geographic
range sizes for most groups of plants and animals (Fig. 13.3). This typical distribu-
tion, consisting of many species with small ranges and a few species with large
ranges, is sometimes called the hollow curve distribution.
What are the ecological controls that produce the hollow curve distribution
of species ranges? If we assume that the species have stable geographic ranges,
the hollow curve indicates that most species are relative specialists in terms of
their ecological niches. Most species are excluded by physical conditions or bio-
logical factors from many geographic areas and habitats. A relatively few species
have broad generalist niches and can survive diverse physical conditions and
compete effectively with other species over a wide range of geographic regions or
habitats. The vast majority of species survive by being adapted to a narrower set
of environmental conditions and perhaps being competitively superior to other
species within the geographic region where those environmental conditions exist.
The widespread occurrence of the hollow curve indicates that the mix of environ-
mental and competitive pressures leading to adaptation for persistence in a select
geographic area is strong.
American biogeographer James Brown has observed that species with
large geographic ranges generally occur in high densities throughout their
range, while species with smaller ranges tend to occur at relatively low densi-
ties. Brown concludes that this is evidence of the key role that niche breadth
plays in determining both geographic range and density. He suggests that gen-
eralist species can use a wide variety of habitat and ubiquitous resources,
which allows these species to be both common on a local scale and widely
382 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

distributed geographically. A contrasting explanation for the relationship ..,

1010
range size and population density comes from metapopulation theory. Accor --
ing to metapopulation theory, the species that develop large geographic ran g _
c\1
and high population densities are superior to other species in terms of disp ;- g 10 8

sal capability and resistance to extinction. In addition, since metapopulatio- Q)

(/J Ol
are by definition geographically separated from other populations of the sa :: Oi c
~
E {.)
species, individual events such as catastrophic disturbances or outbreaks E :.c 10
6
C1l c..
disease will likely not affect all of the populations. It is these properties, parti-- E ~
"0
ularly resistance to large-scale synchronous declines in population, that allo c Ol
0
C1l
for large population sizes. However, since resistance to extinction is higher fc.:- ....J & 104
0
generalist species than for specialists, the importance of large niche breadt" - C1l
in promoting large geographic ranges for metapopulation distributions cann ~
<{
102
be ruled out.
We might ask if there are systematic differences in the range sizes -
different types of organisms? When we compare different groups of organisn:s.
we find that insects and plant species have generally smaller average range s iz~
than vertebrates. Amphibians and freshwater fish tend to have smaller ran=""
sizes than mammals and birds. Birds tend to have larger range sizes than rn a -
1010
mals. North American birds have an average range size of over 3 million sq k
North American mammals have an average range size of around 1.5 million _-:_
km. The relatively large range size of birds may relate to better dispersal capabi:- c\1
ity, broader niches, and or greater ability to take advantage of a wider variety o: E 107
~
microhabitats. Not all flying organisms have range sizes that are particular.. Q)
Ol
large, however. The average range sizes for North and Central American b : c
(/J C1l
species are about the same as for non-flying mammal species. In general, marin:: "E ~ 1Q65
:0 :.c
species have larger range sizes than terrestrial species. "0 c..
c ~
C1l Ol
When we look in more detail at birds and mammals, we find that predato , ....J 0
generally have larger ranges than herbivores. In Africa, large carnivores such as & 10 6
0
lions and hyenas have an average range size of about 9 million sq km. In contra C1l
~
large herbivores such as elephants and zebras have an average range size o:
<{ 105.5
about 4 million sq km. In North America, the average range size of predator:
owls is about 6 million sq km, while for seed-eating sparrows it is about 3.5 ·
lion sq km. The relatively large size of predator ranges likely reflects the fact tha:
the higher trophic levels that predators occupy requires them to utilize a wid
range of different prey species and be able to hunt and survive in a wide range o:
environments in order to obtain adequate food.
Examination of the relationship between the body size of animals an FIGURE 13.4
their geographic ranges shows that both large and small species can posse_: terrestria l mam
large geographic ranges (Fig. 13.4). However, there are very few large-bodie · small ranges, b
species that occupy very small ranges. Species with large body masses requir
greater amounts of foo d and larger browsing, hunting, and foraging areas th an
small-bodied species. In general, the carrying capacity of a given area is less for 6 million sq 1
a large species than for a small one. If a large species has a small geograph i less than 800,
range, it will have a low population size and be particularly prone to extinc- Evoluti'
tion. Thus, a large range size is important for the survival of large-bodie ent range siz(
species. Following from this consideration and our discussion of the differ- streams or sr
ences in the range sizes of carnivores and herbivores, we can see why large car- movement of
nivores in particular require very large range sizes to survive and small gene flow bt
herbivores can persist with small ranges. It is not surprising that the average allopatric ev
range size for predatory cat and canine species in North America is about ranges. Such
 IC DISTRIBUTIONS BIOGEOGRAPHY OF RANGE SIZE AND RANGE SHAPE 383

1 for the relationship o:

·pulation theory. Accord-
' large geographic ranges
c\1
>ecies in terms of disp er-
! 108

...• ."
1, since metapopulation5 Ql

··"
(/) Cl
populations of the same (ii c:
E ~
rbances or outbreaks o: E .>,? 106
Cll .r:
; these properties, partir - E a.
~
in population, that all o -o Cl
c:
g
• •
Cll 104
o extinction is higher fo ~ ...J Cl

: of large niche bread ths 0

tion distributions canno:
Cll
~
••
< 102 ••• •
•
es in the range sizes c:
·ent groups of organi s rr~
•
102 104 106
1aller average range size;: Body mass
d to have smaller ram: ::- (Grams)
=r range sizes than mar:::-
: of over 3 million sq '-
of around 1.5 million s-
)better dispersal capab:..-
.......
• . ....
., r-:.·.• • • ••
..........
..~,~ ••,:.,.,,_,..,'
;,,·~ ......... ...
c\1
tage of a wider variety c· E
.. .:=! :--r., •
zes that are particular._ ==-
Ql

.....,...,..... :.••t•··· •
.. . . ...
Cl
td Central American b.=. (/)
c:
Cll ~ -~....... ••
Jecies. In general, mari;::: :g ;:; 106 5
.c E • • •• • • •• ••
-o a.
•
..
c: ~
1ls, we find that predato. Cll
...J
Cl
0
• • • •
, large carnivores such ::... ~ 106 • •
nillion sq km. In contra5:. 0
Cll
m average range size <
~
• •
: range size of pred a to~ 105.5 • •
mows it is about 3.5 m:..-
ikely reflects the fact tiE
=s them to utilize a \\i ;:::: 100.5 102 104
;urvive in a wide range ~- Body mass
(Grams)
ody size of animals a;:: FIGURE 13.4 The relationship between range size and body mass for North American
nall species can posses. terrestrial mammal and bird species. Small and medium-sized animals can have large or
:every few large-bod:.:: small ranges, but few large animals have small ranges (after Brown, 1995).
rge body masses req ui:-::
, and foraging areas th"-.::.
of a given area is less ~;- ­ 6 million sq km, while for herbivorous small rodents the average range size is
> has a small geograpi:... less than 800,000 sq km.
icularly prone to ext ~.:: ­ Evolution and speciation events also play a role in determining the pres-
mrvival of large-bodi;:-: ent range sizes for plant and animals species. Minor landscape features, such as
jiscussion of the diffe: - streams or small mountain ranges, may be barriers for seed dispersal or the
ve can see why large c2.-- movement of small nonflying animals. The presence of such barriers restricts
s to survive and s m~ gene flow between populations and promotes allopatric speciation. These
prising that the averc.;: allopatric events create new species that initially have small geographic
l"orth America is abo:.. ranges. Such small landscape features are not significant barriers for larger
384 CHAPTE R 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

animals and flying organisms. Biogeographic maps provide some evidence - ~ 10

the role of landscape diversity and physical barriers in the development
small geographic ranges through allopatric speciation. North and Cen t:-_ 8
American mammal species with ranges of less than 10,000 sq km are genera:.. ~
Q)
small bodied and are most commonly found in mountainous regions, isla n '- Ol
c:
and peninsulas. The rugged terrain of mountainous regions and the waterb -- ~ 6
tii

--
ies surrounding islands and peninsulas produce barriers that promo::: c:
i5
allopatric speciation. 2 4
Some very interesting patterns emerge when we examine large-scale gee- §
c:
graphic variations in range sizes. The average range sizes for plants and a- - t1l
Q)
:2 2
mals that live at high latitudes or high elevations tend to be larger than t'-=
range sizes of species that live at low latitudes and low elevations. There a:-::
many more mammal and bird species with very small ranges in southern C - - 0
tral America than in North America. The general pattern of decreasing ran;:: 35
size toward the equator has been observed for many plants and animal gro i='-
and is called Rapoport's Rule. The reasons for the large range sizes at high I :-
itudes and high elevations are still being debated. However, the large ran g : 1800
must in part relate to more generalized and broader niches. The high-latitu ·::
and high-elevation environments have shorter and cooler growing seasons a - 1600
less primary productivity to support species. Animal species may have to us _ g 1400
broader range of food plants or prey species to survive. High latitudes a .:. Q)
Ol
c:
high elevations also have wide differences in seasonal and daily temperature::. ~ 1200
Organisms in such environments must have relatively wide niches in terms c: tii
c:
0 1000
climate and climatic variability. Evidence for increased niche size is fou - i5
when we look at the latitudinal and elevational ranges of high-latitude a .:. iii
a; 800
high-elevation species compared to low-latitude and low-elevation speci - c:
t1l
Plant and animal species found at high latitudes and high elevations typicali: Q) 400
:2
have ranges that extend over a greater range of latitude and elevation than ~ 200
the case for species found at low latitudes and low elevations (Fig. 13.5). Tb ::
greater range of latitudes and elevations in which they can live directly reflec:::
the greater range of temperature conditions that the high elevation and h i~ ;..
latitude species can tolerate.
FIGURE 13,5
The general shapes of geographic ranges reveal additional interesting pa:-
foun d and the
terns that may have ecological significance. In general, the shapes of plant an ·
here are for t re
animal ranges are more elliptical or rectangular than they are circular o::-
square. The tendency for ranges to be elongated along one axis relative to t h ~
other appears to be controlled by climate and landforms. In North Americc.
species that have large geographic ranges typically possess ranges that ar
longer in longitude than they are in latitude. In contrast, animals with smal: of Anna's hl
ranges tend to have latitudinal axes that are long compared to their longitudi- west of then
nal axis. It appears that animals with large ranges are typically distributed par- nia to Washi
( H aliaeetus l
allel to longitude along broad climatic zones. In contrast, rivers and mountaiL
systems, which produce more local climatic regions and also present barriers to Atlantic coa
the distributions of species with small ranges, often run from north to south in excluded fro
North America. Examples of north to south mountain ranges include the Rock- occupies a t
ies and the Appalachians. These landscape scale features and barriers appear two extreme
most important in controlling the range shape for small-bodied species wit h common tha
duced huge
small ranges. This pattern can be seen for many mammals and a number of bird
in the contin
species. For example, the total geographic range (winter and breeding ranges)
 -iiC DISTRIBUTIONS BIOGEOGRAPHY OF RANGE SIZE AND RANGE SHAPE 385

rovide some evidence fo:- 10

·s in the development o~
tion. North and Cent ra. 8
0
0,000 sq km are generalh ~
Q)
ntainous regions, islan ds.. Ol
c
~gions and the waterbod- ~ 6
barriers that promot:: Cii
c
'a
.a 4
examine large-scale geo- ~
c
sizes for plants and an:- ctl
Q)

:nd to be larger than tb:: ~ 2

ow elevations. There ar::
I ranges in southern Ce;::- 0
ttern of decreasing ra ng:: 35 40 45 50 55 60 65 70
plants and animal gro u!'! Degrees of latitude
ge range sizes at high la:-
owever, the large ran g e~
1800
niches. The high-latit uc::
oler growing seasons ar..:. 1600
:pecies may have to us_ _
·vive. High latitudes ac.:.
I
Q)
14oo
Ol
I and daily temperat ures.. ~ 1200
y wide niches in terms c: Cii
1sed niche size is fou;::.:. § 1000
~
ges of high-latitude ac.:. >
Q)
a:; 800
d low-elevation specie;; c
ctl
high elevations typical:: Q) 400
~
1de and elevation th an :..>
200
levations (Fig. 13.5). T;:: ;:
r can live directly refl ec:: 0
high elevation and h i~(- 0 500 1000 1500 2000 2500 3000
Elevation in meters
dditional interesting pa:- FIGURE 13.5 The relationship between the latitude and elevation at which a species is
1, the shapes of plant a::.:. found and the degrees of latitude or meters of elevation that its range spans. The examples
tan they are circular o- here are for tree species in North America and tree species in Costa Rica (after Brown, 1995).

g one axis relative to - ~­

orms. In North Americ-_
possess ranges that a:-::
trast, animals with sma....
tpared to their longituC.:-
typically distributed pa::--
·ast, rivers and mounta=
also present barriers : -
n from north to south :::
ranges include the R oc",:-
lfes and barriers ap p e ~
nall-bodied species ''i:.::
als and a number of b ~.:.
er and breeding range;;
of Anna's hummingbird (Calypte anna) is restricted to a narrow strip of land
west of the main coastal mountains that runs south to north from Baja Califor-
nia to Washington State. In contrast the total natural range of the bald eagle
(Haliaeetus leucocephalus) extends from the western coastline of Alaska to the
Atlantic coastline of eastern North America (Fig. 13.6). The bald eagle is
excluded from desert regions in the south and tundra regions to the north and
occupies a broad band of more temperate climatic conditions between these
two extremes. Prior to European settlement, the bald eagle was much more
common than it is today. European land clearance, hunting, and pollution pro-
duced huge declines in the numbers and geographic distribution of bald eagles
in the continental United States.
386 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

likely to have
important dist

Endemic a
In Chapter 10
plants and an
most contine
endemics. Mo
Rapoport has
butions relati\
occurs and th
endemics are :
istic species a
regions are clc
more regions
km range) wit
tied as macro-
sq km range).
skua (Stercort
America (N ec
region) . This
Humans, who
tied aS COSmO]
also sometime
Endemi(
recently evolv
ranges. These
to the area in
Mauna Kea 01
plant (Argyr.
FIGURE 13.6 The latitudinally elongated geographic range of Anna's hummingbird nowhere else •
(Calypte anna), which is a small-bodied nectar-eating species with a small geographic range . ter. As the isl<
The longitudinally elongated range of the bald eagle (Haliaeetus /eucocephalus), which is a plant is a neoc
large-bodied predator species possessing a large geographic range (after Robbins et al. 1983). of this highly
size today is c
alien plant spt
Some e1
COMMON BIOGEOGRAPHICAL reduced in siz
DISTRIBUTIONAL PATTERNS environmenta
tion or predat
After examining many distributional maps for plants and animals, biogeogra- had much lar
phers have come to recognize a number of geographic distributional patterns coastal redwc
that are often repeated for different taxa. In many cases, study of modern ecolog- demic. Today
ical and dispersal processes, and examination of the fossil record allow us to northern Cali
determine the causes of such distributions. The distributions themselves also pro- the ancestors
vide important evidence regarding the ecology and evolutionary history of the and Eurasia. (
organisms. Of course, in making such inferences we must be careful not to nary caused t
become circular in our reasoning. A rich vocabulary has been developed to graphic range
describe different geographical distributional patterns and the factors that are little later in t
 f>HIC DISTRIBUTIONS
~ important distributional patterns recognized by biogeographers.
l
~ endemics. Most of the time these terms are used in a rather loose fashion .
(:)
Anna's hummingbird
:h a small geographic range.
leucocephalus), which is a
1ge (after Robbins et al. 1983
> and animals, biogeogrc:-
ic distributional patte- -
s, study of modern ecolog-
fossil record allow us rc
ions themselves also pro-
volutionary history of tb::
must be careful not rc
has been developed rc
and the factors that ar~
COMMON BIOGEOGRAPHICAL DISTRIBUTIONAL PATTERNS 387
likely to have created them. In this section, we will investigate some of the more
Endemic and Cosmopolitan Distributions Revisited
In Chapter 10 we saw that biogeographers use the term cosmopolitan to describe
plants and animals that possess very large geographic ranges distributed over
most continents. Taxa with restricted geographic ranges are classified as
Rapoport has fashion ed more precise definitions that classify geographic distri-
butions relative to both the number of biogeographic regions in which the species
occurs and the absolute size of its geographic range. According to this system,
endemics are species that are found in only one biogeographic region. Character-
istic species are those found in two regions. Species that occur in three or four
regions are classified as semicosmopolitan. Those species that are found in five or
more regions are cosmopolitans. Species that have a large range (> 4,400,000 sq
km range) within the biogeographic region(s) in which they are found are classi-
fied as macro-areal. Those with small ranges are classified as micro-areal ( < 1000
sq km range). Meso-areal species have intermediate range sizes. For example, the
skua (Stercorarius skua) is a seabird found in a small area of southern South
America (Neotropical region) on Iceland, and on the Faeroe Islands (Palearctic
region) . This bird would be classified as a micro-areal characteristic species.
Humans, who are found over large areas of all biogeographic regions, are classi-
fied as cosmopolitan macro-areal species. Macro-areal cosmopolitan species are
also sometimes called pandemics. There are very few truly pandemic species.
Endemics can also be classified by their history. Species that have just
recently evolved and speciated often have small population sizes and geographic
ranges. These are referred to as neoendemics. It is assumed that they are endemic
to the area in which they evolved. The higher elevations of the volcanic mountain
Mauna Kea on the island of Hawaii support a beautiful subspecies of silversword
plant (Argyroxiphium sandwicense sandwicense). The subspecies is found
nowhere else on the Hawaiian Islands or anywhere else in the world for that mat-
ter. As the island of Hawaii is only about a million years old, it is likely that this
plant is a neoendemic that evolved on the island. Today, only about 50 individuals
of this highly endemic subspecies grow in the wild. Part of its small population
size today is due to human disruption of its habitat through the introduction of
alien plant species and herbivores such as goats.
Some endemics once had larger geographic ranges that have now been
reduced in size. The ranges of species may be reduced in size because of physical
environmental factors such as climate change, or because of increased competi-
tion or predation from newly arrived or recently evolved species. Endemics that
had much larger geographic ranges in the past are called paleoendemics. The
coastal redwood tree (Sequoia sempervirens) is a good example of a paleoen-
demic. Today the species is only found along a very narrow strip of coastline in
northern California and a small portion of adjacent Oregon. During the Tertiary,
the ancestors of the redwoods were extremely widespread across North America
and Eurasia. Climatic cooling and drying during the late Tertiary and the Quater-
nary caused the extinction of the redwoods over the vast majority of their geo-
graphic range. We will consider the history of this interesting species further a
little later in this chapter.
388 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

Continuous Zonal Biogeographic Distributions fish family Pe1

temperatures ~
Many semicosmopolitan and cosmopolitan plant and animal taxa have geograp - warmer tempe
distributions that form broad latitudinal bands that follow large latitudinal tempe:--
ature belts. Taxa are restricted to these bands because of their physiological sensi ·\-
ity to temperature or possibly in some cases because of their inability to compe:.= Dispersal [
with species that dominate other climatic zones. There are five such zonal distrib> Biogeographe1
tional patterns. Circumpolar taxa have distributions that circle the Arctic or Antar.:- junction is tho
tic regions and lie within the colder climates of the high latitudes. A good example - jump dispersal
a circumpolar taxon is the arrow grass genus (Scheurchzeria) . Species belonging :;: established a l•
the arrow grass genus can be found growing at high northern latitudes from nor0- particularly co
ern Europe to northeastern Siberia and from Alaska to the Atlantic Coast : long-distance
Canada. The terms Boreal (northern hemisphere) and Austral (southern heiL.:- throughout ea:
sphere) are used to describe plants that are distributed in the cooler areas of _ Oregon, is pr•
mid- to high latitudes. The spruce genus (Picea) is an excellent example of a bore ~
taxon. Species of spruce are typical of the northern forests of Alaska, Canada, a.G.::.
northern Eurasia. Temperate plants and animals are found in the intermediate zor::f
typical of the middle latitudes. Maples (Acer) provide an example of a norther:.
hemisphere temperate taxon. Pantropical organisms are found in the tropical z on~
The palm family (Aracaceae) is a good example of a pantropical taxon.

Amphiregional Disjunct Distributions

The term disjunct distribution is used to refer to geographic ranges that ar ~
divided into two or more geographically separate parts. Amphiregional distribu-
tions occur when macro- and meso-areal taxa have two widely separated geo-
graphic ranges. The root "amphi" comes from Greek and can be translated as
meaning "on two sides." Plants and animals that are amphiregional have distribu-
tions that are divided into two distinct ranges that are usually separated b~
biogeographic barriers. Some common distributions of this kind includ
amphitropical, taxa which occur on either side of the tropics but not in the tropi-
cal zone itself. The Chenopodiaceae family of plants, which includes sugarbeets
and spinach, has an amphitropical distribution. It is likely that amphitropical
plants are excluded from equatorial regions by high temperatures or competition
with tropical plant familes. North and South America have about 160 differen
species of temperate plants that have such amphitropical disjunct distribution
One example is the woodland flower species Osmorhiza chilensis, which is fou n · . '
in southern South America and central portions of North America.
Amphioceanic taxa occur on opposite sides of oceans. Amphiatlantic and
amphipacific taxa occur along the western and eastern coastal areas of the Atlanti
and Pacific oceans, respectively. The butterfly species Leptura oblitera, which is
found in eastern Asia and western North America, is an example of a northern
amphipacific species. In the South Pacific, the southern beech (Nothofagus ), which is
found in eastern Australia, New Zealand, and western South America, is an example
of a southern amphipacific taxon. Many shore-dwelling plants and animals, as well
as shallow-water marine species, have amphipacific and amphiatlantic distributions.
Sardines are coastal dwelling marine fish and have an amphiatlantic distribution.
A number of terrestrial and marine species are found in cold polar regions. FIGURE 13.7
but not in the warmer mid- and low latitudes. Such organisms are referred to as {lxobrychus exi
being bipolar. Many members of the whale family Balaenidae and the freshwater (Magnoliaceae)
 IIC DISTRIBUTIONS COMMON BIOGEOGRAPHICAL DISTRIBUTIONAL PATTERNS 389

Jtions fish family Petromyzonidae display bipolar distributions. Intolerance to warm

temperatures and competitive exclusion probably keep such taxa from occupying
imal taxa have geographic
warmer temperate and tropical areas.
N large latitudinal temp e~­
heir physiological sensiti,--
their inability to comper:: Dispersal Disjunctions
:e five such zonal distribe-
Biogeographers often classify disjunct distributions on the basis of how the dis-
;ircle the Arctic or Antarc-
junction is thought to have occurred. Dispersal disjunctions take place when a
titudes.A good example o::
jump dispersal event allows a population of a plant or animal species to become
?ria). Species belonging :c established a long distance from the main population. Dispersal disjunctions are
hem latitudes from no
particularly common for flying animals and for plants that possess seeds prone to
to the Atlantic Coast c:
long-distance dispersal. The least bittern (lxobrychus exilis), which is found
Austral (southern hem:-
throughout eastern North America and as disjunct populations in California and
in the cooler areas of tl::::
Oregon, is probably a dispersal disjunction (Fig. 13.7). Naturally established
ellent example of a boreE...
;ts of Alaska, Canada, ar:.:
din the intermediate zoe::
m example of a northe:::
found in the tropical zo
:ropical taxon.

Jgraphic ranges that a:-::

. Amphiregional distrib:.:-
m widely separated gee-
and can be translated 2.5
)hiregional have distrib:.:-
lre usually separated t ~
!S of this kind incluc::
Jpics but not in the trof':-
lhich includes sugarbee:;
ikely that amphitropiCE...
1peratures or competitio-
have about 160 differe:::
cal disjunct distributiOI:.s.
1 chilensis, which is four::
:hAmerica.
::eans. Amphiatlantic a::
astal areas of the Atlam~:
~eptura oblitera, which ::s

n example of a northe:::
~ch (Nothofagus), which ::s
th America, is an examp:::
•lants and animals, as ,,.e_
nphiatlantic distributim:.s.
)hiatlantic distribution.
md in cold polar regions.
anisms are referred to 2.5
nidae and the freshware:
w
0
Magnolia
FIGURE 13.7 The distribution of a possible dispersal disjunct species, the least bittern
(lxobrychus exilis) The global distribution of a climatic disjunct, the magnolia family
(Magnoliaceae) (after Robbins et al., 1984; Heywood, 1978).
390 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

populations of hydrochores such as coconut palms (Cocos nucifera) foun d t.: - Climatic
isolated tropical islands are also examples of dispersal disjunctions. junctions with
North Americ
in the eastern
Climatic Disjunctions moles from i1
A climatic disjunction occurs when climatic change makes a portion of a taxo:: Mountains. Di
range uninhabitable and splits a once continuous geographic distribution i~: deserts. A nu1
two or more separate parts. A classic example of a climatic disjunction is the m .=- (Eutamius), \\
nolia family of trees (Fig. 13.8) and shrubs (Magnoliaceae ). Fossil evidence sho California, Ne
that this ancient family of plants was distributed over much of the northern he uous ranges 11
sphere during the Cretaceous. The general cooling and drying of the world 's __ climate of tht:
mate that occurred during the Tertiary and Quaternary eliminated magno' elevations in
from most of its range. The family survives today in the Americas and in east :- replacement <
Asia (Fig. 13.7). The alligator genus, which is represented today with one spe i:"' led to the pre:
in China (Alligator sinensis) and one in North America (Alligator mississipiens:... Gorillas
(see Chapter 1) is a similar example of climatic disjunction created by clima _ distribution ir
cooling during the Cenozoic. We can find other species with similar distributio;::..:;. rainforest alo1
For example, the longhorn beetle genus Calloides lives in North America an _ in upland tro1
small area of southern China. Like the magnolia, the disjunct distribution of '-:- tions are sepE
longhorn beetles reflects the cooling and drying of global climate during the la:= dwelling gori
Tertiary and Quaternary that caused the fragmentation of once continuous t Pleistocene, tl
perate forest that stretched across Eurasia and North America. than it is tod
Africa linkin~
was probably

Geologic I
Geologic dis~
movement sp
tribution of t
South Ameri
that this gen
Chapter 10).
to the presen1
ern hemisphe
disjunction. 1
ostriches (Str
extinct moas
dae) of So uti
disjunct durir

Evolution;
Evolutionar)
portions of t
FIGURE 13.8 A sweetbay magnolia (Magnolia virginiana) from the Everglades of
sequent extir
Florida. Magnolias represent one of the most ancient families of flowering plants
(Magnoliaceae) and were once very widely distributed . The three surviving genera
disjunct fror
(Magnolia, Talauma, and Liriodendron) of the magnolia family grow in the wild today only arisen in thi~
in the Americas and in eastern Asia. They are therefore considered to be paleoendemics such as Aca
with a climatically disjunct distribution. south wes ten
 'HIC DISTRIBUTIONS
'Cocos nucifera) found o:.
disjunctions.
akes a portion of a taxor: ·=
agraphic distribution in: -
atic disjunction is the mc:;-
:ae ). Fossil evidence shov- =
mch of the northern he
j drying of the world 's c:_-
nary eliminated magnoL
e Americas and in easte:-
ted today with one sp eci~
L (Alligator mississipiens:_
nction created by cli m a~- ­
: with similar distribut io~ ­
:s in North America anc _
1isjunct distribution of tl::-
bal climate during th e la::-
n of once continuous te;:;-
\merica.
om the Everglades of
>f flowering plants
ee surviving genera
grow in the wild today on I
red to be paleoendemi cs
COMMON BIOGEOGRAPHICAL DISTRIBUTIONAL PATIERNS 391
Climatic changes during the Quaternary caused a number of climatic dis-
junctions within continents. For example, moles (Talpidae) were found throughout
North America during the Tertiary. Today they occur only along the west coast and
in the eastern deciduous and mixed-wood forest region. Cold conditions prevent
moles from inhabiting the northern portion of North America and the Rocky
Mountains. Dry conditions restrict moles from living in the western grasslands and
deserts. A number of other small mammals, such as some species of chipmunks
(Eutamius), which are found on isolated high mountains in the deserts of eastern
California, Nevada, and Utah are climatic relicts. These species had larger contin-
uous ranges in the Great Basin region during the last glacial maximum when the
climate of the region was cooler and moister. Conifer forest extended to lower
elevations in the Great Basin during that time. The onset of dry conditions and
replacement of low elevation conifer woodlands by desert during the Holocene
led to the present disjunct distributions of these small mammals.
Gorillas (Gorilla gorilla) have a particularly interesting climatically disjunct
distribution in Africa. One population of the species is found in lowland tropical
rainforest along the west coast of Africa, and another disjunct population is found
in upland tropical rainforest in the highlands of central Africa. The two popula-
tions are separated by savanna vegetation, which is inappropriate for the forest-
dwelling gorillas. During the early Holocene and other warm periods in the
Pleistocene, the climate of northern portions of Africa was both warm and moister
than it is today, and it is likely that continuous rainforest developed in central
Africa linking the coastal and upland rainforest. The range of the gorilla species
was probably continuous from the coast to the uplands during such periods.
Geologic Disjunctions
Geologic disjunctions occur when geologic processes such as plate tectonic
movement split once continuous ranges into two or more separate parts. The dis-
tribution of the southern beech (Nothofagus) in Australia, New Zealand, and
South America is an example of a geologic disjunction. Fossil evidence shows
that this genus was widespread on the supercontinent of Gondwanaland (see
Chapter 10). The breakup of Gondwanaland in the Cretaceous and Tertiary led
to the present disjunct distribution of the genus. Many plant families in the south-
ern hemisphere share a similar history and show a similar geographic pattern of
disjunction. The large flightless birds of the super order Ratites, including the
ostriches (Struthionidae) of Africa, the emus (Dromaiidae) of Australia, the now
extinct moas (Dinornithidae) of New Zealand, and the now extinct rheas (Rhei-
dae) of South America, evolved from a common ancestral lineage that became
disjunct during the breakup of Gondwanaland.
Evolutionary Disjunctions
Evolutionary disjunctions occur when two new species develop in different
portions of the geographic range of a widespread common ancestor. The sub-
sequent extinction of the common ancestral species leaves the two new species
disjunct from each other. Many amphitropical species of plants may have
arisen in this manner. Species of a number of different tree and shrub genera
such as Acacia and mesquite (Prosopsis) are found in both the deserts of
southwestern North America and the deserts of Chile and Peru. The mesquite
392 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

species Prosopsis juliflora found in the southwestern United States and t :::~ mammal pred;
closely similar species Prosopsis chilensis found in Chile are likely evoluti o~­ of the tuatara 1
ary disjunct species. There ar·
extremely div<
Cretaceous. Fe
Biogeographic Relicts from Greenlm
Biogeographic relicts are taxa that once had larger distributions but have becoiL:- restricted to tr
narrow endemics. Two types of biogeographic relicts are generally recognize.o one species, Z
The first type are climatic relicts, which are species whose geographic ranges ha\-= cycad is relati
been constricted due to recent climatic changes. Warming of the climate at tl:::- tulipifera) of t
end of the Pleistocene created many such climatic relicts. These climatic reli :: more diverse
are sometimes referred to specifically as glacial relicts. One good example is tl::::- America in tl
muskox (Ovibos moschatus), which is found today only in the arctic regions o: remain-the A
North America. Fossil evidence shows that muskox ranged across large portio ' odendron chin
of central North America and northern Eurasia during Pleistocene glacial pen- climates that e
ads. Similar fossil evidence shows that the alpine marmot (Marmota marmoT tree to extinct
ranged across much of Europe during the last glacial maximum, but its nature... classified as be
historic range is restricted today to the Alps. The coas
Many plants have glacial relict distributions. One classic example is t E of China (Met(
Norwegian mugwort (Artemisia norvegica) which ranged across central Euro ;: oendemics the:
during the last glacial maximum but is today restricted to small populations in the these two relic
Scottish Highlands, the coastal mountains of Norway, and the Ural Mountains o::: have seen, the
Russia. In North America, several glacial relict species of pines and cypress e~ extending fror
occur as endemics along the coast of California. These include the Torrey p in ~ occurs as a tin
(Pinus torreyana), which grows today only as small populations near San Diego been known fr
and on Santa Rosa Island, and the Monterey pine (Pinuis radiata), which is fou n extremely wid
in small scattered locations along the central California coast and adjacen and early Tert
islands, as well as on Guadelupe Island of Mexico. Fossil evidence, including ica, from the .
pollen from marine sediments collected off the California coast, suggests tha have also beer
these pines and cypresses formed extensive coastal forests and were widely dis· Shigeru Miki,
tributed along the coastlines of California and adjacent Mexico during Pleis- Kobe, Japan.1
tocene glacial periods. Cooler and moister conditions associated with glaciaJ different from
periods allowed these trees to occupy large ranges along the coast. Warmer and sils. No living
drier conditions became established during the Holocene, and this led to the new genus an
restriction of the pines and cypresses to their present small and disjunct distrib u- tally, during tl
tions. The glacial relicts of the California coast consist of some of the most highly small populat
endemic conifers in the world, including many that are found as tiny populations Szechuan Pro'
on just one site. The pines and cypresses are able to survive on islands and a few now called th
areas of the coast because these sites are typified by unusually large amounts of fog. belonged to t
The fog increases moisture availability and also keeps temperatures and evapora- truly was a liv
tion rates low during the summer months. North Amerie
Evolutionary relicts are paleoendemics that are survivors of formerly more trees and the 1
widespread and diverse evolutionary lineages. The tuatara (Spenodon punctatus) and the QuatE
is a lizardlike reptile that is found only in New Zealand. It is also the only native present status
reptile of that country. The tuatara is the last surviving species of an ancient rep- beautiful gold
tilian order called the Rhynsoles. Prior to the Cretaceous, this order had many are evergreen
species and was geographically widespread. The early separation of New Zealand the California
from the rest of Gondwanaland protected the tuatara lineage from contact and temperate reg
competition with later evolving reptiles and mammals. The lack of reptile and part of its forr
 'HIC DISTRIBUTIONS
rn United States and tb.:
:hile are likely evolutioc-
ributions but have becom::
are generally recognize.:
'se geographic ranges ha\·;;-
ning of the climate at tt::
licts. These climatic relic::;;
. One good example is tl::::
tly in the arctic regions c.:
nged across large portio
g Pleistocene glacial per:-
·mot (Marmota marmm::.
maximum, but its nat ur2...
ne classic example is ti:::
~ed across central Eur o~
:o small populations in tt::
nd the Ural Mountains c:
=s of pines and cypress~
e include the Torrey pi:::-
pulations near San Die5c
:is radiata), which is fo u;:::
)fllia coast and adjace::.-
:;ossil evidence, includi::.;
Jrnia coast, suggests th"--
ests and were widely cts--
=nt Mexico during Pleis-
s associated with glac>
1g the coast. Warmer a;: :
:::ene, and this led to tl::
nail and disjunct distri b:.:-
f some of the most high:_
found as tiny populatio::.s
vive on islands and a fe-
!lally large amounts of fo;.
mperatures and evapoE-
rvivors of formerly mo:-::
tra (Spenodon punctarus
It is also the only na ti\·;;-
pecies of an ancient re;-
US, this order had mac·
paration of New ZealaLt:
neage from contact a- -
The lack of reptile ac:
393
COMMON BIOGEOGRAPHICAL DISTRIBUTIONAL PATTERNS
mammal predators on New Zealand was also an important factor in the survival
of the tuatara there.
There are many evolutionary relicts among plant species. Cycads were an
extremely diverse and geographically widespread group of plants prior to the
Cretaceous. Fossil cycads are found from the equatorial regions to Siberia and
from Greenland to Australia. Today they survive as about 100 species that are
restricted to tropical and subtropical regions. Only one native genus containing
one species, Zamia fioridana, remains in the whole of the United States. This
cycad is relatively common in Florida. Similarly, the tulip tree (Liriodendron
tulipifera) of the southeastern United States is an evolutionary relict of a once
more diverse and widespread family that extended across Eurasia and North
America in the late Cretaceous and early Teritiary. Today only two species
remain-the American tulip tree and a related species in southeastern Asia (Liri-
odendron chinense) . As is the case with the magnolias, increasingly cool and dry
climates that developed during the late Tertiary and Quaternary drove the tulip
tree to extinction across most of its range. Thus, tulip trees and magnolias can be
classified as both evolutionary relicts and climatic disjuncts.
The coastal redwood of California and its nearest relative, the dawn redwood
of China (Metasequoia glyptostroboides), are interesting examples of disjunct pale-
oendemics that are evolutionary and climatic relicts. Unraveling the history of
these two relicts is a very interesting chapter in biogeographical research. As we
have seen, the California redwood is found only along a narrow strip of coastline
extending from central California to near the Oregon border. The dawn redwood
occurs as a tiny population in the Szechuan Province of central China. It has long
been known from fossil evidence that the ancestors of the California redwood were
extremely widespread throughout the northern hemisphere during the Cretaceous
and early Tertiary. Fossils of redwoods have been found throughout North Amer-
ica, from the Arctic to the modern deserts of the Great Basin. Redwood fossils
have also been found throughout Eurasia. In the early 1940s, the Japanese botanist,
Shigeru Miki, described a new fossil species of redwood from fossils found near
Kobe, Japan. The arrangement of the needles and the cones of the fossil species was
different from that of the living California redwoods and previously collected fos-
sils. No living species of trees were like the new fossil redwood, so Miki created a
new genus and species for the fossils-Metasequoia glyptostroboides. Coinciden-
tally, during the 1940s a Chinese forester named Tisang Wang reported finding a
small population of a previously unknown tree species in a remote area of the
Szechuan Province of China. Subsequent comparison of the newly discovered tree,
now called the dawn redwood, and fossil material showed that the living trees
belonged to the fossil species Metasequoia glyptostroboides. The dawn redwood
truly was a living fossil! Fossils of dawn redwood have now been recognized from
North America and Greenland in addition to Asia. As was the case for the tulip
trees and the magnolias, increasing cold and dry conditions during the late Tertiary
and the Quaternary led to the huge decrease in the range of the redwoods and their
present status as evolutionary and climatic relicts. The living dawn redwood turns a
beautiful gold color and loses its needles during the winter. California redwoods
are evergreen. By being deciduous, the dawn redwood can tolerate cold better than
the California redwood. Today dawn redwoods are grown in gardens throughout
temperate regions. A species that was unknown only a few decades ago is regaining
part of its former large range through human intervention.
394 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS BIOGEOGRAPHIC DIST

BIOGEOGRAPHIC DISTRIBUTIONS Centers of

AND THE RECONSTRUCTION The search for
OF EVOLUTIONARY HISTORY long been a pr
proposed to id
How can the modern distribution of organisms be used to reconstruct the evo:_- the 1940s the I
tionary history of plant and animal species? Thi~ question has been a central co- that had been
cern and source of contention for biogeographers for over 150 years. Let's st _ species. One c
back in time and look at the dawn of these debates. In 1843 two British naval shi;:->- greatest numb
the HMS Erebus and the HMS Terror, returned to England from a voyage graphic distrib
exploration around Antarctica and the adjacent continents. Returning with = est number of
expedition was a young English doctor and naturalist named Joseph D. Hook""- declines outw<
He brought back with him a large collection of specimens and a vexing scien ti:_ nated in weste
puzzle. Hooker had noted that many of the same plant families and genera co -
the developm<
be found in the widely separated floras of New Zealand, Tasmania, southern A! '--
evolutionary d
tralia, and southern South America. In addition, he had found fossil wood fro-
of new pheasa
such genera on the remote and treeless Kerguelen Islands off the coast of Antas.:-
is a correct in1
tica. After deciding that long-distance dispersal could not have produced th :~
we map the we
distributional patterns, Hooker concluded that these plants were the survi' i;:=
cult to interpr·
relicts of a once extensive forest that had grown on all of the continents and rn a.; _
equally high c
islands of the southern hemisphere. He suggested that the southern land ar 2..::
Both areas are
must have once been joined together and geologic events must have caused ·1:::
originate in S<
breakup, fragmenting their flora and leading to the evolution of new species.
dence, it is irr
We now know that Hooker was correct. The geographic distributions c-
exactly where
organisms and fossils he studied were the result of the breakup of the Gondwam.-
land supercontinent. As we saw in Chapter 9, geologic events that cause the bre -- with highest di
ing apart of formerly continuous geographic ranges are called vicariance events. - for the taxa. F·
the twentieth century, the primacy of vicariance events in the evolution of n . horse and zeb
species was most forcefully championed by the Venezuelan biogeographer, Leo- evidence that
Croizat. We will learn more about Croizat and his ideas later in this chapter. dispersed to A
Hooker went on to a very distinguished career, and as fate would have it. ~ no surviving S]
co-chaired the 1858 session of the Linnaean Society during which Charles Das- Another
win and Alfred Wallace's papers on natural selection were read. Ironically, bo :. the most evoll
Wallace and Darwin disagreed with Hooker regarding the relative importance o: being studied.
the geologic breakup of landmasses versus long-distance dispersal in generat in~ in form to the
the biogeographic disjunctions seen in the southern hemisphere. Wallace an-' far from the c<:
Darwin believed that dispersal was very effective over geologic time scales an -= Hennig and L
could explain many of the disjunct distributions of plants and animals that ar ~ Many flowerir
present today. In addition, Darwin believed that dispersal was fundamental t South Pacific,
evolution and to the formation of new species. He formulated the center of origir. Australia, and
model to link geographic distributions to evolutionary history. He postulated tha: region as the
evolution and speciation occur as species disperse and spread outward from cen- ever, that fossi
tral points of origin. Darwin suggested that speciation during dispersal is driver; ering plants. Ir
by the new environmental selection pressures encountered when species dispers have suggeste
into new areas. In the twentieth century, famous biogeographers and evolution- advanced spec
ists such as George Gaylord Simpson and Ernst Mayr argued for the importanc These biogeog
of dispersalist evolution in the creation of new species. After 150 years, th promotes evol
debate still continues between vicariance biogeographers and dispersalists-and tive forms, an•
can be extremely heated. With the contentious nature of this debate in mind, let's range of the ta
consider both dispersalist and vicariance approaches to reconstructing evolution- The prec
ary history on the basis of modern distributions. the evolutiona
 'HIC DISTRIBUTIONS
d to reconstruct the evoL.-
ion has been a central co::.-
over 150 years. Let's sr.:--
843 two British naval shi:-:..
:<:ngland from a voyage
inents. Returning with C:".=
named Joseph D. Hookc-:-
ens and a vexing scien!i..;:;,_
families and genera cot:.._
j, Tasmania, southern r\t!._"-
td found fossil wood fro=
tds off the coast of Anta:-:-
not have produced thes.:
plants were the survi \t=
f the continents and maic-
t the southern land are"
:nts must have caused :.::=
ution of new species.
~agraphic distributiom -
·reakup of the Gondwa.:::..::.-
/ents that cause the breh-
~alled vicariance evenrs. --
s in the evolution of nc-
elan biogeographer, L- --
later in this chapter.
d as fate would have it. :::=
1ring which Charles D c:-
vere read. Ironically. bo:::
he relative importance :
:e dispersal in generati:.=-
llemisphere. Wallace a=:
geologic time scales a=.:
mts and animals tha t CC=
:rsal was fundament al :
1lated the center of orig
is tory. He postulated tL
pread outward from ce:::.-
juring dispersal is dri\·c.:
ed when species dispe3=
)graphers and evolurio:::.-
rgued for the importac:::
es. After 150 years. ~::
rs and dispersalists-~.:
this debate in mind. le:
~econstructing evoluric:::.-
BIOGEOGRAPHIC DISTRIBUTIONS AND THE RECONSTRUCTION OF EVOLUTIONARY HISTORY 395
Centers of Origin and the Dispersalist Model
The search for the centers of evolutionary origin of plant and animal species has
long been a preoccupation of biogeographers. Many different criteria have been
proposed to identify the geographic locations in which species have originated. In
the 1940s the biogeographer Stanley Cain listed and critiqued 13 different rules
that had been proposed to indicate the geographic origin for plant or animal
species. One commonly applied criterion is to identify the area in which the
greatest number of related species are found. For example, if we map out the geo-
graphic distribution of pheasant genera (Phasianinae ), we find out that the great-
est number of genera (8) occur in western China and that the number of genera
declines outward from that region. Thus, we might conclude that pheasants origi-
nated in western China and that continued evolution and speciation have led to
the development of many genera and species there, while dispersal and further
evolutionary development by some of these species have led to the development
of new pheasant species that occupy other areas of Asia and Africa. Perhaps this
is a correct interpretation of the evolutionary history of pheasants. If, however,
we map the world distribution of palm genera, we see a pattern that is more diffi-
cult to interpret. In the case of palms, there are two areas in which you have an
equally high diversity of palm genera, South America and southeastern Asia.
Both areas are surrounded by regions of decreasing palm diversity. Did the palms
originate in South America or southeastern Asia? Without fossils or other evi-
dence, it is impossible to say on the basis of modern geographic distribution
exactly where the palms originated. Even if there is only one geographic area
with highest diversity, it does not unequivocably mean that is the center of origin
for the taxa. For example, the highest species diversity for living members of the
horse and zebra genus (Equus) occurs in Africa. However, we know from fossil
evidence that the ancestors of horses first evolved in North America and later
dispersed to Africa. Until the reintroduction of horses by the Spanish, there were
no surviving species of horses in North America.
Another common method for identifying the center of origin is to look for
the most evolutionarily primitive or advanced members of the genus or family
being studied. Some scientists have argued that primitive species, which are close
in form to the supposed ancestral species, are most likely not to have dispersed
far from the center of origin. Two major proponents of this hypothesis were Willi
Hennig and Lars Brundin. The primitive characteristics are called pleisiomorphs.
Many flowering plants with apparently primitive characteristics are found in the
South Pacific, including portions of Fiji, New Caledonia, New Guinea, eastern
Australia, and the Malay Archipelago. This has led some scientists to identify this
region as the center of origin for flowering plants. Other scientists argue, how-
ever, that fossil evidence points to South America or China as the origin of flow-
ering plants. In complete contrast, some biogeographers, such as P. J. Darlington,
have suggested that the centers of origin will possess the most evolutionarily
advanced species. The supposed advanced characteristics are called apomorphs.
These biogeographers contend that high genetic diversity at the center of origin
promotes evolution and speciation. The newly evolved forms outcompete primi-
tive forms, and relegate the latter to survival at the margins of the geographic
range of the taxon.
The preceding discussion shows that it is difficult to confidently determine
the evolutionary center of origin of a species on the basis of modern geographic
396 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS BIOGEOGRAPHIC DISTR

distributions. After a careful review of the 13 different criteria used to ideo...:...

centers of origin, Cain concluded that none of them was completely reliable. E
conclusions have been echoed by many other biogeographers. What then do ~
do with Darwin's theory regarding the evolutionary importance of natural s e l~­
tion during dispersal from the center of origin? Biogeographers such as He - _
and Brundin suggest that Darwin was essentially correct and that there shoul ' ~ _ Ferns c
a gradation from primitive pleisomorphs at the center of origin to more advan ~­
apomorphs at the geographic range boundaries of the taxon. Brundin called 7·-
the rule of deviation, while Hennig coined the term progression rule to descr:_ _
this phenomenon. There is, however, little direct field evidence to support -- -
widespread presence of clear geographic gradients of primitive to advan :::-:.
forms in either plants or animals. The best evidence for the presence of such p::---
gressions comes from the distributions of insects on some island chains. F
example, fruit flies (Drosophila) on the Hawaiian Islands show a progress: =
increase in supposed apomorphic traits as one goes from the oldest islands, s · --
as the outer islands beyond Kauai, to the younger islands, such as Hawaii. S!!--
progressions appear to be somewhat rare, however. In addition, many scient·-:..
have questioned how one can confidently define and identify primitive
advanced traits in the first place. Finally, as Croizat has forcefully argued, Dan _
never presented clear and convincing arguments of exactly how dispersal p;---
motes evolution and speciation at the edges of the expanding ranges, nor did ::::.~
explain why it is that new traits developed at the edges are not introdu :::-:.
throughout the entire range of the species.
FIGURE 13.9 I
Cladistic Biogeography ferns, cycads, pin
number of traits r
One of Brundin's and Hennig's most important contributions was to help link - ;
analysis of geographic distributions directly to phylogenetic reconstructions. Ph.-
logenetic reconstructions of speciation history and the evolutionary link ag~
between species are often presented as branching trees called taxon cladograr-_ together cycad~
(Fig. 13.9). The formulation and study of such diagrams is called cladistics. He:::- from the ferns.
nig pioneered the cladistic method in the 1950s. The endpoints on the taxo;: ing vascular ca
cladograms are modern taxa, such as the different species, genera, or families c: they do not pos
plants or animals that exist today. The construction of the cladogram starts \\i -- do not have fe<
measuring a set of physical traits for a group of species and determining whi ::. off because am
traits are common to all of the species being studied, which traits are shared · : and do not pc
some, and which traits are unique to individual species. It is assumed that tl:: :c roughly similar
traits common to all species are derived from the common ancestor of the plan:.:: they are split ol
or animals. Species that share many traits are assumed to be more recently differ- gram, magnolic
entiated from each other and more closely related than species that share fe tial oils in their
common traits. The species that are similar to each other in terms of physica. not have flowe1
traits are placed close together on the cladogram. Thus, the proximity of the dif- we would infer
ferent taxa to each other infers their evolutionary relationship. The nodes for th; to gnetales and
branches of the cladogram indicate the relative timing of evolutionary diver- The diffe1
gence. The positioning of the nodes corresponds to the timing of the loss o: terms of geogn
shared characteristics or the appearance of new characteristics. The nodes repre- animal will be
sent points of evolutionary divergence between lineages. have many sha1
We can examine a simplified representation of a cladogram for plants to se ist biogeograpl
how taxon cladograms are actually constructed (Fig. 13.9). Hennig produced suet graphic range
a cladogram based on 29 morphological traits. The base of the cladogram groups recently will be
 >HIC DISTRIBUTIONS
BIOGEOGRAPHIC DISTRIBUTIONS AND THE RECONSTRUCTION OF EVOLUTIONARY HISTORY 3 97

nt criteria used to iden

1as completely reliable. E

•
graphers. What then do ~ ~
~l.~
nportance of natural seL:·: .I'~"
~ographers such as He~
'ct and that there shou lc~ ~·-= Ferns Cycads Pines Gingko Gnetales Magnolia
of origin to more adva nc~­
• taxon. Brundin called r"-
rogression rule to descr..::- _
d evidence to support i:..:.=
of primitive to advanc-~..:
r the presence of such p::---
l1 some island chains. F- ~
;lands show a progress. _
om the oldest islands, se:··
mds, such as Hawaii. S12:·
1 addition, many scien··
md identify primitiY
forcefully argued, Da
:xactly how dispersal ~=- -­
Janding ranges, nor die .:::
edges are not introdu:=-

FIGURE 13 .9 A simplified cladogram for magnolias (representing angiosperms) and

utions was to help link :..::
aetic reconstructions. P::
:he evolutionary linkc~::­
s called taxon cladogr~
s is called cladistics. H.:;;: ·
: endpoints on the tax: -
~ies, genera, or families
the cladogram starts w-:· -
=s and determining wt: --
.vhich traits are shared
es. It is assumed that :..::_
aon ancestor of the pic..:.
.o be more recently diE::---
an species that share :::
ther in terms of phys:.-
:, the proximity of the -=
ionship. The nodes for --
1g of evolutionary di,-;e-
the timing of the loss
teristics. The nodes rep:-::
s.
adogram for plants to 5.-"-::
9). Hennig produced St::.::."
• of the cladogram gro'.::'
ferns, cycads, pines, gingkos, and gnetales. The splits in the cladogram are based on a
number of traits related to reproductive structures, leaf shape, and plant chemistry.
together cycads, pines, gingko trees, gnetales, and magnolias and separates them
from the ferns. All of these groups of plants share common traits such as possess-
ing vascular cambium and seeds. Ferns are not included in this group because
they do not possess seeds. The next group to be split off is the cycads because they
do not have features such as ancillary branching and true pollen. Pines are split
off because among other things pines possess needles rather than broad leaves
and do not possess true flowers. The gingkos do have broad leaves that are
roughly similar to those associated with magnolias and other flowering plants, but
they are split off because they do not possess true flowers. At the top of the dado-
gram, magnolias are split from gnetales because the latter do not possess essen-
tial oils in their leaves, have a different method of producing pollen grains, and do
not have flowers similar to magnolia and other angiosperms. From the cladogram
we would infer that magnolias are most similar in form and evolutionary history
to gnetales and most distant from ferns.
The different endpoints (taxa) on the cladogram can also be thought of in
terms of geographic areas. Older species that are similar to the ancestral plant or
animal will be grouped together at one end of the cladogram. These species will
have many shared primitive traits. According to the progression rule of dispersal-
is! biogeographers, these taxa will be found in the central portions of the geo-
graphic range of the taxon (Fig. 13.10). Younger species that have diverged
recently will be found at the other end of the cladogram and will be found in geo-
398 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS BIOGEOGRAPHIC DISTF

Vicariance hypothesis of the role of g

-- --
(Geologic event followed by speciation)
1
A
2
B
3 Sp
c =
-
early voice in t
1950s, he devel

~~
1 11 1 1 11 1 11 1 that is called p:
1 A1 1 A1 plants and anir
3~3
A 1 1
1 11 11 groups, such as
1 1 1 1

- 1

-
Dispersal hypothesis
1

--
(Jump dispersal event followed by speciation)
Cladogram
and disjunctior
capabilities of 1
shared pattern~
dispersal. Like
~~ repeated patter
1 1 1 1 tributions and t
1 forcefully argw
1 A 1 1
1 1 1 tance to the de'
Land areas ing vicariance e
and species isolated geogra
FIGURE 13.10 A three species and three area taxon-area cladogram with vicariance a -
To suppo
dispersalist models of geographic speciation that would produce such a cladogram. that lead to the
taxa and then c
this exercise h1
graphic areas to which the taxa have most recently dispersed. Thus, the endpoiL:_ connecting rei;
on the cladogram can represent not only different species but different ge._ found that mm
graphic areas as well. In some cases, the geographic areas where species a:-:: the Pacific coa~
found are listed on the cladograms rather than the species themselves. Cia c- of Asia, so he d
grams that list and categorize the geographic areas in which the taxa are fo u ;: tracks connect
are called area cladograms. The linking of cladograms with biogeographic histo:-_: appeared for n
is referred to as cladistic biogeography and is an important tool for deducing eYc- As a number c
lutionary history. represented pa
In the 1960s, Brundin produced cladograms for southern hemisphere mi d~;: 13.11). Croizat
species in which he replaced the scientific name for each species on the cl a d~ South Pacific C
gram with the name of the geographic region in which the species was found . l= nections that s
doing so, he produced the first area cladogram. This classic cladogram tied the e\·~ scape, such as
lutionary linkages between different midge species to the geographic regions ·- dismissed cont
which they lived. We now know that the taxon and area cladograms for the south- disjunct populc:
ern hemisphere midges strongly reflect the geologic and geographic history o: nental drift. At
Gondwanaland. Groups of midges such as the genera Afrochlus and Archaeochlu.:; tion, and he
are found only in Africa and are morphologically distinctive from other grou ~ landbridges ac1
because of Africa's early separation from the rest of Gondwanaland and the lo £ with the bioge
period in which midges there have been isolated from other southern hemispher; merged landbr
midges. Genera such as Parochlus and Podonomus occur on New Zealand, Sour.i: South America
America, and Australia, but are represented by different species in each of thE graphic eviden1
three areas. These land areas were joined together longer than was the case f ' dence of the pa
Africa and the rest of Gondwanaland and thus share genera. However, sufficien- intersected eac
time has elapsed since the separation of New Zealand, South America, and Aus- Croizat m
tralia for distinctive species to arise in each region. ing tracks, one
example, fig tre
and Australia. ~
Panbiogeography and Vicariance Models tralasian and A
Deep dissatisfaction with the ambiguous nature of center of origin approaches and are to the Afric
Darwin's model of dispersal and geographic evolution led to a critical revaluation in flora and fal
'HIC DISTRIBUTIONS
2 3 Spec _
8 C AicE
Cladogram
] tributions and that geologic events had split these once continuous ranges. Croizat
Land areas
and species
1
adogram with vicariance a-:.
1ce such a cladogram.
>ersed. Thus, the end poi;:;.:.
;pecies but different ge-:-
: areas where species c..:-::
pecies themselves. Clac:-
which the taxa are fmc_
;vith biogeographic histc::-
:ant tool for deducing e\ ,-_
1uthern hemisphere mi c.;:-:
ach species on the clac:-
the species was foun d.·-
;ic cladogram tied the
the geographic regions -
cladograms for the sou:..:-
md geographic history
rrochlus and Archaeoch.·
inctive from other grot.:;
mdwanaland and the lo:.._
ther southern hemisphe:-::
1r on New Zealand, S o~.::.:.
~nt species in each of C:::
ger than was the case ~2-
~nera. However, sufficie=
South America, and At:..'-
r of origin approaches a;::
~d to a critical revalua t!c
BIOGEOGRAPHIC DISTRIBUTIONS AND THE RECONSTRUCTION OF EVOLUTIONARY HISTORY 399
of the role of geography in reconstructing evolutionary history. The most strident
early voice in this reassessment was the Venezuelan botanist Leon Croizat. In the
1950s, he developed a new method of linking geography and evolutionary history
that is called panbiogeography. Croizat analyzed the distributions of hundreds of
plants and animals and realized that in many cases the species of very different
groups, such as plants and insects, shared similar geographic patterns of endemism
and disjunction. Croizat noted that there were great differences in the dispersal
capabilities of these different species. It did not seem likely, therefore, that these
shared patterns of endemism and disjunction could have arisen by long-distance
dispersal. Like Hooker, Croizat decided that the only logical explanation for such
repeated patterns of disjunction was that these taxa had once had much larger dis-
forcefully argued that long-distance dispersal events were rare and of little impor-
tance to the development of new species. Rather, most speciation occurred follow-
ing vicariance events that split the ranges of species into two or more separate and
isolated geographic regions.
To support his ideas and provide evidence of the past vicariance events
that lead to the splitting of ranges, Croizat amassed data on hundreds of disjunct
taxa and then drew lines connecting the ranges of related disjunct species. From
this exercise he showed that there were a number of frequently repeated lines
connecting related disjunct taxa. He called these lines tracks. For example, he
found that many genera of plants and animals had disjunct distributions along
the Pacific coasts of North America and across the ocean along the Pacific Coast
of Asia, so he drew lines along the coasts and across the Pacific that represented
tracks connecting the distributions of the related taxa. When the same tracks
appeared for many different taxa, Croizat called these lines generalized tracks.
As a number of generalized tracks crossed the oceans, he concluded that they
represented past land connections that had been cut by geologic changes (Fig.
13.11). Croizat found that a number of tracks joined taxa on both sides of the
South Pacific Ocean, the Indian Ocean, and the South Atlantic Ocean. The con-
nections that span oceans or other prominent features of the geographic land-
scape, such as mountain chains, are called baselines. Oddly, Croizat originally
dismissed continental drift as a mechanism for severing connections between
disjunct populations. In fact, he explicitly denounced Wegener's model of conti-
nental drift. At first , Croizat argued that the continents had never shifted posi-
tion, and he hypothesized the creation and destruction of a number of
land bridges across the oceans. The locations of these "lost" connections coincide
with the biogeographic tracks identified by Croizat. He argued for now sub-
merged landbridges across the Pacific, Indian, and Atlantic oceans that joined
South America, Africa, and Australia in the past. He believed that the biogeo-
graphic evidence of past connection was far more reliable than the geologic evi-
dence of the past geography of the globe. Croizat called the points where tracks
intersected each other nodes.
Croizat argued that, by examining geographic distributions and construct-
ing tracks, one could deduce both evolutionary history and geologic history. For
example, fig trees (Ficus) are found in South and Central America, Africa, Asia,
and Australia. Systematics studies based on fig morphology suggest that the Aus-
tralasian and American species are more closely related to each other than they
are to the African species. Croizat's tracks, however, suggest a closer relationship
in flora and fauna and greater past geographic connections between Africa and
400 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS BIOGEOGRAPHIC DISTF

Although
posed by Croiz;
ing geography
strongly agains
speciation even
in allopatric spE
of evolutionary
During tl
have built on
structing evolt
these biogeogr
United States
was to combin(
speciation witl
approach is ca
branches in cla
resenting the c
the history of t
ance event (Fi~
species and the
FIGURE 13. 11 Some ofthe most important tracks identified by pan biogeographic ance events. A<
analysis. The tracks join the disjunct distributions of related plant and animal taxa (after a the history of e
variety of sources including Rosen, 1988; Brown and Lomolino, 1998; Craw et al., 1999). The early
events were raJ
tion. They assu
once continual
Australasia than between America and Australasia. Croizat therefore argued t o tions were a re
the systematics studies were flawed and that the figs of Australasia were m _ grams, the vic;
closely related to those from Africa. In defense of this hypothesis, it has be = similarity betw
shown that a closer relationship exists between Australasian and African wa.:,~ that the fo ssil 1
species that are closely associated with figs as pollinators than there is betwe = cladograms if it
Australasian and American wasp species that serve as fig pollinators. biogeographen
Croizat's ideas were startling to many mid-twentieth-century biogeogr~­ grams and havt
phers. Those who disagreed with him found much ammunition to refute · increasingly rec
claims. Croizat made a number of mistakes in taxonomy and geology. He di>- ate biogeograp
missed the frequency and significance of long-distance dispersal. In addition, ing cladograms
proposed landbridges often had no basis in geologic evidence. Today the e': - results of vicar
dence for plate tectonics and continental drift is too overwhelming and clear : - vicariance ever
be overlooked. Finally, the extreme arrogance and scorn evident in some o: species are ind(
Croizat's writings did little to engender respect or civil discourse on his ideas. species from th
Today, few biogeographers strictly adhere to Croizat's original panbiogeo- An inforn
graphic concept and interpret both evolutionary and geologic history sole _ geologic and '
from the basis of drawn tracks. One exception is in New Zealand, where panbio- allopatric speci
geographic ideas have found favor and panbiogeographers such as R. C. Cra Eucalyptus of t
raise some interesting questions in trying to explain the complex biologi these are ender
affinities of that country's plant and animal species. For example, panbiogeogra- characteristics <
phers argue that certain features of the New Zealand flora and fauna , such as th- cladogram was
presence of Nothofagus trees, appear to be best explained by an ancient lane tributions of th<
connection to New Guinea rather than by a past connection to Australia an - disjunct popula
Antarctica as part of Gondwanaland. drying of Austr
>HIC DISTRIBUTIONS
ld by panbiogeographic
Iant and animal taxa (afte r 2
o, 1998; Craw et al., 1999).
oizat therefore argued ·:.
; of Australasia were II L
his hypothesis, it has b.:::-
ralasian and African \ \ ·_::s-
tars than there is betw.:::-
fig pollinators.
:ntieth-century bioge o~­
ammunition to refute ::.....
tomy and geology. He "'
e dispersal. In addition. ::......
~ evidence. Today the c
werwhelming and cle<L
scorn evident in some
ivil discourse on his ide:
izat's original panbiog::--
ld geologic history so:::
:w Zealand, where pant: -
1phers such as R. C. C :-~
in the complex biolog: ::
)r example, panbioge o~­
]ora and fauna, such as ·-
·lained by an ancient J- -
nnection to Australia ::..:;._
BIOGEOGRAPHIC DISTRIBUTIONS AND THE RECONSTRUCTION OF EVOLUTIONARY HISTORY 401
Although the classical panbiogeographic models and methods initially pro-
posed by Croizat have relatively few adherents today, some of his arguments regard-
ing geography and speciation have been much more influential. Corizat argued
strongly against Darwin's dispersal model of evolution. According to Croizat, all
speciation events arise from vicariance events that split geographic ranges and result
in allopatric speciation. He also showed that many of the dispersalist interpretations
of evolutionary history were ad hoc and lacking in rigorous testing of hypotheses.
During the latter half of the twentieth century, a number of biogeographers
have built on some of Croizat's ideas to formulate a new approach to recon-
structing evolutionary history from geographic distributions. Notable among
these biogeographers are G. J. Nelson, N. Platnick, and D. E. Rosen from the
United States and C. J. Humphries from Britain. The approach they pioneered
was to combine Croizat's concepts on the primacy of vicariance events in causing
speciation with Hennig's systematic organization of taxa on cladograms. This
approach is called vicariance biogeography. Following from this approach, the
branches in cladograms for plants and animals can be thought of as directly rep-
resenting the different geographic vicariance events that have occurred during
the history of the species. Thus, each node where the cladogram splits is a vicari-
ance event (Fig. 13.10). The endpoints of the cladograms represent both modern
species and the geographic areas in which the species developed following vicari-
ance events. According to the vicariance approach, the cladogram presents both
the history of evolution and the geologic and geographic history of the earth.
The early vicariance biogeographers followed Croizat's tenet that dispersal
events were rare and unimportant in producing disjunction and allopatric specia-
tion. They assumed that all disjunction and speciation arose from the splitting of
once continuous ranges by geologic events. They believed that modern distribu-
tions were a reflection of these past events. In producing taxon and area dado-
grams, the vicariance biogeographers argued that only evidence of physical
similarity between taxa or geographic locations should be considered. They felt
that the fossil record could be misleading and would corrupt the production of
cladograms if it was incorporated in their construction. More recently, vicariance
biogeographers have used fossil and geologic evidence as tests of their dado-
grams and have proposed phylogenetic and geographic histories. They have also
increasingly recognized that climatic changes as well as geologic changes can cre-
ate biogeographic barriers and produce vicariance events. In addition, by prepar-
ing cladograms of different plant and animal groups from the same regions, the
results of vicariance biogeographic reconstructions can be further tested. If the
vicariance events that are detected in preparing cladograms for one group of
species are indeed correct, similar cladograms should arise when other groups of
species from the same area are examined.
An informative cladistic study of Eucalyptus species in Australia shows how
geologic and climatic changes can produce vicariance events and lead to
allopatric speciation even in a relatively small area. Today some 29 species of
Eucalyptus of the monocalyptus group are found in southern Australia. Many of
these are endemic to one of eight regions along the coast. Based on 51 physical
characteristics of the tree species and their geographic distributions, a taxon-area
cladogram was constructed. The cladogram suggests that the once continuous dis-
tributions of the ancestral monocalyptus trees were split into increasingly smaller
disjunct populations by incursions of the sea in the Tertiary and then progressive
drying of Australian climate that occurred in the Late Tertiary and Quaternary.
402 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS BEYOND

Cll
.S?
Cll
(.)
biogeographer
Cll Cll
Ql Q; -~ (.)
Cll
(.) many microev
c E E Cll -~
-~ ·s Ql ·~
<1: <1: Ql c E the interpretat:
~ C)
r. r. c. -~ ·s E
t5
::::J
5:
Ql
"5
0
t:
0
e
::::J ~ C)
<1:
r.
<1:
r. c
parallel evolut
<1: z CIJ z LlJ t5
::::J
5:
Ql
"5
0
t:
0 of a clade in sE
N N N F F <1: z CIJ z ings and splits
same physical
graphic areas, i
bility that the t
biogeographic
events but do 1
lineages diverg
cladograms alo
of vicariance e'
Trees Frogs of evolutionar)
N = Nothofagus likely to be im
F =Fagus work of vicaria
In recent
FIGURE 13.12 Simplified taxon and area cladog rams for beech species (Fagus) of th e
the testing of t
northern hemisphere and southern beech (Nothofagus) species and fo r f ro g species (da ta
traditional tax<
from Patterso n, 1981).
which the term
lar genetics an,
research. Much
Allopatric speciation processes led to the development of new eucalyptus spe -~
mtDNA (see C
in these disjunct locations. The cladistic reconstruction of progressive disjuncti -
genetics appro
and speciation through vicariance events is consistent with geologic and paleoc:...-
between croco<
matic evidence of the history of southern Australia.
A classic study comparing the cladograms from a number of different pia::· traits suggest t
and animal groups was conducted by the British paleontologist, Colin Patterso- related to turtl<:
He constructed taxon and area cladograms for several groups of fishes, turtl :. turtles, and birc
birds, frogs, insects, and beech trees. He showed that the area cladograms for th ~ The combinatic
frogs, birds, and fishes were similar, and he consistently divided the groups into _ is an importan t
Australian-New Guinea section and a South America, North America, and Euro- tionary history.
pean section. In contrast, the cladograms for the insects and beech trees divid ~ Phylogeo1
the groups into a North American and European section and an Australia, Ne regarding the d
Guinea, and South American section (Fig. 13.12). The general division of tb-=- analysis (see cr
cladograms between northern and southern continents and between Old Worl::: ariance events 1
and New World continental areas is in agreement with the hypothesis that vicaf.- of new species
ance events caused by the breakup of Pangaea in the northern and souther::: colleagues hav<:
areas of Gondwanaland and Laurasia, and then into the modern continents, is tb- the likely timin
most important factor controlling speciation and the modern distributions o:: species and sut
these different species. Recent interchanges of birds and frogs between Nor United States a
and South America after the two continents were joined by the Isthmus o:: distinct species
Panama has led to greater similarity in the faunas in North and South America. years or so whE
led to expansio
graphic ranges
BEYOND VICARIANCE BIOGEOGRAPHY: ever, detailed 1
THE PHYLOGEOGRAPHIC REVOLUTION revealed s urpri ~
subspecies. The
At present, the vicariance model of speciation and the intepretation of dado- the different sp
grams as records of vicariance events are relatively widely accepted among past and may t
'HIC DISTRIBUTIONS BEYOND VICARIANCE BIOGEOGRAPHY: THE PHYLOGEOGRAPHIC REVOLUTION 403

biogeographers. Vicariance events are undoubtedly important in explaining

ca
(.)
ca many microevolutionary and macroevolutionary events. One weakness is that
(.)
ca ·~ -~
Q)
c E E
the interpretation of taxon and area cladograms can be seriously compromised if
·:; <(
(!l .c
<(
.c
c
c. parallel evolution has occurred and produced morphologically similar members
:;: "5 c
t
0 of a clade in separate areas. In the classic construction of cladograms, all group-
Q)
z
0
(/) z c ings and splits are based on similarities and differences in physical traits. If the
I same physical trait arises in two different species that live in two separate geo-
graphic areas, it will lead to a grouping of the two species and will raise the possi-
bility that the two geographic regions were once joined. Cladistics and vicariance
biogeographic analysis provide information on the relative timing of speciation
events but do not yield direct evidence of the actual number of years since two
lineages diverged. Since the actual age of divergence cannot be determined by
cladograms alone, biogeographers often must speculate on the timing and causes
of vicariance events that may have led to allopatric speciation and the divergence
Frogs
of evolutionary lineages. Finally, even if jump dispersal events are rare, they are
likely to be important over time and are not accommodated within the frame-
work of vicariance biogeography.
In recent years, the development of more refined genetic analysis has led to
1eech species (Fagus) of t he the testing of the proposed evolutionary relationships that have arisen through
es and for frog species (dar.=.
traditional taxomomic and area cladograms. This direct genetic approach, for
which the term phylogeography was coined in the mid-1980s, is based on molecu-
lar genetics and is becoming an increasingly important focus of biogeographic
research. Much of the current work in phylogeography is based on analysis of
t of new eucalyptus spec:::-
mtDNA (see Chapter 11). One example of the potential impact of the molecular
1 of progressive disjuncr: ;:-
genetics approach is provided by a recent examination of the relationship
with geologic and paleo:_-
between crocodiles, turtles, and birds. Classic cladistic studies based on physical
traits suggest that crocodiles are most closely related to birds and less closely
:1 number of different pi::.::.
related to turtles. Comparison of the mitrochondrial DNA of modern crocodiles,
mtologist, Colin Pattersc-
turtles, and birds shows that the turtles and crocodiles are quite closely related.
al groups of fishes, tun.:.::-
The combination of genetic analysis with traditional taxon and area cladograms
he area cladograms for :.=.:
y divided the groups in:c _ is an important means of advancing our confidence in interpretations of evolu-
North America, and E u..:- =- tionary history.
;ts and beech trees diY iC. ~­ Phylogeography can also be applied to explicitly geographic questions
tion and an Australia. \ -:: regarding the distribution and evolution of species. In particular, molecular clock
he general division of =..: analysis (see Chapter 11) has been used to determine the timing of presumed vic-
ts and between Old Wo::- _ ariance events that have split range limits and led to the allopatric development
L the hypothesis that vice.:- ·
of new species or subspecies. Brett Riddle of the University of Nevada and his
he northern and southe::-: colleagues have used mtDNA and the molecular clock approach to determine
e modern continents, is.:.._ the likely timing of vicariance events that have led to the development of new
e modem distribution_ species and subspecies of animals such as mice and lizards in the southwestern
and frogs between No;-...:: United States and adjacent Mexico. It has long been thought that geographically
joined by the Isthmus distinct species and subspecies of these organisms arose during the last 10,000
forth and South Ameri -- years or so when increasing aridity following the close of the last glacial period
led to expansion of dry desert and fragmented the previously continuous geo-
graphic ranges of many mice, lizards, and other plant and animal species. How-
ever, detailed genetic analysis of these different geographic populations has
revealed surprisingly large genetic differences between the different species and
subspecies. The large genetic differences suggest that geographic separation of
he intepretation of claC. =- the different species and subspecies may have occurred millions of years in the
y widely accepted arne=-. past and may be due to environmental and geologic changes that occurred as
404 CHAPTER 13 DESCRIPTION AND INTERPRETATION OF BIOGEOGRAPHIC DISTRIBUTIONS

early as the Tertiary. Riddle refers to vicariance events that have occurred in the :iumphries, C. J. and Parenti
recent geologic past, say the past 10,000 to 20,000 years, as shallow history and ric Biogeography. Clarenc
those that occurred millions of years ago as deep history. On the basis of the :1 mphries, C. J., Ladiges, P.'
newly acquired phylogeographic evidence, he has suggested that such deep his- Zandee, M. (1998). Cladis
A nalytical Biogeography:
tory vicariance events are far more common and important than biogeographers
A pproach to the Study of.
had previously realized. The results of phylogeographic analysis, and molecular
Distributions (ed. A. A . M
clock results in particular, will continue to be the focus of intense research and Giller). Chapman and Ha
lively debate in the years ahead. :-lutchinson, G. E. (1959). He
alia, or why are there so n
mals? American Naturali5
_.linnich, R. A. (1982) . Pseud
KEY WORDS AND TERMS in Baja California? Madn
-!ourelle, C. , and Ezcurra, E.
Amphioceanic Circumpolar Pan tropical rule: A comparative analy
Amphiregional Cladistics Phylogenetic biogeography and North American colu
Amp hi tropical Disjunct Phylogeography can Naturalist 150, 131-1.:1
Areography Hollow curve distribution Range _.!uller, P. (1974). Aspects of;
Biogeographic relicts Macroecology Rapoport's Rule W. Junk Publisher, The H;
Bipolar Panbiogeography Temperate _.!uller, P (1980). Biogeograp
Choropleth maps Pandemic Vicariance biogeography Row, New York .
. !yers, A.A., and Giller, P S.
Biogeography. Chapman ;
O'Brien, S. J., Menotti-Raym
REFERENCES AND FURTHER READING J. , Nash, W. G., Wienberg,
Copeland, N. G. , Jenkins, J
A vise, J. C., and Hamrick, J. L. (1996). Conserva- G. Nelson and D. E. Rosen), pp. 501-523. and Graves, J. A.M. (1999
tion Genetics: Case Histories from Nature. Columbia University Press, New York. comparative genoics in m.
Chapman and Hall, New York. Darlington P. J. , Jr. (1957). Zoogeography: The 458-481.
Bock, C. E ., and Ricklefs, R. E . (1983) . Range Geographical Distribution of Animals. Wiley, Ornduff, R. (1974) . Introduct
size and local abundance of some North New York. Plant Life. University of (
American songbirds: a positive correlation. Gaston, K. J. (1991). How large is a species' geo- Berkeley.
A merican Naturalist 122,295-299. graphic range? OIKOS 61, 434-438. ?atterson, C. (1981). Method
Brown, J. H. (1995). Macroecology. University of Gaston, K. J. (1996). Species-range-size distribu- phy. In Vicariance B iogeo!
Chicago Press, Chicago. tions: patterns, mechanisms and implications. son and D. E. Rosen ), pp.
Brown, J. H., and Lomolino, M. V. (1998). Bio- Trends in Ecology and Evolution 11, 197-201. University Press, New Ym
geography, 2nd edition. Sinauer Associates, Grehan, J. R. (1991). Panbiogeography 1981-91:
Sunderland, MA. development of an earth/life synthesis.
Bush , G. L. (1993). A reaffirmation of Santa Ros- Progress in Physical Geography 15,331-363.
alina, or why there are so many kinds of small Hanski, 1., and Gyllenburg, M. (1997) . Uniting
animals. In Evolutionary Patterns and two general patterns in the distribution of
Processes (ed. D. R. Lees and D. Edwards) , species. Science 275, 397-400.
pp. 229-249. Academic Press, London. Hedges, S. B. , and Poling, L. L. (1999). A molecu-
Cox, C. B., and Moore, P. D. (1993) . Biogeogra- lar phylogeny of reptiles. Science 283,
phy: An Ecological and Evolutionary 998-1001.
Approach, 5th edition. Blackwell Scientific Heusser, L. E ., and Siroko, F. (1997) . Millennia!
Publications, Oxford. pulsing of environmental change in southern
Craw, R. C., Grehan, J. R., and Heads, M. J. California from the past 24 ky: a record of
(1999). Panbiogeography. Oxford University Indo-Pacific ENSO events? Geology 25,
Press, New York. 243-246.
Croizat, L. (1978). Deduction, induction, and bio- Heywood, V. H . (1978). Flowering Plants of the
geography. Systematic Zoology 27, 265-287. World. Oxford University Press, Oxford.
Croizat, L. (1981). Biogeography: past, present, Huggett, R. J. (1998). Fundamentals of Biogeog-
and future. In Vicariance Biogeography (ed. raphy. Routledge, London.
'HIC DISTRIBUTIONS
s that have occurred in C:.::
~ars, as shallow history a:::
istory. On the basis of t1::
~gested that such deep ~
)rtant than biogeographc _
hie analysis, and molecul::
:us of intense research ~=
'an tropical
'hylogenetic biogeograph,-
'hylogeography
~ange
~apoport's Rule
emperate
'icariance biogeography
E. Rosen), pp. 501-523.
rsity Press, New York.
1957). Zoogeography: The
~tribution of Animals. Wi.:e_
How large is a species· ge---
'IKOS 61,434-438.
Species-range-size dist ric __
echanisms and implicatio::s.
y and Evolution 11, 197-::;: _
Panbiogeography 1981- t;:
m earth/life synthesis.
cal Geography 15,331- 36..:
:nburg, M. (1997). Unit in~
:rns in the distribution oi
75, 397-400.
1ling, L. L. (1999). A mole-,::-:_-
reptiles. Science 283,
:iroko, F. (1997). Millelllli.=..
1mental change in sou the::-:.
ile past 24 ky: a record o-
0 events? Geology 25,
8). Flowering Plants of 11>:
Giversity Press, Oxford .
. Fundamentals of Biogeo5-
,London.
Humphries, C. J. and Parenti, L. R. (1986).Cladis-
tic Biogeography. Clarendon Press, Oxford.
Humphries, C. J., Ladiges, P. Y. , Roos, M. , and
Zandee, M. (1998). Cladistic biogeography. In
Analytical Biogeography: An Integrated
Approach to the Study of Animal and Plant
Distributions (ed. A. A. Myers and P. S.
Giller). Chapman and Hall, London.
Hutchinson, G. E. (1959). Homage to Santa Ros-
alia, or why are there so many kinds of ani-
mals? American Naturalist 93, 145-159.
\ finnich, R. A. (1982). Pseudotsuga macrocarpa
in Baja California? Madrono 29, 22-31.
\ fourelle, C., and Ezcurra, E. (1997) . Rapoport's
rule: A comparative analysis between South
and North American columnar cacti. Ameri-
can Naturalist 150, 131-142.
\ fuller, P. (1974). Aspects of Zoogeography. Dr.
W. Junk Publisher, The Hague.
\ fu ller, P. (1980). Biogeography. Harper and
Row, New York.
\ fyers, A. A. , and Giller, P. S. (1988). Analytical
Biogeography. Chapman and Hall, New York.
O 'Brien, S. J., Menotti-Raymond, M., Murphy, W.
J., Nash, W. G., Wienberg, J., Stanyon, R.,
Copeland, N. G., Jenkins, N. A., Womack, J. E .,
and Graves, J. A.M. (1999). The promise of
comparative genoics in mammals. Science 286,
458-481.
Ornduff, R. (1974). Introduction to California
Plant Life. University of California Press,
Berkeley.
?atterson, C. (1981). Methods of paleobiogeogra-
phy. In Vicariance Biogeography (ed. G. Nel-
son and D. E. Rosen), pp. 446-489. Columbia
University Press, New York.
REFERENCES AND FURTHER READING 405
Pielou, E . C. (1979) . Biogeography. Wiley, New
York.
Porsild,A. E., and Cody, W. J. (1980). Vascular
Plants of the Continental Northwest Territo-
ries, Canada. National Museum of the Natural
Sciences, Ottawa.
Rapoport, E. H . (1982). Areography: Geographi-
cal Strategies of Species. Pergamon Press,
Oxford.
Riddle, B. R. (1995). Molecular biogeography in
the pocket mice (Perognathus and Chaetodi-
pus) and grasshopper mice (Onychomys): the
late Cenozoic development of a North Amer-
ica aridlands rodent guild. Journal of Mam-
malogy 76, 283-301.
Riddle, B. R. (1996). The molecular phylogenetic
bridge between deep and shallow history in
continental biotas. Trends in Ecology and
Evolution 11, 207-211.
Robbins, C. S. , Bruun, B., and Zim, H. S. (1983). A
Guide to Field Indentification: Birds of North
America. Golden Press, New York.
Rosen, B. R. (1988). Biogeographic patterns: a
perceptual overview. In Analytical Biogeogra-
phy ( ed. A. A. Myers and P. S. Giller), pp.
23-55. Chapman and Hall, New York.
Stevens, G. C. (1989). The latitudinal gradient in
geographical range: how so many species
coexist in the tropics. American Naturalist 133,
240-256.
Stevens, G. C. (1992). The elevational gradient in
altitudinal range: An extension of Rapoport's
latitudinal rule to altitude. American Natural-
ist 140, 893-911.
Udvardy, M.D. F. (1969). Dynamic Zoography.
Van Nostrand Reinhold, New York.
 CHAPTER 14
THE GEOGRAPHY OF
BIOLOGICAL DIVERSITY

Imagine that you are making a biogeographical fieldtrip and traveling eastwa:-_
from Alaska to Labrador, Canada. The route of your trip will transect the en -=
boreal biome of North America. Being a good biogeographer, you keep ali :
the plants and animals encountered along the way. By the end of the first day. YL _
" ~ .wght agai
have encountered perhaps two species of spruce and no other tree species.
several weeks you have covered hundreds of kilometers, crossed into Cana.: · that is samplec
and have seen only four or five different species of trees. By the time you fin relationship be
arrive at the Atlantic coastline, you have traveled more than 6000 km through · ·- = encountered i~
boreal forest and would be lucky to have encountered more than 10 differ - tionships are r
species of trees. During the thousands of kilometers of your travels, you wo ~ sampling area
have seen at most perhaps 100 different bird species and 30 species of mamm formula for tht
If you had taken a similar field trip across Costa Rica from the Pacific Coast to · ._::
Atlantic Coast, much of your route would have been through a tiny portion of: ·- =
tropical forest biome. Here you might have encountered more than 50 differ - where S is spe
tree species within a distance of a few hundred meters. As you traveled acr :; species per uni
Costa Rica, you would have found several hundred tree species, about
species of birds, and around 140 species of mammals, all in a distance of less th -
200 km from coast to coast! The obvious question that arises from these two fie:.:-
trips is, why is there such a striking difference in the number of plant and ani.n:2.. 600
species in the North American boreal forest compared to the tropical forest
300
Costa Rica?
Documenting and understanding such huge differences in the geograp · _ en
distribution of biological diversity are a central concern for biogeographers. - Q)
"(3 100
Q)
this chapter we will begin by considering exactly what biological diversity is ar:.: a.
en
how it can be measured. We will then examine some geographic patterns in ' :: 0
distribution of biological diversity on continents and the oceans. We will als lii 40
.0
debate which factors control global patterns of biological diversity. As we co- - E 20

(
:J
elude the chapter, we will look at the very interesting distribution of biologi - z
diversity on islands. We will see why the study of island biological diversity he..; 10
long held a particular fascination for biogeographers and ecologists. 6

3
WHAT IS BIOLOGICAL DIVERSITY?
Biological diversity, sometimes called biodiversity, can be broadly defined as t h~ FIGURE 14.1
number, variety, and variability of living organisms. Biodiversity is often simp!: America . The dat
defined as the number of different species in a given geographic area. The great -:: rel atively straight
the number of species present, the greater the biodiversity. For example, th; species found als

406
 WHAT IS BIOLOGICAL DIVERSITY? 4 07

province of British Columbia in Canada has about 70 species of mammals com-

pared to 40 species found in Alaska. Thus, we could say that British Columbia has
a greater mammalian biodiversity than Alaska. This simple measure of biodiver-
sity, based solely on the number of different species in a given area, is also called
species richness by ecologists. In many large-scale biogeographical studies, biodi-
versity is synonymous with species richness.
y When conducting comparative studies of biodiversity in different regions, it
is important that we compare areas that are similar in size. In general, species
richness increases with the size of the area sampled. If, for example, we go into a
forest and conduct surveys of plant species richness, we might start by laying out
a plot that is 1 by 1 m in size. We would identify all of the different plant species
in the plot and determine the species richness of that particular site. In most
trip and traveling eastw:::.: _
instances, if we increase the size of the plot, we will discover additional species.
trip will transect the en:=-:
Thus, if we increased our forest plot to 10m by 10m in size, we might encounter
)grapher, you keep a lis:
twice as many plant species. If we increase our sampling area to 100m by 100m,
the end of the first day. \-" _
we might again double the number of species we discover. The larger the area
no other tree species. Af::::--
that is sampled, the greater the number of species that we will find. The positive
~ters, crossed into Ca n c.~
relationship between the size of the area surveyed and the number of species
ees. By the time you fin L
encountered is called the species-area relationship. In general species-area rela-
ethan 6000 km throu g h :..::;~
tionships are not linear (Fig. 14.1). This typical nonlinear relationship between
·ed more than 10 differ<"-
sampling area and species richness is called the species-area curve. The common
of your travels, you v.-o,___,_
formula for the species-area curve is
md 30 species of rna~::__ .
rom the Pacific Coast to -- S=cAz
b.rough a tiny portion of:_:._
red more than 50 differ.:-- where S is species richness and c is a constant that is based on the number of
ers. As you traveled ac-.: species per unit of area in the geographic region in which the study is conducted.
~d tree species, about '='
all in a distance of less tl:.:..:
arises from these two fie ~.:
600
mmber of plant and a~
ed to the tropical fores ~
300

'ferences in the geogra!='·- (/)

:ern for biogeographe -~Q) 100

t biological diversity is c.- Q_
(/)

geographic patterns in :..:

~ 40
d the oceans. We will C.:., Q)
.0
)gical diversity. As we cc § 20
g distribution of biolog:.:: z
md biological diversity ::.__ 10
and ecologists. 6

3
100 104 106 108 1010
Area (acres)
n be broadly defined as -- FIGURE 14.1 The relationship between sampling area and bird species richness in North
3iodiversity is often siiT.: America. The data are plotted on a log-log graph to render the species-a rea curve into a
eographic area. The grec.::: relatively straight line. As the size of the sampling area increases, the number of bird
diversity. For example. -- species found also increases (after Preston, 1960).
408 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Regions with large numbers of species will have large values of c. If we co-- To capture inJ
ducted our study in Costa Rica, which has very high biodiversity, the value fo::- variety of rna
would be high. If we conducted a similar study in the boreal forest of Alaska. ·'-= richness and e
value for c would be low. The term A is the size of the sampling plot analyzed : same proporti
species richness. If we use a 1 m by 1 m plot, the value for A will be 1 sq m. If : and are domin
use a 10 m by 10 m plot, the value for A will be 100 sq m. Finally, z is a cons ,- diversity valw
that represents the mathematical relationship between area and species richne-s... species presen
It can be thought of graphically as the slope of the line that describes the r ...:..- up. One comr
tionship between area and species richness. For study areas in which species · -::- index (H') wl
ness rises quickly with increasing area, the value for z will be high. In many cas~ index has the J
particularly for animals, the species-area curve can be made into a straight lin ·
a simple logarithmic transformation:
logS= (loge)+ z(logA) where Pi is the
use the index,
There are several reasons why species richness increases as the size of -·- = site, transforrr
sampling plot increases. First, small sampling plots may not include all of ~= index formula
species present in an area because of random chance in the selection of where ::=.:-- 1.5 for system ~
sampling will be conducted. Additional species might be present in the vicic. species richne:
but might be missed because of the small size and choice of location of the sa=- rates both spe
ple site. Second, small sampling sites contain only a small selection of very l -:.. may be obtain
microhabitats and may not provide the right environmental conditions for m--
species. A small increase in the size of the sampling area will increase the varie.
of local habitats and the number of species that are represented. Greatly incr e;:_~ where E is sp1
ing the size of the sampling area will incorporate areas representing wider r a n~ =­
number of spe
of environmental conditions and will in turn increase the number of species :..
Althougl
will be present. The problem with very large sample plots is that it is difficult : adequately me;
completely survey all of the species of plants or animals present. In most cas~ and identify aL
sampling is done by establishing a number of small sample plots or sampling _::-
difficult to cm
tions that are randomly or systematically distributed across the landscape. In '-
species evenne:
manner, it is hoped that both the habitat diversity and the species diversity of -·-=
distributions w
study area will be represented and sampled in an efficient manner. number of ver
When used for comparative purposes, species richness values are of:
species that are
reported as the number of species per hectare or the number of species ;=:-
square kilometer. However, in many cases, censuses of species richness are co:::.-
ducted for political units such as states or countries. The area of the differe=. HOW MANY DIFFI
states or countries will vary, and this will affect the species richness that _ ARE THERE ON Tl
reported. If we compare the species richness for different states, countries. L
continents, it is important to remember the impact that differences in area '-"-:_ An obvious q
have on total species richness. Larger areas will have more species than small=- species are the
ones if all else is equal. present, about
Species richness is a simple and useful means of measuring biodiver i._ from viruses tc
Species richness, however, does not tell a complete story regarding biodiver i _ known species
Simple species richness does not, for example, tell us anything about the rarity c: make up abou
species or the evenness of species richness. What do we mean by the term speci - species are pla1
evenness? Consider that we might find that two areas have exactly the sar:::: known species.
species richness, say 10 species of birds. In one of the two areas, we might disco\·e::- All scien1
that the individuals of the 10 bird species occur in roughly equal numbers. In ti:::: covered. Estim
other area, we might find that 90% of the birds all belong to one species and th::: over 20 millio
the 9 other species of birds are rare. The species richness in the first area is eve!L~ plants, birds, ar
distributed, while the species richness in the second area is unevenly distribute · to be discovere
 HOW MANY DIFFERENT SPECIES ARE THERE ON THE EARTH? 409

trge values of c. If we co:.-
biodiversity, the value fo:-
boreal forest of Alaska. <::=
~sampling plot analyzed :.:-
e for A will be 1 sq m. If " :
sq m. Finally, z is a constc.;:.
m area and species rich n~­
line that describes the r e ~
·areas in which species ric:.-
: will be high. In many cas::-
' made into a straight line ::
To capture information on both species richness and evenness, ecologists use a
variety of mathematical indices of species diversity that include data for both
richness and evenness. Sites that have many species, each occurring at about the
same proportion, will generate high diversity values. Sites that have few species
and are dominated by the individuals of one or two of those species will have low
diversity values. These indices incorporate information on both the number of
species present and the proportion of the flora and fauna that each species makes
up. One commonly applied mathematical measure of diversity is the Shannon
index (H') which takes into account both species richness and evenness. The
index has the form:
H ' = -'f.p; In p ;

where p; is the proportion of the ith species and ln is the natural logarithm. So, to
use the index, one simply counts all the individuals of all the species in a study
increases as the size of :..::..: site, transforms these counts into proportions, and then applies the Shannon
may not include all of :..::.: index formula . In natural systems, the value of H ' has been found to range from
in the selection of where ·- 1.5 for systems with low species richness and evenness to 3.5 to species with high
1t be present in the vici;::- species richness and evenness. The Shannon index for species diversity incorpo-
toice of location of the s.- - rates both species richness and evenness. An estimate of species evenness alone
>mall selection of very l may be obtained from the Shannon index value (H') using the equation
mental conditions for m.:.::
rea will increase the var::: - E= H 'l in S
~presented. Greatly incr ~ ­
where E is species evenness and S is the species richness of the site (the total
ts representing wider mr:~=
number of species identified or collected) .
~ the number of species ~
Although species evenness is an important property of ecosystems, rarely is it
plots is that it is difficul:
adequately measured for natural environments. It is very time consuming to sample
nals present. In most C2S::-
and identify all individuals in a given area. In addition, rare species are extremely
lmple plots or samplin g ~:.
difficult to count adequately and are generally underrepresented in studies of
across the landscape. In :..::;
species evenness. There is no general rule or accepted theory as to how even species
:1 the species diversity o~ ·-
distributions will be in natural environments. Some systems seem to have a large
cient manner.
number of very rare species, while other systems are typified by an abundance of
; richness values are o::"__
species that are neither particularly rare nor overwhelmingly dominant.
the number of species ;::
of species richness are c-.:::
s. The area of the diffe:-:--
HOW MANY DIFFERENT SPECIES
he species richness the:
ARE THERE ON THE EARTH?
ifferent states, countrie·
:hat differences in area
An obvious question we might ask is, how many different plant and animal
~ more species than smc...._
species are there on the earth? The total number of species remains a mystery. At
present, about 1.7 million species of organisms are known to science. These range
. of measuring biodi v e~ -
from viruses to plants, insects, birds, and mammals (Fig. 14.2). About 56% of all
:tory regarding biodive ~ ­
known species are insects. In fact, one group of insects, the beetles (Coleoptra),
anything about the rari _
make up about 24% of all known species. Approximately 14% of all known
ve mean by the term sp eo_
species are plants. Mammals and birds together make up only 2.7% of the world's
·eas have exactly the s.::.=
known species.
wo areas, we might discc· =-
All scientists agree that there are many species that have not yet been dis-
Lighly equal numbers. lc. ·-
covered. Estimates place the total number of species on the earth at 4 million to
long to one species and --
over 20 million. There are thought to be relatively few species of flowering
ess in the first area is e\ ::-
plants, birds, and mammals yet to be discovered. Most of the species that remain
trea is unevenly dis tribe:-_
to be discovered are viruses, unicellular organisms, and invertebrates. The rate of
41 0 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Vertebrates (2 .7%) TABLE 14.1

Nematodes (0.9%) ----
Molluscs (4.2%)
Other invertebrates (4.0%)
Other arthropods (1.2%) Organisms
Fungi (4.2%)
Mammals
Bacteria (0. 2=,
Arachnids (4.5%) Viruses (0. 3° = Birds
Amphibians and
Fish
Other insects (8.9%) Mollusks
Insects
Arachnids

Source: World Con

in Eurasia sho
nia Italy, Israe
Lepidoptera (8.9%)
the basis of tl
FIGURE 14.2 The distribution of known species (in 1990) among different plant and North Americ
animal groups (after World Conservation Monitoring Center, 1992). considered tht
the species. In1
lations is grea1
discovery of flowering plant, bird, and mammal species was relatively hi_: - possible that t
between the eighteenth and early twentieth centuries and has leveled off to _ of the Eurasi<
trickle of new species each year for the past 100 years. In contrast, the rate of ·."- human activiti
covery of new species of invertebrates such as arachnids and crustaceans ~ early stages. H
accelerated in the past few decades On average over 10,000 new invertebr2:.:c measuring the
species are described each year. Of these, most are insects (Table 14.1).
Although the discovery of new species of birds and mammals is rare, it d _
occur. The discovery of some of these new species sometimes occurs under sow:e- LATITU DINAL AN
what amusing circumstances. For example, a new species of bird, the Udzungw-- DIVERSITY GRAC
partridge (Xenoperdix udzungwensis) was discovered by Danish biologist ~
Tanzania in 1991. In this case, the scientists were having dinner and found soc::- If we divided t
unusual looking bird legs in a pot of "chicken" stew that was being served a: ..:. species, mamrr
field camp. Subsequent searching in the surrounding countryside turned up li\i.;:; we would see
members of this previously undescribed species of bird. Comparison of r:.::e sity for all thn
Udzungwaq partridge with fossil material shows it to be a paleoendemic speci ~ ward (Fig. 14.3
that is an evolutionary relict of a group of birds that once ranged from Southeas· north of Cana·
Asia to Africa. In 1994 a new species of tree kangaroo of the genus Dendrolaf!l.L:c than 40 specie:
was discovered by scientists working in New Guinea. This species came to lig : 600 species li\
when a zoologist's dog noticed the animal and began to give chase. groups of orga
In our discussion above, we have focused on species biodiversity. HoweY : of increasing
it is important to remember that large and geographically dispersed populatior:.5 strong increas(
of single species often include many different and distinct genotypes. Biologis.:.: are native to ,
and geographers are increasingly interested in studying genetic biodiversity as Brazil. Michig<
well as species biodiversity. By using genetic analysis, the geographic patterns o:' 1550 species. J
genetic variability within a single species can be recorded, mapped, and analyze The subtropie<
to study the relationship between genetic diversity, geography, the environmen· 7000 known sp
and human impact. For example, analysis of grey wolf populations (Canis lup us similar general

Embryophytes) (14.3%)
Fungi (4.2%)
Bacteria (0. 2°,
Viruses (0.3°,
1mong different plant an d
1992).
)ecies was relatively c.:~·­
~s and has leveled off rc _
In contrast, the rate of C....'-
hnids and crustaceans -..
T 10,000 new inverteb:-.=. .:.
;ects (Table 14.1).
nd mammals is rare, it C .Co:."
etimes occurs under so=:.=
ies of bird, the Udzungv
d by Danish biologisr.;:
ng dinner and found sc-
that was being serve d ~: ~
Juntryside turned up l'- .......=
bird. Comparison of ·-
be a paleoendemic spe.::l
nee ranged from Soutl::.;;::._
'of the genus Dendrol£1?
This species came to t_:-
o give chase.
cies biodiversity. HO\\ e:· c:-
:ally dispersed populac;:
tinct genotypes. Biolo.5_
ing genetic biodiversi[! _
the geographic patter::.;;
led, mapped, and analy::
ography, the environn::e-
populations (Canis lu;
LATITUDINAL AND ALTITUDINAL DIVERSITY GRADIENTS 411
TABLE 14. 1 Number of New Species Described per
Year between 1978 and 1987 for Selected
Groups of Organisms
Organisms Species Described Per Year (1978-1987)
Mammals 26
Birds 5
Amphibians and reptiles 105
Fish 231
Mollusks 366
Insects 7222
Arachnids 1250
Source: World Conservation Monitoring Center (1992).
in Eurasia shows that the remaining small populations in Portugal, Sweden, Esto-
nia Italy, Israel, Iran, and China are all genetically distinctive from each other on
the basis of their mtDNA. The Eurasian populations are also distinctive from
North American populations of grey wolf. Although all of these populations are
considered the same species, they display significant genetic biodiversity within
the species. Interestingly, the genetic biodiversity within the Eurasian wolf popu-
lations is greater than the diversity within the North American populations. It is
possible that the high degree of geographic diversity in the genetic composition
of the Eurasian wolves is due in part to the fragmentation of their range by
human activities. For most groups of organisms such genetic work is in its very
early stages. However, genetic analyses will be an ever more important means of
measuring the true biodiversity of the earth.
LATITUDINAL AND ALTITUDINAL
DIV ERSITY GRADIENTS
If we divided the world into large sample regions and mapped the number of tree
species, mammal species, and bird species found in North and Central America,
we would see a very interesting and relatively consistent pattern. Species diver-
sity for all three groups is lowest in the far north and generally increases south-
ward (Fig. 14.3). Less than 20 mammal species live in the sampling areas in the far
north of Canada, while over 150 are found in Central America. Similarly, fewer
than 40 species of birds make their home in the high arctic of Canada. Well over
600 species live in the rainforest of Central and South America. Many other
groups of organisms, such as reptiles and amphibians, also show a similar pattern
of increasing species diversity toward the equator. Insects show particularly
strong increases in species richness in the tropical regions. Three species of ants
are native to arctic Alaska, 63 species are native to Utah, and 222 species to
Brazil. Michigan has about 134 native species of butterflies, whereas Panama has
1550 species. All of North America has about 750 known species of butterflies.
The subtropical and tropical regions of Central and South America have over
7000 known species. For many other groups of terrestrial organisms, we can find a
similar general pattern of increasing species diversity from high to low latitudes.
412 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

The patt'
of the equator
orchid species
in the tropical
of tree species
ern Scandinav
rial rainforest
in Zaire can cc
species richne:
higher in the S·
The trop
Asia, and Aust
estimated that
rainforests. Th
Tropical rainfc
A typical patcl
as many as 15(
species, and m:
Many gr
species diversi
as foraminifer:
tor. ForaminifE
to the tropics.
about 20 speci
larly, there are
in the far Nortl
species richne~
times as many
areas of comp<
ship between l
ocean. One gn
abyssal zone o
Atlantic to ovE

b TABLE 14.2 Examples

Terrestria
FIGURE 14.3 (a) Tree species diversity, (b) bird species diversity and (c) mammal spec ies
diversity for continental North and Central America (after Currie and Paquin, 1987; Organism Regie
Simpson, 1964; Cook, 1969).
'v1 ammals N. Ame
'v1 ammals S. Ame
_and birds N. Ame
Reptiles N. Ame
Not only does the total number of species increase toward the equator, but the
Amphibians N. Ame
average number of species found per unit of area of land also increases. A
Ants S. Ame
hectare of forest in the boreal forest of Alaska generally supports five or fewer
Orchids S.Ame1
species of trees. A hectare of land in the temperate forest of eastern Nont.
M arine fish N. Ame
America typically supports 10 to 20 species of trees. A hectare of forest in th
M arine mollusks N. Ame
rainforest of Central America or the Amazon may easily contain over 100 dif-
ferent tree species. So urce: Brown and Lomoli no, 198
 LATITUDINAL AND ALTITUDINAL DIVERSITY GRADIENTS 413

The pattern of increasing diversity toward the low latitudes holds up south
of the equator (Table 14.2). South of the equator in South America the number of
orchid species increases from 15 at the southern tip of the continent to over 2500
in the tropical rainforest near the equator. In the eastern hemisphere, the number
of tree species on a typical plot of forest in Europe increases from 2 in far north-
ern Scandinavia to 12 in northern continental Europe. Farther south the equato-
rial rainforest of Africa contains hundreds of different tree species. A 50-ha plot
in Zaire can contain as many as 450 different tree species. In addition, the overall
species richness for some groups of terrestrial organisms has been found to be
higher in the southern hemisphere than in the northern hemisphere.
The tropical rainforests of Africa, South and Central America, southeastern
Asia, and Australasia are the most biologically diverse regions in the world. It is
estimated that 45 % of all plant and animal species on the earth live in tropical
rainforests. The rainforests are particularly rich in plant, bird, and insect species.
Tropical rainforest species likely make up 96% of the world's insect biodiversity.
A typical patch of tropical rainforest that is a few kilometers in area can contain
as many as 1500 plant species, 400 bird species, 100 reptile species, 60 amphibian
species, and many thousands of insect species.
Many groups of marine organisms show a similar pattern of increasing
c species diversity at lower latitudes (Table 14.2). Many planktonic organisms, such
as foraminifera, increase in species diversity from the polar regions to the equa-
tor. Foraminifera species richness increases by a factor of 8 from the polar waters
to the tropics. Small planktonic invertebrates called copepods increase from
about 20 species in the Arctic Ocean to 80 species in the tropical Pacific. Simi-
larly, there are twice as many species of fish in the equatorial Atlantic as there are
in the far North Atlantic. Benthic marine organisms also generally display highest
species richness in the tropics. The Bay of Bengal off India supports about five
times as many species of bottom-dwelling worms and mollusks as do similar sized
areas of comparable benthic habitat off Cape Cod in New England. The relation-
ship between latitude and biodiversity is even seen in the deepest portions of the
ocean. One group of benthic gastropod mollusks, the prosobranchs, found in the
abyssal zone of the ocean, increases in species diversity from 3 in the far North
Atlantic to over 20 in the tropical Atlantic.
414 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Many freshwater organisms also have high species diversity in tropi pattern of spec
regions compared to temperate and arctic areas. The Great Lakes support abo_ sity similar to
172 species of fish. There are over 450 species of freshwater fish known from Ce::- mammals reac
tral America and over 1000 known from the Amazon. All of Europe posses_"'5 alpine environ
about 190 species of freshwater fish, while a single lake in Africa, such as Lake Ta::- species on the
ganyika, may support over 200 species of fish. Other research has shown that th e:-~ plays this pattc
are two to three times more species of aquatic insects in tropical streams in Cent .::... 3000 min elev;
and South America as there are in the temperate streams of North America. upslope towan
Although the pattern of increasing diversity from high to low latitudes ...: Some gn
widely pervasive, there are exceptions. Peninsulas have lower species diversi._ Scolopacidae,
than adjacent mainland areas. The species richness on peninsulas also generall_ of sandpipers
declines toward the tip. The impact of the peninsula effect can be seen in rna - species are fot
mal and bird diversity in Florida and Baja California (Fig. 14.3). In these are<S marine organi
species diversity decreases southward toward the tip of the peninsula. Diversi._ there is no cle
also decreases toward the tip of the Yucatan Peninsula. Islands have low ~ of plants and
species diversity than adjacent mainland areas. We will discuss the biodiversity o: wasps, reach rr
islands in detail below. Desert regions, even those near the equator, have very lo .. diversity towa
species diversity of plants and animals despite their geographic location. ever, to a mon
The species diversity of high-altitude alpine sites in the tropics is low co - the tropics.
pared to that of adjacent lowland forests. On a global basis there is a general pa:- Understa
tern of decreasing species richness at higher elevations. Tundra ecosystems fou ;: diversity is an
at high elevations are poorer in bird, mammal, and plant species than lower ele- diversity gradi
vation forests. For example, in the Himalayan Mountains of Nepal there are O\' ~ important for ·
300 bird species found at 500-m elevation and none above 8000 m (Fig.14.4) . Tt= for a number o
same trend can be seen for Himalayan mammals and vascular plant species.. individualloca
However, high mountains in desert regions often show a different geographi large geograpr
high levels of :
local habitats '
8,000
all levels of spc

7,000
CONTROLS ON Gl
6,000 .
GRADIENTS OF S

Why does biod

~ 5,000 . 5,000 . .
.s higher in tropi
perplexed biof
~ 4,000 · · · 4,000 · - · 4,000
Two broad cat
:;:::;
~ graphic differc
3,000 · - - · 3,000 · - · 3,000
geographic dif
species evoluti
2,000 · - - - · 2,000 · - - ·
ferences in bic
1,000
and biological
1,000 · - - - · 1,000 · - - -
torical theorie!
to as equilibriu
0 0 100 200 300 400 50 100 °1 10 100 the modern p<
Bird species richness Mammal species richness Vascular plant environmental
species richness librium explan
FIGURE 14.4 Declining species richness of birds and mammals at higher elevations in the the present ph
Himalayan Mountains (after Begnn et al. 1996; Hunter and Yonzon, 1992; Whittaker, 1997). torical and equ

ecies diversity in tropia:
}reat Lakes support abou:
rater fish known from Ce:::-
1. All of Europe possesses
n Africa, such as Lake Ta::-
earch has shown that th e:-~
tropical streams in Cent.:-.:...
1s of North America.
m high to low latitudes ::..o
ve lower species divers:._
. peninsulas also genera::
ffect can be seen in mar:-
(Fig. 14.3). In these a re ~
)f the peninsula. Divers:.
1sula. Islands have low::-
discuss the biodiversity -
the equator, have very 1
Jgraphic location.
in the tropics is low co=-
•asis there is a general p.::.:-
.Tundra ecosystems foe:'
mt species than lower e ~ ::­
ns of Nepal there are O\ ::--
we 8000 m (Fig.14.4). T~::
1d vascular plant speC:=-
v a different geograph: -
CONTROLS ON GEOGRAPHIC
GRADIENTS OF SPECIES DIVERSITY
or:::==--
' higher in tropical regions than temperate and polar regions? This question has
o ~s==
0 and biological processes. Explanations of the first category are referred to as his-
01 10
Vascular plant
species richness
s at higher elevations i.. __
1,1992; Whittaker, 1997).
CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 415
pattern of species richness. The lowest elevations typically have low species diver-
sity similar to that of the adjacent desert. Species diversity for plants, birds, and
mammals reaches a maximum in mid-elevation sites and then declines as high
alpine environments and treeline is approached. The diversity of coniferous tree
species on the eastern slopes of the Sierra Nevada Mountains in California dis-
plays this pattern. The greatest coniferous tree diversity occurs between 2000 and
3000 m in elevation and decreases downslope toward the Great Basin Desert and
upslope toward the alpine treeline.
Some groups of organisms, such as whales or sandpiper birds of the family
Scolopacidae, achieve greatest species richness at high latitudes. Only 2 species
of sandpipers are fo und along the Pacific Coast of southern California, and 14
species are found along the Bering Sea coastline of Alaska. For many groups of
marine organisms, such as benthic invertebrates in the southern hemisphere,
there is no clear increase in species diversity toward the tropics. Other groups
of plants and animals, such as pines, aphids, or the Ichneumonidae family of
wasps, reach maximum species richness in the middle latitudes and decrease in
diversity toward the arctic and the equator. These groups are exceptions, how-
ever, to a more general pattern of increasing diversity from the polar regions to
the tropics.
Understanding what causes the large-scale latitudinal gradients in species
diversity is an important topic for research in global biogeography. The global
diversity gradients and the large regional differences in biodiversity are also
important for understanding local species diversity. Analysis of species diversity
for a number of different plant and animal taxa shows that the species richness of
individual local sites is highly correlated with the broader species diversity of the
large geographic region in which the sites are located. Large regions that have
high levels of species richness can be expected to have high species richness in
local habitats when compared to similar habitat in regions which have low over-
all levels of species richness.
Why does biodiversity differ geographically, and in particular, why is biodiversity
perplexed biogeographers since it was raised by Alfred Russel Wallace in 1878.
Two broad categories of explanations have been proposed to account for geo-
graphic differences in biodiversity. Some biogeographers believe that modern
geographic differences in biodiversity may be explained by the past history of
species evolution and extinction. Other biogeographers consider geographic dif-
ferences in biodiversity to be the product of modern environmental conditions
torical theories of biodiversity. Explanations of the second category are referred
to as equilibrium theories of biodiversity. The historical explanations assume that
the modern patterns of biodiversity are not in true equilibrium with modern
environmental conditions but instead reflect the impacts of past events. The equi-
librium explanations assume that the number of species in a given area reflects
the present physical and biological environment of that area. Let's examine his-
torical and equilibrium theories in a little detail.
416 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Historical Theories
Theories that modern global gradients in biodiversity reflect past events in _-_ =
evolution and extinction of species were developed largely to explain the bi_:·
species richness of the tropics compared to the lower biodiversity of the temp.:---
ate and polar regions. This general line of reasoning has a long history that ca n~.
traced back to Wallace. The classic historical theory focuses on the fact that·--=
repeated glacial periods of the Pleistocene resulted in severe disruptions of ·-_ = \....,
environment in polar and temperate regions. According to the theory, ad van ~=
ice and cold during glaciations caused widespread extinctions of plants and a=...-
mals in the higher latitudes. It is surmised that rates of evolution are too slm'> ·
produce signif.cant adaptive radiation and rebuild species richness in the intef\·2.. . .
between glacial periods. So, as long as the earth alternates between glacial a::,.:
nonglacial conditions, species diversity will remain low at the high latitudes.
The classic historical theory further holds that the tropics have not expe~.­
enced significant environmental disruption during glacial periods. Thus, extincti
rates have remained low compared to those of the high latitudes. Long periods
stable environment have allowed for the evolutionary development of a multin:.::=
of new species. This line of general reasoning, that long periods of environmen:_
stability lead to high species richness, is called the stability-time hypothesis. If "- =
earth does not experience another glacial period, it is supposed that decreas::-.:
extinction rates and the evolutionary development of new species will, over til:;:~
lead to higher and higher species diversity at the mid- and high latitudes. Even.--
FIGURE 14.5
ally, the difference in biodiversity between the tropics and other parts of the wo;:--
Southeast Asia d
would disappear. (after Tallis, 1991
Although the premise that long periods of environmental stability redl! -- rainforest was sc
extinction rates and allow for evolution and increased speciation may be corr --
the evidence suggests that the tropical regions were not immune to environm ::-
tal changes during the glacial and nonglacial periods of the Pleistocene. It is no
thought by many scientists that tropical land areas cooled by several degrees a;::.: Contrary
became much drier than present during the last glacial maximum. Some tropi - gested that fra:
lakes dried up, areas of sand dunes and desert expanded, and forested ar 2.... have actually r:
decreased in extent. Records of conditions in the tropics during the last gla · = within the isol<
maximum, about 20,000 years ago, come from the sediments and fossils in dee;:-- In addition, gc
sea cores, fossil pollen from lake sediments, fossil soils, and other lines of e\ .- divergence anc
dence. Many of these records, though not all, suggest that arid conditions led : - forest and sava
the fragmentation of the tropical rainforest in Africa, South America, and Sou :.._ as rainforest fr
east Asia (Fig. 14.5). Fossil pollen records from northeastern Australia shm'> ~ that allopatric
similar replacement of tropical rainforest by dry forest and savanna during tt:e
forests would I
period 26,000 to 10,000 years ago. In some regions, it is likely that tropical fo res:
high levels of ~
survived during the last glacial maximum as small areas of gallery forest al o rr~
Europe where
river valleys. Detailed analysis of sediments from the delta of the Amazon Ri \·~;
forests were si1
suggests that much of the current rainforest in the Amazon Basin may have be ;:.
ecotonal habit
replaced by savanna during the last glacial maximum. Recently obtained foss·
forests is much
pollen evidence from South America shows that in some parts of Bolivia th.:-
Althougr
tropical rainforest only reached its modern geographical extent in the past fe\'
thousand years. Nor were the tropical seas or deep oceans immune from signif.- ents in diversi
cant changes in temperature during the last glacial maximum. For exampl . example, Lake
recent reconstructions of marine conditions based on deep-sea cores indicate sig- glaciated. The
nificant changes in deep-sea environment and crustacean species diversity durin; contains over
glacial and interglacial cycles. These changes show that the stability-time hypoth- these species <
esis cannot be used to explain latitudinal biodiversity trends in the deep sea. tion of the lal
 CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 417

reflect past events in tt::

rgely to explain the hi(-

,_
odiversity of the tempe:--
a long history that can b:: ./ t.
:uses on the fact that tl:::: ~
severe disruptions of tt:-
; to the theory, advancl::;
1ctions of plants and an:-
~volution are too slO\\- ::
:s richness in the inte n -~
ates between glacial a.::.:
1'
'
tt the high latitudes.
tropics have not expe:-:-
1periods. Thus, extinctic-
latitudes. Long periods __ • Rainforest patch
velopment of a multituc::
periods of environm en~
ty-time hypothesis. If t::.::
supposed that decreas.:-.:
:w species will, over tin:.::
1d high latitudes. Even:-:...-
FIGURE 14.5 Suggested extent of tropical rainforest in South America, Africa , and
j other parts of the \\ o~::
Southeast Asia during the last glacial maximum approximately 20,000 years ago
(after Tallis, 1991 ). However, not all biogeographers believe that the fragmentation of
mmental stability redu.:-:: rainforest was so severe.
Jeciation may be correc-_
immune to environme;:-
the Pleistocene. It is nc
:d by several degrees a=c.: Contrary to the time-stability hypothesis, some biogeographers have sug-
maximum. Some tropi.::-.:. gested that fragmentation of the tropical rainforest during glacial episodes may
tded, and forested are.::... have actually promoted high diversity by allowing allopatric speciation to occur
cs during the last glacr.:..._ within the isolated forests that existed along rivers and in areas of moist habitat.
1ents and fossils in dee::-
In addition, genetic studies of modern rainforest species suggest that genetic
;, and other lines of e·.--
divergence and speciation are most likely to occur along ecotones between rain-
tat arid conditions le d :
forest and savanna. The relative area of such ecotonal sites would have increased
,uth America, and S ou ~­
as rainforest fragmented during glacial periods. Although it is entirely plausible
:astern Australia shO\\ _
that allopatric speciation and ecotonal speciation in the fragmented tropical
and savanna during t:::=
forests would lead to increased species diversity, it does not explain why such
likely that tropical fori:5'
high levels of speciation did not also occur in areas such as Siberia and central
ts of gallery forest alo:.=-
lta of the Amazon Ri \ ::- Europe where glacial ice was absent during glaciations and boreal and temperate
:on Basin may have be::- forests were similarly fragmented , leading to an increase in the relative area of
Recently obtained fos~ ecotonal habitat. As we have seen, the biodiversity of boreal and temperate
•me parts of Bolivia t:::= forests is much lower than for rainforests.
al extent in the past fe Although the historical theories may not explain global latitudinal gradi-
ms immune from si g~­ ents in diversity, they may explain more local differences in biodiversity. For
naximum. For exam!"::: example, Lake Baikal in Russia lies in a portion of Siberia that has never been
ep-sea cores indicate s:; - glaciated. The lake has existed there for well over a million years. Lake Baikal
1 species diversity duri=; contains over 580 species of deep-water benthic invertebrates. A number of
he stability-time hypo'-- these species are endemic to Lake Baikal and evolved there after the forma-
:nds in the deep sea. tion of the lake. Great Slave Lake in Canada is a similarly deep and cold
418 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

water body. The modern Great Slave Lake did not appear until about 10. ·
years ago when glacial ice retreated from northern Canada. In contras:
Lake Baikal, Great Slave Lake has only four species of deep-water ben·._ -
invertebrates. CJ
Equilibrium Theories
A number of different equilibrium theories of biodiversity have been prop
L
to explain differences in species richness at small and large scales. To begin L -
study of these theories, let's consider biodiversity within the context of emi; -
mental gradients and niche theory (see Chapters 3 and 4). In theory, areas
high numbers of species reflect one of three conditions.
First, areas of high biodiversity could contain large gradients for res o u:-:-~
2 c
and offer a wide range of habitat for different species to utilize. For exam p ~~
large geographic area, such as the Amazon Basin, may extend across a wide ,-:::-
{11
ety of climatic and soil conditions and thus provide a wide range of habitat :..
can support a large number of different species. A small region, such as C _
Rica, may offer a wide range of environmental conditions because its topogra;::-
creates different environments in the mountains and lowlands. In theory, th~
regions can support a large number of species because each resource gradier.: 3
long and there is a range of habitat that can support different species with d iE~ ­
ent ranges of tolerance and niches (Fig. 14.6).
Second, the range of habitat in two regions may be similar and the reso ·-. .:
gradient lengths may be equal, but the length of the distribution of indivi · -
species along the gradients may be short in one region because they have spe ~­ FIGURE 14.6
ized niches. Thus, if we assume that competition between species precludes : high-diversity ge
much overlap along gradients and between niches, more species are acco~;:­ diversity region I
dated along the resource gradient if each species is a specialist and has a narr~ available niches
are equal, but th
distribution. The development of such specialized niches might reflect the in..:._-
the organisms li•
ence of intense interspecific competition on natural selection. High levels
the resource gra
interspecific competition in one area might produce a large number of specia - abundance is hi~
species with small niches, while lack of competition in another area might allow .: species either bE
few species to persist with very large niches. specialized spec
Third, key resources may be more abundant in one area than in anoth - space, or (b) the
The gradients for the area of high species diversity would be typified by hi ~._ and niche overlc
abundances of resources, while the gradients for the low-diversity area would !::-=
typified by low abundances of resources. The greater abundance of resour c;;
could allow either a relaxation of competitive pressure, thereby allowing ~ generated by <
greater number of generalist species to survive, or it could allow greater numbe;-;: areas with cor
of species with highly specialized niches to survive if interspecific competitio:;: variability of 1
was an important factor. ences habitat c
At present, we do not have enough information to conduct a large-scale gently rolling t
comparative analysis of the gradient distributions and niche sizes for high- c soils, and mois
low-latitude sites. However, we can look at factors that relate to gradient lengt.L and animals th
resource abundance, and the development of specialized niches and see how the: rolling topogn
might relate to latitudinal diversity gradients. sity and have
sented mesic s
Habitat Diversity In some regions, species diversity is positively correlate · wide range of
with habitat diversity. Greater habitat diversity correlates positively with greate:- cooler and me
resource gradient length and greater available niche space. Habitat diversity can ~ cold high-elev

tppear until about lO. OC
n Canada. In contrast ::
es of deep-water ben tb:
~rsity have been prop o~.:-­
large scales. To begin o
tin the context of enviro;::-
d 4). In theory, areas \\;--
·ge gradients for resour.:.::
; to utilize. For example _
extend across a wide ,·c...- -
wide range of habita t (- ~
nail region, such as C<R,--:...
ms because its topogra(:
lowlands. In theory, thtS::
: each resource gradiec:
fferent species with dif-c:-
'e similar and the resou:-:.-
distribution of individ·.:..:
because they have sp eci~
een species precludes : -
Jre species are accornz:..::-
pecialist and has a narrc
tes might reflect the in:._
selection. High levels
large number of specia.:.:.:
mother area might allo. _
one area than in anoti::--
vould be typified by hl_:-
w-diversity area would ::-:::
~ abundance of res ou r~
.sure, thereby allowin~ _
Lild allow greater numb.:-
interspecific competi
L to conduct a large-sc::...::
j niche sizes for high- :
relate to gradient leng-...:..
d niches and see how r.;:_
ity is positively correla:.::_
tes positively with gre2::.:-
:e. Habitat diversity can ::-~:::
CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 419
n=3
C(/\\
,!YYYYf\
roo
n=3
2 I
I
rrcr;&\ I
n=3
1 CYY\ n=6
3
or
ar!YO n=6
(a)
~~(b)
FIG URE 14 .6 Hypothetical resource gradients and niche spaces for species in low- and
high-diversity geographic areas. The first example represents a situation in which the high-
diversity region has greater environmental variability and thus greater gradient length and
available niche space. The second example represents a situation in which gradient lengths
are equal, but the the region with high biodiversity accommodates more species because
the organisms living there possess more specialized niches and require less length along
the resource gradient. The third example represents a situation in which resource
abundance is higher in the area with high biodiversity and thus can accommodate more
species either because (a) the abundance of the resource allows the area to support very
specialized species that exist using a small portion of the resource gradient and niche
space, or (b) they must compete less for the abundant resources and have greater gradient
and niche overlap.
generated by differences in the physical or biological environment. For example,
areas with complex topography present a greater number of habitats and wider
variability of habitat types than do areas with flat topography. Topography influ-
ences habitat diversity at both small and large scales. At the small scale, a field with
gently rolling topography might include relatively dry hilltop soils, mesic hillslope
soils, and moist soils in low-lying areas. Habitat would thus be available for plants
and animals that required both dry soil conditions and mesic or wet conditions. The
rolling topography of the field described above would present more habitat diver-
sity and have a greater species diversity than an adjacent flat field that only pre-
sented mesic soil conditions. At the large scale, mountainous regions can include a
wide range of environments, extending from hot and dry low-elevation deserts,
cooler and moister mid-elevation regions supporting woodlands and forest , and
cold high-elevation tundra. In addition, the diverse topography of mountainous
420 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

regions can lead to the reproductive isolation of species populations and prod _ 3
increased rates of allopatric speciation. In Central America, the mountainous ~ ­
topographically diverse landscapes of Costa Rica support many more tree spe = .i!'
than the relatively flat areas of adjacent southern Nicaragua. In the United St a i.~ '§
(]) 2
mammal biodiversity is highest in the topographically diverse areas of the Pa -= >
:0
coastal mountains, Rocky Mountains, and Appalachian Mountains. rJ)
(])
Although topographic diversity may explain local and regional differe n-~ "ti
(])
a.
in biodiversity, it does not provide an adequate explanation for global biodive _ rJ)

gradients. Both large sections of the Amazon rainforest and the boreal fores: -E
i:C
central Canada are relatively flat areas, with topographic habitat diversity ge =-- /
ated by low uplands, small-scale landscape features, and floodplains. As we he :-
mentioned, the Amazon rainforest, however, has much greater biodiversity , _ 0
0
the Canadian boreal forest. Similarly, tropical mountains have much higher bio- p
versity than the high-latitude mountains of western Canada and adjacent Alas ·
Nor can diversity in topography or physical oceanographic conditions explain .:... : FIGURE 14.7
t he eastern Unii
increasing gradient of marine species richness from the polar areas to the equa -
Vegetation structure is an important biological contributor to habitat d i,-~_
sity. Vegetation structure can vary both horizontally and vertically. The vert:i
structure of the vegetation can greatly influence habitat diversity. A highly srn: -- Large Lano
fied forest provides differences in plant composition and microclimate in each s _ continuous la
tum and presents a wide range of habitats for birds, mammals, reptiles, amphibi - tralasia is an i
and insects. In addition, the different plant species that dominate different st .,_- species divers
provide different food sources. Consider a tropical rainforest with its typically co- - to the large 1
plex stratified canopy. Some animals, such as many species of tropical butterfii~ relative size '
are adapted to life on top of the sunny and relatively dry upper canopy surface. ?:-:- that are typic
mates are often adapted to life within the upper to middle canopy, taking adY -- the area of th
tage of tree limbs for movement and living off fruits. Some amphibians live on ·'-:: to present th<
lower vegetation and moist cool floor of the forest. It has been shown that m - _ relative land
tropical beetle species are adapted to life in a very specific section of the vegeta · that portray t
strata and are absent from all other areas. This helps generate extremely high di,·-~­ see that the 1<
sities of beetles in tropical rainforests. A single Lubea seemannii tree in the P cator projecti
manian rainforest may be host to over 400 species of canopy-dwelling beetles, ,,; -- the vicinity o
the lower limbs and trunks hosting an additional 200 species. Differences in for -:- lies in the vi
strata diversity, even in less complex forests, can have an impact on habitat divers:._ 7,682,000 sq I
and species diversity. A study of bird species diversity in the deciduous forest of :: only 2,175,00(
eastern United States showed a strong positive correlation between diversity L. The pre
foliage heights in the forest and the species richness of birds (Fig. 14.7). such as the A1
Does increasing vegetation stratification and associated habitat divers-•. geographic di:
really explain the latitudinal gradients in species richness from the poles to t :: decreases the
equator? Again, the answer is no. First of all, increased vegetation stratificatio:: population si:
and biological habitat diversity do not explain why biodiversity in tropical deci-'- large areas, sr
uous forest, savanna, and grasslands is higher than structurally similar vegetati ::. gradients and
in mid- and high latitudes. An area of African savanna can contain dozens : populations il
species of large mammal herbivores; a similar sized area of forest-tundra vegetc- problem with
tion supports only two or three large mammal herbivores. Second, attributing rt ::: biome has mu
latitudinal gradient of species diversity to increasing vegetation structure do : ate deciduous
not explain what produces the gradient of increasing plant species diversity froc does not expl
poles to tropics. It is the higher diversity of plant species in the tropics that pro- oceans arounc
duces much of the complex vegetation stratification found in the rainforest. many marine

~s populations and produc=
erica, the mountainous a=-.:
ort many more tree spec!;;:o:
mgua. In the United States.
diverse areas of the Pace:
Mountains.
al and regional differen c_~
ltion for global biodivers:.
st and the boreal fore st -
hie habitat diversity genc:--
rrd floodplains. As we he.· =
h greater biodiversity tl.:.::
rrs have much higher biOC:-
mada and adjacent Al as~
phic conditions explain ~:::
polar areas to the equ ar c~
)ntributor to habitat d i\·c~­
and vertically. The verti.::::..
1t diversity. A highly str-.::.=-
i microclimate in each s:::=-
nmals, reptiles, amphibic..::...
t dominate different srr.::.:..c
orest with its typically cc=-
~cies of tropical butterf.:~
r upper canopy surface. P:- -
ddle canopy, taking adY.:c-
lme amphibians live on :::.-
has been shown that m--
fic section of the veget a::: ~
erate extremely high d i\-~
>eemannii tree in the Pee..:
nopy-dwelling beetles. v.~::
lecies. Differences in fo~::­
impact on habitat diveE..
the deciduous forest of:::.:
ation between diversitY
'irds (Fig.l4.7).
;sociated habitat divers:-;
rress from the poles to ~
~d vegetation stratifica::: :
diversity in tropical dec:
.cturally similar vegeta:i .:
ma can contain dozens :
~a of forest-tundra vegc:..c
res. Second, attributing·-
vegetation structure c co::-
lant species diversity frc
ies in the tropics tha t ~=- -­
und in the rainforest.
CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 421
•
c •• • c
·~ 2 -~
(1)
> >
'5
Cfi
(1) • '5
Cfi
(1)
••
"(3 "(3
(1)
Q.
'f 1
• (1)
Q.
'f
"E "E
i:i5 i:i5
•
0 0
0 1.0 2.0 3.0 0 0.5 1.0 1.5
Plant-species diversity Foliage-height diversity
FIGURE 14.7 The relationship between bird species diversity in the deciduous forest of
the eastern United States and vegetation structure (after MacArthur and MacArthur, 1961)
Large Land Areas of the Tropics It has been suggested that the large
continuous land area of the tropical zone in South America, Africa, and Aus-
tralasia is an important contributor to the development and maintenance of high
species diversity. You may have the impression that the tropics are small relative
to the large land areas of northern Eurasia and North America. However, the
relative size of tropical areas is understated in the Mercator projection maps
that are typically used to show the world. The Mercator projection "stretches"
the area of the high latitudes and "shrinks" the area of the low latitudes in order
to present the spherical earth on a two-dimensional surface. If you look at the
relative land area of the tropics on a globe or on alternative map projections
that portray the relative size of the tropics and high latitudes correctly, you will
see that the land area of the tropics is actually very large. For example, on Mer-
cator projections the island of Greenland, which lies close to the North Pole in
the vicinity of 70° to 80° N latitude, appears to be larger than Australia, which
lies in the vicinity of 12° to 30° S latitude. In reality, Australia covers some
7,682,000 sq km and is over three times larger than Greenland, which covers
only 2,175,000 sq km.
The presence of large areas of continuous habitat in the equatorial zone,
such as the Amazon rainforest, allows species to have large populations and large
geographic distributions. As we have discussed in Chapter 9, large population size
decreases the risk of extinction. Large geographic distributions that support large
population sizes lead to decreased risk of extinction. Thus, if distributed over
large areas, species can survive with narrow ranges of tolerance along resource
gradients and develop specialized niches that would lead to unsustainable small
populations if the geographic area of the species' distribution was small. The
problem with this hypothesis is that it does not explain why the huge boreal
biome has much lower species richness than smaller biomes such as the temper-
ate deciduous forest , Mediterranean, or coastal rainforest biomes. In addition, it
does not explain latitudinal gradients in marine biodiversity. The cold southern
oceans around Antarctica are spatially large but have a much lower diversity of
many marine groups than does the tropical Pacific or Atlantic.
422 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Environmental Stability It has been suggested that environments in whi -- disturbance-se

climate is relatively stable on the short time scale (days, months, seasons, and yea.-:: numbers and <
promote higher biodiversity than unstable environments. Environments with l sity can be e)
amounts of daily, seasonal, and annual variability allow species to become fine quently enoug
adapted and to develop the most efficient form and behavior to take advantage for the surviva
resources without requiring tradeoffs that allow them to cope with variability are not so free
environment. As the species become more specialized and are more efficien bance-sensitiv•
extracting resources, more species can be supported by given resources. Connell'
As we have seen in previous chapters, tropical environments are typified ~ turbance freqt
low amounts of daily and seasonal variability in climate. Mid-latitude enviro:.- Reef of Austra
ments can experience large variations in daily temperatures and huge variatio::... forest fires pre
in seasonal temperatures. The short-and long-term variability of the higher l :..- regimes can p1
tudes requires species to be more generalistic in terms of niche breadth. Becau.s- would becomE
they are generalists, they cannot extract resources as efficiently, but they suni = (Picea glauca)
because they are adaptable to a wide range of conditions. including grasE
One attraction of the climatic stability hypothesis is that it explains both · .__ = aspen (Populu
decrease in species richness at high latitudes and the increasing size of speci -"'" vive in the sha
ranges that are found at high latitudes. The larger ranges found at high latitu ·eo' and longevity, :
are considered to be a reflection of the more generalized niches of the spe ~~ Given time, tht
there. If we assume that competition between species is important in determi ·- = there were no
how many species can be supported in a given environment, we can postula:= throw, and ins•
that since the niches of the higher latitude species are more general, not as rn a::_ allow the shad
species can be supported there. rary high-light
The stability theory has three drawbacks in explaining the geographic :"- extremely freq
tribution of biodiversity. First, it is unclear why short-term climatic instabili._ it takes spruce
should be so limiting to speciation via natural selection. Why could species n sitive long-live,
evolve adaptations to climatic variability without sacrificing their ability to utiliz= The inter
resources? Second, many areas of the earth have very stable environmental co- - the greatest in
ditions but experience low species diversities. For example, the deep ocean ~ tion is low and
extremely stable today in terms of temperature and salinity, but it has lowc. population of s
species diversity of fish and invertebrates than many shallow-water areas whi ._ causes the loca
are far less environmentally stable. Large saline lakes in the desert, such as Mor. takes 30 years
Lake in California or the Great Salt Lake in Utah, have very stable daily or sec.- frequency of 01
sana! conditions compared to adjacent small alpine lakes in the Sierra NevaC2 diversity. Distu
and Rocky Mountains. Yet the saline lakes have much lower diversities of aqua ·:. reproductive rr
organisms. Third, latitudinal diversity gradients are seen in many groups = grasses, herbs, <
species for which there is no evidence for more general niche breadth or wide:- Anumbe
geographic distributions at high latitudes. potential impo
versity. Howev
Disturbance In 1978, the eminent ecologist J. H. Connell proposed a theo _. global gradient
that links biodiversity to disturbance. The idea that Connell proposed is calle would have to
the intermediate disturbance hypothesis. According to this hypothesis, if 2 lower exclusior
ecosystem remains free of disruption by disturbance, the stable homogeneous rates than othe
environmental conditions will favor some species but will lead to the extinctior: proposition. In
of species for which the stable habitat is not favorable or which are prone to com- disturbance rat
petitive exclusion by species that are favored by the undisturbed environmen increasingly cle
The loss of species produces decreased biodiversity. Disturbances cause physica: bance from a n
and biological changes and spatial heterogeneity that can favor species tha: landslides, and
would not survive in a stable undisturbed system. On the other hand, if distur- rates that are 1
bance occurs too frequently and is too severe, it will lead to the extinction o: temperate rainJ
 CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 423

tments in which disturbance-sensitive species that have long generation times or occur in low
sons, and years) numbers and are prone to chance extinction. Thus, maximum levels of biodiver-
ments with low sity can be expected from intermediate levels of disturbance that occur fre-
) become finely quently enough and cause enough disruption and environmental heterogeneity
ke advantage o: for the survival of species which could not persist in an undisturbed system, but
th variability ill are not so frequent or severe as to cause the extinction of long-lived and distur-
wre efficient ill bance-sensitive species.
trees. Connell developed his hypothesis largely on the basis of differences in dis-
s are typified b,- turbance frequency and biodiversity which he observed at the Great Barrier
ttitude environ- Reef of Australia. He initially applied the theory there and to rainforests. Boreal
huge variations forest fires provide a good terrestrial example of how intermediate disturbance
the higher lati- regimes can promote species diversity. If fires were excluded, many boreal sites
readth. Becausi" would become dominated by long-lived conifer species such as white spruce
)Ut they survi' - (Picea glauca) or balsam fir (Abies balsamifera). Other boreal plant species,
including grasses, many herbs, shrubs such as willow, and deciduous trees such as
xplains both tb:: aspen (Populus tremuloides) and paper birch (Betula tremuloides), cannot sur-
\ size of specic vive in the shaded conditions under a spruce or fir canopy. Because of their size
1t high latitudc and longevity, spruce and fir are superior competitors for light and soil moisture.
s of the specic Given time, the shade intolerant species would likely go extinct, at least locally, if
t in determinffi;: there were no disturbances to open the coniferous forest canopy. Fires, wind-
·e can postula:.c throw, and insects create small and large patches in the coniferous forest that
ral, not as man: allow the shade intolerant species to persist by taking advantage of the tempo-
rary high-light intensity in these gaps. If, however, severe fires were to become
geographic dis- extremely frequent, occurring at intervals of say 10 years, (which is less time than
natic instabili:' it takes spruce or fir to reach sexual maturity), then spruce, fir, and other fire-sen-
)Uld species nc: sitive long-lived species would go extinct.
ability to utiliz;: The intermediate disturbance hypothesis predicts that biodiversity will be
ironmental co::- the greatest in systems in which the rate of displacement of species by competi-
~ deep ocean s tion is low and the rate of disturbance is also low. Thus, if it takes 50 years for a
mt it has lowc:- population of slow-growing trees to develop a dense canopy and root system that
lter areas whic::. causes the local extinction of shade intolerant grasses, herbs, and shrubs, and it
rt, such as Mo::: - takes 30 years for the spruce trees to reach reproductive age, then a disturbance
ble daily or sec- frequency of one event every 40 years would be optimal for maintaining species
; Sierra Neva~ diversity. Disturbances once every 40 years would give the spruce time to reach
rsities of aqua:::: reproductive maturity but would not allow the trees to completely displace the
nany groups c· grasses, herbs, and shrubs.
•readth or wirb- A number of field studies and computer modeling experiments support the
potential importance of intermediate disturbance regimes in generating biodi-
versity. However, the hypothesis encounters several problems in explaining
oposed a thea=-: global gradients of species diversity. To explain global biodiversity patterns, it
oposed is call.::: would have to be assumed that tropical ecosystems provide a combination of
hypothesis, if _ lower exclusion rates of species from undisturbed sites and lower disturbance
e homogeneo:..:= rates than other ecosystems. It is difficult to develop evidence for or against this
to the extinctic= proposition. In many areas of the world, both on land and in the ocean, long-term
re prone to co;:;:- disturbance rates and competitive exclusion rates are unknown. It is becoming
~d environme:::.. increasingly clear that even tropical rainforest ecosystems are prone to distur-
!s cause physic.2 bance from a number of sources, including fires, hurricanes, wind throw, flooding,
vor species t h~ landslides, and pathogens. Many tropical environments may have disturbance
r hand, if disn.:.:-- rates that are not much less than areas of the temperate deciduous forest or
he extinction L temperate rainforest. It is also unclear what biological or physical factors would
424 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

ultimately cause tropical species to experience particularly low rates of exclusio- occur in matu
relative to disturbance rates be capable of
that seed pred
Competition It has been suggested that the natural selection and evolutio- members of tl
of species in the mid- to high latitudes is driven by adaptation to physical stress~ tive exclusion
related to climate, such as cold or aridity. In contrast, evolution in the warm a :. predation is p;
moist tropics is thought to be driven more by interspecific competition. The hi ~ ·­ important see
degree of competition in the lower latitudes leads to species developing speci ·- which species
ized niches that restrict their distribution in terms of habitat and along reso ur~ tors are comrr
gradients. Species are competitively superior within narrow ranges of enviro::.- tributed near t
mental conditions. Specialized niches and narrow habitat preferences decreas:: close to the p<
direct competition with other species. This increases the number of species th;;.· from the pare
can survive locally, each species using a narrow range of microhabitats. For exarz.- species. Such a
ple, in the tropics many frog species breed in ponds and other standing water s· · species. Seeds
ilar to the breeding habitat of most temperate frogs. However, other species a;-:: germination.T
adapted to breed in the small pockets of moisture in the forest floor or cano ~ species keeps
including the water that collects in the foliage or flowers of plants such as e .· competitive eJ!
phytic bromeliads. The wide range of specialized breeding areas used by tropi Applyin1
frog species decreases competition for breeding space. versity gradie1
Some evidence of the possible impact of competition on the niche size that large nurr
tropical animals comes from studies of anole lizards (Anolis spp.) in t cases but deer
Caribbean. The niche breadth of the anole species has been measured in term unicellular org
the size of prey they eat. Anoles that hunt prey of only a certain size are consi ""- the pitcher pl
ered specialists, whereas anoles that eat prey of a variety of sizes are conside r ~.:. mosquito larv;
generalists. When a single lizard species occupies an island, it generally will h - · theory does nc
and eat a larger range of prey of different sizes. When several different specie and parasites i
anoles occur together, each species often has a relatively narrow range of pr :
sizes that it will hunt and eat. The narrow ranges of prey size decrease the over! -;: Productivitl
in prey species hunted by each lizard species. Thus, competition between speci : tivity will dev'
for prey is decreased. duces more en
The competition theory suffers serious drawbacks when applied to explai::. high amounts
global biodiversity patterns. First, most plant species, especially trees, are limi t - relatively spec
by a relatively few resources, such as light, water, heat, and soil nutrients. Many 0: consumers, th t
them do not display narrow and nonoverlapping requirements for th es:: environment. -
resources, even in tropical settings. There is in fact little convincing evidence the.: greater diversi
competition and narrow niche specialization can explain tropical tree diversi . pollinators, ant
Second, it still must be explained why competition is such a potent force in th:: lation betweer
lower latitudes, but not in temperate and arctic regions. If competition alone w~ The typical pri
responsible for biodiversity, why wouldn't similar decreases in niche bread rts meter per year
and high rates of biodiversity develop everywhere? meter per yem
These differen
Predation It has been argued that the high numbers of predator and parasi ~ els of plant anc
species, particularly in the tropics, maintain high biodiversity by keeping prey po When ex
ulations low and decreasing the competitive exclusion of one prey species b. ductivity and
another. The reduction in competition in turn allows for more prey species lC' swamps haves
evolve, and this leads to the coevolution of additional specialist and generalis: typically prodt
predators. The predation model has been applied in a very interesting manner fo:- have extremeb
an explanation of tropical tree diversity. D. H. Janzen noted that tropical forests are dominated
contain high numbers of different tree species but that most tree species occur as nia). Studies ol
isolated individuals. In general, stands dominated by one or two species do no: contradicts the

tlarly low rates of exclusio::.
ral selection and evolutio;:
tptation to physical stress~
evolution in the warm ar:::
:cific competition. The hi~-­
species developing speciC.:-
habitat and along resourc=
narrow ranges of envirm:.-
bitat preferences decre2S::
the number of species rt:.·
)f microhabitats. For exa=-
:l other standing water si.;::.-
-Iowever, other species c..:-:'
the forest floor or can o~
;vers of plants such as e;:-i-
:ling areas used by tropi~
titian on the niche sizes ·='
trds (Anolis spp.) in ~=
been measured in terms ,:
y a certain size are cons:::-
ety of sizes are consider.::-::
land, it generally will b··-
;everal different species ~
vely narrow range of p:-::c
y size decrease the ove r ~:­
npetition between spec.::-
:s when applied to explc_
specially trees, are lini :.::-~
md soil nutrients. ManY
requirements for t b~=
~ convincing evidence :.::.:.
ain tropical tree dive~:-'
mch a potent force in :=.-
•. If competition alone "':_
:reases in niche breac:.::..
·s of predator and par~:::
rsity by keeping prey JX"':'-
n of one prey species _
for more prey species ·
1 specialist and generC.:...
~ry interesting manner : _
1oted that tropical for6--::
most tree species occu:-
>ne or two species do ::..
CONTROLS ON GEOGRAPHIC GRADIENTS OF SPECIES DIVERSITY 425
occur in mature tropical lowland forests, which means that no species appears to
be capable of effectively excluding others at a landscape scale. Janzen suggested
that seed predation may explain the spatial isolation of rainforest trees from other
members of the same species. This in turn explains the apparent lack of competi-
tive exclusion of some tree species by others. He suggests that specialized seed
predation is particularly common in tropical forests. Indeed, insects are extremely
important seed predators in the tropics, and many are very specific regarding
which species they will eat. Very high concentrations of these specific seed preda-
tors are common in the vicinity of their host trees. Thus, most seeds that are dis-
tributed near the parent tree are devoured. The loss of these seeds keeps the area
close to the parent tree free of its own offspring. Seeds that are dispersed away
from the parent tree may end up at sites away from other members of their
species. Such areas will, of course, lack the specific predators that prey on the tree
species. Seeds transported to these areas have a greater chance of survival and
germination. The inability of trees to establish close to other members of the same
species keeps the populations of tree species small and dispersed. This decreases
competitive exclusion and allows a greater number of tree species to coexist.
Applying the competition model of species diversity to explain large biodi-
versity gradients raises two issues. First, field and laboratory experiments show
that large numbers of predators lead to increased prey species diversity in some
cases but decreased prey diversity in others. For example, the species richness of
unicellular organisms found in the small pools of water that form in the leaves of
the pitcher plant (Wyeomyia smithii) decreases as the number of predaceous
mosquito larvae living in the pools increases. Second, and most importantly, the
theory does not explain why there should be much greater numbers of predators
and parasites in the low latitudes in the first place.
Productivity It has been suggested that regions with high primary produc-
tivity will develop high biodiversity because the vegetation in such areas pro-
duces more energy that can support more species at the higher trophic levels. The
high amounts of available energy allows for primary consumer species to have
relatively specialized niches in terms of food. The greater the number of primary
consumers, the greater the number of predators that can be supported by the
environment. The large number of different herbivorous species promotes even
greater diversity of the plants through the coevolution of specialized herbivores,
pollinators, and seed dispersers. At a global scale there is indeed a general corre-
lation between primary productivity and latitudinal differences in biodiversity.
The typical primary productivity of tropical rainforests is some 2200 g per square
meter per year, whereas temperate forests produce only about 1200 g per square
meter per year, and boreal forests produce only about 800 g per meter per year.
These differences in productivity can be directly correlated with the relative lev-
els of plant and animal biodiversity in these biomes.
When examined at smaller scales, the relationship between primary pro-
ductivity and biodiversity breaks down. For example, estuaries, marshes, and
swamps have some of the highest rates of primary production in the world. They
typically produce well over 2000 g per square meter per year. Yet these systems
have extremely low diversities of both plants and animals. Many estuary marshes
are dominated by single species of vascular plants such as pickleweeds (Salicor-
nia). Studies of plant biodiversity in terrestrial settings also present evidence that
contradicts the hypothesis that high biodiversity is a result of high productivity.
426 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Comparisons of plant species diversity and primary productivity in Australia an.:

South Africa show that in these regions highest plant biodiversity is correlate.: 180 Trees
with intermediate levels of primary productivity. Areas with very high or very lo>-
160
primary productivity have lower plant biodiversities.
140
en
A Synthesis It appears that no single general rule can be applied to expla·- en
all the differences in biodiversity at large and small scales. Explaining the larg - ~ 120
.s:::
(.)
·;::
scale gradient between the high latitudes and the low latitudes is perhaps o~ ~ 100
greatest interest to biogeographers. The British biogeographer Kevin Gaston h - ·c::; •
Q)
done much analysis of this problem and comments as follows: "The search for :: a. 80
en
single explanation for latitudinal gradients in diversity may be attempting u, • • ••
answer two questions at once, how available energy is converted into individu-
organisms and how these individuals are distributed amongst species" (1996.
Q)
Q)
t!=
60

40
•
.......
•

471). The large difference in biodiversity between the high and low latitud :
likely reflects several factors. 20
First, history cannot be completely excluded as a factor in causing the rel ~­
tively low biodiversity found at high latitudes. Although global glacial periods eli.: 2
result in cooler and drier conditions in the tropics, the magnitude of these chang :
was small compared to the large changes in climate and environment that occurr - 200
in the mid- and high latitudes. During glacial periods, the geographic ranges of man: Vertet
high-latitude plant and animal species were displaced by thousands of kilomete _
The fossil record shows that in both Europe and North America, many specie 100
temperate plants went extinct during the transition from the warm and moist con<E- en
en
Q)
tions of the early to mid-Teritiary to the colder and more variable conditions of tb; c:
.s:::
Pleistocene. We also know that the species composition of mid-and high-latitude (.)
·;:: 50
en
plant and animal communities was markedly different during glacial and nonglaci- Q)
·c::;
periods. The reshuffling of community composition would work against the creati - Q)
a.
(f)
of new species by coevolution or competitive interactions. Finally, the tropical co -
munities, particularly rainforests, were probably fragmented during glacial perio •
but were not dislocated in terms of latitude as was the case for arctic and tempera-;
communities. Environmental factors such as day length or the degree of contras-
between winter and summer climates did not change as much for tropical species •
it did for high-latitute species that experienced large latitudinal shifts in their rang :
during glacial and nonglacial periods. Although they were fragmented, the speci :
composition of the tropical communities may not have changed as much as that o: FIGURE 14.8
temperate and arctic communities. The fragmentation of the rainforests durin; evapotranspirati
glacials likely promoted allopatric speciation. (after Currie, 19f
History alone, however, cannot explain all of the differences in speci !
diversity between high and low latitudes. It is also likely that much of the latit -
dina! diversity gradient is related to climate. At a global scale a very clear rela- vegetative foo
tionship exists between climate, primary productivity, and species diversity. Fo~ and insect spe
example, on continental and global scales, areas with high annual evapotranspira- or arctic settir
tion rates are typified by high species diversity of plants and animals (Fig. 14. food in winter
High evapotranspiration rates occur in regions with high amounts of sol<L ians, and inseci
energy, heat, and moisture. Light, heat, and moisture are key resources for plants birds rely on c
and animals. High amounts of light, heat, and moisture undoubtedly lead to would not sun
higher primary productivity, and at a global scale, this provides energy to suppon There is
a wider range of species. The low-latitude areas of highest biodiversity are also of the tropics
areas where seasonal differences in climate are relatively small. The ability o: water, and 10\
plants to photosynthesize and produce energy throughout the year means tha ment of addit
 ISLAND BIOGEOGRAPHY 427

roductivity in Australia ar..:

tt biodiversity is correlate.: 180 Trees
: I
s with very high or very lo 160

140
(/)
e can be applied to expl2.1.:: (/)

g? 120
cales. Explaining the larg::- .s:::
(.)
·;:
ow latitudes is perhaps c
Jgrapher Kevin Gaston b=....
s follows: "The search for _
~ 100 ~
~ 80
(/)
ch
•

• •
.... ..
• •• •
•• • •
•
•• I
•
•

sity may be attempting :

al • •
s converted into individl.!.=.... t=
60 ••••
•• •
amongst species" (1996. ~ 40
the high and low latitud:- • •
•
:1 factor in causing the re~
~h global glacial periods 6.: 500 700 900 1'1 00
magnitude of these chang::- Annual evapotranspiration (mm)
l environment that occurr::-.:.
~ geographic ranges of rna.::
by thousands of kilomete;:..
200 r v""""'"' •
h America, many species - 100
n the warm and moist con.::..- (/)
(/)
Q)
re variable conditions of --- c
.s:::
Jn of mid-and high-lat i ll.!~ (.)
·;: 50 •
(/)
1uring glacial and nonglac-_ -~
tld work against the crea·· -- Q)
c.
(/)
•ns. Finally, the tropical co=
ented during glacial perio.::
:ase for arctic and temper::.·
I :'1·· • •
:h or the degree of com:-.:...
much for tropical species _
itudinal shifts in their ram:::- 500 1,000 1,500 2,000
'ere fragmented, the spec::- Annual potential evapotranspiration (mm)
changed as much as tha: FIGURE 14.8 The relationship between energy and moisture as measured by
n of the rainforests durc:. evapotranspiration and the species diversity of trees and vertebrates in North America
(after Currie, 1991).
the differences in spec::-
ely that much of the lati __
)bal scale a very clear re vegetative food and fruit are available at all times. Many tropical bird, mammal,
y, and species diversity. F - and insect species, particularly obligate fruit eaters, cannot survive in temperate
tigh annual evapotranspi:-_ or arctic settings because they do not have access to adequate or appropriate
nts and animals (Fig. 1 ..:. .~ food in winter. Warm climates throughout the year also allow reptiles, amphib-
ith high amounts of s o~::_­ ians, and insects to remain active in summer and winter. Many tropical predatory
lre key resources for pia=_ birds rely on consumption of reptiles or insects during summer and winter and
sture undoubtedly lead - would not survive in areas with cold winter climates.
provides energy to sup There is undoubtedly a positive feedback relationship between the ability
ighest biodiversity are a:s of the tropics to support high biodiversity because of available light, heat, and
ttively small. The abili tY water, and low seasonal variability in energy, and the evolutionary develop-
tghout the year means r::_ ment of additional new species. The wide diversity of plant species and the
428 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

resulting complex canopy structure that develops in the tropical rainforests Geograp
produce a wide range of habitat types for animals and smaller plants. As we dis-
cussed earlier, increasing niche specialization in response to competition can be Study after
a strong contributor to biodiversity in areas such as the tropical rainforest lated with i
where resources are abundant. Abundant prey in high-productivity areas than do sm;
allows the evolution of prey-specific predators that can exert pressure on the the Galapa1
population size of their prey species, keeping that species from becoming dom- plant speci<
inant and competitively excluding other species. In summary, we can say that species of L
the high biodiversity of the tropics has defied one single simple explanation and island of H
is more realistically the product of many historical and equilibrium factors. species tod<
in area sup]
the Caribbe
ISLAND BIOGEOGRAPHY port over 5(
and Redon'
Islands have always held a special attraction to biogeographers. The fascination with Jamaica anc
islands extends back to the foundations of the discipline. Darwin's ideas regarding The p.
evolution were very strongly influenced by the biological diversity of the finch found on isl
species that he observed on different islands of the Galapagos Archipelago. So, too,
was Wallace influenced by the similarities and differences in bird species and other
organisms that he observed on the islands of the Malaysian Archipelago. Islands
have been very important in many subsequent studies of the role of geography in
evolution. Examples that we have already discussed in this book include the work of
Mayr on kingfisher species on New Guinea and surrounding islands, and the many
studies of fruit flies, honeycreepers, and plant species on the Hawaiian Islands.
Many biogeographers have focused on intriguing biological attributes and (/l
Q)
phenomena that are characteristic of island organisms and ecosystems. In 1880, Wal- '() 100
Q)
lace published a classic book on this topic called Island Life. In 1965, Sherwin Car- a.
(/)

lquist published an updated synthesis of island biology that was also entitled Island 0
Qj
Life. From these writings we can identify a suite of characteristics that are often .0
found in insular species of plants and animals and distinguish them from mainland E 10
::J

ancestors and relatives. Such features include the loss of dispersability, the develop- z
ment of gigantism or dwarfism, the loss of antipredator defensive features and
behaviors, the development of woody shrubs and trees from species that occur as
soft-stemmed herbs on continents, and the development of highly specialized niches.
The study of species richness on islands has had a long and important history
(a)
in biogeography. In the nineteenth century, the distinguished botanist, Joseph
Hooker, analyzed Darwin's plant collections from the Galapagos Islands and Austra liar
attempted to decipher the cause of interisland differences in plant species richness. 5
In the early twentieth century, 0. Arrhenius studied the biota of islands in the Baltic 4
Sea and produced the first paper on the species-area curve. This paper precipitated 3
heated debate on the meaning of species-area patterns with the noted American 2
ecologist H. A. Gleason (see Chapter 6). The analysis and interpretation of patterns
of biodiversity on islands became a topic of much interest among ecologists and bio-
geographers from that point onward. In 1967, Robert MacArthur and Edward 0.
~
0
Wilson published a seminal book in the field of biogeography. Their book, entitled 0 10
The Theory of Island Biogeography, provides a theoretical construct linking species- (c)
area relationships on islands to dispersal and extinction processes. In the decades
since its publication, MacArthur and Wilson's theory of island biogeography has FIGURE 14.9
remained the subject of intense interest, research, and vigorous debate. Before we amphibians in
pond size in th
examine the theory of island biogeography, let's take a look at the generalized
mo nta ne habi1
patterns in island biodiversity that underlie the theory.
MacArthur an<
 ISLAND BIOGEOGRAPHY 429

in the tropical rainfores:5 Geographic Patterns of Island Biodiversity

d smaller plants. As we di>-
onse to competition can c::
as the tropical rainfore5:
[n high-productivity are2..5
. can exert pressure on tL:
Jecies from becoming doc -
summary, we can say th2.·
1gle simple explanation a- -
md equilibrium factors.
~raphers. The fascination \\i ·"-
ne. Darwin's ideas regardiL;
ogical diversity of the fine:.
Llapagos Archipelago. So, toe
1ces in bird species and othe:-
llaysian Archipelago. Islan-'-
s of the role of geography c
this book include the work c:
unding islands, and the man:
m the Hawaiian Islands.
ing biological attributes an:
Study after study has shown that the species richness of islands is strongly corre-
lated with island size. Larger islands support more species of plants and animals
than do smaller islands. This pattern is seen on local to global scales. Analysis of
the Galapagos Islands clearly shows a strong correlation between island size and
plant species richness. The larger islands in the archipelago support over 300
species of land plants. The smaller islands support a handful of species. The big
island of Hawaii is over 10,000 sq km in area and supports over 30 native bird
species today. Islands in the Hawaiian Archipelago which are less than 400 sq km
in area support fewer than 10 species of native birds. Analysis of biodiversity on
the Caribbean Islands shows that large islands such as Cuba and Hispaniola sup-
port over 50 species of reptiles and amphibians, while small islands such as Saba
and Redonda support less than 10 species. Intermediate-sized islands, such as
Jamaica and Puerto Rico, support an intermediate number of species (Fig. 14.9).
The pattern of increasing species richness on larger islands has also been
found on islands located in lakes. For example, the number of vertebrate species
1000 1- Caribbean reptiles
C/)

and ecosystems. In 1880, wa:- .5!!

(.)
<ll
100

rd Life. In 1965, Sherwin 0.

C/)

y that was also entitled Js[QI ;.; 0

Qj
characteristics that are oft e:::. .0
E
:tinguish them from mainlan.= :::J 10
z
of dispersability, the develor-
dator defensive features an:.
Redonda
~es from species that occur a>
nt of highly specialized nichn 1 ~----~------~------~--------~------~------
1 a long and important histor: 0 10 100 1,000 10,000 100,000
(a) Area of island (km 2 )
istinguished botanist, Josep::
the Galapagos Islands anc Australian fish Great Basin mammals
~nces in plant species richness.
1e biota of islands in the Balti-
curve. This paper precipitatec.
~rns with the noted America::. 0
Qj
and interpretation of patterlli .0
E
:rest among ecologists and bi :::J
z
rt MacArthur and Edward
OL-~L-~L-~~~--~---- 1L-~~~-L--~~~---
;eography. Their book, entitlec 0 10 100 1,00010,000100,000 10 25 50 100 250 5001,000
~tical construct linking species- (c) Area of pool (m 2 ) (b) Area of mountain (square miles)
tion processes. In the decade5
ry of island biogeography ha> FIGURE 14.9 The relationship between island area and species richness for reptiles and
1d vigorous debate. Before \\ :: amphibians in the Caribbean. The relationship between fish species richness and spring-
pond size in the Australian desert, and between boreal mammal species richness and
:~.ke a look at the generalize ~
montane habitat area on isolated mounatins in the Great Basin of North America (after
y MacArthur and Wilson, 1967; Kodric-Brown and Brown , 1993; and Brown, 1971).
430 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

found on islands in Lake Michigan is larger on big islands than on small islauds. Why sh1
In addition to true islands, many other forms of fragmented habitat can The likely rea
thought of as islands. One example is lakes and ponds that are not connected ·c
continents rna
other waterbodies by rivers and streams. These isolated waterbodies are islan •
surrounding e
of aquatic habitat surrounded by land. The analysis of desert spring-ponds i;:;
area, for exan
Australia shows that the species richness of fish generally increases in proportio::
regions. In ad
to the size of the ponds (Fig.14.9). Isolated mountain tops surrounded by warm :-
plants and ani
or drier lowlands are another form of island. For example, a number of bird a .:
ulations from
mammal species typical of the boreal biome can be found on high isolated mou::-
going extinct.
tains in the Great Basin region of the United States. Lowland areas in the Gre :
available to s1
Basin are too hot and dry for these mammal species to survive. Thus, these i o-
supported on
lated mountains can be likened to islands of boreal habitat in a sea of deser_
surrounded b
Analysis of the bird and mammal species richness shows that the number 0:
often difficult
boreal species found on Great Basin mountains is highly correlated with the are""
of alpine habitat provided by the individual mountains and ranges (Fig. 14.9). than small poJ
Larger islands generally have greater numbers of species than smalle:- Differen
islands for all of the same reasons that larger sampling plots on continents h a\-~ variation in sp
more species than small plots. We discussed these reasons in some detail in t h~ the only impo
preceding section. At this point we might then ask two questions. First, are th~ close to conti1
species-area curves for islands essentially the same for different island groups. o:- islands that an
do they differ from one set of islands to another? Second, are species-area cun· .:: Pacific that an
from islands similar to the species-area curves obtained from continents? We ca.::. and Australia I
compare the relationships between island area and species richness with differer.: islands that an
island groups and with continental areas using the species-area curve equatio;:. case of the S01
described previously (S = cA z). If we look at the relationship between island ar "" tant determine
and land plant and animal species richness on the Galapagos Islands, or t been observed
Caribbean islands, the species-area curve equations are as follows: isolated patch(
For Galapagos plants with distance f
species richne
S = 28.6A0 ·32 States is highe:
For Caribbean amphibians and reptiles Before v
the species on
S = 3.3A0 ·30 and islandlike
on the islands
The values for the constant c in the equations is larger for the Galapagos plan·•
mainland. In tl
because there are more plant species on the Galapagos than there are reptile an..:
on smaller isla
amphibian species in the Caribbean. Interestingly, the values of z for both th;
It is rare that 1
Galapagos plants and Caribbean reptile and amphibian fauna are roughly the
or a large isla
same (0.32 and 0.30). This implies that the relative rate of increase in speci .::
mainland or 1:::
related to increasing island area is the same for both systems. In fact, a wide rang.:
of insular com
of island studies have produced values of z that average around 0.30. In contra -
ever, are subse
the species-area equation for land birds in North America is:
many island Sf
S = 40.0A 0 ·17 the number o
island. The isl<
The high value for c relates to the large number of bird species on the continen mainland and
The value for z obtained from the species-area equation for North Americar; California lie c
birds is about half the value obtained for island systems. For species-area equa-
tions derived for different-sized areas on continents or other large landmasse
the value of z tends to range from 0.15 to 0.25. This means that the slope between The Equilit
the number of species and area increases more quickly for islands than for sites Why precisely
of different sizes located on continents. that are near

ds than on small islaud-
~mented habitat can lx:
hat are not connected tc
waterbodies are islanes
f desert spring-ponds l=
y increases in proportio:.
ts surrounded by warme:-
'le, a number of bird aL.:.
don high isolated mou;:-
.vland areas in the Grec.:
survive. Thus, these isc--
tbitat in a sea of dese:-_
)WS that the number e:
correlated with the are.:
nd ranges (Fig. 14.9).
of species than small:c:-
plots on continents hw.:::
ms in some detail in L::.::
questions. First, are C:::
ifferent island groups. ;:-
1, are species-area cu n ~
:rom continents? We c~
:s richness with differe::
:ies-area curve equatic_
ship between island a:-::_
1lapagos Islands, or ~=­
ls follows:
'or the Galapagos pia::::..,
ilan there are reptile c.=.:
values of z for both C::.::
il fauna are roughly ~=
e of increase in spec=--
~ms. In fact, a wide r a::~=
around 0.30. In con trc.o.-:..
:a is:
:pecies on the contine::_
on for North Americ.:..::.
>. For species-area eqe.:-
other large landmas._;;a_
s that the slope ben\·e::
for islands than for si:::
ISLAND BIOGEOGRAPHY 431
Why should species-area curves be different for islands and continents?
The likely reason is that much of the biota that we find on different-sized plots on
continents may be supported by the resources of the plot and resources from the
surrounding environment outside the plot. Birds that nest in a continental plot
area, for example, may forage for food both within the plot area and in adjacent
regions. In addition, a plot on the continent may support a small population of
plants and animals that are prone to local extinction, but immigration from pop-
ulations from just outside the plot boundaries keeps the local population from
going extinct. For island organisms, the resources on the island may be all that is
available to support them, and thus fewer individuals and fewer species can be
supported on an island surrounded by inhospitable ocean than on a plot of land
surrounded by a continent of similar habitat. Finally, immigration to islands is
often difficult, and small island populations are more prone to local extinction
than small populations on continental plots.
Differences in island size appear to explain a considerable amount of the
variation in species diversity between different islands. However, island area is not
the only important control on species richness. When we examine islands that are
close to continents, we find that they tend to have higher species richness than
islands that are located far away from continents. For example, islands in the South
Pacific that are located less than 800 km from the large land areas of New Guinea
and Australia have up to 10 times as many terrestrial and freshwater bird species as
islands that are located more than 3200 km from New Guinea and Australia. In the
case of the South Pacific, both island size and island isolation appear to be impor-
tant determinants of species richness. Low species richness on isolated islands has
been observed in many other studies. For example, the species richness of birds in
isolated patches of montane forest in Venezuela, Colombia, and Ecuador decreases
with distance from the main chain of the Andes Mountains. In a similar manner, the
species richness of boreal mammals on mountains in the southwestern United
States is highest on mountains that are close to other ranges (Fig.14.10).
Before we proceed, we might ask how the species on islands are related to
the species on other nearby islands or continents. In the case of both true islands
and islandlike areas of isolated habitat, it has been found that the species living
on the islands are often subsets of the larger species pool found on the closest
mainland. In the case of groups of islands, the species of plants and animals found
on smaller islands are generally subsets of the species found on the larger islands.
It is rare that the flora or fauna found on a small island located near a continent
or a large island would contain an entirely different group of species than the
mainland or larger island. This phenomenon is referred to as the nested pattern
of insular communities and is very widespread. Not all species on islands, how-
ever, are subsets of the species found on the mainland or larger islands. There are
many island species which have evolved on islands and are endemics. In general,
the number of island endemics increases with the degree of isolation of the
island. The islands of Hawaii are extremely isolated from other islands and the
mainland and have many endemic plants and animals. The Channel Islands of
California lie close to North America and have only a fewer endemic species.
The Equilibrium Theory of Island Biogeography
Why precisely do small isolated islands possess fewer species than large islands
that are near continents? The equilibrium theory of island biogeography was
432 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Theory of Is
bined extinc
sity observec
The th<
biogeograph'
(/)
Q)
121 studies ~
·u years after tr
Q)
0.
(/) MacArthur a
phy. After tt
Wilson rema
attention foe
Isolation of mountain (km) species. Som<
pies to study
FIGURE 14.10 The impact of increasing isolation on the species richness of boreal ure with mar
mammals found on mountains in the southwestern United States. Isolated mountains ha ve his ecologica
fewer species than those located close together (after Lomolino et al., 1989).
he published
tributions of
time. As he \\
Robert Mac;
advanced in 1963 to explain this near consistent relationship between specie= only in his e;
richness and island size and island isolation. It is an exceptional theory in terms o: oped and the
its conceptual simplicity, scope of applicability, and potential implications for he lived long'
species conservation. It is perhaps the most well known theoretical concept to
arise from biogeography in the twentieth century. For over 30 years the theory
has remained at the heart of much research and vigorous debate. Let's start by
looking at the people behind the idea.
In 1953 a young mathematician named Robert MacArthur arrived at Yale
University to begin a Ph.D. degree under the supervision of the noted ecologist.
G. E. Hutchinson. Today, Hutchinson is remembered primarily for his work or.
lake ecology. However, he had broad interests in all areas of ecology and evolu-
tion, including the study of biodiversity. For his part, MacArthur had an avi ·
interest in birds, and this is one factor that brought the young mathematician to
do a Ph.D. in ecology. Under Hutchinson's supervision, MacArthur applied his
mathematical expertise to questions of niche partitioning and community struc-
ture for warblers. He established himself as a leader in ecology by applying th
rigor and logic of mathematics to study the community organization and th
geographic distribution of plants and animals. By 1965 MacArthur was a facul ty
member at Princeton University. During this time, he began to interact with c.
young biologist named Edward 0. Wilson who was a specialist in the study o:
ants. Wilson had been fascinated by ants since his youth. By the 1960s Wilson
had become a faculty member at Harvard. He had a more traditional biolog\·
education than MacArthur and spent long seasons in the tropics collecting ant
Wilson saw the value of mathematics in interpreting ecological data and ha
even taken undergraduate classes in the early 1960s to learn calculus. Both
MacArthur and Wilson felt biogeography was a science that was rich with infor-
mation on species distributions but poor in rigorous theory to explain these dis-
tributions. In 1962 the two men turned their attention to the application of
FIGURE 14 . 11
mathematics to try and explain species-area relationships. By 1963 they ha
the co-ori ginate
published some of their ideas in a paper entitled "An Equilibrium Theory of
tragicall y yo un£
Insular Zoogeography." In 1967 their ideas were expanded in the book, Th e career co nt inue
 ISLAND BIOGEOGRAPHY 433

'cies richness of boreal
:es. Isolated mountains h aY ~
> et al., 1989).
tionship between spe c :~
ptional theory in terms c:
>otential implications fc::-
vn theoretical concep t :c
over 30 years the theor:
ous debate. Let's start )...
Theory of Island Biogeography. The theory they laid out in their book com-
bined extinction, dispersal, and geography to explain the patterns of biodiver-
sity observed on islands.
The theory of island biogeography generated immediate interest among
biogeographers and ecologists. Ten years after publication of the book, at least
121 studies were published that cited the theory of island biogeography. Twenty
years after the book's publication there were over 2000 such published citations.
MacArthur and Wilson quickly became major figures in ecology and biogeogra-
phy. After the book was published, they pursued individual research projects.
Wilson remained a towering presence in the study of biodiversity. Much of his
attention focused on the use of science to aid in the conservation of endangered
species. Some of his positions, such as the use of biological and ecological princi-
ples to study society, have been controversial, but he became a distinguished fig-
ure with many awards to his credit. What of Robert MacArthur? He continued
his ecological work, including a detailed field study of birds, in Panama. In 1972
he published a new and important book, Geographical Ecology: Patterns and Dis-
tributions of Species. By this point, however, MacArthur was battling against
time. As he worked on the 1972 book, he knew he was terminally ill with cancer.
Robert MacArthur passed away shortly after finishing his second book. He was
only in his early 40s. We will never know what new ideas he might have devel-
oped and the role he would have played in the development of biogeography had
he lived longer (Fig. 14.11).

:acArthur arrived at Ya:::

)n of the noted ecologis:
'rimarily for his work c:.
~as of ecology and evoi:.:-
MacArthur had an a'; :
young mathematician :.:-
1, MacArthur applied b
ng and community stru.:-
L ecology by applying t1::-

ity organization and t1::-

MacArthur was a facu::-
began to interact wit h :.
specialist in the study c:
1th. By the 1960s Wilsc:.
more traditional biol o~
1e tropics collecting an· ·
ecological data and he.:
to learn calculus. B o ~
! that was rich with info:-
eory to explain these di-
)n to the application c.
ships. By 1963 they h2:. FIGURE 14.11 Robert H. MacArthur (1930-1972) who along with Edward 0. Wilson was
the co-originator of the equi librium theory of island biogeography. Although he died
1 Equilibrium Theory c:
tragically young, the ecological and biogeographical concepts he proposed during his short
'anded in the book, Ti:!
career continue to stimulate much research by biogeographers and ecologists.
434 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

So, what is the equilibrium theory of island biogeography? The theory is

predicated on the assumption that the species richness of most islands is in a
state of equilibrium with processes of immigration and extinction. Immigration
through dispersal and colonization events brings new species to islands. Extinc- c
0
tion removes species. New species continuously arrive at the island by dispersal, nc
but the addition of these species is counterbalanced by the continuous process
~
of extinction that removes some of the new arrivals and some of the older 0
species. Extinction of species is in turn counterbalanced by further new arrivals, c
0
some of which are successful in colonizing the island. Assuming that no environ- ~Ol
mental change occurs, the species richness of the island will only vary slightly .E
around a stable mean. However, since new species are arriving and older species .~
are going extinct in a continual process, the species composition of the island 0
Q)

will be continuously changing. The changing species composition is called Cti

a:
species turnover.
The best way to consider the theory of island biogeography is to view it
graphically. In a graphical format we can present the number of species along
the horizontal axis and rate of extinction and colonization along the vertical axis
(Fig. 14.12). The number of species on an island can vary from 0 toP, where P
equals the number of species on the mainland from which the island is colo-
nized. The graph presents two curves-one for the rate of immigration by colo- c
0
nizing species and the other for the rate of extinction of species on the island. u
.s
The rate of immigration declines as the number of species on the island xQ)
increases. This is because if there are greater numbers of species on the island, 0
there will be fewer resources and less habitat available for colonization. In other c
0
words, the greater the number of species, the less unoccupied niche space there ~Ol
is. When many species are already present, new colonizing species face stiff com- .E
petition and cannot successfully immigrate. The rate of extinction increases with .~
0
the number of species that are on the island because the more species there are,
the greater the chance of random extinction events occurring to some species. If
more species are competing for a limited amount of resources, the population
sizes of the species will be small due to resource limitations and species with
*
a:

iii TSN --------- .

small population sizes will be more prone to extinction. The equilibrium number
of species that the island can support is the point on our graph where the rates of i:i TLN - -- -- ---- -
E TSF - - - - - - - - - -
immigration and extinction intersect. This is designated as S on the graph. At F! TLF --------- ·

this point, the successful establishment of new immigrants is exactly balanced by

the extinction of other species.
0
The theory of island biogoegraphy states that the equilibrium number of
species on an island is determined by rates of immigration and extinction. What
happens then if we perturb these rates? Suppose that a storm from the mainland FIGURE 14. 1 ~
blows dozens of new bird species to an island. Following this event, the number of MacArthur and
species on the island increases. Because there are now greater numbers of species different sizes
on the island, the rate of extinction will increase and the rate of successful immi- SF= Small Far
gration will decrease. The species numbers will then decline because there will be extinction ev er
more extinctions than immigrations. Over time the number of species will move depending on
back down to the original equilibrium size S. Similarly, imagine if some catastro- mainland and
phe causes massive and sudden extinctions on the island. Then the number of
species will decrease. Now that the number of species is low, competition pres-
sure is relaxed, the rate of immigration will increase, and the rate of extinction
 ISLAND BIOGEOGRAPHY 435

'geography? The theory is

ess of most islands is in c.
1d extinction. Immigratio::.
species to islands. Extinc- c
0
! at the island by dispersa:. t5c
by the continuous process ·~
Q)
ls and some of the old e~ 0
ed by further new arrivals. c
0
Assuming that no environ- ~Ol
and will only vary slightly 'E:
: arriving and older species .£
composition of the islan 0
Q)

ies composition is called ca

cc

biogeography is to view i
e number of species along
ttion along the vertical axis
vary from 0 to P, where P Species richness
1 which the island is colo-
tte of immigration by colo- c
0
'n of species on the island. t5
.~
of species on the island x
Q)
rs of species on the island . 0
e for colonization. In other c
0
Jccupied niche space there ~Ol
izing species face stiff com- 'E:
Jf extinction increases with .£
0
the more species there are. Q)

ccurring to some species. If ca

cc
f resources, the population
mitations and species with T

m. The equilibrium number Q;

>
0
,ur graph where the rates of E
1ted as Son the graph. At ~
rants is exactly balanced by
"'I~ ! ~I p
the equilibrium number of 0 S SF S LF S SN SLN

ration and extinction. What Species richness

a storm from the mainland
ng this event, the number of
'greater numbers of species
the rate of successful immi-
ecline because there will be
umber of species will move
ly, imagine if some catastro-
sland. Then the number of
es is low, competition pres-
!, and the rate of extinction
FIGURE 14.12 A graphical representation of the island theory of biogeog raphy (after
MacArthu r and Wilson, 1972). S equals the equilbrium species number on islands of
different sizes and degrees of isolation (LN = Large Near, LF = Large Far, SN =Small Near,
SF= Sm all Far). S' and S" are nonequi librium species numbers following an extreme
extincti on event (S') or colonization event (S"). Tis the turnover rate on different islands
depending on island size and isolation. Pis the total number of species available from the
main land and is equal to the species richness of the mainland.
436 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

will decrease. Over time, the number of species will increase back to the equili turnover that
rium point S. richness and ·
So far, the theory as we have discussed it explains how biodiversity is main- SLF- SsN > S
tained at a stable size on islands. How does the theory help explain why small iso- have shown ;
lated islands have fewer species than large islands close to the mainland? AgaiL.. islands locate
the best way to consider this question is to look at a graph (Fig. 14.12). MacArthur a
According to MacArthur and Wilson, the size of the island strongly affecc.: A more
rates of extinction. Islands that are small have fewer resources and less hab i t ~: prediction th<
diversity. Therefore, small islands cannot support either large populations o:- to island size
large numbers of species. The species that are on these islands must live in clo_" MacArthur a
proximity, share limited habitat, and experience stiff intraspecific and interspe- much influen
cific competition for resources. The intraspecific competition keeps populatio;::. located in th•
sizes low. The small size of the islands and the small size of the plant and animc... eruption dest
populations mean the species are prone to random extinction events due rc tung,Anak K
events such as fires, hurricanes, or flooding. The small resource base and sma: of any life im
populations also means that extinction due to competitive exclusion is mor" plants and an
likely on small islands. km away fro n
The proximity of islands to the mainland largely influences rates of immi- the Krakatoa
gration. Islands that are near the mainland are more likely to receive colonis· - nizing specie1
by jump dispersal than are islands that are far away from the mainland. Th" Islands. Cem
model predicts that the equilibrium number of species is controlled by isla c 1932-1934 sh
size and island isolation as follows. Islands that are small (S) and far (F) fror::. then stabilize
the mainland will experience the highest rates of extinction and lowest rates o: increase signi
immigration and thus will have the lowest number of species (Ssp) . Islands tha: on the island
are large (L) and near (N) the mainland will experience the lowest rates o: species went
extinction and the highest rates of immigration and will have the highest num- species turno·
ber of species (SLN) · Islands that are large and far or small and near the mai - island biogeo
land will have intermediate numbers of species (SLF and SsN)· Thus, the Some o
relationship between species numbers and island geography can be summa- ducted in atte
rized as: and island si
These tests c:
and animal c
experimental
According to island biogeography theory, equilibrium species numbers ar made by islar
determined by the impact of island size and isolation on rates of extinction an · Some o
immigration. If a small island near the mainland has higher rates of extinctio:: regarding tur
and immigration than a large island that is far away from the mainland, then th
smaller island must experience higher rates of species turnover (T). Becau
species are arriving and going extinct at a relatively high rate on the small islan
close to the mainland, the species composition must be changing relatively ra -
idly (TsN)· The inverse is true for a large distant island. It will have a low rate of TABLE 14.3
turnover (TLF) because extinctions will be rarer and immigration events fewer.
Thus, according to the theory of island biogeography, turnover rates are relate
to island size and island isolation in the following manner:
Year

1908
1919-1921
The theory of island biogeography is especially powerful because it not only
1932- 1934
offers a simple explanation for why species diversity varies according to island
size and isolation, but it also provides a set of hypotheses on species richness and Source: Ma cArth

:rease back to the equili 0-
how biodiversity ism ~­
elp explain why small is0-
~to the mainland? A ga:-
tph (Fig. 14.12).
the island strongly affe c
·esources and less h a b i~·
her large populations .:·:
islands must live in clos.::
ntraspecific and intersr=-
>etition keeps populatic-
:e of the plant and an~
extinction events due : -
resource base and sm::.
etitive exclusion is mo::-::
influences rates of im::::...-
ikely to receive colon:.S:.
from the mainland . T.::. ::
~s is controlled by isl2..:._
nall (S) and far (F) frc-
tction and lowest rat es
;pecies (Ssp). Islands tt..:
ence the lowest rates -
ill have the highest me -
small and near the mc_:_::-
)LF and SsN)· Thus. ::.=__
ography can be surnc::.:.
-ium species numbers c..:-::
m rates of extinction <:::..:.
1igher rates of extin c t~.:=
m the mainland, then ~::
:s turnover (T). BecaG::
h rate on the small isle..:..:.
~ changing relatively rc.;:-
It will have a low rat
nmigration events fe\\ ::-
urnover rates are rel2:::_
er:
.verful because it not o=-
·aries according to isl- -
~s on species richness ~ -
ISLAND BIOGEOGRAPHY 437
turnover that can be used to test the theory. The first hypothesis is that species
richness and island size and isolation are related in the following manner; SLN >
SLF - SsN > Ssp. A s we have noted, many studies of islands and islandlike habitat
have shown that small isolated islands have lower species richness than large
islands located close to the mainland. Indeed, it was this observation that first led
MacArthur and Wilson to develop the theory.
A more crucial test of the island biogeography theory is presented by the
prediction that species turnover is a continuous process on islands, and is related
to island size and isolation in the following manner TsN > TLN - Tsp > TLF·
MacArthur and Wilson 's ideas on continuous species turnover on islands were
much influenced by evidence from the islands of Krakatoa. The islands are
located in the Malaysian Archipelago and were created when a huge volcanic
eruption destroyed the original island of Krakatoa in 1883. The islands of Ser-
tung,Anak Krakatoa , Panjang, and Rakata that were created were almost devoid
of any life immediately after the eruption. The islands were quickly colonized by
plants and animals from the large islands of Sumatra and Java, which lie about 40
km away from the Krakatoa group. Smaller islands that lie between Sumatra and
the Krakatoa group also provided colonists or acted as stepping stones for colo-
nizing species. By 1935 tropical rainforest had been established on the Krakatoa
Islands. Censuses of bird populations carried out in 1908, 1991-1921, and
1932-1934 showed that bird species richness increased quickly until 1921 and
then stabilized somewhat (Table 14.3). Although bird species richness did not
increase significantly after 1921, there were turnovers in the bird species present
on the islands. After 1921 some new colonists became established, and other
species went extinct. The data from Krakatoa regarding seemingly continuous
species turnover on the islands were central to MacArthur and Wilson 's theory of
island biogeography.
Some of the most interesting field studies in biogeography have been con-
ducted in attempts to test the hypothetical relationship between species turnover
and island size and isolation presented in the theory of island biogeography.
These tests can be broadly divided into two types: (1) the examination of plant
and animal census data from unmanipulated environments; and (2) the direct
experimental manipulation of island ecosystems to observe if the predictions
made by island biogeography theory hold true.
Some of the earliest attempts to test the island biogeographic predictions
regarding turnover rates were conducted by Jared Diamond and his colleagues.
TABLE 14.3 Changes in Freshwater and Terrestrial Bird Species on
Two Islands of the Krakatoa Group (Rakata and Sertung)
Following the 1883 Eruption
Year No. of Species Found No. of New Species No. of Extinctions
1908 14 14
1919-1921 60 48 2
1932-1934 64 11 7
Source: MacArthur and Wilson, 1967.
438 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

Their analyses were based on bird census data from the Channel Islands of C - be different,
ifornia, small islands near New Guinea, and the Farnes Islands of Britain. In t :: all of the isla
case of the Channel Islands, land bird census data from 1917 were compar ·.:: Such small r
with census data from 1968. The census data suggest that a number of specie> large degree
disappeared from the islands during the twentieth century, while other n of the small
species colonized. Consistent with island biogeography theory, the total nu - ment and p<
ber of bird species on the different islands did not change greatly between 19> biogeograph
and 1968. However, the exact species of birds that were present on each islar:.::
had changed significantly. On some islands, 62% of the land bird species fou;:;.::
in the 1960s were different from the species found in 1917. In general, Jar:§::: The Theo
islands had lower rates of turnover than smaller islands. For example, tt.::
largest island, Santa Cruz, had a turnover rate of 17%, while the tiny island : In the deca<
Santa Barbara had a 62% turnover rate. Interestingly, turnover rates did spawned a I
appear to be strongly influenced by distance from the North American con·- achieved par
nent. However, all of the islands are relatively close to the mainland, and fc- vation of enc
birds the different distances involved may not be great enough to be a signiS- raised man;
cant barrier to dispersal. More detailed analysis of year-by-year species couc:..:: attempts to 1
on the Channel Islands and the Farnes Islands also provided evidence of coi:- nor universa
tinuous turnover of species. The annual species turnover rates ranged from : ory has, how.
species to over 20 species per year. Similar evidence of continuous sp eci~ factors that <
turnover was reported from other island studies, including an examination ~ initial posi ti'
birds on the island of Mona near Puerto Rico. the addition.
A classic experimental test of island biogeography theory was conduct - generate. As
in the 1960s by Edward Wilson and his student, Daniel Simberloff. They co;::- can be imprc
ducted the experiment by manipulating the environment of small islands in tb"'
Florida Keys. The islands are located in shallow waters and are created by tb:: Classic Te:
growth of salt-tolerant mangrove trees (Rhizophora mangle). The four islan -- that differen
they studied are all very small in size, ranging from 75 to 250 sq min area and arE of species tu
located about 0.002 to 1.2 km from the mainland. Since all the islands are sm a~ early studies
size is not an important variable in the experiment, but distance from the main- ducted by I
land is. The islands support terrestrial arthropod faunas that are comprised of , However, ot
to 40 species of insects and arachnids. The mainland species pool is over 10 , by humans r
species of arthropods. Simberloff and Wilson used the insecticide methyl bro-- bird species 1
mide to exterminate all the arthropods in the islands. They then conducted care- the Californi
ful repeated censuses of the islands' arthropod fauna for over two years. peared from
census allowed them to document the return of the arthropods to the islands predators sw
and to determine whether arthropod species diversity was consistent with t h~ aetus), and p
patterns predicted by island biogeography theory. Consistent with island bio-- tims of pesti
geography theory, they found that species turnover was a relatively constan: starlings (Stz
process on all of the islands. They also found that the lowest rates of immigratior: recently intn
and turnover occurred on the most distant island. The highest species richness bird fauna o
and the highest rates of species turnover and immigration were found on th has been am
island closest to the mainland. non breeding
After less than a year, the species richness of the islands had returned to its on the island
pre-experiment levels. This result suggests that island biogeographic processes of (Aphelocom,
immigration and extinction had returned the islands to their equilibrium state o · no evidence
species richness. However, only about 19 to 40% of the arthropod species on the been caused
islands before the experimental defaunation were present in the new fauna of the the island of
islands. Why would the pre-experiment and post-experiment species composition caused by go
 ISLAND BIOGEOGRAPHY 439

rmel Islands of C a ~ ­ be different, and is this finding consistent with island biogeography theory? Since
js of Britain. In the all of the islands are small, they can only support small populations of arthropods.
'17 were compare.: Such small populations are prone to random extinction events. Thus, there is a
number of specie;; large degree of chance determining exactly which species will be present on one
y, while other ne of the small islands at a given time. So, the difference between the pre-experi-
ory, the total nun - ment and post-experiment species composition is not inconsistent with island
eatly between 19 biogeography theory.
sent on each islan.:.
bird species fa un.:
7. In general, l arg~
The Theory of Island Biogeography Today
For example. tt=
e the tiny island c: In the decades since its introduction, the theory of island biogeography has
tover rates did n0: spawned a large amount of research and been applied in many areas. It has
h American con::- achieved particular attention in regard to its potential application for the conser-
mainland, and fc ~ vation of endangered species. This intensive research on island biogeography has
Ligh to be a signi::- raised many questions about the assumptions of the theory and previous
rear species co ur. :.> attempts to test it. At present, the theory has been neither universally accepted
d evidence of co;::- nor universally rejected. The research prompted by MacArthur and Wilson's the-
ttes ranged from : ory has, however, led to a much more extensive and deeper understanding of the
:ontinuous specie-5 factors that affect species richness on islands. Let's look at some areas where its
an examination c: initial positive tests of the theory have been criticized and then examine some of
the additional insights that study of island biogeography theory has helped to
:)ry was conducte.:. generate. As we shall see, some of these findings suggest that the original model
tberloff. They co::.- can be improved.
;mall islands in
are created by t ::~ Classic Tests Revisited At the heart of island biogeography is the premise
). The four isl a - ~­ that differences in immigration and extinction rates lead to a continuous process
l m in area and ~~ of species turnover that maintains species diversity at an equilibrium size. The
=islands are sm · early studies of bird species turnover of islands off California and England con-
tee from the mai::.- ducted by Diamond and his colleagues appeared to support this contention.
re comprised of ~ However, other scientists have pointed out that environmental changes caused
pool is over 1():) by humans may have been largely responsible for the differences between the
ticide methyl br.:-- bird species found in early census studies and the species found more recently on
m conducted c ~=- the Californian and English islands. For example, some of the birds that disap-
1er two years. T::.:: peared from the Californian islands between the different census periods were
·ods to the islan.:... predators such as the bald eagle (Haliaeetus leucocephlus), osprey (Pandion hali-
onsistent with t!:: :: aetus), and peregrine falcon (Falco peregrinus). These species were likely the vic-
tt with island bic- tims of pesticide poisoning. Some of the new immigrants were species such as
·elatively constc..::.- starlings (Sturnus vulgaris) and house sparrows (Passer domesicus) that were
tes of immigrat ic:: recently introduced to North America by humans. The most recent analysis of the
st species rich ne,_ bird fauna of the Channel Islands has shown that most of the species turnover
<ere found on t:-:. = has been among species of birds from the continental North America which are
nonbreeding visitors or species that maintain low, unstable breeding populations
h.ad returned to ::_ on the islands. Endemic island species and subspecies, such as the island scrub jay
aphic processes cc (Aphelocoma insularis), have had relatively stable populations and have shown
1uilibrium state c no evidence of turnover. Some changes in bird species composition may have
Jod species on r:: ~ been caused by changing land cover on islands. For example, the bird fauna on
e new fauna of r::::- the island of Mona near Puerto Rico was likely impacted by vegetation changes
'ecies composi tic- caused by goat herds that were introduced there. Finally, studies of several island
440 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

systems, such as research on the avifauna of islands off Mexico in the Sea o: Target Are
Cortez and in the Pacific Ocean, have been unable to detect clear evidence o: gration rates
species turnover as predicted by the MacArthur and Wilson model. land. The mo
The experimental defaunation of the Florida mangroves by Simberloff an · to disperse t,
Wilson yielded results consistent with island biogeography theory. However, the size will also
turnover of species on these small islands was so rapid that reliable curves for get for dispe
immigration and extinction could simply not be calculated and compared witt immigration
the hypothetical immigration and extinction curves of MacArthur and Wilson. Ir; that travel bt
addition, subsequent work has shown that many of the arthropods collected oc crossing ban
the islands represent highly transitory species. Individuals of these species c smaller islan
move rapidly between sites even on the mainland. The appearance and disap- snow-covere'
pearance of these species may not represent true immigration and extinctioc. more likely 1
events. Later work by Simberloff led him to conclude that the rates of extinctio versed the ic
islands in Fir
suggested by his original study were overestimates of the true rates.
dispersers su
cept waterbc
Small Island Effect Island biogeography theory assumes that species rich-
target. Studi1
ness will decrease with decreasing island size in a continuous manner. The smalles:
richness of t
island should have a smaller number of species than the next largest island and so
islands. Thus.
on. However, MacArthur and Wilson themselves suggested that this relationshi
reflect highe1
might not hold on very small islands. Indeed, it has been shown in several cases tha:
if only the smallest islands of island groups are studied, there is often no relation- Biological
ship between island size and species richness. The reason for this may be that thes general as pc
islands are so small that none of them can support large enough population sizes to essentially e'
produce significantly decreased extinction rates. It may also be the case that a important di
these small sizes differences in habitat diversity related to island area are nil. clades and gu
example, the
Rescue Effect The MacArthur-Wilson model assumes that the distance o: mainland is n
an island from the mainland affects species diversity by controlling immigra- tiles than it i~
tion rates. Immigration rates impact on extinction rates by influencing compet- likely to eros:
itive exclusion. According to the model, islands that are near the mainlan logs and othe
should have higher rates of immigration, prompting higher rates of extinction various relati
and thus higher rates of species turnover. However, more recent studies hav these animal~
shown that the proximity of an island to the mainland also may serve to that many dis
decrease extinction rates. In some studies, it has been shown that, contrary to and wind-dis
theory, islands that are near the mainland have significantly lower turnover richness for l
rates than distant islands. Examples of such studies include the species richnes logical facto r
of arthropods on isolated patches of thistles, the distributions of small mam- is not in equ
mals on isolated piles of rocky habitat, and land birds on small islands in Lake sioned by isla
Gatun, Panama. The Lake Gatun study shows that islands near the shore of the The thE
lake have very low turnover rates, islands an intermediate distance from th e that plants a
shore have the highest turnover rates, and islands located far from shore again For example
have low turnover rates. The decreased turnover rates observed in the near insect sp e ci e
islands have been attributed to a phenomenon called the rescue effect. The res- diversity. Bir
cue effect occurs when small populations of a species are rescued from the nesting or th
brink of extinction by the arrival of new immigrants of the same species. Iso- dependent 01
lated islands are less likely to receive frequent immigrants, so small populations tion and est
on these islands are more likely to go extinct. The rescue effect has been shown immigrate to
to be important in maintaining some populations of continental bird species on thus promot
the California Channel Islands. MacArthur and Wilson did not anticipate the The th
rescue effect when they produced their model. should be fo

)ff Mexico in the Sea o:
detect clear evidence o:
tlson model.
groves by Simberloff ar,.:
phy theory. However, tl::.o
j that reliable curves fo::-
ated and compared \\it:.
1acArthur and Wilson. I.=
arthropods collected o:.
Jals of these species ca:.
e appearance and disa;:-
nigration and extinctio::
.at the rates of extinctio:.
e true rates.
ssumes that species riL~­
ous manner. The smalJQ-
aext largest island and s-
ted that this relationsh.::::
wwn in several cases t!L:.-
here is often no relatio:.-
:or this may be that the5e
wugh population sizes : -
also be the case that ::.·
island area are nil.
mes that the distance c:
Jy controlling immigr::.-
by influencing compc:-
are near the mainla:..:
gher rates of extinctic:.
ore recent studies ha· ~
md also may serve :c
;hown that, contrary :-
'icantly lower turn o\·:::
1de the species richnes!
butions of small mac-
n small islands in L a,;;,~
js near the shore of :i::~
liate distance from r::::
~d far from shore age:.:=
; observed in the ne::.:
~ rescue effect. The res-
are rescued from rt::
· the same species. Isc-
ts, so small populatio:...:
effect has been sho\<-::
inental bird species c:.
did not anticipate tL:
ISLAND BIOGEOGRAPHY 441
Target Area Effect The theory of island biogeography assumes that immi-
gration rates are controlled solely by the distance of the island from the main-
land. The more isolated an island is, the less likely it is that organisms will be able
to disperse to it from the mainland. However, it has been suggested that island
size will also influence rates of immigration. Larger islands provide a bigger tar-
get for dispersing organisms. The positive relationship between island size and
immigration rates is called the target area effect. For actively dispersing animals
that travel between the mainland and islands by flying, swimming, or otherwise
crossing barriers, larger islands are more likely to be seen and colonized than
smaller islands. For example, an analysis of the travel of land mammals across
snow-covered ice in the Saint Lawrence River showed that these animals were
more likely to head toward the larger islands than smaller islands as they tra-
versed the ice. Similar results have been seen in the dispersal of shrews to small
islands in Finland. The target area effect also influences immigration by passive
dispersers such as plants. Islands with large areas are simply more likely to inter-
cept waterborne and airborne seeds because they offer more surface area as a
target. Studies along the northeastern coast of Australia show that the species
richness of the seaborne seeds is positively correlated with the beach area of
islands. Thus, the relatively high species richness observed on large islands may
reflect higher rates of immigration in addition to lower extinction rates.
Biological Factors and the Nonequilibrium Effect In order to be as
general as possible, the theory of island biogeography treats all species as being
essentially equal in terms of immigration and extinction. However, the many
important differences between different groups of organisms and even within
clades and guilds will substantially impact on immigration and extinction rates. For
example, the decrease in species richness observed as one moves away from the
mainland is much more pronounced for terrestrial mammals, amphibians, and rep-
tiles than it is for birds and bats. This is of course because flying animals are more
likely to cross water barriers and arrive at distant islands. However, by floating on
logs and other flood debris, many species of terrestrial animals have been carried to
various relatively distant islands. Thus, although it will take long periods of time,
these animals do have the potential to reach distant islands. Therefore, it is possible
that many distant islands may have achieved equilibrium species richness for flying
and wind-dispersed organisms, but may not yet have reached equilibrium species
richness for less efficiently dispersed plants and animals. This is one of many bio-
logical factors that can cause a nonequilibrium effect in which the species richness
is not in equilibrium with the processes of immigration and extinction as envi-
sioned by island biogeography theory.
The theory of island biogeography does not take into account the impact
that plants and animals might have in facilitating immigration by other species.
For example, many insects can feed on only one or two species of plants. Thus, the
insect species diversity of an island will be highly dependent on the plant species
diversity. Birds can be relatively specific about the vegetation they require for
nesting or the fruit and insects they eat. Thus, bird fauna richness can also be
dependent on certain plant species being present in order for successful immigra-
tion and establishment to occur. The inability of some species to succesfully
immigrate to an island may make it impossible for other species to immigrate and
thus promote nonequilibrium conditions.
The theory of island biogeography predicts that periods of extinction
should be followed by a rapid increase in immigration because of the relaxation
442 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

of competitive exclusion. However, it is also possible that the extinction of som ~ Historical J
species may make it impossible for other immigrants to become established. Fo:- state betweer
example, if a hurricane were to cause the extinction of most large fruit trees on :o number of h
small tropical island, the successful immigration of insects, birds, or bats th : extremely imJ
depend on those species of fruit trees for food would be precluded until a ne continents WE
population of fruit trees became established. If such disturbances were freque : about 20,000 :
enough, there might never be enough time between them for the island biota tc Such islands <
ever reach equilibrium. to the mainla
The English biogeographer Robert J. Whittaker (not to be confused wit during the la~
the American ecologist Robert H . Whittaker) and his colleagues have conductec have never b<
considerable analysis of the biota of the Krakatoa island group to study how dis- islands were c
persal and biological interactions have affected species richness there. Their date. organisms, an
extend forward in time from the 1931 records from Krakatoa that MacArthu:- With the rise ,
and Wilson considered when they formulated the theory of island biogeograph became separ
The more recent work on Krakatoa shows that the species richness of plants tha: of plants and
disperse their seeds in sea water leveled off before 1920 and has remained rela- their small p(
tively stable. The species richness of wind-dispersed plants continues to increas . rates would h
but the rate of increase slowed considerably after the 1920s. In contrast, th species richnt
species richness of animal-dispersed plants has continued to increase at a rapic from a larger
rate. Bird species richness on the Krakatoa islands is just slightly below tba: How qt
found on other nearby islands that did not suffer from the catastrophic volcani - rium with isla
eruption. However, the species richness of nonflying mammals on the Krakatoa is good evide1
islands remains far below that of nearby similar islands. Thus, after more than 10C lated mountc
years, the Krakatoa island group may be at or near equilibrium species compo i- United State~
tion for some well-dispersing groups of plants and animals, but far from equilib- cial maximun
rium species richness for poorly dispersing groups such as nonflying mammals. Guinea may
Some animals may be absent from the island because of poor dispersal capabili- during the la
ties, whereas others may be absent because their host plant species have not ye: Guinea, speci
become established on the islands. How long will it take for islands of Krakatoa species relax:
to reach equilibrium species richness for all groups of plants and animals? Whit- became isolat
taker and his colleagues predict that this will take thousands of years. the Pleistoce1
Other studies support the contention that the development of true equilib- ness. In contr
rium in species richness on islands can take very long periods time and perha s find evidence
never be achieved. A survey and comparison of the bat and nonflying mamm ~ For example,
fauna of many ancient islands located in the Pacific far from mainland areas sug- was dammed
gest that even after tens of thousands of years or more the species richness o: Canal. The ca
nonflying mammals remains far below what would be predicted as equilibrium b,- ated islands, 1
the theory of island biogeography. The dispersal of nonflying animals is such a nested on the
slow process that islands may never have a true equilibrium fauna . Given th may now be c
slow rate at which islands reach equilibrium and the relatively high frequenci es species relax<
of disturbances such as fires or hurricanes that can cause extinctions, many of islands. In
islands probably never reach a state of steady turnover rates and equilibrium small islands
species numbers as predicated by the theory of island biogeography. When a
A final biological factor we might consider in the case of nonequilibrium ens because i
island biotas is the impact of humans. As we have seen in Chapters 8 and 12, th e diversity on
arrival of humans on remote islands has generated pronounced peaks in extinc- endemic hon
tions of native fauna and introductions of new species. In some cases, the original finches on tt
species richness in terms of native plants and animals may never be truly known. process in ge
All modern island data sets pertaining to species richness are to a greater or less island biogeo
extent influenced by the impact of humans. while evoluti

at the extinction of some
' become established. Fo:-
nost large fruit trees on 2.
tsects, birds, or bats tba:
be precluded until a ne
;turbances were frequ er.:
~m for the island biota t;:
~not to be confused \\-it::
>lleagues have conducte.:
i group to study how dis-
richness there. Their d a:~
rakatoa that MacArt h··-
y of island biogeograph:
[es richness of plants tb::::
J and has remained rei:::-
Its continues to increas"=
~ 1920s. In contrast. tl::~
ed to increase at a rapl.:
just slightly below t h~­
the catastrophic volcac.L
1mmals on the Krak atc_
rhus, after more than 1)
librium species compO:->
als, but far from equilic-
1 as nonflying mamm ~
:poor dispersal cap a b~­
iant species have not ~ :'
~ for islands of Krakat c_
ants and animals? V. 1- ,. -
mds of years.
:lopment of true eq uili --
eriods time and perhc.,..
and nonflying rna
rom mainland areas S'..!~­
~ the species richn ess
:dieted as equilibrium ::-
tflying animals is suet _
brium fauna. Given L:.::
atively high frequ enCJ:-<-
;ause extinctions, me.;: r
r rates and equilib-: . -
)geography.
case of nonequili bri:_-
1 Chapters 8 and 12 . ;_:=
ounced peaks in exti.L.:-
some cases, the origi::
y never be truly k n o'~'­
s are to a greater or ::--
ISLAND BIOGEOGRAPHY 443
Historical Factors and the Nonequilibrium Effect A nonequilibrium
state between species richness, island size, and island isolation can be caused by a
number of historical factors. The recent geologic history of the island can be
extremely important in this regard. Many modern islands that are located close to
continents were actually joined to the mainland during the last glacial maximum
about 20,000 years ago. This is because sea level was about 80 m lower at that time.
Such islands are called landbridge islands. Islands that have never been connected
to the mainland are called oceanic islands. Tasmania was connected to Australia
during the last glacial maximum and is a landbridge island. The Hawaiian Islands
have never been a part of the mainland and are oceanic islands. When landbridge
islands were connected to the mainland, there was no dispersal barrier to terrestrial
organisms, and they likely possessed flora and fauna similar to that of the mainland.
With the rise of sea level between 15,000 and 6000 years ago, the landbridge islands
became separated from the mainland. When they became islands, their populations
of plants and animals became cut off from the larger mainland populations. Since
their small populations were now restricted to a limited area of land, extinction
rates would have increased and caused a decline in species richness. The decline in
species richness that occurs when a region, such as a landbridge island, is cut off
from a larger landmass or area of similar habitat is called species relaxation.
How quickly does species relaxation bring island biodiversity into equilib-
rium with island size? The answer varies from island to island. For example, there
is good evidence that some of the species richness in boreal mammals seen on iso-
lated mountains in the desert areas of the Great Basin and the southwestern
United States is a reflection of species richness that developed during the last gla-
cial maximum. It is also thought that the bird species richness on islands off New
Guinea may have developed when the islands were connected to New Guinea
during the last glacial period. In the cases of both the Great Basin and New
Guinea, species richness may still be declining very slowly due to the process of
species relaxation that began when these mountain environments and islands
became isolated at the end of the Pleistocene. In the 10,000 years since the close of
the Pleistocene, these regions may not yet have reached equilibrium species rich-
ness. In contrast to these potentially slow rates of species relaxation, we can also
find evidence of extremely fast declines in biodiversity on newly created islands.
For example, a number of very small islands were created when the Chagras River
was dammed, and Lake Gatun was created during the building of the Panama
Canal. The canal was completed in 1914, and by 1980 the largest of the newly cre-
ated islands, Barro Colorado, had lost 45 of the 108 species of land birds that had
nested on the area of the island prior to the creation of Lake Gatun. The island
may now be close to equilibrium species richness for birds. It is likely that rates of
species relaxation vary for different groups of organisms and on different groups
of islands. In general, rate of species relaxation might be expected to be fastest on
small islands because they support small, and thus extinction-prone, populations.
When applied to long-time scales, the theory of island biogeography weak-
ens because it excludes the impact of evolution and speciation in generating bio-
diversity on islands. We know from much evidence such as the diversity of
endemic honeycreepers on Hawaii or the many endemic species of Darwin's
finches on the Galapagos Islands that island speciation can be an important
process in generating island biodiversity. It might be argued that the theory of
island biogeography was designed to address time scales of years to millennia,
while evolutionary processes occur over periods of millions of years. However, not
444 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

all speciation events are so slow. Some differences in the modern biodiversity of been demon:
islands and islandlike habitats may result from differences in rates of evolution target effect,
and not from equilibrium processes of island biogeography. The over 300 endemic trolled solei)
species of cichlids found in Lake Victoria in Africa is far greater than the total fish by island siz<
diversity of comparable size lakes in the temperate zones. As Lake Victoria was target effect
completely dry during part of the last glacial maximum, much of this diversity revisions are
arose from speciation events that occurred over the past 10,000 years or so. the rescue e
We still know little about contemporary rates of evolution on islands, par- island size ar
ticularly those in the tropics, and how island species richness might relate to spe- located near
ciation rather than processes of immigration and extinction. We might speculate. rescue effect
however, that speciation rates will be highest on relatively large and isolated were control
islands such as those of Hawaii. Large islands offer a wider variety of habitats and be postulate<
available niche space that promote adaptive radiation. Larger islands can also small isolate<
support larger populations, which increases the probability of new traits arising and near the
through chance mutations and also increases the probability of species escaping extinction. T'
extinction. Isolated islands are not as likely to receive immigrant species that will rates, we nee
be capable of filling all available niches. This leaves niche space available for fill- curves to rep
ing through adaptive radiation. The large number of endemic species found on islands, and 1:
the large remote oceanic islands of Hawaii supports this contention. The im
In recent years, detailed analysis of species turnover rates on islands such effect can alE
as Jamaica, the California Channel Islands, and the Revillagigedo Islands of we can postu
Mexico, shows that the endemic species that have evolved on these islands have lowest on sm
very low rates of turnover or extinction unless they are disturbed by human grants and p
land-use changes and the introduction of alien species. In contrast, species that
have recently dispersed from the mainland areas to an island exhibit relatively
high rates of extinction and turnover. We should recognize that island species
dynamics reflect two very different groups of species. One group can be called Revised me

continental taxa because they represent species that have their main popula-
tions and centers of origin on continents. The other group can be called insular
taxa because they have their evolutionary origins on the island. The continental
taxa experience high turnover rates because they are often unable to maintain
viable population sizes on islands inasmuch as they are not well adapted to con-
ditions on the island, unable to compete with the endemic island taxa, or unable
to persist at the small population sizes dictated by the limited carrying capacity
of the island. In contrast, unless the island environment is altered physically or
biologically, the insular taxa are more resistant to extinction because they are
well adapted to the island and to maintaining themselves, even if population
sizes are low. It has been suggested that once an endemic island biota has c:
0
·.;::;
evolved, it occupies the available niche space, and this leads to the resistance of '6
"0
successful colonization by plants and animals dispersing from the continent. <ll
0
The apparent high turnover rates and seeming inability of island biota to resist Q)
Cil
colonization by new species today are most likely a result of the large changes cr:
humans have caused to island environments.

Adjustments to the Theory of Islan d Biogeography In view of the

many shortcomings that have been pointed out over the past 30 years, it is not
surprising that several revisions have been suggested for island biogeography FIGURE 14. 1:
theory. In particular, it is difficult to conclude that the flora and fauna of remote takes into aceo
islands are in a true state of equilibrium between size, isolation, and species rich- the fact t hat thE
ness of plants and animals. Leaving the equilibrium questions aside, it has also evolution .
 ISLAND BIOGEOGRAPHY 445

te modern biodiversity o:
1ces in rates of evolutio:.
,hy. The over 300 endem;.:
greater than the total fis:::
tes. As Lake Victoria we:.'
n , much of this diversi:-c
10,000 years or so.
~volution on islands, pa:--
mess might relate to sp.:-
tion. We might specula!.:
tively large and isolate.:
ler variety of habitats ar..:
. Larger islands can aJs_
ility of new traits ari s i::.~
bility of species esca p i::.~
1migrant species that \\-:...._
te space available for fi:..-
ndemic species fou nd c-
: contention.
ver rates on islands sl!.:-
:{evillagigedo Islan ds -
·ed on these islands ha' ~
ue disturbed by hu iL~
In contrast, species t l~
island exhibit relati\·c.
gnize that island spec::-
)ne group can be caL::-.:
wve their main p op ~~­
mp can be called insul- ·
e island. The continer;:
been demonstrated tha t because of factors such as the rescue effect and large
target effect, we cannot accept the assumption that immigration rates are con-
trolled solely by relative isolation or that extinction rates are controlled solely
by island size. H owever, we might try to incorporate the rescue effect and large
target effect into a slightly revised version of island biogeography theory. The
revisions are best considered in their graphical form (Fig. 14.13). In considering
the rescue effect, island biogeography theory might take into account both
island size and island isolation when predicting relative extinction rates. Islands
located near the mainland experience lower rates of extinction by virtue of the
rescue effect. Remember the original theory postulated that extinction rates
were controlled solely by island size. In a revised version of the theory, it could
be postulated that large near islands experience the lowest extinction rates and
small isolated islands suffer the highest rates of extinction. Islands that are small
and near the mainland, or large and faraway, experience intermediate rates of
extinction. To convey the relationship between island isolation and extinction
rates, we need to revise the familiar island biogeography graph to include three
curves to represent extinction rates on small far islands, small near and large far
islands, and large near islands.
The impact of island size on immigration rates due to the large target
effect can also be incorporated with a slight revision to the theory. In this case,
we can postulate that immigration rates will be highest on large near islands and
lowest on small far islands. Large near islands are close to the source of immi-
grants and present a large target. Small far islands are far from the dispersal
Revised model
c
0

,ften unable to ma in
s
~Q)
not well adapted to cc:.- "0
c
lic island taxa, or u n a~ · =· co
c c
imited carrying capa •. 0 0
:;:;
[!:! t5c
t is altered physically -- Ol
nction because the\· ~=
.E ~
lves, even if popu la:: c-
.s 0
~
Q)

tdemic island biota :._::_ c 1il

a:
eads to the resistance : 2
i5
"0
ing from the conti n.:.- co
, of island biota to res 0
Q)

.ult of the large ch ar;~ ::- 1il

a:

raphy In view ot :_:._ SF SN/LF LN

e past 30 years, it is :. -
:or island biogeogra~ .:
Jra and fauna of rerr:.:-- _
)lation, and species r:.:-
estions aside, it has 2-
Species richness
FIGURE 14.13 A revised view of the theory of island biogeography by the author that
takes into account the rescue effect on near islands, the target effect of large islands, and
the fact that the addition of species to island biota occurs both through immigration and
evolution .
446 CHAPTER 14 THE GEOGRAPHY OF BIOLOGICAL DIVERSITY

center of immigrants, and present small target. Islands that are large and far, or Themo
small and near, will have intermediate rates of immigration because they either ing the theor~
are not close to dispersal sources but offer a large target area, or they are close more such at
to dispersal centers but provide only a small target for dispersing organisms. The done on colle
inclusion of three immigration curves, for large near islands, large far and small these ideas. p,
near islands, and small far islands expands the island biogeography theory to the memory c
incorporate the impact of target size on immigration rates.
What impact does evolution have on species richness? Evolution might be
thought of as a factor that combined with immigration produces the rate of
species additions to the island. On short time scales of years to millennia, species KEY WORDS ANI
additions will mainly reflect immigration. Over longer time scales, the evolution
Biodiversity
of endemic island species will be an increasingly more important factor in deter-
Continental taxa
mining island species richness. Taking both immigration and evolution together, it Equilibrium theories of
is plausible to speculate that the large near islands will have high rates of immi- biodiversity
gration because they are near a dispersal source and moderately high rates of Equilibrium theory of islan
speciation because they offer a range of habitat and niche space for adaptive biogeography
radiation. In addition, large islands support large enough population sizes to gen- Historical theories of
erate a reasonable chance of introducing new genotypes through mutation. Small biodiversity
and isolated islands will have lower rates of immigration and low rates of specia-
tion because they offer little niche space for adaptive radiation. Because small
islands can only support small populations, the chance occurrence of new geno-
types by mutation decreases, and the chance that endemic species will go extinct REFERENCES AN
due to chance events increases. Therefore, we might predict that the rate of
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islands, lowest on small far islands, and intermediate on large far and small near Ecology 9, 95-99.
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(1996). Ecology: Individ~
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Communities, 3rd editior
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Oxford.
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(Fig. 14.13). relationship between gee
The revised model of island biogeography predicts that equilibrium species latitudinal gradient in sp<
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field studies that originally presented evidence of species turnover rates consis- Highly structured fish co
tralian deep springs. Eco1
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Brown, J. H., and Lomolino.
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Biogeography, 2nd editio
the differences in turnover rates predicted by the original theory of island bio- Associates, Sunderl a nd ,~
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 REFERENCES AND FURTHER READING 44 7

large and far, o:- The model presented here is only one of a number of approaches for refin-
:mse they eithe:- ing the theory of island biogeography. Many attempts have been made, and many
~r they are close
more such attempts will be made in the future. Also, much more work will be
~ organisms. The
done on collecting field data on the plants and animals that live on islands to test
5e far and small these ideas. Perhaps this continued research is the best compliment we can pay to
·aphy theory tc the memory of Robert MacArthur.

1lution might be
tees the rate c::
illennia, speci"" KEY WORDS AND TERMS
~s, the evolutim:
Biodiversity Insular taxa Species relaxation
factor in deter-
Continental taxa Intermediate disturbance Species richness
Ltion together. i: Species turnover
Equilibrium theories of hypothesis
1 rates of il1ll1li-
biodiversity Nested pattern of insular Stability-time hypothesis
ly high rates o: Equilibrium theory of island communities Target area effect
ce for adap ti\ ;:' biogeography Peninsula effect
ion sizes to ge;:- Historical theories of Rescue effect
mutation. S m ~ biodiversity Species evenness
rates of specie.-
. Because smc:=
;e of new geno-
s will go exti nc: REFERENCES AND FURTHER READING
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15
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from Nicaragua after Hurricane Joan. useful for the study of tropical montane
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bird communities of the Channel Islands. In alternative explanation of species coexis- a synthetic d i ~
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specifically or
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Landscape ecc
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'· 307-314. U.S. Department
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onservation Genetics: Case
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1ing plant species richness: an
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Ecology and Evolution 12,
C H APTER
BIOGEOGRAPHY AND THE
CONSERVATION CHALLENGE
When Europeans first established permanent settlements in California in the
eighteenth century, a small endemic butterfly called the bay checkerspot
(Euphydryas editha bayensis) was relatively common in grassland habitat devel-
oped on serpentine soils adjacent to the San Francisco Bay. As the city of San
Francisco and nearby communities grew and expanded during the nineteenth
and twentieth centuries, the bay checkerspot became less and less common. By
1987 the butterfly had become so rare that it was listed as a threatened species by
the Environmental Protection Agency. Things did not look completely bleak for
the bay checkerspot, however. One population of the butterflies lived on Stan-
ford University's Jasper Ridge Biological Preserve. The lands of the reserve were
maintained as natural areas by the university and were under no direct threat
from development. Yet, despite the best conservation efforts, by 1996 the butter-
fly had gone extinct at Jasper Ridge. The butterfly had also disappeared or suf-
fered population declines in many of the other remaining areas in which it was
found. We have seen in earlier chapters that human activities, such as hunting,
land-use changes, and the introduction of alien species, have caused many extinc-
tions over the past several centuries. For most of these species, no attempts were
made to conserve their habitat and save them from extinction. With regard to the
bay checkerspot, however, an effort was made to preserve a relatively large pop-
ulation at Jasper Ridge, and that effort failed. The story of the bay checkerspot
raises two important questions for biogeographers. First, why did the butterfly
species go extinct at Jasper Ridge despite habitat conservation efforts to save it?
Second, what can be done to save the remaining bay checkerspot populations at
other sites? The answers to these two questions are not important just for the
preservation of the bay checkerspot; they are also important for the survival of
literally tens of thousands of plant and animal species. Finding the answers to
these questions in order to help conserve endangered species and habitat is the
most important challenge facing biogeography in the twenty-first century.
In efforts to address issues of conservation, biogeography finds itself closely
aligned with the fields of conservation biology and landscape ecology. These
three fields overlap considerably. Conservation biology can be broadly defined as
a synthetic discipline that applies principles from biology and other sciences to
the maintenance of biodiversity. Conservation biogeography might be differenti-
ated from conservation biology by the fact that most biogeographers concentrate
specifically on how spatial distributions or temporal patterns and historical
events affect the present abundance and potential conservation of biodiversity.
Landscape ecology is concerned principally with spatial patterns of landscapes,
including the distribution of species and habitat on local to regional scales, and
451
452 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

with how these patterns are developed and maintained by nature and human the largest sol
actions. In general, the spatial and temporal scales analyzed by biogeography are African coun1
somewhat greater than those analyzed for conservation biology and landscape elephants, gir;
ecology. In the end, however, it is difficult to draw firm lines between the three and attractivt
fields. Because the work of conservation is so important, biogeographers, conser- intense conse
vation biologists, and landscape ecologists should put their effort into working sale of new iv
together rather than trying to define and accentuate differences. tions. The bm
In this chapter, we will briefly consider why the conservation of nature is so (Loxodonta a
important, and then we will look at the challenges that face us as we try to pre- are also chari!
serve endangered species and habitat around the world. We will then investigate nia, which at
the role of biogeography in developing conservation strategies to protect species intense prese1
and habitat. Finally, we will examine the challenge that is presented to conserva- set aside as
tion biogeography by the future environmental changes associated with global (Sequoia semj
climate warming. fornia. Some
tourism will gl
trees. In recer
THE VALUE OF CONSERVATION fornia have a·
visits to Sequ·
Why is the conservation of habitat and species important and worth so much over 800,000.
effort? This is a legitimate question with many different potential answers. It to the local ec
would take an entire book to do justice to this issue. Let's just summarize some of Ecologic
the possible answers which we may divide into two categories: economic and world. In thes
spiritual-ethical. positive contr
In many cases, the preservation of biodiversity and natural habitat has an and the recre<
economic value. The view that conservation is an important issue for the provi- Costanza and
sion of resources such as timber was championed by people such as the American ecosystems d
forester Gifford Pinchot at the start of the twentieth century. Pinchot was head of crops, and pel
the early forest service and an original author of its philosophy of multiple land that each yea
use on forest land. In many countries, loss of natural resources such as timber due value. That is
to habitat deterioration could exact a huge and direct economic cost to society. In 194 nations.
Canada, for example, during the 1990s the forest industry accounted for 14% of We can 1
the nation's exports, employed directly or indirectly some 800,000 people, and oping ecotour
generated about U.S. $26 billion of revenue annually. seemingly ins
Ethnobotanists and pharmacologists point out that wild plants may provide some cases, tb
many unknown and potentially life-saving drugs. For example, a compound from valuable ecos
the bark of the Pacifc yew plant (Taxus brevifolia) found in old-growth forests changes in co
along the Pacific Coast of North America can be used to produce a potent drug argued that bi
called taxol, which fights ovarian cancer. The Amazon plant, Chonodrodendron that the more
tomentosa, is traditionally used by native peoples to poison the tips of arrows. The imental manij
chemical tubocurarine from this plant is now widely used as a muscle relaxant in gest that plot~
surgical anesthesia. Today close to 100 species of plants are used to produce more resistan
pharmaceutical products. The value of such natural clinical compounds that still species divers
remain unknown to science could tally in the billions of dollars. There may be used to suppo
important medicines that we will never discover if we do not protect biodiversity. debated amon
Social scientists studying the economic value of recreation point out that In contr
ecotourism in natural areas generates large amounts of money in many parts of tion of specie~
the world. During the peak bird-watching month of May, Point Pelee National thetic enrichrr
Park in Ontario, Canada, generates over $3 million in revenue for the local econ- is a spiritual \
omy. A decline in the numbers and diversity of migrating birds that visit the park such as the p
each year would produce a large decline in visitors and revenues. Ecotourism is Although pea
GE
[ned by nature and hum a::.
alyzed by biogeography a:-e
tion biology and landscape
rm lines between the three
mt, biogeographers, cons
1t their effort into workins:
lifferences.
conservation of nature is sc
at face us as we try to pre-
"ld. We will then investiga::-
;trategies to protect sp ec ie~
1t is presented to conserYc-
lges associated with globe...
ortant and worth so muc::
erent potential answers. ::
;t's just summarize some c:
categories: economic aa.::
and natural habitat has a::
Jortant issue for the pro\-:-
;ople such as the Americc.=
;ntury. Pinchot was head c::
>hilosophy of multiple Jar..::.
:sources such as timber du:-
;conomic cost to society. I=
lstry accounted for 14 % c:
some 800,000 people, an.::.
lat wild plants may pro\ iCe
!Xample, a compound fr oc=
mnd in old-growth foresL.>
I to produce a potent drui
1 plant, Chonodrodendror:
~ison the tips of arrows. Th;e
sed as a muscle relaxant i::.
ants are used to produc_;e
inical compounds that st L:
> of dollars. There may b;
io not protect biodiversir:.
· recreation point out tha:
)f money in many parts o::
\1ay, Point Pelee Nationa.:
·evenue for the local eeoc·
ng birds that visit the par',;
ld revenues. Ecotourism i.5:
THE VALUE OF CONSERVATION 453
the largest source of foreign revenue for Kenya. The interest of tourists in visiting
African countries such as Kenya would be greatly diminished if animals such as
elephants, giraffes, and lions were no longer found in the wild there. These large
and attractive animals, called charismatic megafauna, are often the object of
intense conservation measures for this reason. In 1989 a worldwide ban on the
sale of new ivory was put into place to preserve Africa's native elephant popula-
tions. The ban was supported in part because of the importance of elephants
(Loxodonta africana) as a tourist attraction for poor countries in Africa. There
are also charismatic flora such as the giant redwood and sequoia trees of Califor-
nia, which attract tourists from around the world and are also the object of
intense preservation efforts. Large areas of land with valuable timber have been
set aside as parks and conservation areas in order to protect the redwood
(Sequoia sempervirens) and sequoia (Sequoiadendron giganteum) trees of Cali-
fornia. Some conservationists argue that the income generated by long-term
tourism will greatly exceed the income that would be generated by harvesting the
trees. In recent years, Redwood National Park and adjoining state parks in Cali-
fornia have averaged between 300,000 and 400,000 visits each year, and annual
visits to Sequoia National Park in the Sierra Nevada Mountains have averaged
over 800,000. Ecotourism to these parks has added millions of dollars each year
to the local economies.
Ecological economists have tried to place a monetary value on the natural
world. In these estimates, they weigh factors such as the value of resources, the
positive contribution natural areas make to world water resources and climate,
and the recreational value of natural ecosystems. In one such calculation, Robert
Costanza and his colleagues recently looked at the economic value that natural
ecosystems delivers in producing freshwater, regulating climate, pollinating
crops, and performing other economically important functions. They concluded
that each year the earth's natural ecosystems produce $33 trillion in economic
value. That is nearly twice the combined gross domestic product of the world's
194 nations.
We can understand economic arguments for preserving resources or devel-
oping ecotourism attractions, but why preserve small, economically valueless, and
seemingly insignificant plants or animals such as the checkerspot butterfly? In
some cases, these organisms may serve as keystone species within economically
valuable ecosystems, and their removal can cause large disruptions due to
changes in competitive interactions and trophic structure. Some ecologists have
argued that biodiversity in and of itself helps to maintain ecosystems. They argue
that the more species that are present, the more stable the system will be. Exper-
imental manipulations of grasslands by G. David Tilman and his colleagues sug-
gest that plots with high plant species diversity produce more biomass and are
more resistant to disturbances such as droughts than are plots with less plant
species diversity. The view that biodiversity imparts stability and the evidence
used to support this hypothesis are very contentious, however, and remain hotly
debated among ecologists and biogeographers.
In contrast to economic valuations, many people believe that the conserva-
tion of species and natural habitat is important because of the spiritual and aes-
thetic enrichment that nature provides. In North America, the concept that there
is a spiritual value of conserving nature owes much to nineteenth-century writers
such as the philosopher Henry David Thoreau and the naturalist John Muir.
Although people like Thoreau and Muir put an emphasis on conservation that
454 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

was divorced from monetary values, they still viewed conservation from the p :--
spective of what nature could provide to people rather than in any sense th - 80
humans have an ethical responsibility to preserve nature. In the twentieth ce;:-
tury, the wildlife ecologist, Aldo Leopold, espoused the concept that hum ar;s 70
en
have an ethical responsibility to conserve and preserve the natural environmen- Q)

He articulated his views in an influential book called A Sand County Alman _ -~ 60

Q_
en
(1949). Leopold called this responsibility to nature the land ethic. He argued thz: '0
human communities were part of a greater community that included the nature... ~ 50
Q)

world. We should extend the same ethical treatment to nature that we exten.: co
~ 40
within human communities. Leopold's views were fundamental to the develo :5
ment of the modern environmental movement. ~c 30
Today, the basic philosophy that humans have an ethical responsibility to tb:E Q)
f::
natural world is being further developed by thinkers such as Robert Cahn \\-b cf. 20
argue that progress within society depends on humans developing a broad en\i -
ronmental ethic with regard to nature. In this vein, the cofounder of the Socie,_. 10
for Conservation Biology, Michael Soule, has articulated four postulates regardin§:
0
the ethical value of nature: (1) Diversity of organisms is good; (2) complexity i;:
ecosystems is good; (3) natural evolutionary development is good; and (4) biolo=-
ical diversity should be valued for and protected for itself regardless of utilitari -
values. We might summarize the spiritual-ethical view by simply saying that \\-;
should work to preserve other species and their natural habitat, not because o: § 50
economic values, but because it is the right thing to do. In doing so, we become 2. ii
ct!
.s:::
better people and create a better world for humans and all other life. 0)
c
-~ 40
~
(.)
(.)

ENDANGERED AND THREATENED SPECIES 0

en
Q)
Ti
Q)
30
In earlier chapters, we discussed some very specific definitions for the terms loca. Q_
en
species extinction and global species extinction. What exactly do we mean wbe '0
Q)
c
we use terms such as threatened species or endangered species? The governm en ~ Q)
co 20
of the United States tackled this question when an act of Congress was passed tC' ~
:5
protect species that were in danger of extinction. The Endangered Species Act o: 0
1973 defines an endangered species as a species or subspecies that is at risk o::" cQ)
10
extinction throughout all or a portion of its range. From a biogeographical per- f::
Q)
0...
spective, we might define an endangered species as a species that is no longer su::-
ficiently abundant and reproducing to ensure its survival. Species can 0
endangered at both the local and global level. A threatened species is a speci ~
that is likely to be endangered in the foreseeable future. It takes much research to FIGURE 15.1
demonstrate that a species is endangered or threatened. In general, conservatior.. bird spe cies. Th
biologists consider the total population size, rate of mortality, reproductive rate. and birds. The t
rate of overall population decline, and frequency of disturbances that produc from North AmE
significant population declines. Species that have too small a population size anc Center, 1992).
too low a reproductive rate to offset normal mortality and the impact of popula-
tion losses caused by disturbance are threatened or endangered.
The human activities that are endangering plant and animal species fall into
and 60 % of tt
seven broad categories: habitat destruction; hunting, introduction of alien specie
condition to
international trade in items such as ivory or animal and plant parts used in tradi-
responsi ble fo
tional medicines, drainage of wetlands, pollution, and the incidental destruction
Tropical rai nfc
or removal of species. Habitat destruction is by far the leading cause of species
rate of ab out C
endangerment (Fig. 15.1). Over 70% of the threatened or endangered mammals
are from tropi
E
ENDANGERED AND THREATENED SPECIES 455

onservation from the per-

er than in any sense tha: 80
ure. In the twentieth cer;-
the concept that huma115 70
: the natural environmen: "'
Q)
• Mammals
A Sand County Almanac -~ 60
Cl. Hu nting Birds
land ethic. He argued tha:
-g"'c 50
' that included the nat ure__ Q)

to nature that we exter;c cr;

~ 40
damental to the develo .;::;
~c 30
:thical responsibility to tht' Q) Introduced
~
:uch as Robert Cahn \\ h.:- rf. 20
developing a broad en\-:- International Wetland
10 trade drainage Pollution
cofounder of the Socie-
1four postulates regard i;::~
0
is good; (2) complexity i.:.
Threat type
ntis good; and (4) biol o~­
~lf regardless of utilitaria.::.
by simply saying that '-'=
al habitat, not because c: cr;
15 50
In doing so, we become .:: ctl
.<::
all other life. Ol Oceanic
c Islands
• Mammals
-~ 40
::::>
(..)
Birds
(..)
0

"'
Q)
-~ 30
nitions for the terms loc. Cl.

xactly do we mean \\ h== "'

"0
Q) Seasonal
c woodlands
species? The governme=· 2 20
ctl Fresh
>f Congress was passec :- ~ water Mountains
.;::;
ndangered Species Act.::: 0
bspecies that is at risk c: cQ)
10
m a biogeographical pe:- ~
Q)

~cies that is no longer s~­ 0...

0
urvival. Species can -:::--::
tened species is a spec:- Habitat type
It takes much research: FIGURE 15.1 The major causes of threat and endangerment to global mammal and
. In general, conserva tic-::- bird species. The habitat distributions of threatened and endangered species of mammals
rtality, reproductive ra::: and birds. The bird species represent global totals, while the mammals are based on totals
isturbances that prodtc::: from North America, South America, and Australia (after World Conservation Monitoring
tall a population size a.::.~ Center, 1992).
nd the impact of po p~­
angered.
td animal species fa ll i;:::-
and 60% of the bird species that are endangered worldwide owe their present
oduction of alien spec:-.:
condition to habitat destruction. In the United States habitat destruction is
plant parts used in tra;:.-
responsible for about 50% of bird, mammal, and plant species endangerment.
he incidental destructi.::-
Tropical rainforests are being cleared throughout the world at a global average
leading cause of specc:--
rate of about 0.9% per year, so it is not surprising that many endangered species
or endangered marnm.::._,
are from tropical forest regions. About 45% of the world's threatened bird and
456 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

mammal species are found in tropical forest habitat. Tropical rainforests and species extin
deciduous forest are crucial for the wintering of many birds that breed in the high destruction a
latitudes. For example, many species of flycatchers (Empidonax), which are com- rate of extin1
mon during summer months in the United States and Canada, spend their win- larger than o
ters in southern Mexico, Central America, and South America. The loss of habitat Habita1
and species diversity in the tropics will undoubtedly cause the extinction of some fragmentatio
of the bird species that breed in the temperate latitudes and winter in the tropics. ance of fore~
Systematic bird surveys of the North American avifauna show a general decline the eastern 1
over the past 30 years in the number of many migrant songbirds that winter in (Fig. 15.2). 1
Central and South America. smaller meta
As we have seen in previous chapters, many endemic species of birds may to remain vi:
be found on islands such as Hawaii and the Galapagos Islands. Because of the tion of habi
high number of endemic island bird species, the destruction of oceanic island forests have
habitat is just as significant a cause of bird endangerment as is the destruction of America mu
the rainforest. From 1600 to the mid-twentieth century, the number of animal of varying si:
such as road:
restrict the rr
Prehistoric natural vegetation -1900 Vegetation A gooc
serve it is pn
the most end
on the conti1

-1980 Vegetation

Forest
• Urban
Agricultural/other
-Main roads

FIGURE 15.3
America , the C.
FIGURE 15.2 Habitat loss and fragmentation caused by the clearance of land for type of char isn-
agriculture and urbanization in southern Ontario (after MacDonald, 1987). ex situ and in s

Tropical rainforests an ·
rds that breed in the hig
1idonax), which are com-
::::anada, spend their win-
lerica. The loss of habita:
se the extinction of some
and winter in the trop i~~
a show a general declin e
songbirds that winter i::.
mic species of birds rn a:.
; Islands. Because of th :-
uction of oceanic islan.:
1t as is the destruction o:
y, the number of anum:.:
·allother
js
learance of land for
ld, 1987).
ENDANGERED AND THREATENED SPECIES 45 7
species extinctions on islands was higher than on continental areas. As habitat
destruction and the introduction of alien species have increased worldwide, the
rate of extinctions on continental areas has increased to the point that it is now
larger than on islands.
Habitat destruction leads to both a decline in the total area of habitat and a
fragmentation of remaining habitat into geographically separated patches. Clear-
ance of forest in the nineteenth and twentieth centuries to develop farmland in
the eastern United States and Canada is a good example of such fragmentation
(Fig. 15.2). The fragmented patches separate once continuous populations into
smaller metapopulations. In many cases, the remaining populations are too small
to remain viable, and they go extinct in the isolated patches. Similar fragmenta-
tion of habitat has occurred throughout the world. In recent times, tropical
forests have become increasingly fragmented, so that in areas such as Central
America much of the rainforest and dry tropical forest exists as isolated patches
of varying sizes (see Chapter 6). Even small corridors of human modified land,
such as roads and powerline cuts, can be important forms of fragmentation that
restrict the movement of many plants and small animal species.
A good example of an endangered species and the efforts made to pre-
serve it is provided by the California condor (Gymnogyps californianus), one of
the most endangered birds in North America (Fig. 15.3). It is also the largest bird
on the continent, and it can obtain a wingspan of 3 m. The condors require the
FIGURE 15.3 The largest and one of the most endangered bird species in North
America, the California condor (Gymnogyps californianus). This huge bird represents the
type of charismatic megafauna that has been the object of an intensive combination of
ex situ and in situ conservation programs.
458 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

carcasses of large animals to feed upon and the protection of rocky ledges for no nearby p
nest building. The females only breed every second year and produce one egg. <: population f
reproduction rate that is very low. The settlement of southern California b,- on Vancouv(
Europeans led to the loss of habitat for condor feeding and the introduction o: gered speciE
damaging pollutants such as DDT. By 1985 only five condors remained alive iL wild. Today,
the wild. The threat of extinction to this endangered species was so great that the tion is down
remaining wild condors were trapped in 1987 and placed in special breeding being bred i1
facilities at the Los Angeles and San Diego zoos. The condors were successfulh- to save the r
bred in captivity, and by the 1990s their offspring were being released into a spe-
cial preserve to try to develop new wild breeding populations.
The Vancouver Island marmot (Marmota vancouverensis) provides anoth e;- BIOGEOGRAPH,
good example of an endangered species and the efforts being made to preserve iL
The Vancouver Island marmot is one of the most endangered mammal species iL The study oJ
North America. Even prior to endangerment by recent human activity, the Van- of species th
couver Island marmot was not extremely abundant. It has a low reproductiY marmot sho
rate, and it lives in small, scattered populations. The recent decline of the marmo: very typical
can largely be attributed to the impact of logging on marmot habitat. Once ulation num
regional populations began to decline and go extinct, even those located in pro- collapsed. D
tected areas such as parks could not escape local extinction because there wer many sites lc
at additiona
collapse has
In the
Port
last remain1
• Active sites range. Popul
o Inactive sites than at peri
• Solitary marmots higher pote1
* Urban centers strategies an
concentrate
many cases,
the peripher
lapse are pr'
Scale red wolf (CG
t---+--+-1
0 50km (Dasy urus g,
periphery is
by human ac
and vertebr;
Lomolino ar
that left the
In total. 98 ~
periphery of
Vancouver periphery of
Island
Why dl
tend to surv
popul at i ons ~
hunting are
The periphE
regions that
FIGURE 15.4 The past historical breeding sites of the Vancouver Island marmot tions of red '
(Marmota vancouverensis) and its modern distribution (from Marmot Recovery Foundatio n the Gulf of ~
data). The population decline of the marmot has occurred in tandem with a range collapse. tains of soutl

~tion of rocky ledges for
r and produce one egg. a
· southern California b,.
~ and the introduction o:
mdors remained alive i_
cies was so great that the
1ced in special breeding
ondors were successfuU,·
leing released into a sp"'-
ations.
~rensis) provides anot b e~
BIOGEOGRAPHY AND ENDANGERED SPECIES
1eing made to preserve i:
?;ered mammal species i.;:
human activity, the Van-
has a low reproducti\·;:
nt decline of the marmo:
1 marmot habitat. One::
ren those located in pro-
:tion because there we~ ;
ver Island marmot
rmot Recovery Foun dafc-
lem with a range coll aps:o
BIOGEOGRAPHY AND ENDANGERED SPECIES 459
no nearby populations to replenish park populations during periods of natural
population fluctuation s. Today the marmot is found on only a few scattered sites
on Vancouver Island (Fig. 15.4). The Canadian government declared it an endan-
gered species in 1980 when there were still several hundred individuals in the
wild. Today, despite protective measures, the Vancouver Island marmot popula-
tion is down to around 100 animals. Some marmots were captured and are now
being bred in captivity. The offspring are being released into the wild in an effort
to save the noncaptive populations.
The study of historical changes in geographic distributions can provide evidence
of species that are threatened or endangered. The history of the Vancouver Island
marmot shows a linkage between population decline and range decline that is
very typical of endangered species. In the case of the marmot, not only have pop-
ulation numbers fallen precipitously, but the geographic range of the species has
collapsed. Documentary evidence shows that the marmots had once existed at
many sites located hundreds of kilometers north of their present populations and
at additional sites to the south (Fig. 15.4). The phenomenon of geographic range
collapse has been documented for most endangered animal species.
In the case of the Vancouver Island marmot, range collapse has led to the
last remaining populations surviving in the central area of the former natural
range. Population size in the central portions of a species range are often higher
than at peripheral sites, so we might assume that endangered species have a
higher potential to survive near their range centers. In selecting conservation
strategies and refuge areas to save endangered species, we might be tempted to
concentrate our efforts at the centers of the species' natural ranges. However, in
many cases, range collapse has led to the species surviving only in small areas at
the periphery of their original ranges. Good examples of this type of range col-
lapse are provided by the giant panda (Ailuropoda melanoleuca) in China, the
red wolf (Canis rufus) in the southeastern United States, and the western Quoll
(Dasyurus geoffroii) in Australia (Fig. 15.5) . This pattern of range collapse to the
periphery is extremely common. The geographic pattern of range collapse caused
by human activity have been collected for over 250 different plant, invertebrate,
and vertebrate species and have been analyzed by the biogeographers M.V.
Lomolino and R. Channel. Only 2% of these species experienced range collapses
that left the surviving populations at the center of the former geographic ranges.
In total, 98% of all the species studied had some surviving populations at the
periphery of their ranges. About 36% of the species studied survived only at the
periphery of their former ranges.
Why do species that are declining in population size and geographic range
tend to survive as geographically peripheral populations rather than as central
populations? The likely answer is that human activity such as land clearance and
hunting are often concentrated in certain areas and are less practiced in others.
The peripheral sites where species survive human endangerment are often
regions that are not heavily used by humans. For example, the remaining popula-
tions of red wolf in the United States survive in swampy and wooded areas along
the Gulf of Mexico coastline, while the giant panda survives in the rugged moun-
tains of southern China. Much of the former range of these two animals has been
460 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

this diffe
savanna ;
islands ty
species ra
pattern h
fauna. Ra
Original range coastal ar
0 Surviving range The
degree to
small pop
Giant Panda
tion than
very small
tors such
as a resul
threatene'
erred spec
direct cor:
the degret
sis of wat<
500 sq km
over 1000
Geo;
tion on p<
produce n
Mace-Lan
based on
decline, ge
small pop1
Western Quoll

TABLE 15.1 A Sur

Red Wolf Threa

FIGURE 15.5 Examples of geographic range collapses with survival on the periphery of
Observations
the former range; giant panda (Ailuropoda melanoleuca) in China, the red wolf (Canis rufus )
in the southeastern United States, and the western Ouoll (Dasyurus geoffroii) in Australia Range
(after Lomolino and Channel, 1995; Brown and Lomolino, 1998).

Population size

extensively cleared for farming or otherwise impacted by human land uses. Declining
Analysis of the history of range collapse in North American and Australian mam- Population
mals shows that many species went extinct in a east to west direction. The direc-
tion of extinction and range collapse paralleled the westward expansion of Projected decline
European settlement and land-use patterns. For example, the black-footed ferret
(Mustela nigripes) disappeared from most of the intensively farmed portions of
the Great Plains of North America and survived only along the western edge Extinction probability
near the Rocky Mountains.
There are some other interesting patterns in the geography of range col-
lapse. In North America, Eurasia, and Australia, about 75% of all range collapses
led to peripheral survivors, while in Africa only about 58% did so. The reasons for Sources: Mace and Lande, 1
 BIOGEOGRAPHY AND ENDANGERED SPECIES 461

this difference are unclear but may relate to the lack of habitat refuges for
savanna species at the periphery of the African savanna. Unlike continents,
islands typically display a pattern of central range collapse. The collapse of island
species ranges tends to occur first along the coast and then to proceed inland. This
pattern has been observed for many native species of the Hawaiian flora and
fauna. Range collapse occurs in this manner on islands because it is usually the
Original range
coastal areas that are first occupied and modified by humans.
~
D Surviving range The degree of endemism exhibited by a species can provide evidence of the
j degree to which it is endangered by human activity. As shown in earlier chapters,
small populations restricted to small geographic areas are more prone to extinc-
:? ~ tion than large populations that are widely dispersed. Species that persist only in
very small geographic areas are very prone to complete extinction because of fac-
tors such as disease or large disturbances. Thus, species that are highly endemic,
as a result either of natural factors or range collapse, are likely to be the most
threatened or endangered by human activity. Analysis of endangered and threat-
ened species of plants and animals in many geographic locations shows that a
direct correlation exists between the present geographic range of a species and
the degree to which it is threatened with global extinction. For example, an analy-
sis of waterfowl found that all species with total geographic ranges of less than
500 sq km were threatened or endangered. In contrast, no waterfowl with ranges
over 1000 sq km were considered endangered.
Georgina Mace and Russ Lande have attempted to incorporate informa-
tion on population size, rates of population decline, and geographic range to
produce more analytic definitions for species endangerment (Table 15.1). The
Mace-Lande criteria classify species as vulnerable, endangered, or critical
based on present population size, observed and predicted rates of population
decline, geographic range size, and degree of edemicity. Critical species have very
small population sizes ( <250 individuals) , high rates of observed and projected

\] TABLE 15.1 A Summary of Mace and Lande (1991) Criteria for Assessing
Threatened Species

Degree of Threat
;urvival on the periphery o'
Observations Critical Endangered Vulnerable
1a, the red wolf (Canis ru fL:>
1rus geoffroii) in Austra lia Range < 100 sq km < 5000 sq km < 20,000 sq km
1 location <5 locations < 10 locations
Popu lation size < 250 tota l <2500 <10,000
< 50 at each location < 250 at each location <1000 at each location
:d by human land us es. Declining 80% decline 50% dec line 20% decline
can and Australian mau:.- Population per-decade o r per-3 pe r-decade or per-3
per-decade or per-3
vest direction. The dire-.:- generations generations ge nerations
westward expansion c: Projected decline >20%
>25% >20%
e, the black-footed fe rr~ = per-3 years or per-1 per-5 yea rs or per-2 per-10 yea rs or per-3
ively farmed portio ns c: generations generations generations
along the western e dg~ Extinction proba bility >50% >10%
>20%
per-10 years or per-3 per-20 years or per-5 per-100 years
geography of range cc.- generations generations
5% of all range collaps::-::
% did so. The reasons :·;::~ Sources: Mace and Lande, 1991; Dobson , 1996.
462 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

TABLE 15.2

Continent

Africa
Asia
Australia
··-.Hawaii Europe
N. America
S. America
Western Ecuador
Uplands of Source: World Co
Western
Amazonia
Central
Chile

to India , Chi
FIGURE 15.6 Global biodiversity hotspots (after Myers, 1988; 1990).
many specie!
unknown to :
Today t
population declines (80% in 10 years and 25% in 3 years), and single small ge ~ and endange1
graphic ranges of less than 100 sq km. According to Mace and Lande, criti servation of
species have a 50% probability of going extinct within 10 years. A number of con- Endangered :
servationists have accepted the Mace-Lande criteria and applied them to varia - mental and r
species. One such analysis applied the Mace-Lande criteria to the study of pri- endangered s
mates and concluded that 12% of the world's primate species were in a criti · gered species
state and at high risk for extinction. on a continer
Based on the premise that endemic species with relatively small ranges an · animal speci(
small population sizes are more prone to be threatened or endangered, we caL threatened 01
see some global patterns in the distribution of potentially endangered plant an · over 8000 enc
animal species. The tropical regions and the southern hemisphere in general pos- mals. The sm<
sess larger numbers of endemic animals and plants than the higher latitudes o:- also the sma
the northern hemisphere. Brazil has about twice as many endemic amphibia . species alrea
bird, and mammal species as the United States. Globally, a number of biodiver· includes over
sity hotspots have been identified which contain very large numbers of endemi species, 22 arr
plant and animal species that are prone to threat by human activity (Fig. 15 .6). high numbe r
The hotspots represent areas where, due to the large numbers of endemic specie- Zealand has c
human activity has the potential for producing a large number of extinctions and
bird species. l
significant declines in overall global biodiversity. Not surprisingly, most of these
species. The ~
hotspots are in the tropics in regions such as equatorial South America, equato-
erred or enda
rial Africa, and southern Asia. Important extra tropical hotspots are found in cen-
the oceans ar
tral Chile, southern Africa, and southern Australia. Two hotspots, Hawaii and . .
spec1es are su
California, are located in the United States. Because of their contribution to
Now.le
world biodiversity, the world's biodiversity hotspots are receiving high priority
learned. it is
for conservation efforts.
population si;
Taken globally, different groups of organisms appear to be facing differ-
ent levels of endangerment. It is estimated that about 3.3% of the world's mam- in the San Fn
mal species and 1.5% of the bird species can be classified as endangered. Only ingly small ar
about 0.75% of the reptile species and 0.25% of the known amphibian and combination
invertebrate species are classified as endangered. Although these percentages turbance that
may sound small, it should be stressed that the extinction of 3.3% of all known of habitat rec
mammals represents hundreds of species in countries ranging from Madagascar tion has been
E BIOGEOGRAPHY AND ENDANGERED SPECIES 463

TABLE 15.2 Threatened or Endangered Species on a Continental Basis

Continent Plants Mammals Birds Reptiles Amphibians Fish

Africa 3308 688 453 89 8 49

Asia 6608 497 918 146 9 124
Australia 2024 38 39 9 3 16
Europe 2677 66 396 16 15 48
N. America 5747 145 219 88 27 277
S. America 2061 239 535 58 2 14

Source: World Conservation Monitoring Center, 1992.

3; 1990).
trs), and single small geo-
Mace and Lande, criti
0 years. A number of cou-
d applied them to varioc
iteria to the study of pr..-
species were in a critica.
~latively small ranges ar..:
d or endangered, we c<:=
lly endangered plant ar;-
~misphere in general p05-
m the higher latitudes o::-
any endemic amphibia;;.
ly, a number of biodher·
trge numbers of endern.::
uman activity (Fig. 15.
nbers of endemic speci e-s..
umber of extinctions ar..:
1rprisingly, most of th e5::
South America, equaro-
otspots are found in ce;:-
vo hotspots, Hawaii an..:
of their contribution :
e receiving high prior. .
pear to be facing diffe::--
.3% of the world 's rna,.,.-
ied as endangered. 0 _
known amphibian ac
tough these percentage-5
on of 3.3% of all kn O\\-
mging from Madaga sr ~-
to India, China, Australia, and the United States. In addition, it is likely that
many species, particularly invertebrates, are going extinct but are currently
unknown to science.
Today two international organizations are working to identify threatened
and endangered species throughout the world: the International Union for Con-
servation of Nature (IUCN) and the Convention on International Trade in
Endangered Species (CITES). The IUCN is now composed of over 600 govern-
mental and nongovernmental organizations working to identify and preserve
endangered species around the world. Estimates of the total number of endan-
gered species in the world are tabulated in the IUCN Red Books. When viewed
on a continental basis for the entire globe, the number of endangered plant and
animal species is staggering (Table 15.2). The highest total number of known
threatened or endangered species is found in Asia. On that continent there are
over 8000 endangered species of plants, fish, amphibians, reptiles, birds, and mam-
mals. The smallest number of endangered species is found in Australia, but that is
also the smallest continent and one that has lost many of its native mammal
species already. The threatened or endangered species in the United States
includes over 2262 plant species, 27 mammal species, 43 bird species, 25 reptile
species, 22 amphibian species, and 164 species of fish. Many islands also have very
high numbers of endangered plant and animal species. For example, New
Zealand has over 323 threatened or endangered plant species and 26 endangered
bird species. This represents about 10% of the native New Zealand plant and bird
species. The global total of known plant and vertebrate species that are threat-
ened or endangered on all continents and islands is well over 25,000. Species in
the oceans are not immune to the impact of human activity, and so many marine
species are similarly threatened or endangered.
Now, let's return briefly to the checkerspot butterly. Given what we have
learned, it is not surprising that the Jasper Ridge population went extinct. The
population size of the butterfly had been in decline since the arrival of Europeans
in the San Francisco Bay area. The habitat of the butterfly was becoming increas-
ingly small and fragmented. During the time of its extinction at Jasper Ridge, the
combination of a drought followed by very high rainfall served as a climatic dis-
turbance that catastrophically reduced the butterfly population size. This pattern
of habitat reduction, habitat fragmentation , population decline, and then extinc-
tion has been the local and global fate of thousands of species.
464 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

One of
BIOGEOGRAPHY AND ing sure that
CONSERVATION PLANNING genetic vari~
studies gene
We have seen that thousands of species are at risk of extinction as a direct result of tance of gen
human activity. Many more such species will undoubtedly be identified in the futur . have large g1
These species are not yet extinct, however, and efforts can be made to preserYe play high de
them. Many conservation efforts are aimed at preserving certain targeted endan- protect or br
gered species. In these cases, planning is tightly focused on the goal of preserving th gered specie
identified species. The advantage of such species-based conservation is that it pro- geographic 1
vides simplified objectives and focuses resources on those clear objectives. genetic vari<
In some cases, it is impossible to ensure the survival of a species in the wild. ent habitats.
It may be that too much habitat has been destroyed or that the remaining popu- sure that clo
lation size is too low to sustain the species any longer. In these cases, species can also leac
preservation must be done by maintaining the species in botanical gardens an Theim
zoos. In some cases, such as the California condor, the remaining wild individuals of the H awa
may even be captured and bred in captivity. These efforts, which rely on the which is an;
preservation of the species in zoos and botanical gardens, are called ex situ con- species, plan
servation. Over 800 botanical gardens and a smaller number of zoos are direct[\· to recoloniD
involved in ex situ conservation. In some cases, it is not the actual plant or animal very low lev
that is preserved in ex situ facilities. Many botanical gardens have banks of pre- the maj ority
served seeds for endangered species, while animal facilities may preserve frozen
ent. In this SJ
sperm, egg cells, and embryos. Despite the importance of ex situ conservation. i· relatives cau
cannot be regarded as the answer to the conservation of biodiversity. For one to outplanti
thing, it is simply not possible from an economic standpoint to try to preserve Because of i
huge numbers of endangered species in botanical gardens and zoos. Second, ex rapidly grO\\
situ conservation requires that scientists know that a species exists, recognize that Specie
it is endangered, and then collect it. Thousands of species, particularly inverte- ecosystem ir
brates, remain unknown to science but are endangered. Third, in many cases it is species. \Vh1
difficult, if not impossible, to generate successful reproduction in captive popula - species to e}
tions. Passenger pigeons (Ecopistes migratorius) and the Carolina parakeet which speci<
(Conurpsis carolinensis) both went extinct in North America despite efforts by based conse
zoos to maintain the species. Finally, it would be a rather dreary world if the on ly serve the sp
place we could see real biological diversity was in controlled environments such individual SJ
as botanical gardens and zoos. preserving t<
Most conservation efforts are aimed at preserving species in their natural Almost all e
environment. This approach is called in situ conservation. Attempts to save the success of b
remaining 350 peninsular bighorn sheep (Ovis canadensis cremnopbates) in south- censuses of'
ern California is an example of an in situ conservation strategy. Fences have been Today
built to keep the sheep from crossing sections of highway where they have aim is to con
frequently been killed. Areas of sheep habitat have been removed from potential gered specie
development, and access has been restricted for hunting. The United States· serve not or
attempts to save the California condor and Canada's effort to save the Vancouver known and
Island marmot are examples of hybrid programs that combine ex situ breeding based cons<:
efforts with in situ efforts to maintain a selected species in the wild. A similar endangered
effort of linking captive breeding with the release of offspring has been used to The p
reintroduce black-footed ferret to the Great Plains and the Hawaiian goose involve det
(Branta sandvicensis) and Hawaiian crow (Corvus hawaiiensis) to the wild in ures to use f
Hawaii. The ferret and goose reintroductions have been moderately successful. In protect and
the case of the Hawaiian crow, between 1993 and 2000, 24 were hatched in captiv- gauge whet
ity and released. As of 2000, only 3 of the released birds had survived.

ction as a direct result of
e identified in the future.
m be made to preserw
certain targeted endan -
he goal of preserving th
nservation is that it pr
lear objectives.
of a species in the wild.
tat the remaining popu-
In these cases, species
L botanical gardens an ·
1aining wild individuals
=>rts, which rely on the
:, are called ex situ con-
ber of zoos are direct!
~ actual plant or anima:
.ens have banks of pre-
es may preserve froze
·ex situ conservation . i:
>f biodiversity. For one
•oint to try to presen ·.:
1s and zoos. Second. e:r
es exists, recognize tha:
~s, particularly inven e-
hird, in many cases it is
:tion in captive popula-
he Carolina parakee:
!rica despite efforts
lreary world if the o
led environments su
;pecies in their natur~
. Attempts to save the
remnopbates) in soutb-
tegy. Fences have bee =
way where they h a,-~
~moved from poten tia:
g. The United States
to save the Vancou,·e:-
nbine ex situ breedim:
in the wild. A simile:
>ring has been used rc
l the Hawaiian goo~
iiensis) to the wild i=
>derately successful. L:
vere hatched in capti'"-
1 survived.
BIOGEOGRAPHY AND CONSERVATION PLANNING 465
One of the biggest challenges facing all species conservation efforts is mak-
ing sure that ex situ and ex situ strategies preserve as much as possible the natural
genetic variability of the endangered species. The field of conservation genetics
studies genetic variations within endangered species and examines the impor-
tance of genetics in conservation practices. Many species, particularly those that
have large geographic distributions and occur as separated metapopulations, dis-
play high degrees of geographic variability in their genetic composition. If we
protect or breed in captivity only a small sampling of the population of an endan-
gered species, particularly if that sample is from only one of the environments or
geographic regions in which the species lives, we may not preserve all of the
genetic variability that allows the species to survive across its range and in differ-
ent habitats. In captive breeding programs, genetic analysis is crucial to make
sure that close relatives are not bred, for this promotes low genetic diversity and
can also lead to congenital abnormalities and illness.
The importance of conservation genetics can be illustrated by the example
of the Hawaiian silversword subspecies, Agroxiphium sandwicense sandwicense,
which is a narrow endemic on Mount Mauna Kea. In order to preserve the sub-
species, plants were grown from seeds in botanical gardens and then outplanted
to recolonize wildland sites. Unfortunately, the outplanted populations displayed
very low levels of seed production. A subsequent genetic analysis showed that
the majority of the outplanted individuals descended from the same female par-
ent. In this species, the plants generally cannot fertilize themselves, and such close
relatives cannot successfully reproduce with each other. Genetic screening prior
to outplanting would have increased the success of the conservation effort.
Because of its clear importance, it is not surprising that conservation genetics is a
rapidly growing area of scientific interest.
Species-based approaches may provide little help to the other species in the
ecosystem involved and indeed may have a negative impact on some of the other
species. When we apply species-based approaches, how do we choose which
species to expend resources on and which species to ignore, or how do we know
which species we may possibly harm in our efforts? An alternative to species-
based conservation is biodiversity conservation in which efforts are made to pre-
serve the species richness of the entire ecosystem rather than focusing on one
individual species. In biodiversity management we often must choose between
preserving total biodiversity or just the biodiversity of native plants and animals.
Almost all ecosystems now contain species introduced by humans. Gauging the
success of biodiversity preservation programs is also difficult without frequent
censuses of species.
Today many efforts are focused on habitat-based conservation in which the
aim is to conserve the entire biological and physical environment in which endan-
gered species live. Habitat-based conservation has the benefit of working to pre-
serve not only those species that we can recognize as endangered, but all of the
known and unknown species in the ecosystem as well. In most cases, habitat-
based conservation is the only way to ensure the maintenance of important
endangered species and biodiversity in general.
The problem of biodiversity and habitat-based conservation programs
involves determining what management strategies to implement and what meas-
ures to use for gauging the program's success. It is impractical to try specifically to
protect and take a continuous census of all species in a protected ecosystem to
gauge whether biodiversity is being preserved. An alternative approach is to
466 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

focus on the maintenance of key selected species that, because of their resourc can help add1
requirements and role in the ecosystem, provide evidence of biodiversity, habita: aside to pres(
diversity, and healthy ecological functioning. These indicator organisms are calle · ration of that
umbrella species because it is thought that managing the ecosystem to presen· Conser'
them provides an umbrella strategy that preserves the biodiversity and habita· from the stan
integrity of the area in which they live. The northern spotted owl (Strix occiden- viable popul:
talis caurina) of the Pacific Northwest of North America is one such umbrella reserve areas
species. By preserving the spotted owl through protection of the old-growth for- lations have
est stands in which it lives, the Unites States has protected a number of other. more likely il
often less charismatic, animal and plant species with similar habitat requiremen species. The r
The continued presence of the spotted owl in the protected stands is a relatively study of griZ2
reliable indication that the ecosystem is functioning well and is providing prope by Mark Sha
habitat for the other species associated with it. species as the
Habitat has sometimes been so profoundly disturbed by human activity existence ovc
that conservation efforts are no longer a matter of preservation but of restora- his concept. l
tion. The practice of environmental restoration requires rebuilding the biological the south we~
and physical environment from damaged ecosystems, and in some cases in areas fewer were n
where the natural habitat has been completely destroyed. Environmental tion is, what
restoration should be differentiated from environmental reclamation in which a viable popul c:
severely damaged environment is modified to develop an ecosystem that is quit It takes
different from the one that existed prior to human modification. For example, sta- bance regime
bilization of slopes and planting of lawn grasses and horticultural varieties o: tat are a fo r
trees to cover up open mine tailings are examples of environmental reclamation. population si
not restoration. mammals in
Biogeography plays an important role in developing in situ conservation of Yellowstm
strategies and specific plans to save both endangered species and endangere support a mi1
habitat. In particular, biogeographical analysis can help to determine the optimal Yet field ce n ~
size, spatial configuration, and geographic location of preserves and conservation years. In othE
areas. In addition, because biogeographers also work on issues of temporal in the Natim
changes in the environment, they can help provide the information required to mammal bioc
manage conservation areas over long periods of time and to restore damaged viable popul;
habitat. In the next section, we will explore some of the ways in which biogeo- one species.
graphical approaches can be applied to in situ species conservation, biodiversity which are qu
conservation, and habitat restoration. lation size b<
extinction pr
that the resul
General Strategies for Species Conservation and
base consen·
Biodiversity Conservation
is too low. \H
In order to provide the best possible plan for the development of conservation In rece
areas, it would be ideal to know all of the physical resource requirements, demo- size have b(
graphic attributes, and biological interactions that are important to the species method olog~
we wish to conserve. In many cases, however, such detailed autecological and tion viabilit~Y
synecological information is not available, and there may not be enough time and a given area
resources to gather this information. Most of the time we cannot preserve all of PYA are 9 -c
the habitat that was naturally occupied by the species we wish to protect. Indeed, Mace and L
much of the original habitat may already be occupied by urban areas or farm- bance rates. <
land. In these circumstances, biogeographical analysis can provide some guide- conservation
lines in establishing reserve areas to maximize the chances of success in our bution of th a
conservation efforts, despite our lack of detailed ecological information and the ity are deter
limitations we might face in setting aside large areas for preserves. Biogeography woodpecker

because of their resouw
ce of biodiversity, habit at
:a tor organisms are callec
he ecosystem to preserYe
biodiversity and habita:
10tted owl (Strix occiden-
ica is one such umbreilc.
on of the old-growth for-
:cted a number of other_
ilar habitat requirement
:;ted stands is a relative!,·
1 and is providing prope~
trbed by human actlVL .
servation but of resto ra-
rebuilding the biologi
1d in some cases in areas
strayed. Environmen tC..:
I reclamation in which 2.
n ecosystem that is qu ite
ication. For example, stc.-
wrticultural varieties of
ironmental reclamat i o ~ .
·ing in situ conservatio;:;
>pecies and endangerec
o determine the optimC..:
serves and conservatio;::
on issues of tempore.:
information required rc
md to restore damagec
: ways in which biogeo-
'nservation, biodiversir:·
on and
>pment of conserva tio:::
ce requirements, dem
nportant to the specie-5
liled autecological anc
not be enough time aLe
: cannot preserve all c::
wish to protect. Indee ·
y urban areas or farc-
n provide some gui
nces of success in ol!:
:al information and tl:c
reserves. Biogeograpt:-
BIOGEOGRAPHY AND CONSERVATION PLANNING 467
can help address two general questions. What is the minimum area we should set
aside to preserve a species or set of species, and what is the best spatial configu-
ration of that reserve area?
Conservation biogeographers approach the question of the reserve area
from the standpoint that the reserve must be large enough to support a minimum
viable population size (MVPS) for the species they wish to preserve. Larger
reserve areas can support larger populations than small areas. Since large popu-
lations have a lower probability of suffering extinction, the larger the area the
more likely it is that it will support a minimum viable population size of a given
species. The minimum viable population size concept was first outlined during a
study of grizzly bears (Ursus horribilis) conducted in Yellowstone National Park
by Mark Shaffer (1981). He defined the minimum viable population size for a
species as the smallest isolated population having a 99% chance of remaining in
existence over a 1000-year period. Many conservation scientists have embraced
his concept. In one classic study focused on bighorn sheep (Ovis canadensis) in
the southwestern United States, it was found that populations of 50 sheep or
fewer were not able to persist more than 50 years. For biogeographers, the ques-
tion is, what is the minimum area of habitat required to support the minimum
viable population for a given species?
It takes many years of data collection on birth rates, death rates, and distur-
bance regimes to determine minimum viable population size and minimum habi-
tat area for most species. The difficulties in calculating minimum viable
population sizes are highlighted by the population sizes of grizzly bears and other
mammals in the national parks of the United States. Some demographic models
of Yellowstone grizzlies suggest that the area of the current park is too small to
support a minimum viable population and that the grizzlies should be in decline.
Yet field censuses have shown a slow increase in grizzly numbers over the past 30
years. In other studies, empirical data show a decline in mammal species richness
in the National Park system that is at odds with calculations of their potential
mammal biodiversity based on park area. The main problem with some minimum
viable population size estimates is that they are often using empirical data from
one species, such as the bighorn sheep, and applying the data to other species
which are quite different. In addition, models that predict minimum viable popu-
lation size based on general relationships between birth rates, death rates, and
extinction probabilities are sometimes applied without sufficient effort to verify
that the results are applicable to the species we are attempting to preserve. If we
base conservation efforts on an estimate of minimum viable population size that
is too low, we can doom the species to extinction.
In recent years, the techniques for estimating minimum viable population
size have become increasingly sophisticated and explicitly geographic. The
methodology of estimating minimum viable population size is known as popula-
tion viability analysis (PVA). PYA specifies a survival probability for a species in
a given area for a given number of years. In many cases, the target values used in
PYA are 95% probability of survival for 100 years. Following the approach of
Mace and Lande that we discussed earlier, the birth rates, death rates, distur-
bance rates, and geographic distributions of endangered species are examined by
conservation planners. In addition, a target population size and geographic distri-
bution of that target population required to meet the specified survival probabil-
ity are determined. For example, a recent PYA focused on the red-cockaded
woodpecker (Picoides borealis). Historical records show that the woodpecker
 --- - --- -----------------
468 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

had a natural range extending from central Texas and Oklahoma to the west an - Better

0
Missouri and Kentucky north all the way to the tip of Florida. Due largely to Ian ·
clearance, the range of the woodpecker has retreated southeastward, so that it is
now absent or rare in Missouri, Kentucky, Tennessee, and Maryland. Its distribu-
tion within its remaining range has become increasingly fragmented. The P -~
study found that in order for the red-cockaded woodpecker to escape extinctio;:
in the next 100 years, the birds would have to reach a population size of 500 indi-
viduals in each of 15 different geographic regions in the southeastern Unite '
States. The red-cockaded woodpecker is listed as an endangered species in th
United States, and some steps, including captive breeding and release program
0
are being taken to meet the PYA goal. PYA is still an evolving discipline tha:
must be applied on a species by species and geographic area by geographic are2. C8
basis. Thus, it remains a very time-consuming process.
Recognizing the difficulties in determining minimum viable populatioc 0
size, we might consider whether the conservation area required for preserva-
tion of a given species can be estimated using empirical biogeographic data tha:
link the body size of species to their geographic ranges. If we assume that th
oc
present population size of the species is large enough to ensure viability, we can FIGURE 15.i
areas incorpo
assume that its present geographic range represents a very conservative esti-
continuous h<
mate of the minimum area required to support a minimum viable populatio
and Will is, 19
size. If we look at data on the body size of North American land mammals an
birds relative to their geographic ranges, we can see that this is not a very pre-
cise aid but does provide some insights (see Chapter 13). Mammals have many
diversity is J-
species with small body masses that have large ranges (> 10,000 sq km). Ma ny
lated islan d~
others of these small-bodied mammals have small ranges ( < 1000 sq km) . How-
and isolated
ever, almost no large mammal species (> 10 kg) have ranges that are less th an
produce lov
at least 1000 sq km. For bird species there is not much difference in the range
island bioge
sizes for large or small species. Few bird species, however, have ranges less th an
elude that a
about 10,000 sq km.
smaller sep
The sobering message from these data is that very few species of large
together cor
mammals or of birds appear to be sustained with very small ranges. Most individ-
Diamc
ual parks and nature reserves are far smaller than the natural ranges that species
conservatior
appear to require for survival. This is not to say that many species might not be debate is be
sustainable in smaller geographic areas, particularly with intensive managemen
areas and th
efforts, but in nature few of these species are found to be restricted to such small
biodiversity
areas. At this stage, it would be fair to say that we have little unequivocal evi-
Large Or Se
dence of the true minimum sustainable population size or area requirements for
several stror
most species. It is better to err on the side of caution and to make our reserves as
habitat dive
large as possible. Unfortunately, the majority of national parks and wildlife
porate. Tim
refuges, even in the United States, are relatively small, often encompassing less
species diYeJ
than a few hundred square kilometers. Other semiprotected areas such as gov- Secane
ernment forest lands may be larger in area but are often subject to habitat distur-
area is nonl
bance by logging, mining, or grazing.
Most natural preserves and conservation areas exist today as fragments sur-
rounded by large areas of land that is used and modified by humans. Established
where 5 is S
national parks and wildlife areas are often like islands in a sea of developed and
species per
modified land. In 1975 Jared Diamond suggested that the theory of island bio-
total area of
geography may provide a general model for assessing the ability of conservation
tionship ben
areas to preserve biodiversity. According to island biogeography theory, species
 iE
Oklahoma to the west an d
'lorida. Due largely to land
;outheastward, so that it is
md Maryland. Its distribu-
gly fragmented. The PYA
ecker to escape extinctio
opulation size of 500 indi-
the southeastern Unite
!ndangered species in th
ing and release programs.
n evolving discipline th a
c area by geographic area
timum viable population
!a required for preserva-
1biogeographic data th a:
es. If we assume that the
o ensure viability, we car.
a very conservative esti-
timum viable populatio::
rican land mammals an
wt this is not a very pre-
3). Mammals have m a n ~
(> 10,000 sq km). Man ~
;es ( < 1000 sq km). H ow-
ranges that are less th ar:.
1 difference in the rang -
er, have ranges less th a~
=ry few species of large
1all ranges. Most indivic-
ttural ranges that species
my species might no t be
h intensive managemen:
! restricted to such sma1
e little unequivocal e,·i-
)r area requirements fo~
to make our reserves as
)nal parks and wildlife
Jften encompassing less
!cted areas such as go' -
mbject to habitat distur-
today as fragments su;--
by humans. Establish ·
a sea of developed anc
1e theory of island bio-
! ability of conserva tio::
ography theory, speciQ
BIOGEOGRAPHY AND CONSERVATION PLANNING 469
Better Poorer
0 00
0 ~
GO 00
0
00 000
FIGURE 15.7 Different shapes and configurations of conservation areas compared. All
areas incorporate the same amount of area . Shapes that provide the maximum amount of
continuous habitat with the lowest ratio of area to perimeter are optimal. (Based on Wilson
and Willis, 1975.)
diversity is highest on large islands close to the mainland and lowest on small iso-
lated islands. The theory holds that rates of extinction are highest on small islands
and isolated islands experience low rates of immigration. Together these factors
produce low biodiversity on small isolated islands. If these simple rules from
island biogeographic theory are applied to conservation areas, we would con-
clude that a large single conservation site would be preferable to a number of
smaller separate and isolated conservation sites even if the separate sites
together comprised the same total area as the large site (Fig. 15.7).
Diamond's conjectures about the application of island biogeography to
conservation sparked a controversy in biogeography that continues today. The
debate is between those who favor the establishment of single large conservation
areas and those who maintain that several smaller areas are better for preserving
biodiversity. This difference of opinion is known as the SLOSS Debate (Single
Large Or Several Small). Opponents of the island biogeography approach have
several strong arguments. First, a single large area may not incorporate all of the
habitat diversity that several small and strategically placed reserves might incor-
porate. Thus, a single large area might not provide the range of habitat and
species diversity that several smaller reserves might.
Second, the increase in species numbers that occurs with increasing island
area is nonlinear, having the form:
S= cAz
where S is species richness; c is a constant that is based on the overall number of
species per unit of area geographic region in which the site is located; A is the
total area of the island; and z is a constant that represents the mathematical rela-
tionship between area and species richness. It can be thought of graphically as the
 "--------====-------------
4 70 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

slope of the line that describes the relationship between area and species rich- decrease tr
ness. The value of z ranges from 0.15 to 0.40 (see Chapter 14). What this means is perimeter 1
that an increase in island area of 10 times will generally lead to an increase in example, aJ
species richness of roughly only < 2 to 4 times. Thus, since biodiversity increases increased '
in a less than 1 to 1 relationship with proportional increases in area, several forest at th
smaller reserves could have the same or greater total biodiversity than one large been a sig1
reserve. Indeed, some empirical data have shown several instances in which small stands of o
reserves have a combined total species richness that is greater than the species logging. La
richness of single large reserves of approximately the same area as the total area terns and a
of the smaller reserves. This result has been demonstrated for communities of animal spe•
arthropods, lizards, birds, and mammals. extra light<
Third, a single conservation site might be more prone to the extinction of tances of b(
species owing to a large disturbance such as fire or disease than would be the case significant '
for several separate sites. Fourth, some biogeographers argue that the biota on edges is on•
most islands, particularly oceanic islands, provides a very poor model to use in ridors such
trying to develop conservation strategies for fragments of continental habitat . the road . Ir
They point out that much of the species richness on remote islands can be pro- and animal
duced by endemic insular species that are adapted to survival on small and iso- it is necess;
lated areas of habitat. We have seen that many island species have special including ir
adaptations for limited dispersal or specialized feeding that aids their survival on The r
small islands. Continental species, particularly birds and mammals, often do not ing extincti
have such adaptations and appear to rely on relatively large ranges and popula- more diffic1
tion sizes. In short, they are not evolutionarily adapted to survival on small iso- If a species
lated fragments of habitat. through im
There is one area in which the study of island biogeography has provided a are optima
widely accepted lesson for species conservation. That lesson concerns the reduction that are pi
in biodiversity that is seen when once continuous areas of habitat are fragmented species dive
into smaller units. We can recall from Chapter 14 that the decrease in biodiversity Biage
that occurs when such new islands or fragments of habitat are created is called developme1
species relaxation. When a small habitat area is preserved, while the surrounding (GIS) a nd ~
landscape is modified, the species richness of the newly isolated fragment of pre- making sur
served habitat will likely decline over time. There are many reasons for this decline. of a region.
For some species, the populations remaining in the small preserve will be below the analysis. a 1
minimum viable population size and will go extinct due to chance factors. The frag- ment are pi
ment may not provide sufficient resources for larger animals that require large on topograJ
areas for hunting, browsing, or grazing. In very small fragments of forest, because of veget ation
edge effects, the physical environment in terms of light penetration, wind, evapora- conservatio
tion rates, and other factors may be so altered as to cause some forest-dwelling overlay spe
plants and animals to go extinct. When one species becomes extinct in the frag- butions of
ment, this might cause a trophic cascade and generate additional extinctions. In cover (Fig.
addition, if a keystone species becomes extinct, this may cause changes to competi- can be OYe
tive balances that lead to additional extinctions. Time and time again it has been areas are i
shown that the preservation of only small isolated fragments leads to a decline in Often this
the native species diversity of the fragments. This has been shown in many studies serva tion si
of tropical birds, primates, and insects in small fragments of Amazonian rainforest conserYatio
in Brazil that have been preserved experimentally following large-scale clearance gered bi rd
of surrounding forest. Thus, preserves that are too small may not provide enough conserYatio
land area and resources to support the species we wish to protect. 6693-h a H a
Aside from area, other features of spatial configuration can be important to completed
the success of conservation areas (Fig. 15.7). First, conservation area designs that with conse

en area and species rich-
er 14). What this means is
lly lead to an increase in
nee biodiversity increases
ncreases in area, several
iodiversity than one large
J instances in which small
greater than the species
tme area as the total area
ated for communities of
rone to the extinction of
;e than would be the case
. argue that the biota on
ry poor model to use in
s of continental habita t.
note islands can be pro-
lrvival on small and iso-
Jd species have special
1at aids their survival on
mammals, often do nor
1rge ranges and popula-
to survival on small iso-
!Ography has provided a
n concerns the reduction
'habitat are fragmented
decrease in biodiversitY
tat are created is called
I, while the surrounding
alated fragment of pre-
reasons for this decline.
eserve will be below th
;hance factors. The frag-
mals that require large
:nts of forest, because of
=tration, wind, evapora-
e some forest-dwellin g
nes extinct in the frag-
lditional extinctions. In
tse changes to competi-
time again it has been
tts leads to a decline in
shown in many studies
Amazonian rainforest
g large-scale clearance
1y not provide enough
otect.
m can be important to
1tion area designs tha:
BIOGEOGRAPHY AND CONSERVATION PLANNING 471
decrease the ratio of perimeter to area are better than shapes that produce a large
perimeter relative to the preserved habitat area. The edges of forest preserves, for
example, are subject to higher wind speeds than the interiors of closed forests. The
increased wind stress often causes windthrow and the destruction of old-growth
forest at the edges of the conservation areas. Wind throw due to this edge effect has
been a significant problem in portions of northern California where very small
stands of old-growth redwoods (Sequoia sempervirens) have been set aside from
logging. Large edges also allow greater penetration of light into forest floor ecosys-
tems and also permit easier invasion of the conservation area by alien plant and
animal species. The edge effect has been demonstrated to cause tree blowdowns,
extra light and wind penetration, and other physical and biological changes for dis-
tances of between 100m and 500 min North American and Australian forests. The
significant expansion of environmental changes inward to patches of forest from
edges is one reason why the fragmentation of landscapes by relatively narrow cor-
ridors such as roads can have significant impacts beyond the immediate vicinity of
the road. In addition, roads often provide corridors that allow invasive alien plant
and animal species to expand into landscapes. For many types of protected habitat,
it is necessary to have buffer zones around the core habitat so that edge effects,
including invasion by alien species, will not compromise our conservation efforts.
The rescue effect (see Chapter 14) can be extremely important in prevent-
ing extinction within a preserve. The more isolated a conservation area is, the
more difficult it is for species to disperse to it from other areas of similar habitat.
If a species goes extinct in an isolated reserve, it is less likely to be resurrected
through immigration. Reserves that are connected by corridors of similar habitat
are optimal for promoting the rescue effect. Alternatively, conservation areas
that are placed close to other similar conservation areas can help maintain
species diversity through the rescue effect (Fig. 15.7).
Biogeographic approaches to conservation are being improved by the
development of newer analytic tools such as geographic information systems
(GIS) and satellite remote sensing. In particular, GIS can be an immense help in
making sure that conservation networks adequately capture the habitat diversity
of a region. The approach to conservation planning is called gap analysis. In gap
analysis, a number of variables relating to the physical and biological environ-
ment are placed in GIS databases. The variables commonly include information
on topography, landforms, waterbodies, geology, hydrology, soils, climate, general
vegetation cover, species distributions, and the locations of current and planned
conservation lands. GIS allows maps to be made of these features, and we can
overlay species maps to find areas of high biodiversity or to compare the distri-
butions of endangered species to factors such as soil conditions or vegetation
cover (Fig. 15.8). Areas of high biodiversity or the ranges of endangered species
can be overlaid on maps of current or planned conservation areas to see if those
areas are in the right location to preserve biodiversity or endangered species.
Often this type of analysis identifies gaps in the geographic distributions of con-
servation sites where areas of high biodiversity are not protected by any existing
conservation area. Gap analysis was pioneered in Hawaii to help preserve endan-
gered bird species. The analysis showed that there were indeed major gaps in the
conservation network in Hawaii and that this resulted in the establishment of the
6693-ha Hakalau Forest National Wildlife Refuge. Gap programs are now being
completed or are ongoing in many areas, particularly in the United States, to help
with conservation planning.
4 72 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

Maps soils to tra•

Distribution of plant speci<
endangered species

Habitat I
Environme
Soils map
ing the typ<
Private land and how to
ownership map project by c
many so-ca
Conservation thousands c
area map human-mQ(
of wildland
human land
Polynesian-

s( at restoring
ate a landsc
In rna
to provide .
regimes, an<
America WI
"Gap"
.r-- Area requiring to 200 year
Conservation Land to landscape 2
preserve species and activities of
associated habitat
use patt e rn ~
FIGURE 15.8 A schematic diagram of the application of gap analysis using a geograph ic of what the
information system to overlie species ranges, environmental conditions, and land disturbance
ownership information. conserve na
change: unl<
changes and
A ri ch
In many instances, it is difficult to adequately monitor habitat conditions in
tat restorati
large conservation lands. Remote sensing, particularly from satellites, is an
theory and
extremely important biogeographical tool for conservation. Estimates of trop i-
summarized
cal forest habitat loss and fragmentation in areas such as the Amazon are
mine th e id
dependent on data from satellite imagery. Sequences of images collected over
measured. I
different years provide quantitative data on rates of habitat change. Attemp t
such as t h e~
are being made to use satellite imagery to estimate plant diversity in a number
and feed. In
of biomes. In addition, since bird diversity is often correlated with the diversity
populations
of vegetation structure, efforts are being made to estimate bird diversity on the
ulation prio
basis of satellite imagery. By using GIS, the satellite-derived data on habitat dis-
design our e
tribution can be mapped and combined with ground-based measurements to aid
populat ion
in gap analysis. The resulting spatial distributions of habitat or conservation
Restoration
areas can be analyzed using an array of specialized landscape statistics that pro-
been very i
vide measures of the degree of clumping between patches, the connectivity
such as res
between patches, the ratio of edges to perimeter of habitat patches, and the
restoring th
homogeniety of habitat within patches. All of these features are important in
Durin
conservation planning. In analyzing such landscape metrics, biogeography and
against eco
landscape ecology are closely allied. Finally, by using GIS, time series of air pho-
peoples an
tographs or satellite images can be linked with features such as topography or
regions whe
 E
Maps
>istribution of
:mgered species
)agraphic map
Soils map
>rivate land
vnership map
onservation
area map
"Gap"
,a requiring
vation Land to
te species and
:iated habitat
malysis using a geograp hic
1ditions, and land
tor habitat conditions C.
y from satellites, is a;:;
ion. Estimates of tropi-
::h as the Amazon are
f images collected 0\·e:-
tbitat change. Attemp ~
.t diversity in a numbe:-
lated with the diversi _
te bird diversity on rb.:
ved data on habitat dis-
:d measurements to ai""
abitat or conservatio1:
cape statistics that pro-
tches, the connecti \·ir:-
tbitat patches, and tb.:
tures are important i;:
rics, biogeography an--=
time series of air pho-
such as topography o:-
BIOGEOGRAPHY AND CONSERVATION PLANNING 473
soils to trace the large-scale patterns and controls of the progress of invading
plant species or other vegetation changes caused by humans.
Habitat Restoration and Conservation
Environmental restoration is often a very difficult process. It involves determin-
ing the type of ecosystem we wish to "restore", how to go about the restoration,
and how to assess its success. In addition, although we may start on an restoration
project by declaring that our goal is to restore the landscape to a "natural state,"
many so-called natural landscapes have been modified by human land use over
thousands of years and our idealized view of the natural state may actually be a
human-modified system. For example, the sparse and shrubby vegetation typical
of wildland areas in Mediterranean portions of southern Europe is the product of
human land use, as is much of the lowland forest that Europeans encountered on
Polynesian-occupied islands such as Hawaii. Should restoration efforts be aimed
at restoring this earlier human modified landscape, or should the aim be to recre-
ate a landscape that may not have existed for many thousands of years?
In many cases, conservation managers must rely on short historical records
to provide information regarding the original species composition, disturbance
regimes, and population dynamics of the habitats they wish to restore. In North
America written accounts of landscapes and ecosystems often go back only 100
to 200 years. In many instances, these historical accounts commence after the
landscape and environment have already been significantly impacted by the
activities of European settlers. Information on natural conditions or native land-
use patterns is difficult to interpret from historical records. Without a clear idea
of what the earlier environment was like, as well as what natural variability and
disturbance regimes it was subject to, it is impossible to know how to restore or
conserve natural habitat. In addition, many environments are in natural states of
change; unless we recognize this fact, our management policies may retard those
changes and artificially alter the natural environment.
A rich scientific literature has developed on the theory and practice of habi-
tat restoration and conservation. The term restoration ecology is applied to the
theory and practice of restoring ecosystems. The restoration approach can be
summarized quite clearly in a simple graph (Fig. 15.9). The first step is to deter-
mine the ideal restoration goal and how success in reaching that goal will be
measured. In one instance, the goal may be to preserve an endangered species,
such as the giant panda, by restoring the bamboo groves in which the pandas live
and feed. In this case the success of the project is determined by the size of panda
populations in the restored habitat. If we can estimate the size of the panda pop-
ulation prior to human impact, we have an ideal target to shoot for and we can
design our efforts accordingly. However, we may have to compromise on a lesser
population size that will still be viable for long-term survival of the species.
Restoration of the giant panda habitat is a project in which biogeographers have
been very involved. In many cases, the restoration goal may be more complex,
such as restoring the total biodiversity of native plant and animal species or
restoring the primary productivity of an ecosystem.
During restoration attempts, the restoration goals must often be balanced
against economic and cultural concerns. In many parts of the world, indigenous
peoples and their cultures depend on land use and resources from the same
regions where we might wish to protect endangered species or general biodiversity.
474 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

times called
Optimum goal- attempt to ga
Complete restoration

/
--- torical docurr
of what the 1
I when historic
I mining what
I
I
study desigm
Compromise goal - cisco Bay, rna
Viable population
Spanish land
topographic r
Worst case outcome - photos, and
Extinction geographic lc
Past Present Future resolution. 11
pretation , an
Time tions. Often s
FIGURE 15.9 A graphical representation of restoration aims and tradeoffs (after conditions ca
National Research Council , 1992). In this case the goal is restoration of habitat to support One ex
giant pandas. determine th'
from researcl
United State
Such problems are particularly acute in portions of Central America, South throughout P
America, Asia, and Africa. For example, to restore all of the pandas' original ized by large
habitat in China would be impossible because it would mean displacing millions understories
of people and expending huge amounts of money. In most restorations a compro- large fires th.
mise range of goals is identified. The compromise goal may not be to bring panda trees. The reg
populations completely back to natural levels, but to bring the population up to a 100 years of
level higher than it is today that will ensure the survival of the species in the wild . pine stands <
In general, a time limit is set as to when the goal should be met and different writings of e
restoration strategies are developed. and for est m
In setting restoration goals and strategies, it is essential to know what the widely scattl':
state of the ecosystem was in the past so that we can know the ideal state of the understory w
system we wish to recreate. We must also be able to recognize if long-term natu- of twen tieth-
ral trends, such as increases or decreases in panda populations, are affecting the progressive i
ecosystem. For example, there is very good evidence that the geographic range coniferous tr
and population size of the California condor have been shrinking since the end of and photogr<
the Pleistocene. European settlement and land-use changes may have simply lyzed. These
accelerated the movement of the species toward extinction. In this case, our con- quent. but lo
servation efforts would have to go beyond restoration of condor habitat if we and fire supr:
wished to save the species. This raises an ethical issue regarding whether or not kept the unc
we should intervene to save species that are on a natural road to extinction. allowed gras~
Because biogeographers are concerned with changes that occur over rela- and this prod
tively long periods of time, they have been very active in producing long environ- pines protect
mental records that can cast light on the earlier natural state of ecosystems. Becaus
Biogeographers have also been very active in developing records of the long-term enced very I<
variability and disturbance patterns of different ecosystems. The techniques that and den e un
biogeographers use in such studies include analysis of fossil pollen, fossil diatoms, large ponder
and other plant and animal remains from lake sediments, peatlands, and packrat fires that des
middens; analysis of tree-rings and fire scars from forested sites; and study of his- National Par
torical records such as written material, old maps, paintings, and photographs. The record an d
application of these techniques to aid in restoration and conservation is some- to restore th(
 BIOGEOGRAPHY AND CONSERVATION PLANNING 475

times called applied historical ecology. In most cases, applied ecology studies
attempt to gather records from a number of different sources, ranging from his-
torical documents to fossil evidence, and synthesize these into a coherent picture
of what the past environment was like. This work can be quite difficult. Even
when historical documents such as texts and maps are available, the task of deter-
mining what the past environment was like can be difficult. For example, in a
study designed to aid in the restoration of coastal marshes along the San Fran-
cisco Bay, maps and documents ranging from eighteenth- and nineteenth-century
Spanish land grant papers to nineteenth- and twentieth-century coastal surveys,
topographic maps, and aerial photographs were available. In most cases, the maps,
photos, and documents provided evidence of past conditions at different
geographic locations in the marshes at different times and at different scales of
resolution. These different sources required careful geographic reference, inter-
pretation, and synthesization to provide a meaningful overview of past condi-
tions. Often specific information regarding species presence or details on habitat
d tradeoffs (after
conditions can only be inferred from such data sources.
n of ha bitat to support
One example of the application of biogeographical research in helping to
determine the natural state of an ecosystem prior to European settlement comes
from research on ponderosa pine (Pinus ponderosa) forests in the southwestern
United States. Today extensive areas of ponderosa pine forest may be found
:ntral America, South throughout Arizona and New Mexico. In many cases, these stands are character-
f the pandas' original ized by large dominant trees that are several hundred years old and by dense
~an displacing millions understories of smaller trees and shrubs. Such stands are extremely prone to
restorations a com pro- large fires that spread upward to the canopies and destroy both young and old
. not be to bring panda trees. The region has been impacted by the grazing of sheep and cattle, as well as
the population up to a 100 years of fire suppression, and it has been unclear if the present ponderosa
the species in the wil d. pine stands are a natural vegetation type. Studies of old photographs and the
be met and different writings of early travelers suggest that prior to extensive European settlement
and forest management, many of these ponderosa pine stands were typified by
tial to know what th widely scattered pine trees that were extremely large and generally old. The
v the ideal state of the understory was dominated by grasses. Small trees and shrubs were rare. Analysis
1ize if long-term natu- of twentieth-century aerial photographs further showed that there has been a
:ions, are affecting the progressive invasion of grassy meadows throughout much of the Southwest by
the geographic range coniferous trees, including ponderosa pines. To supplement the written evidence
inking since the end of and photographic records, tree-rings and fire scars have been collected and ana-
ges may have simply lyzed. These records show that the ponderosa pine forests were prone to fre-
L. In this case, our con- quent, but low-intensity, fires until extensive sheepherding European settlement
· condor habitat if v..-e and fire suppression began in the late nineteenth century. These frequent burns
uding whether or not kept the understory of the pine stands clear of most shrubs and saplings and
>ad to extinction. allowed grasses to dominate. In some areas the fires kept trees from establishing,
; that occur over rela- and this produced grassy meadow. In other areas, the thick bark of the ponderosa
oducing long environ- pines protected the larger trees from ground fires that spread through the stands.
state of ecosystems. Because of sheep grazing and fire suppression the twentieth century experi-
cords of the long-term enced very low rates of burning, allowing the conifers to invade some meadows
s. The techniques th at and dense understories of small trees and shrubs to become established under the
pollen, fossil diatoms. large ponderosa pines. The current vegetation is now prone to very large, intense
>eatlands, and packrat fires that destroy timber and structures. The United States Forest Service and the
sites; and study of his- National Park Service have used the information provided by analysis of historical
and photographs. The records and tree-rings to institute an aggressive prescribed burn policy to attempt
~onservation is some- to restore the ponderosa pine stands and other southwestern ecosystems to their
476 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

natural state. The prescribed burn program is extremely expensive and will take documents
many decades to complete. If the natural state of the forests had been studied and often very c
taken into account in the early twentieth century, fire suppression might have At th
been limited to the protection of structures and townsites, and the costly restora- result of lo1
tion efforts that are being conducted today using prescribed burning might have of fire intol<
been avoided. Similar studies of natural fire regimes and the impact of fire sup- reintroduct
pression are being conducted by biogeographers in large portions of North Amer- Should this
ica, including many national parks in the United States and Canada. This is an Here the st•
economically important area of research. In 1999 the U.S. Department of Agricul- provided b~
ture and the U.S. Department of the Interior practiced prescribed burning and Howes Prai
other forms of fuel reduction on over 2 million acres of federal land in the United teenth cent
States. The annual budget requested for such work has grown to $380 million. nearby are<
A particularly interesting study that sought to reconstruct a pre-European tural fields.
ecosystem and examine options to restore it was conducted at the Indiana Dunes ragweed (A
National Lakeshore on Lake Michigan. The site is of special interest because it is something e
where the ecologist Henry Cowles conducted his classic studies on plant succes- park. The c
sion during the nineteenth and early twentieth centuries (see Chapter 5). The site almost 2000
is also close to Chicago and affords a large urban population the chance to see a by increasec
"natural" ecosystem. The current upland forest of the Indiana Dunes National sition from
Lakeshore is dominated by black oak (Quercus velutina) and white oak (Quercus shoreline of
alba) with open areas of wet grasslands and prairie. However, the site had been site is becon
impacted by European land-use practices, and the Park Service wished to see pines. In adc
what the natural vegetation was and if the park could be returned to that condi- ing, while tl
tion. In order to address this problem, Ken Cole of the United States Geological lished, frequ
Survey and his colleagues collected data from a number of sources, including Lakeshore c:
ground and aerial photographs, government documents, tree-rings, and fossil by Europea1
pollen records from small ponds and marshes. The ground photographs included As mi~
a systematically collected time series of photos from survey points in the park. ing a restor<
The systematic ground photos were taken regularly from 1978 to present. Aerial given to re:
photographs for the region span the period 1929 to present. excessively c
The historical ecological evidence shows that the National Lakeshore has sion, which \
indeed been significantly changed by recent human activity. The evidence from oak stands a
the survey photographs shows that there has been a steady increase of fire intol- vice is to ap
erant tree and shrub species such as black cherry (Prunus serotina) and witch and limit the
hazel (Hamamelis virginiana). The increase in these species is directly attributa- The ex
ble to fire suppression. The aerial photographic evidence shows that there has plex the issu
been a steady encroachment of oaks and other trees onto dry prairie areas. example alsc
Again, it is likely that fire suppression has allowed this expansion of woody vege- ment al cond
tation onto the prairie. cannot be cc
Documentary evidence of the vegetation of the park in the year 1834 is to return site
available from General Land Office (GLO) surveys conducted by the U.S. gov- ecosystems.
ernment. The GLO surveys provide data on the dominant tree cover encountered natural shift
in 1834. The GLO records were compared with tree surveys conducted in 1985. habit at shou
Between 1835 and 1985, there has been a significant increase in the density of oak artifici ally p
trees and a significant decrease in the density of pine trees. Today there are small
numbers of white pine (Pinus strobus) and jack pine (Pinus banksiana) in the
area. The survey records indicate that in 1835 these pines were extremely abun- The Glob<
dant and were probably the dominant tree, or codominant with oak. Other histor- As we saw i
ical documents show that the southern shores of Lake Michigan were heavily the burning
logged for pines in the mid-nineteenth century. In addition to logging, historical in global lir
 BIOGEOGRAPHY AND CONSERVATION PLANNING 4 77

expensive and will take

documents reveal th at fires set by Europeans in the nineteenth century were
sts had been studied and
often very extensive and likely contributed to the destruction of pine stands.
suppression might have
At this point, we might conclude that the shift from pine to oak was the
s, and the costly restora-
result of logging, while subsequent oak expansion onto prairies and the increase
bed burning might have
of fire intolerant plants in oak stands was the result of fire suppression. Would the
j the impact of fire sup-
reintroduction of fire lead to reestablishment of a pine-dominated vegetation?
portions of North Amer-
Should this be the restoration goal for the Indiana Dunes National Lakeshore?
and Canada. This is a
Here the story becomes more complex and requires the longer time perspective
Department of Agricul-
provided by fossil pollen and charcoal records. Such a record from a site called
prescribed burning an
Howes Prairie Marsh shows that the onset of European land use in the nine-
deralland in the Unite
teenth century produced a regional decrease in both oaks and pines. Some
::Jwn to $380 million.
nearby areas that were dominated by these trees were transformed to agricul-
'nstruct a pre-Europear.
tural fields. The pollen record shows increases in weedy field plants such as
ed at the Indiana Dun e ~
ragweed (Ambrosia) during this time. However, the pollen records also show
;ial interest because it is
something else that is very important in formulating a restoration strategy for the
studies on plant succes-
park. The decline in pines relative to oak and prairie grasses actually began
:see Chapter 5). The site
almost 2000 years ago. It appears to coincide with increasing fires, as represented
tion the chance to see
by increased amounts of charcoal in the pond sediments. The reason for the tran-
ndiana Dunes National
sition from pine to oaks and prairie likely reflects the fact that the southern
and white oak (Quercus
shoreline of Lake Michigan is rising relative to the water level of the lake and the
vever, the site had bee
site is becoming progressively drier. The dry conditions favor oaks and grass over
: Service wished to see
pines. In addition, the oaks and prairie grasses are well adapted to frequent burn-
returned to that condi-
ing, while the pines are not. Once the dry grass and oak vegetation are estab-
nited States Geologi c~
lished, frequent fires serve to exclude pines. The decrease in pines in the National
~r of sources, including
s, tree-rings, and fo ss.. Lakeshore appears to have been a long-term trend that was simply accelerated
by European logging and burning in the nineteenth century.
j photographs include
As might be anticipated from the information provided above, in formulat-
vey points in the park.
ing a restoration plant for the National Lakeshore no serious thought has been
1978 to present. Aeria:
lt. given to restoring pine densities. Not only would it be somewhat unnatural,
excessively difficult, and very expensive, but it would require intense fire suppres-
~ational Lakeshore has
sion, which would in turn cause an artificial increase in fire tolerant tree species in
rity. The evidence from
oak stands and the invasion of prairie by oaks. The course taken by the Parks Ser-
ly increase of fire intol-
vice is to apply prescribed burns to limit the densities of fire intolerant species
~ us serotina) and witct
and limit the invasion of prairies by oaks.
ies is directly attributa-
The example from the Indiana Dunes National Lakeshore shows how com-
~ shows that there has
plex the issues associated with habitat restoration and conservation can be. The
mto dry prairie areas.
example also demonstrates that without adequate long records of past environ-
>ansion of woody veg"'-
mental conditions, disturbance regimes, and natural ecosystem trajectories, we
cannot be confident that restoration and conservation strategies will truly serve
Lrk in the year 1834 is
to return sites to a natural condition. Finally, the Indiana Dunes region, like many
lucted by the U.S. go\--
ecosystems, was in a natural state of change prior to the European impact. These
tree cover encountere
natural shifts will, and should, continue into the future. Attempting to restore a
~ys conducted in 1985.
habitat should not be considered the equivalent of trying to freeze it in time and
se in the density of oak
artificially preserving some idealized "natural" state .
. Today there are smal
;nus banksiana) in the
were extremely abun- The Global Warming Challenge
with oak. Other histor-
As we saw in Chapter 7, the increasing levels of C02 in the atmosphere caused by
vfichigan were heaYil_
the burning of fossil fuels are likely to result in significant warming and changes
n to logging, historic ·
in global climate over the next century. The impact of greenhouse warming on the
l
4 78 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

biosphere could be dramatic and is the subject of much study. This problem is of
major interest to biogeographers. One approach to anticipating the biogeo-
graphic impacts of greenhouse warming is to examine how the geographic distri-
bution of biomes might be affected. The strength of this approach is that we have
long studied the link between climatic conditions and the present distribution of
biomes. By assuming that vegetation formations will move to adjust to geo-
graphic shifts in climate, we can project the future distributions of the biomes.
There are two weaknesses to this approach. First, we do not know how long it will
take species to shift their range boundaries. The rate of climatic warming in the
next 100 years could be faster than any natural climatic changes since the end of
the last ice age. Biomes may remain out of adjustment with new climatic condi-
tions for some time, and some species may go extinct. Second, it is possible that
combinations of temperature and precipitation that do not have a modern coun-
terpart could develop and create biomes that have no exact modern counterpart.
In any case, a number of attempts have been made to predict the changes in the
global distribution of biomes due to greenhouse warming. In addition, some of
the more recent studies have also incorporated the direct impact of increased
C0 2 on rates of photosynthesis and plant growth. One of the most recent projec-
tions indicates that large areas of the arctic tundra will be replaced by boreal for-
est (Fig. 15.10). In turn, large areas presently occupied by boreal forest will be
occupied by temperate deciduous forest and grassland. Grasslands and the grass-
land-temperate forest ecotone will expand and replace large areas of temperate
deciduous forest in North America. Rainforest will expand in South America,
Africa, and Asia.
As mentioned earlier, the shifts in species ranges and biomes that might
accompany anticipated global warming will not occur instantly. Rather, some
species will adjust their ranges quickly, while those that have lower dispersal rates
may lag behind the climatic changes. Biogeographers are now addressing the
questions of how fast species will be able to adjust to rapid climatic changes by
using a variety of approaches.
Some biogeographers are looking at fossil records to see how quickly flora ..
and fauna responded to the natural climatic changes that occurred over the Qua-
ternary. Fossil pollen evidence shows that the rate at which tree species dispersed
northward following the close of the last age ranged from < 100 m per year to >
1000 m per year, and this provides some evidence of the rate at which these
species might adjust their ranges in response for future global warming. Unfortu-
nately, these past dispersal events do not provide a perfect analogue for future
warming because most natural changes in global climate occurred at a much
slower rate than anticipated greenhouse warming. Other biogeographers are
FIGURE 15. 1
looking for evidence of recent shifts in ecosystems and species range boundaries that m ig ht res
that might indicate that organisms are already responding to warmer tempera-
tures. It has already been found that southern species of coastal invertebrates are
extending their ranges northward along the Pacific Coast of North America, bird sensitive to
species are extending their ranges northward in Europe, and the size of tree pop- already com
ulations is increasing at the northern edge of the boreal forest biome. Biogeogra- Much
phers are using tree-ring analysis to see if increasing levels of C02 in the might adjust
atmosphere are serving as a fertilizer and producing higher growth rates of trees. from these
Evidence remains ambiguous. Remote sensing and GIS are being combined to First, the rat
help detect large-scale shifts in ecotones. such as treeline, that are difficult to require unu
resolve from ground-based studies. Treeline location should be particularly habita t have
 BIOGEOGRAPHY AND CONSERVATION PLANNING 479

>roblem is of
the biogeo-
"# .-!..
~
raphic distri-
that we have
stribution of
!just to geo-
' the biomes.
w long it will
rming in the
:e the end of
matic condi-
possible that
-.
wdern coun-
counterpart.
1anges in the
ion, some of
of increased • Tundra, forest-tundra, ice ~'t
ecent projec- • Boreal forests
'Y boreal for- • Temperate forests
·orest will be • Tropical forests
nd the grass- Savannas, dry forests, woodlands
of temperate
1th America,

~s that might
~ather, some
ispersal rates
!dressing the
c changes by

quickly flora
-- ,
1ver the Qua-
:ies dispersed
per year to >
which these
ing. Unfortu-
ue for future /
:data much
graphers are
FIGURE 15.10 One estimate of the changes in the location of major terrestrial biomes
e boundaries
that might result from global warming (after Nielson and Marks, 1994; Watson et al., 1996).
ner tempera-
:rtebrates are
\merica, bird sensitive to future climate warming. There is some evidence for northward shifts
=of tree pop- already commencing for some major vegetation zones in northern Eurasia.
e. Biogeogra- Much work is being done to develop computer models of how organisms
C02 in the might adjust their ranges in the face of changing climatic conditions. The results
rates of trees. from these models suggest that many species will face two difficult obstacles.
combined to First, the rates of climatic warming are faster than natural rates of change and
·e difficult to require unusually rapid dispersal capabilities. Second, because many types of
: particularly habitat have become extremely fragmented due to human activity, in many areas
T

480 CHAPTER 15 BIOGEOGRAPHY AND THE CONSERVATION CHALLENGE

species will have to cross large barriers, such as extensive farmlands or urban REFERENCES J.
areas as they migrate. The reliance on jump-dispersal events to cross such barri-
ers will slow or even preclude the ability of some species to adjust to new climatic Avise, J. C., and Hamric!
conditions. In these cases ex situ conservation measures, such as maintaining tion Genetics: Case H
breeding animal populations in zoos or holding plants and seed banks in botani- Chapman and H all, !'
cal gardens, may be required to ensure the survival of some species during the Baker, W. L. (1992) . Effe
period of rapid climatic change. suppression on lands<
73, 1879-1887.
Baker, W. L. (1993). Spat
multi-scale response <
SOME REFLECTIONS pression. Oikos 66, 6t
Baker, W. L. (1994) . Rest
Throughout this book we have examined and celebrated the great diversity of structure altered by fi
life that exists on our planet today. We have seen that geography and history vation Biology 8, 763-
Baker, W. L. , H onaker. J.
have worked together to produce the incredibly complex and beautiful modern
(1995). Using aerial p
biosphere that we are a part of. We have also learned that humans have played map the forest-tundra
a lamentable role in the extinction of a great many species. In this final chapter, Mountain National p<
we have observed that we face many challenges in preserving the remaining change research. Phoz
biodiversity and natural habitats of the world. Our efforts must take on a spe- ing and Rem ore Sensil
cial urgency because as human population grows, the rate of habitat and species Bartlein, P. J. , Whitlock. C.
loss also increases. The specter of global warming means that no species or Future climate in the 'l
environment can be counted on as completely secure in the years ahead. We Park region and its pot
tion. Conservation Bio,
have seen how the study of biogeography and allied fields can help us to both
Berger, J. (1990). Persiste
understand and preserve the natural life of the planet. The application of bio- populations: an empir.
geography for conservation is still in an early stage, and many exciting discov- extinctions in bighorn
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learning more about biogeography, ecology, paleontology, and the other life Brown, J. H. (1995 ). Macr
and earth sciences. You may actually take up the challenge of working as bio- Chicago Press, Chicag•
geographers to understand and protect the biosphere. If so, this book has been Brown, J. H. , and Lomoli1
more than successful. For others, I will consider this book successful if a trip geography, 2nd editior
through a national park or even a simple walk through autumn leaves is just a Sunderland, MA.
Bush, M. B. (1996). Amaz
little more enjoyable because you are now more aware of how you, and all the changing world. Biola~
plants and animals you encounter, are part of the marvelous and continuing 219-228.
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Wildlife Ecology and J
Science, Cambridge. 'v
Cole, K. L. (2001 ). A mult
past and ongoing \·ege
KEY WORDS AND TERMS the Indiana dunes. In
Handbook (ed. D. Ega
Applied historical ecology Environmental restoration Population viability analysis pp. 391-412. Isl and Pre
Biodiversity conservation Ex situ conservation (PVA) Costanza, R. (1991 ). Ecole
Biodiversity hot spots Gap analysis Range collapse Science and Ma nagenu
Charismatic megafauna Habitat-based conservation Restoration ecology Columbia University P
Charismatic flora In situ conservation SLOSS debate Costanza, R. , d' Arge. R ..
Conservation biogeography Land ethic Species-based conservation Grasso, M. , H annon . B
Conservation biology Landscape ecology Threatened species S. , O'Neil, R. V.. Parue
Conservation genetics Minimum viable population Umbrella species ton, P. , and van den Be
Endangered species size (MVPS) of the world 's ecoS\'ste
Environmental reclamation capital. Nature 387. 25

sive farmlands or urban
rents to cross such bani-
to adjust to new climatic
res, such as maintaining
nd seed banks in bot ani-
some species during the
~d the great diversity of
. geography and history
~x and beautiful modern
1at humans have played
ies. In this final chapter.
·eserving the remaining
)rts must take on a spe-
te of habitat and species
~ans that no species or
in the years ahead. We
~Ids can help us to both
The application of bio-
d many exciting discov-
>tart you on the road to
ogy, and the other life
~nge of working as bio-
f so, this book has been
ook successful if a trip
autumn leaves is just a
of how you, and all th e
rvelous and continuing
tlation viability analysis
>VA)
~e collapse
oration ecology
SS debate
ies-based conservation
atened species
1rella species
REFERENCES AND FU RT HER READING
A vise, J. C. , and Hamrick, J. L. (1996). Conserva-
tion Genetics: Case Histories from N ature,
Chapman and Hall, New York.
Baker, W. L. (1992). Effects of settlement and fire
suppression on landscape structure. Ecology
73, 1879-1887.
Baker, W. L. (1993). Spatially heterogeneous
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GLOSSA
AND TE~
Acheulean Tradition The star
Homo erectus.
Adaptive Radiation Th e de vel
from a single founding spec
occupy the different range <
ent resources that are prese
adaptive radiation occurs.
Adventitious Roots Roots th a
other plant parts th at are n
buried by soil due to floods
events.
Agriculture The deliberate cui
mals for hum an use.
Alleles The gene form s that ex.
Allelopathy Competition bet\\
chemical compounds releas
the germination or growth <
Allen's Rule The general obser
in cold environments have sl
limbs or ears, than related sr
ronments.
Allopatric Speciation The fo rn
geographic isolation.
Alpha Diversity The degree of
found on different sites of tl
nity type.
Amphioceanic Having a goegr
divided by an ocean.
Amphiregional Having a bioge
is divided by a barrier into 1
Amphitropical Having a bi oge
is divided by the tropics.
Analytical Biogeograph) The d
rules that explain how geogr
and distribution of plants an
distributions and evolutiona
modern distributions.
Anemohydrochores Organisms
water.
Aneomochores Organisms disp
Angiosperm Floweri ng plan ts._
which are developed " it bin<
onomies refer to angiosperm
Annual Plants that germinate. g
in one growing season.
tlyses of the distribution of
e expansion of the human
rzce 44, 592-604.
That is conservation biol-
i, 727-734.
iable Populations for Con-
lge University Press, Cam-
rrberton, R. W. (2000). Ecol-
trol of invading species-
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'cology and Evolution of
vi. L. Cody and J. M. Dia-
. Belkmap Press, Cam-
GLOSSARY OF KEYWORDS
AND TERMS
Acheulean Tradition The stone tool making tradition of
Homo erectus.
Adaptive Radiation The development of many species
from a single founding species. The new species evolve to
occupy the different range of habitats and use the differ-
ent resources that are present in the region in which
adaptive radiation occurs.
Adventitious Roots Roots that develop from stems or
other plant parts that are near the soil surface or get
buried by soil due to floods or other depositional
events.
Agriculture The deliberate cultivation of plants and ani-
mals for human use.
Alleles The gene forms that exist for a given locus.
Allclopathy Competition between plant species in which
chemical compounds released by one species inhibits
the germination or growth of competing species.
Allen's Rule The general observation that animals that live
in cold environments have shorter extremities, such as
limbs or ears, than related species that live in warm envi-
ronments.
Allopatric Speciation The formation of new species by
geographic isolation.
Alpha Diversity The degree of difference between species
found on different sites of the same habitat or commu-
nity type.
Amphioceanic Having a goegraphic distribution that is
divided by an ocean.
Amphiregional Having a biogeographic distribution that
is divided by a barrier into two distinct regions.
Amphitropical Having a biogeographic distribution that
is divided by the tropics.
Analytical Biogeography The development of general
rules that explain how geography affects the evolution
and distribution of plants and animals and how past
distributions and evolutionary history are reflected in
modern distributions.
Anemohydrochores Organisms dispersed by wind and
water.
Aneomochores Organisms dispersed by wind.
Angiosperm Flowering plants. Angiosperms have seeds
which are developed within an ovary. Some newer tax-
onomies refer to angiosperms as the Magnoliophyta.
Annual Plants that germinate, grow, reproduce and die all
in one growing season.
/r-
Anthropochores Organisms dispersed by humans.
Applied Historical Ecology The use of paleoecological
techniques such as pollen analysis and dendrochronol-
ogy to help in ecosystem restoration.
Areography The ecological analysis and interpretation of
the large scale geographic distributions of plant and
animals.
Ark A geologic event that leads to the movement of a
land mass and the geographic transport of flora and
fauna.
Artificial Selection Conscious and unconscious genetic
selection practiced by humans on species of plants and
animals.
Association Groupings of plant species that are typically
found growing together in similar habitats.
Atmosphere The envelope of gasses that surrounds the
earth.
Australopithecus The earliest known genus of hominids.
Found only in Africa and now extinct.
Autecology Ecological research which focuses on one
species.
Autotrophs Photosynthetic plants that produce chemical
energy using sunlight, water, and carbon dioxide.
Background Mortality The natural mortality rate for
plants and animals caused by the death of individual
organisms and not associated with large disturbance
events.
Barrier Geographic features that block dispersal and colo-
nization.
Batesian Mimicry Occurs when one species, which is not
poisonous or unpalatable, has the same coloring or shape
as a species which is poisonous or unpalatable. The palat-
able species benefits from coexisting with the unpalat-
able species because predators mistake it for the other
species and avoid it. This can be considered a form of
commensalism.
Benthic Organisms Plants or animals that live on the bottom
surfaces of water bodies.
Beringia The region of eastern Siberia and Alaska that
links North America with Asia during periods of low-
ered sea level.
Beta Diversity The degree of difference in the species
composit ion between two different habitats or commu-
nity types.
485
486 GLOSSARY AND KEY WORDS AND TERMS

Biodiversity Can be broadly defined as the number, vari- Carrying Capacity The number of individuals of a given Colonization The expansion
ety, and variability of living organisms. Biodiversity is species that a given area of the environment can sup- ment of a self-sustaining ~
often simply defined as the number of diffe rent species port. graphic region.
in a given geographic area. The greater the number of Catena A gradient in soil characteristics and types across Commensalism A symbiotic
species present, the greater the biodiversity. a landscape. species benefits and the ot
Biodiversity Conservation Efforts that are made to pre- Centers of Domestication Discrete areas of the world Community All the different
serve the species richness of an entire ecosystem rather where a number of different plant and animal species that live and interact with
than focusing upon one individual species. were first domesticated. scribed area.
Biodiversity Hot Spots Areas that contain high numbers of Charismatic Flora Showy, large, or otherwise interesting Community Type Th e coll ect
endemic species that are threatened by extinction due to plants that are often the obj ect of intense conservation ally found in a specific typ<
human activities. measures. Competition A s the interactit
Biogeographic Line A geographic boundary dividing two Charismatic Megafauna Large, attractive and/or otherwise same species or indi v id ual~
biogeographic regions. interesting animals which are often the object of the growth and/or fertilitv
Biogeographic Province Subcontinental areas that contain intense conservation measures. is increased for th e interac
similar flora and faun a. Chloroplasts Photosynthetic bodies within plant cells. Competitive Exclusion PrinciJ
Biogeographic Realm Supercontinental areas that contain with identical niches cann a
Choropleth Maps Maps on which the geographic distribu-
similar flora and faun a. graphic region because om
tions of species (or other fe atures and phenomena) are
Biogeographic Region Continental and supercontinental other to extinction.
delineated by continuous shading or circumscribed by
areas that contain similar flora and fauna. lines. Conifer Cone bearing gym no'
Biogeographic Relicts Taxa that once had larger distribu- Chromosome Threadlike structures within cells upon
spruces.
tions, but have become narrow endemics. which genes are arranged. Conservation Biogeography 1
Biogeographical Provincialism The tendency for different Chronosequence Approach A technique for constructing
graphical data and principl•
regions to possess unique species, genera, or familie s. seres by studying many different patches created by and management.
Biogeography The study of the past and present geographic disturbances that occurred at differe nt times in the Conservation Biology A synth
distributions of plants and animals and other organisms same type of environment. This type of approach is principl es from bi ologv and
and the environmental and evolutionary forces that pro- sometimes called space for time substitution. tenance of biodiversity.
duce those distributions. Circumpolar Having a geographic distribution that circles Conservation Genetics The ap
Biomass The mass of living material in an ecosystem. Gen- the polar regions. and techniques of genetics t

erally considered the dried weight of all living matter in Clade The lineage of different related species that arise
versity.
a prescribed area. from a common ancestor. Continental Drift A wide ly act
Biome Very large areas of the earth's surface that have a Cladistics, Cladograms The reconstruction of evolutionary logic plates on which th e co
similar climate and vegetation. history and linkages between species by constructing their relative geographic po
branching diagrams (cladograms) based upon the num- over geologic time.
Biosphere All life on the earth.
ber of shared physical traits observed between different Continental Taxa Species that
Bipedal The ability to walk on two legs.
taxa. Convergent Evolution Th e de'
Bipolar Having a biogeographic distribution with popula-
Cladogenesis (see Speciation). phological or physiological t
tions located in both the North and South Polar
Class A taxonomic category that includes groups of living in geographicall v sepa
regions.
related orders. similar environm ents.
Blitzkrieg Theory of Pleistocene Overkill P.S. Martin's
Climatic Disjunction A disjunct distribution caused by cli- Cope's Rule A general trend tc
hypothesis of how a geographically expanding front of
matic change. eage evolves. This pattern oc
prehistoric human colonists led to the extinction of
history of many species.
many large mammals in North and South America. Climatic Relict Species that have had their geographic
ranges constricted due to climatic changes. Corridor Geographic features
Bottleneck The decrease in genetic diversity that results
from a significant decrease in population size such as colonization.
Climax The idealized (and often unrealized or unrealistic)
might occur after a natural catastrophe. end point of succession. Cosmopolitan Having a global
Buttress Trunks Tree trunks that are wide and splaying at Cline A geographic gradient in a genetically controlled Crassulacean Acid Metabolism
the base. Typical of tropical rain forests. trait. CAM plants, C0 2 is absorbe
C3 and C4 Photosynthesis Different photosynthetic path-
malic acid. During the ligh t
Clovis Points An elongated stone point with a fluted base
ways. In C3 plants the carbon dioxide from the atmos- conducted by th e C3 pathwa
that is accepted by many archaeologists as representing
phere is converted into a 3-carbon molecule called synthesis.
the earliest tradition of tool making in North America.
3-phosophoglyceric. C4 plants convert carbon dioxide Most points of this type date from around 11 ,000 years Deciduous Perennial plants tha
into two 4-carbon molecules, malic and aspartic acid. The ago (10,000 radiocarbon years BP). annual cold or dn· seasons.
C4 pathway is common in grasses from warm regions. Coefficients of Similarity Mathematical measures of how Density The num ber of indi,·idu
Canopy Fire A fire that burns in the canopy cover of a for- similar the flora and fauna of one geographic area are uni t of area.
est. to the flora and fa una of another area. Dicotyledon A cl ass of angiosp
Canopy The uppermost layer of vegetation cover. Coevolution Occurs when two unrelated species evolve leaf-like photosyn thetic stru
Carnivore An organism that consumes animals. traits that are tied to th eir interactions. develop as a pair. DicotYledo

1ber of individuals of a given
of the environment can sup-
1aracteristics and types across
)iscrete areas of the world
:ent plant and animal species
nge, or otherwise interesting
object of intense conservation
rge, attractive and/or otherwise
1 are often the object of
1sures.
c bodies within plant cells.
which the geographic distribu-
features and phenomena) are
; shading or circumscribed b,-
ructures within cells upon
l Conservation Biogeography The application of biogeo-
A technique for constructin g
lifferent patches created by
!d at different times in the
Jt. This type of approach is
or time substitution.
rap hie distribution that circles
ent related species that arise
r.
reconstruction of evolution af\
teen species by constructing
lograms) based upon the num-
tits observed between differen:
n).
· that includes groups of
met distributi on caused by eli-
have had their geographic
climatic changes.
•ften unrealized or unreali stic
tin a genetically controlled
tone point with a fluted base
trchaeologists as representing
Jl making in North America.
Lte from around 11,000 years
ears BP).
>thematical measures of how
a of one geographic area are
mother area.
vo unrelated species evolve
r interactions.
Colonization The expansion of a speci es and its establish-
ment of a self-sustaining population in a new geo-
graphic region.
Commensalism A symbiotic relationship in which one
species benefits and the other is not affected.
Community All the different populations of organisms
that live and interact with each other within a pre-
scribed area.
Community Type The collection of species that is gener-
ally fo und in a specific type of habitat.
Competition As the interaction betwee n individuals of the
same species or individuals of different species in which
the growth and/or fertility is decreased or the mortality
is increased for the interacting organisms.
Competitive Exclusion Principle The theory that species
with identical niches cannot exist in the same geo-
graphic region because one would eventually drive the
other to extinction.
Conifer Cone bearing gymnosperms such as pi nes and
spruces.
graphica l data and principles for conservation planning
and management.
Conservation Biology A synthetic discipline that applies
principles from biology and other sciences to the main-
tenance of biodiversity.
Conservation Genetics The application of the principles
and techniques of genetics to the conservation of biodi-
versity.
Continental Drift A widely accepted theory that the geo-
logic plates on which the continents lie have shifted
their relative geographic positio ns very significantly
over geologic time.
Continental Taxa Species that have evolved on continents.
Convergent Evolution The development of similar mor-
phological or physiological traits in unrelated species
living in geographically separated regions that have
similar environments.
Cope's Rule A general trend towards larger size as the lin-
eage evolves. This pattern occurs in the evolutionary
history of many species.
Corridor Geographic features that promote dispersal and
colonization.
Cosmopolitan Having a global geographic distribution.
Crassulacean Acid Metabolism (CAM) Photosynthesis In
CAM plants, C0 2 is absorbed at night and stored as
malic acid. During the light of day, photosynthesis is
conducted by the C3 pathway. Cacti use CAM photo-
syn thesis.
Deciduous Perennial plants that Jose their leaves during
annual cold or dry seasons.
Density The number of individuals of a given species per
unit of area.
Dicotyledon A class of angiosperms in which the earliest
leaf-like photosynthetic structures (cotyledons)
develop as a pair. Dicotyledons also have leaves with
GLOSSARY AND KEY WORDS AND TERMS 487
net-like venation. Some newer taxonomies classify
dicotyledons as the class Magnoliopsida.
Diffusion The expansion of the range of a species along a
discrete front.
Dimorphic Leaves Different sized or shaped leaves on the
same plant that may vary by season or by the high on
the plant at which the leaf grows.
Diploid A cell o r organism containing both sets of chro-
mosome pairs.
Disjunct Having a biogeographic distribution in which
two or more populations of the same taxon are sepa-
rated by large geographic areas in which the taxon is
absent.
Dispersal Disjunction Taxa which have a disjunct distribu-
tion due to jump dispersal events.
Dispersal The movement of an organism away from its
point of origin. Active dispersal involves locomotion
by the organism. During passive dispersal the organ -
ism is transported by gravity, wind, water, or other
organisms.
Disturbance A discrete and relatively rapid physical or
biological event that disrupts ecosystem, community, or
population structure and changes resources, substrate
availability, or the p hysical environment.
DNA (Deoxyribonucleic acid) The DNA molecul es are
made up of various combinations of sugars and phos-
phates that are joined together by nitrogenous com-
pounds consisting of adenine, thymine, cytosine and
guanine bound by hydrogen. The DNA structure
resembles a twisted ladder with the sugars and phos-
phates on the sides and the nitrogenous compounds
forming the rungs. Genes are formed of DNA.
Domestication The process of artificial selection by which
plant and animal species come to depend upon humans
for survival while in turn providing humans with practi-
cal or other benefits.
Dominance Type Plant associations that are defined an d
described on the basis of the one or several plant
species th at structurally dominate them.
Ecological Biogeography The biogeographic study of th e
modern relationships between organisms and the envi-
ronment.
Ecological Equivalents Widely separated and unre lated
species th at are sim ilar in morphology and behavior.
Widely separated, but physiognomically and struc-
turally similar vegetation formations.
Ecosystem All of the interacting biological and physical
components of a prescribed area.
Ecotone The geographic boundary between two different
adjacent communities.
Endangered Species A species or subspecies that is at risk
of extinction throughout all or a portion of its range.
Endemic Restricted to one geographic area.
Environmental Determinism The premise that human cul-
tures and changes in culture are driven mainly by the
natural environment and changes in environment.
Environmental Reclamation The replacement of a
severely damaged or destroyed ecosystem with a land-
488 GLOSSARY AND KEY WORDS AND TERMS

scape such as a lawn that does not attempt to replicate Facilitation The process by which the establishment of one Geologic Time Scale An int
the original ecosystem. species changes the environment and allows the subse- that serves as the standar
Environmental Restoration The rebuilding of the original quent establishment of other species many millions of years ot
biological and physical environment in damaged Family A taxonomic category that includes taxonomically GIS (Geographic Informatic
ecosystems, and in some cases in areas where the natu- similar and related genera. terns used to combine an•
ral habitat has been completely destroyed. Fertile Crescent A crescent of land that extends from Glaciation A period of regie
Epiphyte So called air plants that grow on the branches the mouth of the Tigris and Euphrates rivers, north to and expansion of glaciers
and trunks of host plants, but do not tap into the vascu- eastern Turkey and then south to the coastal regions Gondwanaland A large cont
lar tissue of the host plant for water or nutrients. of Lebanon and Israel. Some of the earliest evidence million years ago and inch
Equilibrium Theories of Biodiversity Theories which of plant and animals domestication comes from this Africa, Australia, South A
assume that modern biodiversity is in equilibrium with region.
Great American Exchange 1
modern environmental conditions. Filter Avenues of dispersal and colonization that are not fauna that occurred foll m
Equilibrium Theory of Island Biogeography The theory equally favorable for all species. Isthmus of Panama.
advanced by R.H. MacArthur and E.O. Wilson that the Food Web The complex flow of energy between different Greenhouse Effect The pher
biodiversity on islands is governed by rates of coloniza- trophic levels. dioxide, methane, and cer
tion and extinction, which are in turn controlled by Formation A classification of vegetation based upon struc- phere allow the down wan
island isolation and island size. ture. solar light energy and trar
Erosion The transport of weathered rock and other mate- Fossils The preserved and usually lithified remains of Increases in carbon di oxic
rials. organisms found in geologic deposits. gasses due to the burni ng
Euryhaline Organisms that can withstand a wide range of human actions ma y lead t<
Founder Principle The idea that populations founded by a
salinities. perature and global sea le·
very small number of individuals generally contain a
Euryphagous An organism that can consume a wide vari- small subset of the total genetic variability of the main Growth Form A classificatior
ety of different foods. population and are prone to allopatric speciation. their morphology.
Fundamental Niche The niche space of a species if there Guilds Groups of species witi
Eurythermic Organisms that can withstand a wide range
was no competition with other species that decreases similar forms, habitat, and
of temperatures.
the range of resources conditions it could exist within. Gymnosperm Seed bearing p
Eustatic Sea Level Change Global changes in sea level
Gap Analysis The use of GIS to overlay species maps to produced outside of a prot
caused by increases and decreases in the amount of
water in the world's oceans. The storage of water in gla- find areas of high biodiversity or map endangered Habitat Based Conservation
cial ice is a major cause of eustatic sea level declines. species and to compare the distributions of biodiversity serve the entire biological
or endangered endangered species to factors such as which endangered species
Eutrophic Lake Shallow, warm lakes with high amounts of
nutrients that have high primary productivity. These soil conditions or vegetation cover. Areas of high biodi- Habitat The explicit spatial. p
lakes generally have low amounts of oxygen due to versity or the ranges of endangered species are then ronment in which species c
high rates of respiration and aerobic decomposition. overlayed on maps of current or planned conservation Haploid A cell or organism cc
areas to see if those areas are in the right location to the chromosome pairs.
Evapotranspiration Water that is added to the atmosphere
preserve biodiversity or endangered species. Harmonization, Harmony Tw
by both evaporation from waterbodies and soils, and by
transpiration from plants. Gaps Openings in vegetation cover caused by large and harmony when they conrail
small disturbances. Heliophytes Plants that grow
Evolution Genetically controlled changes in physiology,
anatomy and behavior that occur to a clade over time. Gene Basic unit of heredity carried and transmitted by Heliotropic Morphologi cal fe<
Microevolution refers to evolution at the scale of the chromosomes. The genes of plants and animals consist are influenced by the rel ati·
species. Macroevolution refers to evolutionary changes of molecules of deoxyribonucleic acid (DNA).
Herbivore An organism that c
viewed at higher taxonomic units such as genera and Generalist A species which has a wide range of environ-
Heterotrophs Non-photos,·mr
families. mental tolerances and a large niche.
upon consuming other orga
Evolutionary Disjunction A disjunction that occurs when Genetic Drift Stochastic changes in the genetic composi-
Heterozygous Refers to a locu
species evolve at two different peripheries of the geo- tion of a population that occur over time as new genes
associated with it
graphic range of a population. Extinction of the ances- arise via mutation and other genes are lost through
chance processes. Historic Extinctions The recen
tral population causes the related species to be disjunct
(starting around 1600 AD )
from each other. Genome The complete range of genes present in a species.
of European colonization aJ
Evolutionary Relict Survivors of once more widespread Genotypic Variations Differences in the genes between sion, and the rapid global in
and diverse evolutionary lineages that are now narrow different species or members of the same species. Not tion.
endemics. all genotypic variations may be observable as pheno- Historical Biogeography The t
Ex situ Conservation Attempts to save species from typic variations. past distribution s and e\·olu
extinction by maintaining them outside of their natural Genus A taxonomic category that includes groups of taxo- Historical Theories of Biodivers
habitat in areas such as zoos or botanical gardens. nomically similar and closely related species. The plural that modern biodi versitY is n
Exponential Population Growth The geometric growth of of genus is genera. ern environmental condition
a growth of a population in an environment where Geographic Range The entire geographic area in which a events of the evoluti on and
there are no limits to population size- dN!dt = rN species can be found living. Hollow Curve Distribution .-\
Extinction The loss of all individuals of a species, genus, Geologic Disjunction A disjunct distribution caused by species ranges, consisting of
family, or order. Extinctions may be local or global. geologic events such as continental drift. ranges and few species with

which the establishment of one
ronment and allows the subse-
•ther species
•ry that includes taxonomically
ra.
t of land that extends from
and Euphrates rivers, north to
1 south to the coastal regions
)orne of the earliest evidence
mestication comes from this
and colonization that are not
species.
N of energy between different
Jf vegetation based upon struc-
tsually lithified remains of
1gic deposits.
that populations founded by a
lividuals generally contain a
genetic variability of the main
~ to allopatric speciation.
:he space of a species if there
other species that decreases
mditions it could exist within.
rs to overlay species maps to
~rsity or map endangered
he distributions of biodiversitY
ed species to factors such as
:ion cover. Areas of high biodi-
ndangered species are then
rrent or planned conservation
s are in the right location to
~ndangered species.
•n cover caused by large and
carried and transmitted by
of plants and animals consist
onucleic acid (DNA).
has a wide range of environ -
arge niche.
mges in the genetic composi-
occur over time as new genes
her genes are lost through
:e of genes present in a species.
·ences in the genes between
bers of the same species. Not
1ay be observable as phen o-
·y that includes groups of taxo-
sely related species. The plura:
e geographic area in which a
~-
unct distribution caused bv
mtinental drift.
Geologic Time Scale An internationally accepted system
that serves as the standard means of subdividing the
many millions of years of earth's hi story.
GIS (Geographic Information Systems) Computer sys-
tems used to combine and map spatial data.
Glaciation A period of regional or global development
and expansion of glaciers.
Gondwanaland A large continent that existed around 150
million years ago and included the modern continents of
Africa, Australia, South America, and Antarctica.
Great American Exchange The movement of terrestrial
fauna that occurred following the establishment of the
Isthmus of Panama.
Greenhouse Effect The phenomenon in which carbon
dioxide, methane, and certain other gasses in the atmos-
phere allow the downward transmission of incoming
solar light energy and trap outgoing heat energy.
Increases in carbon dioxide and other greenhouse
gasses due to the burning of fossil fuels and other
human actions may lead to significant increases in tem-
perature and global sea level over the next 100 years.
Growth Form A classification of plants on the basis of
their morphology.
Guilds Groups of species within a community that have
similar forms, habitat, and resource requirements.
Gymnosperm Seed bearing plants in which the seeds are
produced outside of a protective ovule.
Habitat Based Conservation Efforts which attempt con-
serve the entire biological and physical environment in
which endangered species live.
Habitat The explicit spatial, physical, and biological envi-
ronment in which species can be found.
Haploid A cell or organism containing only one half of
the chromosome pairs.
Harmonization, Harmony Two geographic areas are in
harmony when they contain the same flora and fauna.
Heliophytes Plants that grow best in full sunlight.
Heliotropic Morphological features or movements that
are influenced by the relative position of the sun.
Herbivore An organism that consumes plant matter.
Heterotrophs Non-photosynthetic organisms that depend
upon consuming other organisms to acquire energy.
Heterozygous Refers to a locus that has different alleles
associated with it
Historic Extinctions The recent episode of extinctions
(starting around 1600 AD) caused by the combination
of European colonization and socioeconomic expan-
sion, and the rapid global increase in human popula-
tion .
Historical Biogeography The biogeographic study of the
past distributions and evolution of life.
Historical Theories of Biodiversity Theories that assume
that modern biodiversity is not in equilibrium with mod-
ern environmental conditions because it reflects past
events of the evolution and extinction.
Hollow Curve Distribution A typical distribution of
species ranges, consisting of many species with small
ranges and few species with large ranges.
GLOSSARY AND KEY WORDS AND TERMS 489
Homeotherm Animals that maintain a relatively steady
body temperature through the metabolic generation of
heat.
Hominid Moden human beings and their closest extinct
relatives of the genera Homo and Australopithecus.
Homo erectus An extinct hominid.
Homo habilis An extinct hominid.
Homo sapiens sapiens The genus, species and subpecies
classification of all modern humans.
Human Revolution A rapid expansion 50,000 years ago of
Homo sapiens sapiens within and out of Africa. These
modern hominids created art, practiced ritual burials,
and displayed other indications of sophisticated culture.
Hunters and Gatherers Human populations who rely
upon naturally occurring vegetation, fruits, nuts, car-
rion, fish, and game for subsistence.
Hybridization Sexual reproduction between two different
species.
Hydrochores Organisms dispersed by water.
Hydrophyte Plants that require high levels of soil mois-
ture.
Hydrosphere All of the water in the atmosphere, on the
surface, and below the surface of the earth.
Igneous Rocks Formed by the cooling and solidification of
molten material called magma. Intrusive igneous rocks
are formed when the cooling occurs deep underground.
Extrusive igneous rocks from when cooling occurs at
the surface.
In situ Conservation Attempts to save species from
extinction by maintaining them within their natural
habitat.
Individualistic Community Concept Gleason's concept
that species are distributed in the environment accord-
ing to their individual resource requirements and not
due to interactions with other species.
Industrial Revolution The rapid development of new tech-
nologies and the shift of the socioeconomic system
away from an agrarian base towards a techno-industrial
base that has occurred since about 1800.
Inhibition Model A model of succession that is based
upon the premise that the first species that arrive fol-
lowing disturbance come to dominate the site and com-
petitively inhibit the establishment of later arriving
species.
Insular Taxa Species that have evolved on islands.
Intermediate Disturbance Hypothesis According to this
hypothesis, if an ecosystem remains stable and free of
disruption by disturbance, the stable homogenous envi-
ronmental conditions will favor some species but will
lead to the extinction of species for which the stable
habitat is not favorable or which are prone to competi-
tive exclusion by species that are favored by the undis-
turbed environment. Disturbances cause physical and
biological changes and spatial heterogeneity in the sys-
tem that can favor species that would not survive in a
stable undisturbed system. On the other hand, if distur-
bance occurs too frequently and is too severe, it will
lead to the extinction of disturbance sensitive species
 -------- ~=--==-=--=-=~-------

490 GLOSSARY AND KEY WORDS AND TERMS

that have long generation times or occur in low num- ulations to the interaction between predators and prey: traits are reproductively r
bers and are prone to change extinction. Prey populations: dN/dt = rN- a'PN and Predator pop- individuals.
Invasion The colonization of a new region by a species ulations dP/dt = fa'PN- qP
Neanderthal Homo sapiens'
that has been introduced to that region by humans. Macroecology The ecological analysis subspecies of Homo sape1
Irruption Episodic explosions in the population size and and interpretation of the large-scale geographic distri- humans.
geographic ranges of insects of animals. butions of plant and animals.
Nekton Organisms such as fi
Isostatic Sea Level Change Changes in regional sea level Megafauna Large birds and mammals that are greater through the water.
caused by the uplift or downward motion of land sur- than about 44 kilograms in weight.
Neoendemic Species that ha
faces. Mesoamerica An archaeological term that refers to south- speciated and have small ]
Isotopes Forms of an element that have the same atomic ern Mexico and northern central America. geographic ranges.
number, but differ in the number of neutrons. Mesophyte Plants that require intermediate levels of soil Nested Pattern of Insular COJ
Jump Dispersal The dispersal of a species across a geo- moisture. dency for the biota on isla
graphic area that is not occupied by the species. Metamorphic Rocks Formed when existing rocks are flora and fauna found on r
Keystone Species A species that due to its presence or altered by high temperatures and pressures. nents.
absence can greatly change the productivity, species Metapopulations Members of the same species that are Niche The multidimensional
composition, or species diversity of an ecosystem. found in different and separated locations. Members of species exists.
Krummholtz Individual trees that have a low shrubby to different metapopulations may not interact with other Oasis Theory The original the
mat-like growth form due to cold and sind found at meta populations on a regular or frequent basis. that conditions in south we~
high elevation and high latitude treelines. Milankovitch Orbital Theory of Glaciation The widely warmer at the close of the
Land Ethic The concept espoused by A. Leopold that accepted theory that natural changes in the orbital development of agriculture
humans have an ethical responsibility to conserve and geometry of the earth produced the cycles of glaciation able plants and animals bee
preserve the natural environment. and non-glaciation that typify th e late Quaternary. oases.
Landscape Ecology The study of the spatial patterns of Minimum Viable Population Size (MVPS) The popula- Old Growth Forest Forest th<
landscapes, including the distribution of species and tion size required to keep a species from extinction disturbances or human Ian
habitat on local to regional scales, and how these pat- with the foreseeable future. years.
terns are developed maintained by nature and human Mitochondrial DNA (mtDNA) DNA found in the mito- Oldowan Tradition The oldes
actions. chondria. The mitochondria are small bodies found in tion, dating from 2.5 to abc
Last Glacial Maximum (LGM) The period of maximum the cytoplasm of cells and only contain maternal DNA. associated with Homo hab
ice advance and minimum global temperatures associ- Molecular Clock Analysis Th e use of differences in Oligotrophic Lake D eep. cole
ated with the last glacial stage. The LGM occurred mtDNA between different species or populations in oxygen and low amounts c
around 20,000 years ago. order to estimate the length of time they have been nutrients. They typicall y h<
Laurasia A large continent that existed around 150 mil- reproductively isolated. ity.
lion years ago and consisted of the modern ~ontinents Monocotyledon A class of angiosperms in which the first Omnivore An organism th at c
of North America, Europe, and most of Asia. leaf-like photosynthetic structures (cotyledons) animals.
Law of Superposition An assumption made in geology develop as a single shoot. Monocotyledons also have Optimal Foraging Theory The
that overlying material is younger than the underlying leaves with parallel venation. Some newer taxonomies evolve to focus their fo ragic
material. classify monocotyledons as the class Liliopsida. species that provide th e hig
Law ofUniformitarionism Am assumption made in geol- Monophyletic A group of species that descended from a relative to the energy requi
ogy that the natural processes of rock formation, common ancestor. Order A taxonomic category t
weathering, erosion, and deposition that occur today Moustarian Tradition The stone tool making tradition related families.
also occurred in the past and can be applied to other associated with the Neanderthals. Ordination Analysis The stud1
historical studies in natural history. The law is often
Muellerian Mimicry A type of mutualism that occurs lions relate to th e environrr
stated as "The present is the key to the past".
when a species, that is poisonous or unpalatable to ranges and densities are dis
Liana Woody vines typical of the tropical rainforest. predators, possesses the same coloring or shape as tal gradients.
Life Form A classification of plants on the basis of phys- another species that is also poisonous or unpalatable. Orogeny The form at ion of mo
iognomy and life history. Both species benefit since predators avoid either processes.
Life Zone Elevational and latitudinal bands of similar cli- species. Out of Africa Model A model
mate and vegetation Multiregional Model A model of human evolution that suggests most homin id spec
Lithosphere The geological structure of the earth. It suggests that modern humans, and perhaps some other humans, first evolved in Af
includes both solid rock and liquid forms such as species of Homo evolved independently through paral- from there to ot her parts 0
magma. lel evolution in different geographic regions. Out of Asia Model A model o
Locus The point at which a gene is located on a chromo- Mutation A change in the genes or chromosomes of a cell gests that modern humans e
some. or organism that creates a genotype that is unlike the persed from there.
Logistic Population Growth The S-shaped growth of a parent type. Overkill Extinction of a prev S
population in an environment in which there is a limita- Mutualism A symbiotic relationship in which both species ing by predators. and part ic
tion on the ulitmate size of the poplaution that can be benefit. Paleoendemic Older species th
supported-dN! dt =rN(K-N). Natural Selection The process by which the genes for graphic ranges in the past.
Lotka-Volterra Model A mathematical model the relates genetically controlled traits become more common in a Palynology The analysis of fos
cyclic fluctuations in the size of predator and prey pop- population over time because individuals with those used to reconstruct past veg

ion between predators and prey:
t = rN - a'PN and Predator pop-
- qP.
~ical analysis
he large-scale geographic distri-
Jimals.
1d mammals that are greater
•s in weight.
logical term that refers to south-
rn central America.
1uire intermediate levels of soil
Jed when existing rocks are
atures and pressures.
·s of the same species that are
;eparated locations. Members of
Jns may not interact with other
egular or frequent basis.
•ry of Glaciation The widely
tlural changes in the orbital
>roduced the cycles of glaciation
typify the late Quaternary.
m Size (MVPS) The popul a-
ep a species from extinction
ture.
NA) DNA found in the mito-
tdria are small bodies found in
nd only contain maternal DNA.
The use of differences in
ent species or populations in
ngth of time they have been
·angiosperms in which the first
: structures (cotyledons)
>t. Monocotyledons also have
ation. Some newer taxonomies
s as the class Liliopsida.
species that descended from a
stone tool making tradition
nderthals.
e of mutualism that occurs
oisonous or unpalatable to
same coloring or shape as
!so poisonous or unpalatable.
ce predators avoid either
>del of human evolution that
1mans, and perhaps some other
d independently through paral -
t geographic regions.
:enes or chromosomes of a cell
a genotype that is unlike the
ationship in which both specie>
:ess by which the genes for
aits become more common in a
cause individuals with those
traits are reproducti vely more successful than oth er
individuals.
Neanderthal Homo sapiem neanderrhalensis- an extinct
subspecies of Homo sapeins close!'' related modern
humans.
Nekton Organisms such as fis h that can propel themselves
through the water.
Neoendemic Species that have just recently evolved and
speciated and have small popul ation sizes and small
geographic ranges.
Nested Pattern oflnsular Communities The general ten-
dency for the biota on islands to contain a subset of the
flora and fauna found on nearby larger islands or conti-
nents.
Niche The multidimensional resource space in which a
species exists.
Oasis Theory The original theory by V.G. Chi ide postulated
that conditions in southwest Asia became drier and
warmer at the close of the Pleistocene. and this led to the
development of agriculture as people and domesticat-
able plants and animals became concentrated around
oases.
Old Growth Forest Forest that has not experienced large
disturbances or human land cl earan ce for hundreds of
years.
Oldowan Tradition The oldest known tool making tradi-
tion, dating from 2.5 to about 1.5 million years ago. It is
associated with Homo habilis in Africa.
Oligotrophic Lake Deep, cold lak es with high amounts of
oxygen and low amounts of phosphorus and other
nutrients. They typically have low primary productiv-
ity.
Omnivore An organism that consumes both plants and
animals.
Optimal Foraging Theory The theory that predators
evolve to focus their foraging activities upon prey
species that provide the highest ratio of food energy
relative to the energy required for foraging.
Order A taxonomic category that includes groups of
related families.
Ordination Analysis The study of how species distribu-
tions relate to the environment, including how species
ranges and densities are distributed along environmen-
tal gradients.
Orogeny The formation of mountains by geologic
processes.
Out of Africa Model A model of human evolution that
suggests most hominid species, including modern
humans, first evolved in Africa and then dispersed
from there to other parts of the world.
Out of Asia Model A model of human evolution that sug-
gests that modern humans evolved in Asia and dis-
persed from there.
Overkill Extinction of a prey species caused by overhunt-
ing by predators, and particularly by humans.
Paleoendemic Older species that had much larger geo-
graphic ranges in the past.
Palynology The analysis of fossil pollen and spores. Often
used to reconstruct past vegetation and climate.
GLOSSARY AND KEY WORDS AND TERMS 491
Panbiogeography A method of reconstructing evolutionary
history based upon geographic distributions and that vic-
ariance events were the overwhelming most important
way in which speciation occurred. Developed by L.
Croizat.
Pandemic Having a widespread distribution between and
within biogeographic regions.
Pangea A super continent that included most of the land
masses of the modern continents and existed around
200 million years ago.
Pantropical Having a biogeographic distribution across
the tropical regions.
Parallel Evolution Occurs when geographically isolated
populations derived from the same ancestor evolve into
morphologically and physiologically similar descendent
species.
Parapatric Speciation Speciation caused by the evolution-
ary divergence of populations that occupy different
habitat or niches in the same geographic area.
Parasitism A symbiotic relationship in which one species
benefits at the expense of the other.
Paripatric Speciation Speciation via founder effect.
Pastoral Nomadicism A form of subsistence in which
herders migrate and move along with their herds dur-
ing seasonal changes in grazing areas.
Patches Areas at different stages of succession due to
being impacted by disturbances at different times, or
different disturbances. Patches may also relate to differ-
ences in local environmental conditions.
Peninsula Effect The general observation that species
richness on peninsulas also generally declines towards
the tip of the peninsula.
Perennial Plants that live for several growing seasons.
Peripatric Speciation Occurs when peripheral popula-
tions become geographically isolated from the main
population and undergo genetic divergence and
speciation.
Permafrost Soil A soil in which the temperature does not
reach above freezing even in the summer months.
PhAR Photosynthetically active radiation. This normally
corresponds to visible light.
Phenology The seasonal timing of changes in the growth
and reproduction of plants and animals.
Phenotypic Variation Observable differences in the physi-
ology, anatomy, or behavior of different species or indi-
viduals of the same species.
Phloem Cells in vascular plants that are specialized for the
flow of food.
Photoperiodism The influence of seasonal or daily
changes in light intensity on organisms.
Photorespiration The release of carbon dioxide by plants
without photosynthetic production having occurred.
This occurs under bright light and high temperatures in
c3plants.
Photosynthesis The process by which plants combine
water and carbon dioxide in the presence of sunlight to
capture visible light spectrum electromagnetic energy
in the sunlight and transform it into chemical energy in
the form of simple sugars.
492 GLOSSARY AND KEY WORDS AND TERMS

Phyla A taxonomic category that contains groups of Primary Succession Occurs when a previously lifeless Rhizomes Horizontal root ~
related classes. environment is first colonized by plants and animals. grasses and other plants :
Phyletic Gradualism A slow and gradual process of evolu- Primates Members of the Primate Order that includes Riparian The environment:
tion during which new traits arise by mutations and lemurs, tarsiers, monkeys, apes, and humans. Scarification A physical or bi
traits which infer greater reproductive success are Propagule The stage in the life cycle (a plant seed for exam- must undergo prior to ger
selected for and eventually become dominant over ple), part of an organism (a piece of marram grass which Sciophytes Plants that grow
many generations. can develop a new set of roots and grow into a full sized Sclerophyllous Leaves Stiff
Phylogenetic Biogeography The analysis of modern geo- grass plant), or group of organisms (a male and female cuticles and stomata th at
graphic distributions to infer evolutionary history. rabbit of mating age) that is required to establish a new decrease moisture loss.
reproducing population.
Phylogeny The evolutionary history of a taxon. Seasonal Migration The ann'
Punctuated Equilibria A model of evolution in which new
Phytogeography The reconstruction of past biogeographic from one regularly occupi
genotypes and species arise as small isolated populations
history on the basis of modern geographic distributions another for purposes of a'
or populations at the edges of the main species popula-
of genetic characteristics. and/or for feeding and rna
tion. These small populations increase and very rapidly
Phytogeography The study of the biogeography of plants. become dominant when environmental changes cause Second Growth Forest The I
Phytophagous An organism that consumes plants. them to be better adapted to new environmental condi- develops followin g a larg'
tions. clearance.
Plankton Organisms that exist by passively floating in the
waters of an ocean, lake, or other water body. These Radiocarbon Dating (14C Dating) The dating of organic Secondary Succession OccUJ
include many microscopic plants (phytoplankton) and remains using the amount of radioactive carbon (14C) recovers from a disturban
small invertebrate animals (zooplankton). in them as a guide to their age. Radiocarbon ages are Sedimentary Rocks Formed
reported as years before present (yrs BP) with present accumulates and lithifies.
Plate Tectonics The modern theory of how continental
taken to be AD 1950. Radiocarbon years do not Sere An idealized or typical
drift occurs due to movement of the mantle and out-
directly equal calendar years. An age of 10,000 BP occurs in an ecosystem fo·
flows of magma that cause movement of the overlying
equals a little over 11,000 calendar years ago.
crust. Serotinous Cones The cones
Random Model A model of succession that suggests that
Pleistocene Overkill The theory that the large mammal that remain closed and ho
the patterns of species abundance that occur following
extinctions that typify the late Pleistocene in Australia released following disturb
a disturbance are highly variable and largely influenced
and the Americas were caused by over hunting by the parent trees and/or he
by the random chance dispersal and establishment of
humans. post disturbance colonizers. Sixth Extinction The widesp1
Pluvial Lakes Lakes that form during long periods of over the past 10,000 years
Range As used in this book the term refers to the geo-
increased precipitation and/or decreased evaporation. humans.
graphic area permanently occupied by a plant or ani-
The formation and evaporation of pluvial lakes are mal taxon. SLOSS (Single Large Or Sevt
often linked to global glacial cycles. Range Collapse The linked phenomena of population entific and political debatE
Pneumatophores Elongated vertical root structures that decline and geographic range decline that is very typi- reserve is better than seve
rise above water saturated soils and the level of stand- cal of endangered species. servation of biodiversit y.
ing water and allow roots to obtain oxygen. Rapoport's Rule The general pattern of decreasing range Soil The uppermost laye r of 1
Poikilotherm Organisms that assume the temperature of sizes of plant and animal taxa that live near the equator found on the earth 's surfa<
their environment. compared to those that live in higher latitudes. Specialist A species with a na
Polymorphism Genetically controlled variation within a Realized Niche The reduced niche space occupied by a ances and a small niche.
population or species. species due to the impact of competition with other Speciation The development
Polyploid Organisms that have twice the chromosome species. differentiable species frorr
number of either parent. This is a common mutation in Red Queen Hypothesis States that because all of a species. Speciation results
plants. species' competitors are continually evolving and but not all evolutionary ch
becoming more competitive, if a species cannot evolve ment of two or more speci
Population All individuals of a given species in a pre-
quickly enough to keep pace with the evolution of com- Speciation is also referred
scribed area. Usually members of the same population
peting species, it will become extinct. Species There are several diff
are assumed to be in close enough proximity to be able
to interact and interbreed frequently. Regolith Weathered rock and mineral matter from which Biological species can be t
soil is developed. egory that includ es all indi
Population Viability Analysis (PVA) The methods used
Releve A rapid, semi-quantitative way of conducting a cen- reproduce and produce fer
for estimating minimum viable population size.
sus of a plant community. netic species concept ident
Predation A biological interaction in which one organism sexually reproducing or ga
Rescue Effect The rescue effect occurs when small popu-
consumes another. diagnostic character that is
lations of a species are rescued from the brink of
Prehistoric Extinctions Extinctions caused by the initial extinction by the arrival of new immigrants of the same the species, but absent in o
prehistoric geographic expansion of human beings from species. species are organisms that
Africa and Eurasia. descendent relationship th
Respiration The oxidative chemical reaction that breaks
Primary Forest Forest which has not experienced signifi- the high-energy bonds of carbohydrates to release record.
cant human modification or land clearance. energy for an organism's metabolism. Species Area Curve The t,·pic
Primary Productivity The energy fi xed in an ecosystem by Restoration Ecology The application of ecological principles between sampli ng area and
photosynthesis. Rates of biomass accumulation are and techniques to restore ecosystems that have been the species-area curYe. The
sometimes used to determine primary productivity. damaged by human activity. species-area curve is S=cA

s when a previously lifeless
mized by plants and animals.
Primate Ord er that includes
'S, apes, and humans.
life cycle (a plant seed for exam-
(a piece of marram grass which
·roots and grow into a full sized
organisms (a male and female
1t is required to establish a new
ode! of evolution in which new
ise as small isolated populations
:es of the main species popula-
:ions increase and very rapidly
environmental changes cause
d to new environmental condi-
lating) The dating of organic
nt of radioactive carbon (14C)
:ir age. Radiocarbon ages are
' present (yrs BP) with present
1diocarbon years do not
tears. An age of 10,000 BP
0 calendar years ago.
,f succession that suggests that
bundance that occur fo llowing
variable and largely influenced
ispersal and establishment of
:ers.
~ the term refers to the geo-
ly occupied by a plant or ani-
phenomena of population
·ange decline that is very typi-
:s.
·al pattern of decreasing range
I taxa that Jive near the equator
ive in higher latitudes.
d niche space occupied by a
t of competition with other
tes that because all of a
continually evolving and
tive, if a species cannot evolve
Jace with the evolution of com-
orne extinct.
nd mineral matter from whi ch
tative way of conducting a cen-
ffect occurs when small popu-
'scued from the brink of
of new immigrants of th e same
:hemical reaction that bre aks
' carbohydrates to release
metabolism.
plication of ecological principles
ecosystems that have been
ty.
Rhizomes Horizontal root structures typical of many
grasses and other plants such as cattails.
Riparian The environment al ong rivers an d streams.
Scarification A physical or biol ogical event that some seeds
must undergo prior to germination.
Sciophytes Plants that grow best in shade.
Sclerophyllous Leaves Stiff leaves with hard and waxy
cuticles and stomata that lie in small pits. These features
decrease moisture loss.
Seasonal Migration The annual movements of organisms
from one regularly occupied geographic region to
another for purposes of avoiding harsh conditions,
and/or for feeding and mating.
Second Growth Forest The forest vege tation that initially
develops following a large disturbance or human land
clearance.
Secondary Succession Occurs when an ex isting ecosystem
recovers from a disturbance such as fire or flood.
Sedimentary Rocks Formed when the weathered material
accumulates and lithifies.
Sere A n idealized or typical set of successional stages that
occurs in an ecosystem following a disturbance.
Serotinous Cones The cones of certain species of conifers
that remain closed and hold viable seeds. The seeds are
released following disturbances such as fires which kill
the parent trees and/or heat the cones.
Sixth Extinction The widespread episode of extinctions
over the past 10,000 years or so that has been caused by
humans.
SLOSS (Single Large Or Several Small) Debate The sci-
entific and political debate over whether a single large
reserve is better than several small reserves for the con-
servation of biodiversity.
Soil The uppermost layer of mineral and organic matter
fo und on the earth's surface.
Specialist A species with a narrow range of resource toler-
ances and a small niche.
Speciation The development of two or more genetically
differentiable species from a single common ancestor
species. Speciation results from evoluti onary change,
but not all evolutionary change leads to the develop-
ment of two or more species from a common ancestor.
Speciation is also referred to as cladogenesis.
Species There are several different definitions for species.
Biological species can be thought of as a functional cat-
egory that includes all individuals that can sexually
reproduce and produce fertile offspring. The phyloge-
netic species concept identifies a species as a group of
sexually reproducing organisms that share at least one
diagnostic character that is present in all members of
the species, but absent in other organisms. Evolutionary
species are organisms that have a direct ancestor-
descendent relationship that is traceable in the fossil
record.
Species Area Curve The typical non-linear relationship
between sampling area and species richness is called
the species-area curve. The common formula for the
species-area curve is S=cAz.
GLOSSARY AND KEY WORDS AND TERMS 493
Species Based Conservation Conservation efforts that are
aimed at the preservation of an individual species.
Species Eveness The degree to which the number of indi-
vidual organisms are evenly divided between the di ffer-
ent species of the community. An estimate of species
eveness alone is obtainable from the Shannon Index
value (H') using the equation E = H '/In S.
Species Relaxation The decline in species richness that
occurs when a region, such as a land bridge island, is
cutoff from a larger landmass or area of similar habitat.
Species relaxation also occurs when land clearance pro-
duces small reserves in place of once extensive areas of
natural habitat.
Species Richness The number of different species in a
given area. In many large-scale biogeographical studies,
biodiversity is synonymous with species richn ess.
Species Turnover Changes in the species represented at
diffe rent sites or between different times periods at the
same site.
Stability-Time Hypothesis The hypothesis that long peri-
ods of environmental stability lead to high species rich-
ness.
Stand An individual plot of plants, particularly forest
trees.
Stenophagus An organism with a very narrow range of
foods.
Stenothermic Organisms that can withstand only a narrow
range of temperature conditions.
Stepping Stones Chains of closely distributed islands or
discrete areas of similar habitat, such as mountains sur-
rounded by low elevation deserts, that aid in the disper-
sal of some species.
Stolon A prostrate stem that runs just at or below the sur-
face. Common for many species of grass.
Stomata Specialized cells on the surfaces of plants that
open to allow the exchange of gasses between the plant
interior and the atmosphere.
Succession The changes in the physical and biological con-
ditions of an ecosystem that follow disturbances.
Succulent Plants that have enlarged leaves and stems that
contain water holding tissue.
Super Tramp Species that are particularly well suited to
long distance dispersal and successful colonization.
Superorganism Community Concept Clements' concept
that the species in communities had evolved together
over long periods of time and were highly dependent
upon each other, much like the individual organs in an
animal depend upon the functioning of all the other
organs.
Surface Fire A fire that is restricted to the ground level
and lower vegetation and does not rise into the tree
canopy if trees are present.
Sweepstakes Routes Geographic routes with low proba-
bility of successful dispersal , but which are occasionally
crossed by species.
Symbiosis The close associati on between two species that
generally develops through coevolution. In many cases,
the symbiotic association is obligatory for the survival
of one or both species.
494 GLOSSARY AND KEY WORDS AND TERMS

Sympatric Speciation The development of new species Troposphere The lowest stratum of the atmosphere that
within the same geographic area as the parent species. extends from the surface to about 9-17 km height. The
Synecology Ecological research which focuses on the troposphere consists of roughly 78% nitrogen, 21% oxy-
interactions between different species in communities. gen, 0.93% argon , and 0.036% carbon dioxide, and a
small amount of other gases and water vapor.
Systematics Taxonomic studies that are directed at deter-
mining the evolutionary relationship between different
groups of organisms.
Umbrella Species Species, which because of their resource
requirements and role in the ecosystem, provide evi-
PHOTO (
Target Area Effect The observed positive relationship dence of biodiversity, habitat diversity, and healthy eco-
between island size and immigration rates due to the logical functioning.
fact that larger islands provide a bigger target for dis- Vicariance Event Geologic events or environmental
persing organisms. changes that divide the ranges of species into geograph-
ically isolated distributions. Chapter 2 Page 11 (left): M
Taxon Any biological taxonomic category such as a family,
genus, or species. The plural of taxon is taxa. Vicariance Biogeography Reconstruction of evolutionary Researchers. Page 11 (right): (
Taxonomy The subdiscipline of biology concerned with histories that assumes that splits in cladograms repre- Researchers. Page 17: Dr. Ken
the classification and naming of organisms. sent vicariance events that split the geographic ranges Researchers.
of taxa.
Temperate Having a geographic distribution in the tem- Chapter 3 Page 44: Courtes
perate mid-latitude regions of the world. Waif Dispersal Successful long distance jump dispersal
52: Gregory Ochocki/Photo R<
events.
Threatened Species A species that is likely to be endan- Christian Grzimek/Photo Res<
gered in the foreseeable future. Wallace's Line The boundary between the Austalasian and
Oriental biogeographic regions. First described by Chapter 4 Page 88: Stuart V
Tillers The photosynthetic stems and blades of grass. Alfred Russell Wallace. Researchers.
Tills (Tillites, Glacial Tills) The poorly sorted mineral mat-
Weathering The physical, chemical, and biological Chapter 5 Page 110: Step he
ter that is weathered, eroded, and deposited by glaciers.
processes that cause rocks to break down into fine par- Researchers. Page 114: David
Tolerance Model A model of succession that assumes that ticles and soluble chemicals. Researchers. Page 121: Fred
the species that will dominate the ecosystem following Xerophyte Plants that require dry soils. Researchers. Page 123: Louisa
a disturbance are all established very quickly following
Xylem Cells in vascular plants that are specialized for the Page 125: Pat & Tom Leeson!f
the disturbance. All species can tolerate the physical
and biological conditions that occur after the distur- flow of water.
Chapter 6 Page 142: Photo
bance. As time progresses some species disappear Xylopdia Large woody subsurface organs found on some Gregory G. Dimijian/Photo R<
because they cannot tolerate the environment of later plants, particularly in the tropical savanna. Courtesy Glen MacDonald. P
successional stages. Younger Dryas A brief period of sharp Degginger/Photo Researchers.
Transpiration The release of water vapor to the air by cooling that affected the northern Atlantic region and Lepore/Photo Researchers. P:
plants. many other parts of the world around 12,000 years ago Suzio/Photo Researchers. Pag
Trophic Cascade Occurs when the loss of an important
(11 ,000 radiocarbon years ago). MacDonald. Page 168: Jim Ste
prey species causes further ecosystem disruptions and Zoochores Organisms dispersed by other organisms. Page 172: George Ranalli!Pho
extinctions because of the loss of food for higher preda- Zoogeography The study of the biogeography of animals. Art Wolfe/Photo Researchers.
tors. Krasemann/Photo Researcher
McHugh/Photo Researchers.
Trophic Levels The hierarchical levels of the food chain
through which energy flows from primary producers to
primary consumers, secondary consumers and so on.