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EBOOKS Information in the Environment
FOR THE Christopher J. Paradise A. Malcolm Campbell

APPLIED BIOLOGY COLLECTION
This book identifies the commonalities between communication
SCIENCES
within a species and communication between species. Behav-
LIBRARY ior and exchange of non-heritable information occurs between
Create your own individuals of different species, in animals and plants, in order
Customized Content to exploit other species and compete for resources. Several
Information in
Bundlethe more examples of adaptations of one species to exploit the informa-
books you buy, tion passed between individuals of another species are given.
the greater your This book describes how animals make decisions while gath-
discount! ering information and resources, selecting habitat, and inter-
acting with potential competitors. Plants grow in response to the Environment
nutrients in soil, which may require gene regulation in response
THE CONTENT
to information in the environment. Information is also exhibited
Energy Physics in biodiversity, in the number and types of species present, and
Engineering this information is used by other organisms as they assess their
Biotechnology surroundings. The information content of ecological systems
Biology changes when species are added or lost.
Mathematics
Christopher J. Paradiseis professor of biology and environ-
Chemistry
mental studies at Davidson College. He teaches introductory
biology, ecology, entomology, and topical seminars on ecotoxi-
THE TERMS cology and renewable natural resources. He also occasionally
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A. Malcolm Campbell
Information in the
Environment
Information in the
Environment
Christopher J. Paradise, PhD

A. Malcolm Campbell, PhD
Information in the Environment
Copyright Christopher J. Paradise and A. Malcolm Campbell. 2016.
All rights reserved. No part of this publication may be reproduced, stored

in a retrieval system, or transmitted in any form or by any means
electronic, mechanical, photocopy, recording, or any other except for
brief quotations, not to exceed 250 words, without the prior permission
of the publisher.
First published in 2016 by

Momentum Press, LLC
222 East 46th Street, New York, NY 10017
www.momentumpress.net
ISBN-13: 978-1-60650-969-2 (print)

ISBN-13: 978-1-60650-970-8 (e-book)
Momentum Press Biology Collection
Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

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Printed in the United States of America.

Abstract
This book identifies the commonalities between communication within
a species and communication between species. Behavior and exchange of
non-heritable information occurs between individuals of different spe-
cies, in animals and plants, in order to exploit other species and compete
for resources. Several examples of adaptations of one species to exploit the
information passed between individuals of another species are given. This
book describes how animals make decisions while gathering information
and resources, selecting habitat, and interacting with potential competi-
tors. Plants grow in response to nutrients in soil, which may require gene
regulation in response to information in the environment. Information is
also exhibited in biodiversity, in the number and types of species present,
and this information is used by other organisms as they assess their sur-
roundings. The information content of ecological systems changes when
species are added or lost.
Keywords
communication, information, resource, natural enemies, predators,
parasites, host, optimal foraging theory, diminishing returns, m
arginal
value theorem, nutrients, d ensity, competition, coral settlement,
biodiversity, composition, global extinction
Contents
Preface...................................................................................................ix
Acknowledgments....................................................................................xi
Introduction.........................................................................................xiii
Chapter 1 Organisms Have Evolved to Exploit Communication
Between Individuals of Other Species.................................1
Cricket Songs Are Exploited by Natural Enemies...............2
Frog Choruses Attract Predators.........................................6
Chapter 2 Organisms Assess Their Environment When
Searching for Resources....................................................13
Optimal Foraging Behavior of Lizards..............................13
Plants Assess their Environment In Search
of Nutrients..................................................................18
Chapter 3 Chemical Communication May be Used to Block
Competition....................................................................25
Egg-Laying Moths Detect Larvae of Other Moths............25
Information Is Used by Corals During Settlement...........30
Chapter 4 A Change in the Number of Species Affects
Information Content of an Ecological System..................37
Ethical, Legal, Social Implications: We Have an
Obligation to Preserve Biodiversity...............................41
Conclusion............................................................................................45
Glossary................................................................................................47
Index....................................................................................................49
Preface
This book about information in the environment is part of a thirty book
series that collectively surveys all of the major themes in biology. Rather
than just present information as a collection of facts, the reader is treated
more like a scientist, which means the data behind the major themes are
presented. Reading any of the thirty books by Paradise and Campbell
provides readers with biological context and comprehensive perspective
so that readers can learn important information from a single book with
the potential to see how the major themes span all size scales: molecular,
cellular, organismal, population and ecologic systems. The major themes
of biology encapsulate the entire discipline: information, evolution, cells,
homeostasis and emergent properties.
In the twentieth century, biology was taught with a heavy emphasis
on long lists of terms and many specific details. All of these details were
presented in a way that obscured a more comprehensive understanding.
In this book, readers will learn about information in the environment and
some of the supporting evidence behind our understanding. The historic
and more recent experiments and data will be explored. Instead of believ-
ing or simply accepting information, readers of this book will learn about
the science behind information in the environment the way professional
scientists dowith experimentation and data analysis. In short, data are
put back into the teaching of biological sciences.
Readers of this book who wish to see the textbook version of this
content can go to www.bio.davidson.edu/icb where they will find
pedagogically-designed and interactive Integrating Concepts in Biology
for introductory biology college courses or a high school AP Biology
course.
Acknowledgments
Publishing this book would not have been possible without the generous
gift of Dr. David Botstein who shared some of his Breakthrough Prize
with co-author AMC. Davids gift allowed us to hire talented artists (Tom
Webster and his staff at Lineworks, Inc.) and copyeditor Laura Loveall.
Thanks go to Kristen Mandava of Mandava Editorial Services for project
management and guidance. In particular, we are indebted to Katie Noble
and Melissa Hayban for their many hours and attention to detail.
Kristen Eshleman, Paul Brantley, Bill Hatfield and Olivia Booker
helped us with technology at Davidson College. We are grateful to ad-
ministrators Tom Ross, Clark Ross, Carol Quillen, Wendy Raymond,
Verna Case, and Barbara Lom who had confidence in us and encouraged
us to persist despite setbacks along the way.
Thanks to my wife Amy Brooks for her constant support during the
development of this textbook, and my daughter Evelyn for her endless
energy. Thanks to Malcolm Campbell for his steadfast resolve and opti-
mism. Without him, this book would not exist. Thanks to collaborator
Laurie Heyer for taking my sometimes half-baked math ideas and turn-
ing them into powerful and elegant Bio-Math Explorations. I learned a
lot from both of them. While the math is largely absent from this book,
our collaboration with her made this a better book. Nancy Stamp at
Binghamton University, and Bill Dunson and Richard Cyr at The
Pennsylvania State University influenced me greatly in how I think as
a scientist and approach my teaching. Finally, I thank my students in
Integrated Concepts in Biology II, who enthusiastically participated in
our experiment to redesign introductory biology, starting with the text
and ending with a new approach to teaching biology.
Introduction
Communication is important to everyday life, and allows people to gather
and distribute information. Handwritten private notes, emails or cell
phones are often used to exchange information with only certain people,
with potentially drastic consequences of those private exchanges being
intercepted by someone else. Imagine how much more difficult and dan-
gerous life would be if the only way to communicate was to broadcast
information by shouting out loud or posting on an open forum on the
Internet in hopes that the intended recipient got the message. That is what
life is like for many species.
The book Behavior and Information Exchange, part of this series of
digital books, explored several examples of information exchange be-
tween individual animals and plants within the same species. What hap-
pens when these communications are intercepted? Ecological systems,
which are environments in which organisms live and interact with each
other and with nonliving components, are teeming with information.
Interactions between species, communications between individuals, and
substances in the environment all contribute to the information in the
system. In this book, how plants and animals exploit the information that
they gather from their environment and prevent being exploited them-
selves is explored. It will also show how sensitive an ecological system can
be to the presence or absence of a single species and show how to measure
the effects of extinction or the introduction of a species.
CHAPTER 1
Organisms Have Evolved

to Exploit Communication
Between Individuals
of Other Species
The mechanisms and purposes of communication between individuals

of the same species include species recognition, mate/sex recognition,
mate attraction, defense against threats, facilitation of cooperation, and
warning of danger, to name just a few. These signals are sent into the
environment with some risk. One risk in communicating to find a mate
or to announce location is that another species could perceive and re-
spond to the communication and use the communicator as a resource,
say for a meal. A resource is a natural feature or organism used as a
supply of energy or nutrients to other organisms. A risk to male fireflies
when flashing their double pulse of light is that a predator could use
the light to locate the male. The interceptors of the signals are natu-
ral enemies of the signaler and can be classified as either predators or
parasites. Natural enemies are organisms that use other organisms as a
resource. Predators are organisms that kill and feed on other organisms
and include lions, hawks, snakes and dragonflies. Parasites are organisms
that live on or in another organism, a host and cause harm to the indi-
vidual by using them as a resource. Parasites include tapeworms, ticks,
and mosquitoes. In this chapter, the ability of predators to locate prey
and parasites to identify hosts by intercepting communications will be
investigated.
2 INFORMATION IN THE ENVIRONMENT
Cricket Songs Are Exploited by Natural Enemies

Prey and hosts of predators and parasites are often insects. Crickets and
grasshoppers are insects found in many areas around the world, being es-
pecially common in tropical, subtropical, and temperate zones (Figure 1).
Males of many species of crickets and grasshoppers make vocalizations to
attract mates, much like male fireflies flashed light to attract their mates.
They can be heard chirping all day and night at certain times of the year.
Mole crickets are a particular type of cricket with large shovel-like fore-
legs adapted for burrowing into the ground (Figure 1). These crickets are
nocturnal and spend much of their time underground. They are known
to fly during the mating season, which is often the only time people ever
see them. Both males and females can fly, but only males emit mating
vocalizations. Several scientists have studied vocalizations of males and
responses of females.
In one study, S. Ulagaraj and Thomas Walker studied two species of
mole cricket, the southern mole cricket (Scapteriscus borellii) and the tawny
mole cricket (S. vicinus) in Florida. They recorded male vocalizations of
these two cricket species and broadcast the songs through speakers in a
playback experiment. Each speaker was mounted in the center of a large
funnel and faced skyward (Figure 2). One funnel contained a speaker
playing southern mole cricket songs, another funnel contained a speaker
playing tawny mole cricket songs, and a the third funnel contained no
speaker.
Figure 1 A katydid (left) and a mole cricket (right). Note the

variation in the legs of these two species.
Source: A, http://upload.wikimedia.org/wikipedia/commons/5/57/Katydid_tx.jpg; public domain.
2007 B, http://en.wikipedia.org/wiki/File:Scapteriscus_vicinus.JPG; Author: Ilona Loser, 2009.
This file is licensed under the Creative Commons Attribution-Share Alike 3.0 Unported license.
ORGANISMS EXPLOIT COMMUNICATION OF OTHER SPECIES 3
funnel with
no speaker
funnel with
speaker
funnel with
speaker
collecting jar
speaker wire
Figure 2 Experimental design for testing mole cricket response to

vocalizations.
Source: Based on Ulagaraj & Walker, 1973, Figure 1.
600
number of individuals captured at traps
= southern males
500 = southern females
= tawny males
400 = tawny females
300
200
100
0
southern tawny control
male call male call (no sound)
Figure 3 Responses of southern and tawny mole crickets to broadcast

recordings of male calls of both species and a control broadcasting no
sound. Bars indicate crickets captured in traps broadcasting male calls.
Source: Modified from Ulagaraj & Walker, 1973, Figure 2A.
At the base of each funnel was a jar, where adults that flew into the
funnel were collected. Adults collected from each funnel were brought
back to the laboratory, identified, and their sex was determined (Figure 3).
Male crickets use sound to attract mates, and this communication is

used for species and mate recognition. Male mole crickets have vocaliza-
tions that are responded to by members of their own species, as the data
show and as is known for many other species. Most of the individuals that
came to a particular speaker were individuals of the species being broad-
cast from that speakermore than 80% in every case. This indicates that
the vocalizations are specific to a species.
In addition, most of the mole crickets trapped at speakers were females,
suggesting that the function of the males vocalizations is to attract females.
Other males may approach the vocalizations of males to check them out.
A scientist might hypothesize that vocalizing males have a location that
is good for vocalizing or that a vocalizing male may have already brought
several females near to his location, which increases the chances of a second
male finding a mate if that second male approaches the vocalizing male.
The sounds that crickets and grasshoppers make are quite loud, and
when a person walks by a chirping cricket it will stop making those vocal-
izations. The benefit of making the vocalizations is that, hopefully, they
will attract mates. The cost is that they will attract predators or compet-
ing males. From the vocalizing male crickets perspective, a person is a
potential predator, so their pause in vocalizing is a mechanism to avoid
detection by passing predators.
Just as crickets have adaptations to detect animals approaching them,
other animals have adaptations to detect and respond to the males mat-
ing vocalizations. A variety of parasitic insects use crickets and grasshop-
pers as hosts for their developing offspring. These insects lay their eggs
on crickets and grasshoppers, often on particular species. Then the eggs
hatch and the larvae, the immature, wingless, and often wormlike forms
of many insects, burrow into the host. The larvae live off the host tissue,
usually killing it by the time they finish developing. For example, the
larvae of tachinid flies, which resemble small house flies, live inside an-
other animal as larvae, usually another insect. Many of these flies parasit-
ize crickets and grasshoppers.
In order to successfully reproduce, parasitic flies require hosts on
which they lay eggs. The particular species of flies that lay eggs on mole
crickets have evolved in such a way that adults are active at the same time
of year that male mole crickets are vocalizing to attract females. Another
requirement for success is to lay eggs quickly and fly away before the mole
cricket has an opportunity to defend itself or escape from the fly under-
ground. Once the fly recognizes an appropriate host by its call, it makes
the decision to land and lay eggs, which it does quickly.
A simple playback experiment was designed by Harold Fowler to de-
termine the extent that one species of fly (Ormia depleta) was attracted to
the vocalizations of the southern and tawny mole crickets, along with the
imitator mole cricket (S. imitatus). Fowler set up playback devices with
speakers, spaced 50 meters apart, and each night for a full year, he played
the songs of the three species (Figure 4).
100
90
= tawny
80 = southern
number of flies captured
= imitator
70
60
50
40
30
20
10
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
month
Figure 4 Monthly captures of female parasitic flies at speakers

playing the male calls of three species of mole cricket, the tawny,
southern, and imitator.
Source: Data from Fowler, 1987, Table 1.
The speakers trapped flies that landed to investigate the sound. Fowler
recorded the number of flies landing according to the month and the spe-
cies of male mole cricket vocalization. He also observed the behavior of
flies around speakers and noted that flies that landed deposited eggs and
left in 3 seconds or less.
Fowler next played the vocalizations of five species of mole cricket dur-
ing the peak activity period of the fly (Table 1). He included the southern,
tawny, and imitator mole crickets, and he added two more species, the
Table 1 Numbers of flies captured at traps playing vocalizations from

one of five species of mole cricket.
mole cricket call number of tachinid flies
southern 24
tawny 51
imitator 33
changa 0
northern 0
Source: Data from Fowler, 1987, text p. 476.
changa (S. didactylus) and the northern mole cricket (Neocurtilla hexa-
dactyla). He played recorded vocalizations of each of these species for ten
nights during the peak annual activity period of the parasitic fly and col-
lected all flies attracted to the mole cricket vocalizations.
The mole cricket/tachinid fly interaction is one example among many
of a natural enemy that intercepts the communication of another spe-
cies, illustrating how information is collected from the environment to
obtain resources. The particular species of parasitic fly discussed here has
evolved to recognize vocalizations of at least three species of mole cricket,
but it did not appear to recognize the vocalizations of two other closely
related mole cricket species. All five species of mole cricket may be found
in Florida, but the three to which the flies are attracted arrived in Florida
from South America. The parasites may have evolved a mechanism to find
their hosts if the parasites and hosts coexisted in the same habitat, but if
there is no shared evolutionary history, there would be no natural selec-
tion for flies that could recognize the vocalizations of the host. Therefore,
it might be speculated that the flies have also invaded North America.
Alternatively, perhaps the changa and northern mole crickets have adapted
their vocalizations to be unrecognizable to the flies and have escaped from
that natural enemy.
Frog Choruses Attract Predators

Other natural enemies, such as predators, may also exploit communication
signals to locate prey. To better understand these interactions among ver-
tebrates, animals with an internal skeleton made of bone and a backbone,
investigators led by Merlin Ryan studied frogs in Panama. Male frogs,

like male mole crickets, vocalize to attract mates, and they often do so in
large choruses where many males are located along the shores of a wetland
or pond. Once a female chooses a male, mating takes place in the pond.
The benefits of vocalizing are that males will win mates and reproduce.
The potential cost, as for insects, is discovery by natural enemies. The
biologists were interested in the abilities of predators to find vocalizing
prey, and whether predators were adapted to cue in on particular species.
Ryan and colleagues studied two predators of a community of frogs
that congregate around ponds in South America, the philander opos-
sum (Philander opossum, a small 30 cm long opossum with distinctive
white patches above its eyes) and the fringe-lipped bat (Trachops cirrhosus;
Figure 5 inset).
In the first study, the researchers observed opossums near a pond that
contained a breeding male chorus of Tungara frogs (Physalaemus pustulo-
sus), which are small brown frogs that resemble toads. Because the frogs
vocalize after sunset and the opossum is nocturnal, they used a night
vision scope to observe predator-prey interactions. They observed ponds
at night and recorded thirty-nine captures. Typically, the opossum ap-
proached the pond, stopped, and then turned toward the vocalizing frogs.
As the opossum got closer, it stopped again, often turning its head from
side-to-side and rotating its ears. Then it ran forward and pounced, at-
tempting a capture.
The researchers performed a playback experiment where they played
recordings of Tungara frog vocalizations from a speaker located 2 meters
from the edge of the ponda location where opossums did not nor-
mally venture and frogs were not found. The recordings were played only
when an opossum was nearby but facing away from the speaker and no
frogs were vocalizing. In five out of five trials, the opossum turned toward
thespeaker and performed the head tilting and ear rotating behaviors.
The opossum then approached the speaker and pounced on the speaker
three out of five times. Even after an opossum attacked the speaker with-
out reward of a meal, it continued to circle the speaker if the playback was
left on. In the two trials where the opossum did not attack the speaker, it
had spotted the researchers, which scared the predator away.
60
leading to capture of a frog 50

percentages of bat visits
40
30
20
10
0
full chorus partial chorus few calling none calling
A
100
approaches to vocalizations
80
percentages of
60 = cage
= field
40
20
0
edible poisonous small large
frog toad frog frog
B prey vocalization type
Figure 5 Results of experiments on fringe-lipped bats preying on

frogs. A, Percentage of bat visits resulting in a frog capture for four
categories of Tungara frog vocalization frequency. Inset, A fringe-
lipped bat. B, Responses of frog-eating bats to vocalizations of edible
versus poisonous or small versus large frog or toad prey.
Source: From Tuttle and Ryan, 1981, Table 1. Photo inset: Author: Karin Schneeberger. This file
is licensed under the Creative Commons Attribution 3.0 Unported license.
Although the sample sizes for this experiment were small, the results
offer compelling evidence that nocturnal predators exploit acoustic com-
munication within other species to locate prey. Even more impressive data
would be evidence that a predator could distinguish vocalizations from
different species of frog. Because some frogs that vocalize are poisonous,
Ryan and his colleagues set up an experiment to test the reactions of a

predator, the fringe-lipped or frog-eating bat, to vocalizations from differ-
ent frog species; some poisonous and some not. Most bats use echoloca-
tion, or sonar, to locate prey, but this species does not. The fringe-lipped
bat locates prey by the sound that the prey makes, and they specialize in
eating frogs.
Ryan and colleagues visited frog breeding ponds between January and
June, where they observed the fringe-lipped bat hunting. Half of the ponds
were breeding ponds of the Tungara frog. At these locations the research-
ers categorized their observational times by the amount of vocalizations
coming from the male frogs (Figure 5A). The categories included a full
chorus when many males were vocalizing simultaneously, a partial chorus
when several males were vocalizing, times when only a few were vocalizing
intermittently, and finally, when no males were vocalizing. The researchers
determined the number of bats that visited during time periods in each
category and tracked how successful they were at capturing frogs.
The researchers then exposed captive and wild frog-eating bats to vo-
calizations of an edible versus a poisonous frog, or a frog small enough
to eat versus one too large for a bat to capture (see Figure 5B). Data were
recorded for captive bats in cages and free (wild) bats in the field.
The observations of opossum behavior and the data in Figure 5 indi-
cate that both opossums and frog-eating bats are using vocalizations to lo-
cate prey. It was long hypothesized that there is a potential predation cost
to vertebrates that advertise their presence to attract mates, but Ryan and
his colleagues demonstrated it first in 1981. The confirmation of this hy-
pothesis should not be very surprising to careful observers of hunting ani-
mals. A predator that preys on frogs may be cueing in on the sound, which
is exploitation of information transfer, but the exploiter is not using the
information to identify the species, only to locate potential prey. Opos-
sums could not locate their prey in the dark without the frogs vocalizing,
and bats were much less successful during visits to ponds when no frogs
were vocalizing. Success was much higher when frogs were vocalizing.
One surprising result of Ryans experiments was that fringe-lipped
bats could discriminate between vocalizations and knew the vocalizations
of their preferred prey. There were strong preferences for the edible frog
vocalization over the vocalization of the poisonous toad. When given a
choice of frogs of different sizes, bats preferred the vocalizations of frogs

that were not too large, presumably because very large frogs would be
hard for a relatively small bat to carry.
The overly large and the poisonous frogs incur little cost in making
themselves as conspicuous as possible, because this will make it easier for
females to find them. The other frogs and toads do not make themselves
very conspicuous, hiding in the bushes and ceasing their vocalizations
when they detect a nearby threat. One of the advantages of communica-
tion within species is that individuals can successfully find a mate. How-
ever, if they are not protected by toxins, large size, or some other defense,
then the advantages are balanced by predation risk.
Experiments like these are important to carry out in the laboratory as
well as the field. In the laboratory, extraneous variables can be controlled,
whereas field studies are crucial in order to corroborate and increase the
validity of the controlled laboratory studies. The preferences seen in cap-
tive bats could have been caused by some behavior exhibited or learned
by the bats either in the wild or in captivity, so it was important to carry
out the experiment in both locations. The fact that both groups responded
in the same manner added to the credibility of the study. Yet a person may
wonder whether fringe-lipped bats are born with the ability to recognize
different species of frogs or if they learn it from other bats.
Continuing with his studies of the frog-eating bats, Ryan and another
colleague sought to determine how bats learn which vocalizations are
made by frogs that they can eat and which are made by frogs that are too
big or poisonous. The fringe-lipped bat lives in social groups, which the
researchers predicted would be important for learning in the bats. In their
study, the scientists played the vocalization of cane toads (Bufo marinus)
for the bats. This species of toad is too large for the fringe-lipped bats to
eat and is very poisonous.
The biologists studied caged bats to determine the conditions under
which bats could learn to associate the vocalizations of the cane toad with
a palatable food reward, that is, it was agreeable or acceptable to consume.
First, they tested the caged bats and found that none were initially at-
tracted to cane toad vocalizations. Next, they trained a group of bats to fly
to and land on a speaker that was broadcasting the cane toad vocalization.
When they landed on the speaker, they were given a reward of raw fish.
Once these bats became experienced with cane toad vocalizations, they
were paired up with bats that had never experienced these vocalizations.
In each pair, the inexperienced bat observed the experienced bat. The
researchers also presented cane toad vocalizations to pairs of two inexpe-
rienced bats and to single inexperienced bats with no other bats present.
Bats were subjected to a number of trials. In each trial, between one
and five speakers were set up with one randomly chosen one playing the
cane toad vocalization. The researchers determined the average number
of trials needed for each inexperienced bat to associate cane toad vocaliza-
tions with food (Table 2). The criterion for learning was the bat flying
to and landing on the speaker broadcasting cane toad vocalizations and
getting a piece of raw fish three trials in a row. In all trials, the bats ap-
proached only speakers that were broadcasting vocalizations, and they
never ate fish from a location other than a broadcasting speaker.
By setting up appropriate controls, the biologists reduced bias in their
experiment. By randomly shuffling the location of the cane toad vocaliza-
tion speaker, the researchers ensured that bats were not learning that a
food reward was in a particular place. Other speakers were used to control
for potential bias of the speaker itself, and the various treatments were
used to demonstrate that inexperienced bats were learning from experi-
enced bats.
This study showed that fringe-lipped bats, which live and forage in
small social groups, can learn from their neighbors, and they can associate
a new frog vocalization with edible prey by watching what experienced
neighbors eat. Rates of learning were faster for inexperienced bats when
they were paired up with experienced bats that had learned that cane toad
vocalizations were associated with edible prey. This begs the question of
Table 2 Number of trials needed for fringe-lipped bats to learn to

associate the cane toad vocalization with palatable prey.
treatment mean number standard sample
of trials error size
inexperienced bat
5.3 1.7 10
with experienced bat
two inexperienced bats 96.8 3.2 5
one inexperienced bat 96.2 3.8 5
Source: From Page and Ryan, 2006, Figure 2.
how bats learn to recognize new edible prey in the first place. In the wild,
bats will occasionally approach new frog vocalizations and attempt to eat
the new prey type. This species of bat most likely has an innate, or present
from birth, ability to find frogs through their attraction to frog vocaliza-
tions, but they also have the ability to learn to discriminate among vocal-
izations of palatable and unpalatable frogs.
Information from other species, much like the information that passes
between members of the same species, can lead to changes in the behav-
ior of the receiving organism. Both parasites and predators can intercept
information exchanged between members of other species and use it to
their advantage. The interception of information may even be species-
specific, in that the natural enemy is using the information for recognition
of particular prey or host species. While many natural enemies intercept
information sent within a species, many other organisms obtain food and
other resources without perceiving other species communications. These
organisms use other types of information. In the next chapter, how some
organisms gather information from their environment while searching for
resources and how they attempt to maximize benefits and minimize costs
will be examined.
Bibliography
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dae): phonotactically orienting parasitoids of mole crickets (Orthop-
tera: Gryllatalpidae: Scapteriscus), J NY Entomol Soc 95(4):474480,
1987.
Page RA, Ryan MJ: Social transmission of novel foraging behavior in bats:
frog calls and their referents, Curr Biol 16(12):12011205, 2006.
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philander opossums, Biotropica 13:233234, 1981.
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and female crickets to male calling songs, Science 182(4118):1278
1279, 1973.
CHAPTER 2
Organisms Assess Their

Environment When
Searching for Resources
Animals can use the communications of other species to help them find
prey, as shown in the previous chapter. But there is more to information
than just intercepting signals of other species. Many organisms find prey
or resources by gathering other information from the environment. In-
formation is crucial to the survival of all organisms. The better informed
an individual organism, the better it can meet the demands of a vari-
able world where resources can be difficult to obtain. Uncertainty in the
environment poses significant problems for life, but uncertainty can be
reduced by the acquisition of information. Gathering and processing in-
formation on a continuous basis can allow exploitation of resources and
avoidance of potential harm, and this is what will be explored in this
chapter.
Optimal Foraging Behavior of Lizards

Prey do not want to signal their presence to foraging animals searching
for food. Potential prey benefit if they keep their presence hidden when a
predator is nearby. This extra complication to foraging requires predators
and prey to make decisions about where, when, and how to search for
food, and what to eat. For example, prey are often found in discrete habi-
tat types, or patches, and animals must gather information to decide how
long to forage within a patch. Many studies of how animals search for
food have tested hypotheses of optimal foraging theory, the theory that
evolution of animals results in their foraging in a manner that maximizes
their rate of energy intake per unit time spent feeding. Optimality implies
that the animal is attempting to maximize something relevant to the ani-
mals success, in terms of survival and production of offspring. Decisions
that animals make when foraging will affect their success.
In one study of optimal foraging in patches, James Munger worked
with two species of horned lizards (the Texas horned lizard, Phrynosoma
cornutum, and the round-tailed horned lizard, Phrynosoma modestum,
that live in the deserts of the southwestern United States. The lizards feed
on ants, which are found concentrated mostly at their nests. Each lizard
in the study was followed for an entire day. Munger knew from the previ-
ous day where an individual had sought shelter the previous night, and
he was at the shelter in the morning before the lizard became active. He
followed a lizard from a distance and observed its foraging activity, often
with binoculars, until it sought shelter during the heat of the day.
Munger resumed observation when the lizard came back out in the
cooler, late afternoon. Munger meticulously recorded all feeding activity
at each nest by speaking into a recorder, which allowed him to note every
ant eaten, how quickly they were eaten, and how long the lizard remained
at a nest. He created a model of foraging that described the cumulative
number of ants captured in a patch. The foraging data helped Munger
test his hypothesis that lizards were exhibiting optimal foraging behavior.
Munger used a model of foraging given by the general form of the equa-
tion y = 118.6*(1 e0.00084x). Munger found a polynomial to fit the data
for each lizard, based on their foraging at an ant nest; the one shown here
is one example from one lizard.
Each exponential equation depicts the relationship between the cu-
mulative number of ants eaten (the dependent variable, y) and the time
the lizard spent at the nest (the independent variable, x). To better un-
derstand the relationship between ants eaten and time spent at the nest,
consider a hypothetical nest. In a fixed amount of time, say 60 seconds,
the lizard can search a fixed percentage of the nest and capture a fixed
percentage of the ants in the area it searches. Therefore, assuming the
susceptible ants are uniformly distributed around the nest, the lizard
would capture a fixed percentage of the total number of remaining sus-
ceptible ants every 60 seconds. The number of ants captured per minute
steadily decreases over time, which is a concept sometimes referred to as
ORGANISMS ASSESS THEIR ENVIRONMENT WHEN FORAGING 15
diminishing returns. As the number of ants remaining decreases, it gets

more and more difficult for the lizard to capture them.
Once Munger had this model for the foraging process, predictions can
be made of when the lizard should leave the nest to optimize its resources.
The longer a lizard stays at a nest, the slower the rate of capture becomes.
Under these conditions, the marginal value theorem states that the opti-
mal time for a lizard to leave a nest is precisely when the rate of capture at
that nest slows down to the overall average rate of capture at all nests in the
habitat. The marginal value theorem predicts that optimal foragers leave
a food patch when the rate of capture at that patch falls below the overall
habitat rate. Leaving earlier or later than the optimal time would decrease
the total number of ants caught each day. Leaving earlier than the optimal
time, while the rate of capture is still above the overall average, would de-
prive the lizard of some of its best foraging; spending less time foraging at
a higher than average rate would lower the overall average. On the other
hand, leaving later than the optimal time lowers the overall average, because
all foraging after the optimal leaving time is below the average capture rate.
Therefore, the optimal behavior for the lizard is to do all of its forag-
ing at an above average rate. This seems impossible until it is realized that
because the lizard forages in patches, the travel time between patches is
completely unproductive, but is still included in the overall average rate
calculation. Because the capture rate is zero during travel between nests,
the lizard has to make the most of its time at each nest, catching enough
ants so that the overall average (considering time at the nest plus time in
transit) stays as high as possible. The marginal value theorem states that to
keep the overall average as high as possible, the lizard should stay at each
nest until the current capture rate decreases to the overall average rate,
but no longer. The most accurate estimate of the capture rate at any time
is the slope of a line that touches, but does not cross, the feeding curve at
the point (i.e. the slope of the tangent to the curve at that point).
Optimal foraging behavior is modeled by mathematical equations,
but lizards are not doing a bunch of calculations while they roam around
their habitat. However, because there is a selective advantage for lizards
with optimal behavior, evolution has resulted in a set of behaviors that
are nearly optimal. Mathematical models can be used to predict foraging
behavior and its evolution.
The marginal value theorem predicts the optimal time for a forager
to leave a patch based on its rate of feeding in a patch and its expectation
of whether it could find more food by moving to another patch. The
expectation of doing better somewhere else is based on information that
the individual has collected over time while feeding in different patches.
The quality of a patch is thus shown to affect the optimal leaving time.
Munger tested predictions of the marginal value theorem of how long
a forager should remain in patches with various amounts of resources. He
converted the number of ants consumed by each lizard at each nest to
mass by determining the average biomass of ants. For each lizard in the
study, he calculated its overall habitat rate of energy intake, calculated as
the mean mass of ants consumed (in milligrams) per 1,000 seconds.
Munger then plotted the consumption rate at departure for each
lizard at each nest against that lizards overall habitat rate. Because the
researcher wanted to determine whether lizards were optimal foragers,
he plotted the data for all lizards but noted that some lizards left a nest
for reasons apparently unrelated to maximization of energy intake. For
instance, some were observed to move immediately to shade, presumably
to cool off. Munger determined the best fit line when considering all ant
nest visits and when considering the subset of visits in which the lizard
did not leave early due to heat or other factors (Table 3).
He then calculated the average consumption rate at the time of depar-
ture for all ant nest visits in which the lizard did not leave the nest early
for reasons unrelated to energy maximization. The best fit line for those
eight lizards had a slope of 0.84 (Table 3).
The data show that the rate of prey capture decreased the longer the
lizard remained at an ant nest. If lizards forage optimally, then the mar-
ginal value theorem predicts that they will leave a nest when their in-
stantaneous capture rate at a nest is equal to the mean capture rate for
the entire habitat, which was averaged across all nests. Foragers gather
information as they move through the habitat, but they only have infor-
mation on patches they have visited: They cannot know the size of ant
nests that they have not visited. But the marginal value theorem predicts
that the slope of the line for rate at departure versus overall habitat rate
(Table 3) should be 1. Munger found that the capture rate at the time of
departure for each lizard at each ant nest was approximately the same as
Table 3 Slopes of rate of departure for horned lizards leaving ant nests
versus the overall habitat rate of each lizard. The capture rate at the
time of leaving a nest is compared to the average capture rate during
foraging for each lizard.
condition slope sample size
rate of consumption at departure
1.04 36
from ant nest versus overall habitat rate
rate of consumption at departure from
ant nest versus overall habitat rate for 0.76 19
all visits in which lizard did not leave early
average individual capture rate at
departure from an ant nest for each lizard 0.84 8
versus overall habitat rate for each lizard
Source: From Munger, 1984, Figures 3 and 5.
the overall average capture rate for that lizard, supporting the marginal
value theorem.
Munger noted two explanations for the slopes that are less than 1 in
Table 3. First, they were not statistically different from 1. In any experi-
ment there is variation in the data caused by experimental error; there was
enough variation in the data for the statistical analysis to conclude with
95% confidence that the slopes were not different from 1.
The second explanation Munger noted was that he only calculated
the overall habitat rate while lizards were at an ant nest, and the marginal
value theorem makes predictions for rates of energy intake for foragers
at the patch as well as while traveling between patches. If the researcher
had included the time traveling, the overall habitat rate of intake would
have been lower. That would shift all the values on the x-axis slightly to
the left, which would have a net effect of raising the slope of the best fit
line. The higher slope for the full data set can be explained by the fact that
individual visits that were interrupted were included. If a lizard began
foraging at a nest and had to leave early, it would likely have a high forag-
ing rate upon departure, which would shift values up on the y-axis and
therefore increase the slope of the line.
Lizards not only gather information about the number of ants and
how quickly they ate them, but they also collect information over a lon-
ger period of time about the average rate of ant capture and the expected
time to travel between nests. If the overall capture rate was low, lizards
did not expect to find many ants at any one nest, and generally left a nest
relatively quickly. There was no point in a lizard staying at a low-quality

nest if it could possibly do better by leaving and searching for another
nest that might have more ants. The foraging lizard was integrating two
pieces of information: the current rate and the long-term average rate.
The long-term average rate may be assessed on a daily basis by lizards, or
perhaps over a longer period of time, although the current rate was close
to meeting predictions of the theory.
Lizards that left an ant nest for reasons other than having a lower than
average capture rate were also using information to make decisions about
behavior. Munger cited three reasons why a lizard left early: 1) it went
immediately to shade, implying that its body temperature was too high,
2) it stopped eating altogether, implying that its stomach was full, and
3) another lizard arrived on the scene. In each of these cases, information
from either the lizards internal or external environment was integrated
into the decision-making process. Lizards are ectotherms and use behav-
ior to regulate their body temperature. They seek shade when their body
temperature is too high, and this requires the ability to assess information
about their body temperature and the temperature of the environment.
If they are full, their stomachs send information to their brain, which
causes them to stop eating. Most readers would be familiar with this sen-
sation. Full lizards may leave an ant nest early, whereas extremely hungry
lizards may stay longer than one would expect. Finally, if another lizard
arrives, the lizard must assess its own status: Is it big enough to fend off
the intruder, or should the feeding lizard abandon the nest? Decisions are
based on information gathered from the environment. Other animals use
similar information and decision-making processes for foraging, finding
mates, and performing a range of behaviors.
Plants Assess their Environment In Search of Nutrients

All organisms assess their environment in some fashion. The mechanisms
are quite varied. For example, plants assess their environment and gather
information about the location of nutrients, food or chemicals that or-
ganisms obtain from their environment and need to live and grow. But
the manner in which information is gathered and resources are acquired
is very different for plants than for animals. Like resources for animals,
plant resources are often located in patches. Consider the soil in which a
plant is growing. When a handful of soil is examined, it probably looks
the same to the average observer, but a persons perception of the soil is
much different than that of a plants. The plant did not choose to be in
that particular place; the seed from which it grew found its way there on
the wind or perhaps on the fur of some animal. To survive where it lands,
a seedlings roots must have ways to sense where nutrients are within the
soil and then gather them. But unlike animals that can move to a new
patch when resources are depleted, plants are stuck in one place, and they
must deal with soil that may be resource-poor on one side and resource-
rich on the other.
Two scientists, Rebecca Farley and Alastair Fitter, assessed the varia-
tion in soil in a forest in England. They selected four small plots that were
2 meters apart along a straight line. Each month for a year, they removed
a core of soil from four randomly selected 10 cm2 locations within each
plot. For each sample, Farley and Fitter analyzed the nutrients nitrate
(NO32), ammonium (NH4+) and phosphate (PO432). Ammonium is
an ion of NH4+ derived from ammonia by combination with a hydrogen
ion. Nitrate is an ion of NO32 and has a negative one charge. Phosphate
is an ion of PO432 derived from phosphoric acid H3PO4. For a shorter
period of time, Farley and Fitter also examined nutrients in smaller plots
closer together. With these two spatial scales, they examined key plant
nutrients over time and space.
Nutrients such as nitrogen and phosphorus can enter soil with rain,
when an animal urinates or defecates, or when an organism dies and de-
cays. Nutrients are released during decay and are washed into the soil.
A decaying animal carcass causes soil nutrient variation in one spot over
time. Two meters away, there may be no carcass. The decaying animal car-
cass also leads to soil nutrient variation from place-to-place and can lead
to patches of high nutrients. Farley and Fitter showed high levels of varia-
tion at a time scale of 1 month as well as a spatial scale as small as 20 cm.
Data for phosphate and ammonium showed similar levels of variability.
Once natural temporal and spatial variation was shown for nutrients
in the soil, the next logical experiment for Farley and Fitter was to deter-
mine how plants respond to these variations. Their experimental setup
involved seven flowering plant species. The plant species were all small,
ranging in height from 5 to 100 cm. They grew these harvested plants in
a greenhouse and allowed them to reproduce. The offspring were then
transplanted to the center section of rectangular pots that measured
20 centimeters long x 5 centimeters wide x 4 centimeters deep.
Pots were subdivided into sections by coarse netting to help maintain
distinct soil patches. The netting allowed roots to grow through but did
not allow for soil mixing. Seedlings were placed in the 4 cm wide center
section. Each pot had a nutrient-enriched treatment patch on one side
of the center section and a non-enriched control patch on the other side.
These two patches were always the same size, such that the pots were sym-
metrical on either side of the center section.
The researchers tested small (2 cm), medium (3.5 cm), and large (8 cm)
nutrient-enriched patches. Non-enriched areas and areas outside the test-
ing zone were filled with sand, and nutrient-enriched areas were filled
with one of two treatments: 100% soil, or 50% soil and 50% sand. Four
pots were set up for each combination of species, nutrient level, and patch
size (4 pots x 7 species x 2 nutrient levels x 3 patch sizes = 168 pots).
Concentrations of ammonium, nitrate, and phosphorus were deter-
mined in each type of growth medium. The researchers found that all
three nutrients increased in concentration as the soil type went from sand
to 50% soil to 100% soil. The nutrient concentrations in sand were all
close to 0 and the concentrations in 50% soil were about half that of
100% soil, as one would expect.
Farley and Fitter let the seedlings grow for 6 weeks and then harvested
the plants by cutting out each pot section and extracting the root frag-
ments. They washed the soil and sand off the roots and made digital im-
ages of the roots. A computer program determined the total root length
in each section of the pot and the root length density, equal to length di-
vided by section volume. The researchers defined the response of the plant
to be the difference between root length density in the treatment and root
length density in the control (non-enriched) patch. They also measured
the biomass that was in the roots relative to the total plant biomass.
Biomass is the total amount of organic matter in an organism, popula-
tion, or other ecological system after water is removed. They compared
the response and relative biomass measurements for the seven species
across the two nutrient-enrichment levels.
On average, across all 7 species, the researchers found that root bio-
mass as a proportion of the entire plant biomass was 1.8 to 3 times as
great in the 50% soil treatments as in the 100% soil treatments. Plant
nutrients as already shown are found in variable concentrations in the
soil. Plants gather information from their environment and respond to
patches of low nutrient concentrations by growing a higher fraction of
their total biomass as roots, presumably to maximize nutrient acquisi-
tion. In high nutrient soils, plants do not have to allocate as much of their
growth to roots because nutrients are easy to capture with fewer roots.
However, when examining the responses of the plants as the difference
between root length density in the treatment and root length density in
the control patch, two species had no response to increased nutrients.
Two other species had equal responses and increased their root length
density the same amount whether an individual was exposed to 50% or
100% soil, relative to the control. Several species had an increased re-
sponse to higher nutrient concentrations over their response to low nu-
trient concentrations. Even if there is a lower proportion of biomass as
roots in the high nutrient soil, as indicated by the root biomass data, the
roots are denser for these species in the nutrient-enriched patches, which
isrelative to the control patches. The response to soil enrichment is not
the same for every species of plantsome have a more sensitive and im-
mediate response than others.
The researchers also compared the responses of the seven species
across the three patch sizes averaged across the two nutrient levels. For
each species and patch size treatment, Farley and Fitter determined the
average proportion of the root system that was in each patch. In the small
and medium sized patches, the total proportions did not always add up to
100% because roots had grown beyond the treatment and control patches
on either side of the central experimental sections. In general, there were
three types of responses. Some species never responded to nutrient en-
richment in any patch size. Some always responded positively to nutrient
enrichment, but the response was equivalent across patch sizes, and then
some responded with greater root density (relative to the control) in the
medium and large nutrient patches than in the small nutrient patches.
In terms of the total root system, two species always had a large per-
centage of their total root system in the center section, as they had small
root systems and did not allocate much of their roots to either the control
patch or whichever nutrient patch they had. For other species, when their
total root system went beyond the center section, more of the root system
was in the nutrient-enriched side than in the sand control side, and that
amount typically grew larger as the patch size increased.
Plants, like animals, have different abilities to assess their environment
to determine where nutrients are, and they then respond accordingly. Farley
and Fitter found several types of responses to the size of nutrient patches.
Some species did not really have a response to nutrient patches, mostly
because its roots were small and confined mostly to the center patch. Some
species have a similar response to nutrient patches in terms of root density,
no matter the size of the patch. However, as patches get larger, some spe-
cies grow a greater percentage of their total root system in the nutrient
patch than in the control patch; they are responding to information in the
environment and sending their roots toward where they detect nutrients.
The mechanisms that plants use to assess their environment are not
well understood; however, the induction of genes has been studied to de-
termine the role of gene induction and transcription of messenger RNA
(mRNA) in response to nutrient exposure.
In one such experiment, Zhang and Forde studied gene expression in
Arabidopsis thaliana. They looked for the mRNA of ANR1, a gene that
they hypothesized was important for root growth. The method used to
detect mRNA was the northern blot, a technique to study gene expres-
sion by detection of specific mRNA sequences on an electrophoresis gel.
The researchers used tubulin mRNA as a loading control to confirm that
mRNA loading is the same across the gel. The expression levels of the
control should not vary between the different sample lanes, and tubulin is
an essential gene expressed at roughly equal levels in all cells. First, the re-
searchers exposed plants to different levels of nitrate and collected samples
over 7 hours. They found that the expression of ANR1 increased over time.
The researchers then extracted mRNA from different plant tissues to
determine where ANR1 was expressed. They found expression only in the
roots. The researchers also wanted to know whether ANR1 responded to
other nutrients in the soil. To answer this question, the biologists exposed
Arabidopsis tissues to nitrate but starved them of either potassium (K+) or
phosphate (PO432); at time zero, the nutrient was resupplied. As before,
the scientists extracted mRNA to determine presence and abundance of

ANR1. They found high expression regardless of lack of potassium or
phosphorus, suggesting that it was the nitrate that the plants were re-
sponding to.
Zhang and Forde demonstrated that ANR1 was expressed rapidly
upon exposure to nitrate, at least in controlled laboratory conditions;
subsequent research has shown that under some conditions exposure to
nitrate does not always induce ANR1. The researchers also demonstrated
that ANR1 expression occurred only in roots. Again, under certain other
conditions, ANR1 expression can be detected outside of the roots, al-
though the levels in roots are much higher. ANR1 transcription remained
high when exposed to nitrate but starved of potassium or phosphorus.
This indicates that ANR1 is sensitive to presence of nitrate and not to
those other nutrients. The results supported the hypothesis that ANR1
plays a role in root growth, specifically in response to nitrate detected in
the environment.
No one knows exactly how root growth is stimulated by the presence
of the ANR1 gene product, but one hypothesis about the function of this
product or protein is that ANR1 regulates the transcription of a set of
genes that together control root growth. Nitrate detected in the environ-
ment stimulates lateral root growth when ANR1 is present.
Plants forage for resources in patches just as many animals do, al-
though animals and plants sense their environments in very different
ways. Scientists do not yet have a good understanding of how nitrate is
sensed by plant roots, but they do know that it is sensed. In both animals
and plants, information is gathered from the individuals internal envi-
ronment, from other organisms, or from the external environment. Soil
nitrate information obtained by plants is used to switch on a gene that
produces a gene product that controls the direction of root growth. The
gathered information leads to changes in behavior and subsequent growth
of the entire plant. Information can also be used by one species to change
the behavior of another species. In the next chapter, how chemicals are
used in communication and how individuals of one species use chemicals
to alter the behavior of individuals of another species is explored. The
function of such chemical communication is to better monopolize re-
sources such as food and space.
Bibliography
Farley RA, Fitter AH: Temporal and spatial variation in soil resources in a
deciduous woodland, J Ecol 87:688696, 1999.
Farley RA, Fitter AH: The response of seven co-occurring woodland
herbaceous perennials to localized nutrient-rich patches, J Ecol
87:849859, 1999.
Munger JC: Optimal foraging? Patch use by horned lizards (Iguanidae:
Phrynosoma), Am Nat 123(5):654680, 1984.
Zhang H, Forde BG: An Arabidopsis MADS box gene that controls
nutrient-induced changes in root architecture, Science 279(5349):
407409, 1998.
CHAPTER 3
Chemical Communication
May be Used to Block
Competition
Foraging organisms assess their environment and take in information

from ecological systems, including where and when resources can be
found. Organisms that are food for other species do not benefit if they
are used as resources, and organisms that are exploiting a resource do not
benefit if they have to share it with other organisms. Over time, organ-
isms have evolved adaptations to help reduce the frequency of becoming
the food of the forager or to minimize other organisms from monopoliz-
ing a potentially shared resource. One class of adaptation includes pro-
duction of chemicals that deter or prevent other organisms from using a
resource. These chemical signals are a way of communicating information
that may warn away other species or inform them that a resource patch
is already occupied. In this chapter, the use of such chemical information
in the environment by animals to make decisions about resource use will
be described.
Egg-Laying Moths Detect Larvae of Other Moths

Females of many animals lay their eggs on a resource that the offspring
will feed on as they develop. These animals may have evolved very spe-
cific feeding requirements, like the flies that parasitize mole crickets in
Chapter 1. Often, the offspring will feed on only one or a few types of
food, but their small size restricts their mobility and prevents them from
moving from one patch to another. The females choice of where to lay
eggs will determine the success of her offspring because they are trapped
in a patch. Females of some species have evolved the ability to locate

resources for their offspring and minimize competition between their
offspring by sensing chemical information, as will be shown in this sec-
tion. Competition is the demand by two or more organisms for limited
environmental resources. Offspring or females may provide information
to others that a patch is occupied, but what kind of information is it, and
can other species collect and correctly interpret this information?
To better understand the communication of chemical information,
Sarah Corbet studied larvae of the Mediterranean flour moth, Ephestia
kuehniella. These moths are small and their larvae live on a variety of
grains and grain products, including flour and rice. Moth and butterfly
larvae are also known as caterpillars. Corbet discovered that larvae of the
Mediterranean flour moth secreted droplets of a liquid from glands near
their mouths. Moth larvae enter a cocoon phase called pupation, the
process in many insects of transforming from a larva to an adult, prior to
becoming adults, and Corbets experiment measured the time it took for
50% of a test population to pupate when exposed to different numbers
of other larvae or larval.
Corbet performed pupation tests in three conditions. In the first,
she placed different numbers of larvae in small boxes to simulate various
population densities. In the second, she placed variable numbers of larvae
in boxes for a period of time and removed them before the test larvae
were added. In the third, she extracted the secretion from the mandibular
glands of larvae, spread the secretion on a coverslip, and exposed test lar-
vae to the extract. In a related experiment, she also determined the num-
ber of eggs laid by females in the presence of larvae or their secretions.
Corbet showed that secretions from glands of an individual larva led
to responses by other larvae and egg-laying females of the same species.
The Mediterranean flour moth uses the glandular secretions of larvae to
help reduce competition; the presence of high densities of larvae inhib-
its both pupation and egg-laying. Low densities of larval secretions also
led to longer pupation times, but intermediate densities actually reduced
pupation time and increased egg-laying, which may indicate that the
presence of some larvae is information that a patch is a potentially good
resource for offspring.
Chemical Communication May be Used to Block Competition 27
The Indian meal moth, Plodia interpunctella, shares the same habitat
and larval food resources as the Mediterranean flour moth, making these
two species potential competitors. Indian meal moth larvae also secrete
droplets from glands with similar results within their species. P. Anderson
and J. Lofqvist (1996) wanted to determine whether the Mediterranean
flour moth and the Indian meal moth could sense each others glandular
secretions. If the information about density of larvae were restricted to
one species, females would not be able to determine the total number of
potential competitors in a resource patch. Inability of females to estimate
total density of all possible competitors for her offspring would decrease
her reproductive success. As shown earlier, competition causes larvae to
grow slower, because pupation time increased under conditions of high
larval density.
Anderson and Lofqvist set up three different choice experiments in
which a mated female of either species was released into a small screened
cage (150 x 110 x 60 mm) containing two cups filled with a wheat germ
mixture, which represented patches of resources where females might
choose to lay eggs. A choice experiment is an experiment used to deter-
mine the choices organisms make under certain conditions, usually be-
tween two alternatives. Some of these cups had been filled with a variable
number of larvae (1, 5, 20, or 40) of one of the two species for 24 hours
prior to the test and then removed.
The biologists let each female inhabit the cage for 48 hours, during
which time she could choose to lay eggs in either or both cups. In the first
experiment, one of the cups had contained larvae of the same species, and
the other cup was not infested (Figure 6A and B). In the second experi-
ment, one of the cups had contained larvae of the other species, and the
other cup was not infested (Figure 6C and D).
The two bars for any density treatment in Figure 6AD representa
choice between a cup that had contained the indicated density and a
cup that had contained no larvae. In the third experiment, one cup had
contained larvae of the same species, and the other cup had contained
an equal number of larvae of the other species (Figure 6E and F). The
height of each bar represents the percentage of adults that grew from eggs
females laid in a cup, a measure of choice, or preference, for a cup.
100 Med moth laying in 100 Indian moth laying in

Med or noninfested cups Indian or noninfested cups
80 80
% choice
% choice
60 60
40 40
20 20
0 0
A B
100 Med moth laying in 100 Indian moth laying in

Indian or noninfested cups Med or noninfested cups
80 80
% choice
% choice
60 60
40 40
20 20
0 0
C D
Med moth laying in
100 Med or Indian cups 100 Indian moth laying in
Med or Indian cups
80 80
% choice
% choice
60 60
40 40
20 20
0 0
1 5 20 40 1 5 20 40
E density of larvae in pretreatment F density of larvae in pretreatment
Figure 6 Results of choice experiments by female Mediterranean

flour and Indian meal moths. Females chose between cups that had
contained larvae of the same species or were not infested (A and B),
between cups that had contained larvae of the other species or were
not infested (C and D), and between cups that had contained larvae
of one or the other species (E and F). Med = Mediterranean, light
gray bars = Med moth cups, dark gray bars = Indian moth cups, and
open bars = noninfested cups.
Source: Data from Anderson and Lofqvist, 1996, Figures 1 and 2, courtesy of P. Anderson, Dept.
of Plant Protection Biology, Swedish University of Agricultural Sciences, Sweden.
When given a choice between a non-infested cup and a cup that pre-
viously held one larva (of either species), Mediterranean flour moth fe-
males did not exhibit a clear preference, choosing cups that held larvae
between 45% and 55% of the time (Figure 6A and C). However, the
Mediterranean flour moth females can detect the prior presence of lar-
vae of Mediterranean flour and Indian meal moth larvae, as evidenced
Table 4 Mean mass of adult female and male Mediterranean flour

moths (A) and Indian meal moths (B) when grown at different larval
densities of larvae of the same species or half one species and half the
other species. MFM = Mediterranean flour moth; IFM = Indian
flour moth; s.e. = standard error of the mean.
density male mass ( 1 s.e.) male mass ( 1 s.e.) female mass ( 1 s.e.) female mass ( 1 s.e.)
of larvae when grown with when grown in when grown with when grown in
other MFMs 50:50 mix other MFMs 50:50 mix
5 11.09 0.59 13.92 0.67
10 10.38 0.45 10.50 0.51 13.18 0.62 13.41 0.56
20 8.69 0.34 9.51 0.56 10.21 0.37 11.79 0.59
A Mediterranean flour moth adults
density male mass ( 1 s.e.) male mass ( 1 s.e.) female mass ( 1 s.e.) female mass ( 1 s.e.)
of larvae when grown with when grown in when grown with when grown in
other IFMs 50:50 mix other IFMs 50:50 mix
5 5.92 0.08 9.28 0.25
10 5.78 0.08 5.30 0.20 10.49 0.14 9.51 0.56
20 5.69 0.11 6.06 0.11 9.45 0.17 10.01 0.37
B Indian flour moth adults
Source: From Anderson and Lofqvist, 1996, Figures 4 and 5.
by their avoidance of cups that previously held high densities of either

species when a non-infested cup was available. When given a choice to
lay eggs in cups that held larvae of their species versus the other species,
Mediterranean flour moth females prefer to lay eggs in cups that had
previously held larvae of the other species (Figure 6E). This indicates that
the glandular secretions of each species are not the same, and the Mediter-
ranean flour moth can distinguish between the two signals.
Indian meal moth females showed some avoidance of Mediterranean
flour moth larvae only when cups held 40 larvae per cup, indicating
that they could detect the presence of the other species. Densities below
20 larvae per cup of either species seemed to attract Indian meal moth fe-
males to lay eggs as indicated by Figure 6B and D. The preference was mod-
erate, between 60% and 75%. Low amounts of secretion from larvae could
indicate to a female looking for resources that a patch is suitable for her off-
spring, because other larvae are surviving on the resources at that patch. At
the same time, the concentration of the chemical signal is not high enough
to indicate to her that densities are excessive, which would result in too
much competition for resources between existing larvae and her offspring.
In another set of experiments, Anderson and Lofqvist tested the rate
of growth and the total size attained by larvae of each species growing in
containers with different population densities (Table 4). They used 5, 10,
or 20 larvae of the same species, or 10 or 20 larvae total, with half of them

being Mediterranean flour moth larvae and the other half being Indian
meal moth larvae. As moths completed development, they were collected,
examined to determine their species and sex, and then weighed.
Female moths that can detect high densities of larvae have a distinct
advantage. High larval densities caused a decrease in mean weight for
the Mediterranean flour moth (Table 4A). Attaining a large size increases
the success of insects like these moths, because larger females can lay more
eggs than smaller females, and they may also live longer. This decrease
in mean weight of the Mediterranean flour moth was more pronounced
when it was with high densities of larvae of the same species compared
to high densities of the Indian meal moth, indicating that it competes
well for resources against Indian meal moth larvae but less well against
other Mediterranean moth larvae. Perhaps Mediterranean flour moths are
more sensitive to the signal of their own species than the other species
because of this competitive disadvantage. The Indian meal moth typically
develops faster and does not get as large as the Mediterranean flour moth.
Adult size does not seem to be affected by varying densities of either spe-
cies (Table 4B), so Indian meal moths may not have evolved mechanisms
to reduce competition.
The two moth species do not have the same competitive ability, but
they are each successful. The Mediterranean flour moth grows more slowly
in a crowded environment and is sensitive to the presence of other larvae.
The Indian meal moth has a weaker response to secretions of the Mediter-
ranean moth but survives by growing faster and maturing at a smaller size.
Both species have evolved strategies to deal with limited resources and the
competition that often occurs for those resources.
Information Is Used by Corals

During Settlement
Many species emit signals that indicate occupation of a resource. Often
these signals are intended for individuals of the same species, but when
a resource is shared with other species, there is an advantage to species
that can detect the presence of individuals of any type. The remainder
of this section investigates the communication of such signals in coral,
which, like plants and moth larvae, cannot move from patch to patch to
avoid competition. Individuals may communicate their presence through
chemical signals, and this information may have effects on the receivers of
these signals. How do nonmotile organisms, such as corals and the algae
that often encrust coral reefs, use such signals?
Coral reefs are large structures produced by living organisms, mainly
coral animals, which are animals that secrete a hard skeleton. Many dif-
ferent species together make up a reef. A coral grows as individuals repro-
duce asexually, that is, reproducing without sex or union of gametes. But
coral animals can also release sperm and eggs into the ocean, which is a
process called spawning, a process that is usually synchronized within a
species. Coral spawning usually occurs for different colonies of the same
species at the same highly specific time. For instance, individuals of Mon-
tipora verrucosa release their gametes each June, July, and August, three
days after the full moon, between 8 and 10 PM at night. Sperm and
egg unite to form a small organism called a larva. The larva has some
swimming ability, but most of its mobility is controlled by ocean cur-
rents. Coral settlement, the landing and attachment of coral larvae that
precedes maturation into adults, occurs when a larva lands on a surface.
These surfaces may be reefs or some other hard facade and may be in
limited supply. Once a larva settles, it grows into a non-moving adult, so
it only gets one chance to select its location for life.
Larval corals can settle on other corals, bare mud, beds of small
soft algae, or coralline algae that are hardened with calcium carbonate
(CaCO3). Mud and soft algae are not good surfaces for settlement, be-
cause they are too soft for successful colony development. Corals that
settle on coralline algae may have the best chance at survival. Scientists
have noted that coral animals differ in the distribution of colonies on
surfaces of coralline algae versus other surfaces. Researchers set out to
determine what information coral larvae use to decide on a place to settle
and how that decision affects their ability to thrive.
The Great Barrier Reef is a coral reef that lies off the coast of Queensland,
Australia. Katharina Fabricius and her Australian colleagues studied two
species of coral animal common on the Great Barrier Reef, Acropora mil-
lepora, called milli staghorn coral, and A. tenuis, known as the finger coral
(Figure 7; Harrington et al., 2004).
Figure 7 Two coral reef species (staghorn coral on left and brain coral
on right) and coralline algae.
Source: Staghorn coral, http://commons.wikimedia.org/wiki/File:Staghorn-coral-1.jpg: Author:
Adona 9 at the English language Wikipedia. This file is licensed under the Creative Commons
Attribution-Share Alike 3.0 Unported license. Brain coral, http://upload.wikimedia.org/
wikipedia/commons/5/56/Brain_coral.jpg: Public domain. 2007. Corraline algae, http://
en.wikipedia.org/wiki/File:Coralline.1.jpg: Author: FalsePerc. This file is licensed under the
Creative Commons Attribution-Share Alike 3.0 Unported license.
They collected five different species of coralline algae from the Great
Barrier Reef and placed them in several outdoor aquaria. To determine
settlement patterns of coral larvae, they used fragments of live and dead
coralline algae as well as clay tiles and pieces of dead coral.
Fabricius and her colleagues placed one of each type of surface (five
species, both dead and alive, plus two abiotic surfaces = 5 x 2 + 2 = 12
treatments) into each of five different tanks. They added about 8,000 fin-
ger coral larvae to each tank, waited for 3 days, and counted the numbers
of settled corals on each type of surface.
The process of selecting an appropriate habitat is essential for most
organisms. Larvae that can collect information to determine the best lo-
cation for settlement will have a selective advantage over individuals that
cannot utilize such information. Coral settlement was much greater on
live coralline algae than dead. It may be hypothesized that coral larvae are
avoiding dead coralline algae fragments. They may either be attracted to
live algae or inhibited by dead algae. The various species of coralline algae
have different properties that increase or decrease their suitability as sur-
faces for coral larvae. The scientists found that settlement was highest for
one species (A), which had about 15 times more settlement than the spe-
cies with the lowest settlement (E). The other three algae species (BD)
had coral larvae settlement that was in between the highest and lowest
species. Because coral settlement on the two worst coralline algae species
(D and E) is also lower than settlement on the bare tile surface, these two
species may be inhibiting coral settlement.
Fabricius and her colleagues next wanted to determine the survival
of coral larvae on each of these surfaces. They transferred the fragments
with settled coral larvae to another tank. They observed the larvae for
22 days and determined the proportion of coral animals surviving at
the end of each day. Finger coral has survival patterns that vary with the
type of surface on which they settle. In general, survival was correlated
with settlement in that when settlement was high, resulting survival over
22 days was also high, but when settlement was low, survival was low. For
instance, for the species with the lowest settlement (E), coral larva survival
was 100% after 1 day, 50% had died on the next day, and all were dead
by day 3.
The researchers next ground up a known mass of each type of coral-
line algae, plus their clay tile and coral controls, and extracted chemicals
from each ground up sample. The biologists mixed the seven extracts with
filtered seawater to make five different concentrations of each residue and
placed a portion of each concentration in small dishes. Fabricius and her
colleagues added ten to twenty coral larvae to each dish. After 36 hours,
they determined the percentage of larvae that had settled and grown at
each concentration, and they determined which concentration produced
the maximum survival and growth of settled coral larvae (Figure 8). A
low maximum concentration of any extract means that coral larvae were
sensitive to chemicals from that species. The extract controls of clay tile
and coral produced no growth into adults of settled larvae.
Fabricius and her colleagues found strong correlations between the
ability for coralline algae extract to induce coral growth and the amount
of settlement and survival of coral larvae that occurred on those species.
The researchers hypothesized that there are chemical signals that affect
growth of the coral larvae into adults and that the species of coralline
algae produce different chemical signals. Coralline algae species A and B
induced the most settlement in corals and were the ones on which cor-
als had high growth and survival. The low maximum concentrations of
extracts from those species meant that coral larvae were more sensitive to
chemicals from those species than to other coralline algae. The chemical
produced by species A and B could have been different or the same.
35
maximum survival and growth of coral

extract concentration that produced
30
25
20
15
10
0
A
l
ra
til
e
e
e
co
ga
ga
ga
ga
ga
ay
al
al
al
cl
al
al
Figure 8 Extract concentrations that produced maximum survival and

growth of settled coral larvae in laboratory experiments (with 95%
confidence intervals). Concentrations that were both lower and higher
than indicated produced less coral growth and maturation.
Source: Data from Harrington et al., 2004, Figure 4.
Algal species D and E, on which settlement and survival of corals were

lowest, still produced chemicals that affected growth, but not as strongly
as other species. Coralline algae had other mechanisms to prevent corals
from settling and growing on them. For example, the biologists observed
that settled coral larvae died when the algae shed their outer layers. The
shedding removed and subsequently killed settled larva. Coral larvae also
died when they were overgrown by algae.
High survival was related to high sensitivity to the corresponding
chemical extract. This suggests that coral larvae may be choosing surfaces
on which to settle based on information they gather from the environ-
ment. It appears that coral larvae are able to recognize and choose coral-
line algae species on which to settle that are least able to kill the coral.
Although the larvae are mostly at the whim of ocean currents, they have
some ability to reject a surface and remain in the current. The strategy of
releasing chemicals may have evolved to deter settlement of corals, but
corals have evolved to recognize different species based on the chemical
information they emit. They use that information to accept or reject a
settlement location.
The data suggest that information is transmitted by species to sig-

nal prior occupation of a resource. These signals can prevent use of a
resource by others. Different species assess their environment to make
decisions about where to lay eggs, where to settle, or where to feed. Some
species have evolved the ability to assess the information transmitted by
other species, allowing them to more efficiently use resources and avoid
resources where competition may be fierce or the supply of resources may
be limited. Information is an intrinsic part of the interactions among
individuals of the same and different species. There is a great deal of in-
formation in the environment for organisms to assess, such that the loss
or gain of species in an ecological system can change the quantity and
quality of information that is available.
Bibliography
Anderson P, Lofqvist J: Asymmetric oviposition behaviour and the
influence of larval competition in the two pyralid moths Ephestia
kuehniella and Plodia interpunctella, Oikos 76(1):4756, 1996.
Corbet SA: Mandibular gland secretion of larvae of the flour moth,
Anagasta kuehniella, contains an epideictic pheromone and elic-
its oviposition movements in a hymenopteran parasite, Nature
232(5311):481484, 1971.
Corbet SA: Oviposition deterring pheromone in mandibular glands of
Ephestia kuehniella, Nature 243(5409):537538, 1973.
Harrington L, Fabricius K, Death G, et al.: Recognition and selection
of settlement substrata determine post-settlement survival in corals,
Ecology 85(12):34283437, 2004.
CHAPTER 4
A Change in the Number

ofSpecies Affects
Information Content
of an Ecological System
Ecological systems are made up of many species and environmental fac-

tors, and are full of information. Species assess ecological systems by col-
lecting this information and using it to obtain resources, avoid danger, or
find mates. Information is an inherent part of ecological systems.
Humans measure the information content in ecological systems in
a variety of ways. Biological diversity, or biodiversity, is the variety of
life at all of its levels, from genes to ecological systems, and the ecologi-
cal and evolutionary processes that affect biodiversity. Biodiversity in an
ecological system is defined as the number of species living in that system
or as all of the species, their genes, populations, abundances, and interac-
tions. The interactions often require the transfer of information, making
information a fundamental aspect of biodiversity. Each species occupies
a unique niche; therefore, the loss of any species is a loss of biodiversity
and the unique information in the DNA of that species. An ecological
niche is where a species lives, how the environment affects it, and how it
affects the environment. All of the effects of that species on other species
will now be absent from the ecological system, and this represents a loss
or alteration of information in the system.
In some ecological systems that exist as discrete patches, such as lakes or
ponds, different species may exist in different locations such that the spe-
cies and the information content vary from location to location. A study
on the interactions and biodiversity of small forest ponds was performed
by David Chalcraft and William Resetarits who were interested in how

interactions between species were affected in ecological systems with dif-
ferent predators. Specifically, they wondered whether a predator could
have a large impact on some prey species but not on others. This study
can be used as a simple example of how the addition or subtraction of
a species can change the information content of the environment. The
prey they studied were three species of tadpole, the larvae of frogs, and
toads: the southern toad (Bufo terrestris), the southern leopard frog (Rana
sphenocephala), and the spring peeper (Pseudacris crucifer).
Chalcraft and Resetarits tested for effects of six common predators,
three fish and three salamanders, on the tadpole prey community. The fish
were the banded sunfish (Enneacanthus obesus), mosquitofish (Gambusia
holbrooki) and eastern mud minnow (Umbra pygmaea). The salamanders
were the marbled salamander (Ambystoma opacum), two-toed amphiuma
(Amphiuma means) and red-spotted newt (Notophthalmus viridescens).
Chalcraft and Resetarits created experimental ponds using large tanks.
They set up six treatments (one for each predator) and a control with
each treatment and control repeated in four tanks. The tanks were filled
with pond water, some leaves, and 175 tadpoles of each prey species. The
scientists randomly selected which tanks would receive each treatment,
and added two individuals of the designated predator. They checked the
tanks daily and collected any tadpoles that had grown into frogs or toads.
After 60 days the scientists emptied the tanks and searched for remaining
tadpoles. Their experimental design allowed them to quantify the effect
of each predator on prey species diversity and the relative proportions
of each prey species. The investigators quantified biodiversity using the
Shannon diversity index.
For each predator condition, including the control, the researchers
quantified the proportion of each tadpole species remaining. For the con-
trol with no predator, 56% of the tadpoles were southern toads, 41%
were leopard frogs and 3% were spring peepers. About 56% of the tad-
poles were southern toads in the amphiuma treatment, but leopard frogs
were a bit less at about 32% and spring peepers were at about 12%. In
addition, when mosquitofish were predators, the proportions were about
the same as the control. Thus, these two predators had effects on the
tadpole communities that were fairly similar to the no predator control.
Number ofSpecies Affects Information Content 39
With newts and salamanders as predators, southern toads nearly or

completely disappeared from the tanks, with relative increases in leop-
ard frogs and spring peepers. When minnows were the predator, almost
100% of the tadpoles that survived were southern toad tadpoles, and
when sunfish were the predators 100% of remaining tadpoles were leop-
ard frog tadpoles.
Chalcraft and Resetarits next took those proportions and quantified
the impact of different predators on prey communities with a measure of
biodiversity called the Shannon diversity index. This measure is based on
the number of species and the number of individuals of each species in an
ecological system. An ecological system with many species in which most
of the individuals in the community belong to one species will have lower
diversity than a system with the same number of species but an equal
number of individuals of each species.
Biodiversity was affected both in the number of species and their in-
dividual proportions in the system, depending upon the identity of the
predator. The researchers determined that for most predators and the con-
trol, when two species survived and the third made up less than 10% of
the prey community, the Shannon diversity index was between 0.65 and
0.9. Both minnows and sunfish reduced diversity to nearly zero, which is
a consequence of eliminating or nearly eliminating all but one prey spe-
cies. There is zero diversity and a dramatic decline in information content
in a prey community with only one species.
A predator in that situation will encounter only one type of prey.
If the predator does not eat that type of prey, or cannot locate it using
information in the environment, then the predator population may ulti-
mately go extinct, further reducing the diversity of the system. As the data
illustrate, the ability to assess information varied among specieseither
predators were able to locate only certain species of prey, or the prey were
able to detect the presence of only certain predators. Predators and prey
were collecting and assessing information in their environment, although
some are better than others at utilizing the information they collect.
When new species enter or existing species disappear from an ecologi-
cal system, information is gained or lost and the flow of information is
altered. Species present initially are now subject to a different quantity
and quality of information. As the composition, that is, the variety, or
types, of species existing at a particular point in time of an ecological

system changes, interactions between species change. The spring peeper
did not do well in ponds without predators, suggesting that they may
not compete well against the other tadpole species. However, when other
tadpole species were reduced by newts or marbled salamanders, the peep-
ers remained at high proportions in the community, indicating they can
process amphibian predator information successfully. The competitive in-
teractions between species of tadpole were affected by the species of preda-
tor present, allowing for greater success of spring peepers. Spring peepers
were extremely susceptible to fish predators, suggesting that they lack the
ability to assess information about those predators. However, other exper-
iments have shown that adult female spring peepers avoid laying eggs in
ponds where certain fish predators are present, so spring peeper tadpoles
may not normally be subjected to fish predators as in this experiment.
In the same manner that spring peepers lack the ability to detect fish
predators, when a non-native predator is introduced into an ecological
system, all existing prey species likely lack an ability to detect or assess
information about this new predator. However, any prey that avoids the
predator may flourish, whereas susceptible prey become rare or extinct,
further altering the information content. Even though the addition of a
new species increases the information content in an ecological system,
a new predator may ultimately cause the reduction of biodiversity and
information as it causes extinctions of existing species.
In a manner similar to the addition of a predator, the loss of the
southern mole cricket from Florida would alter the information avail-
able in the ecological system. This change in information might affect the
tachinid flies, and they would have to find new hosts to parasitize. More
flies might parasitize tawny and imitator mole crickets, and reduce their
population sizes.
The species interactions in ponds and other ecological systems require
the transfer of information, making information a fundamental aspect of
biodiversity. In one sense, we can measure the amount of information in
an ecological system by measuring its biodiversity, and in another sense,
an understanding of how species interact with one another demonstrates
the multiple ways that species transfer information from one individual
to another.
Ethical, Legal, Social Implications: We Have an

Obligation to Preserve Biodiversity
Each species is unique; it has a unique evolutionary history, ecological
niche, and set of genes. The loss of any species represents a loss of bio-
logical information. The permanent loss of any species from the planet is
global extinction, and is a permanent loss of the unique storehouse of in-
formation of that species. Extinction is natural; approximately 99.9% of
all species that have ever existed on Earth are now extinct. Yet, human ac-
tivity has caused the extinction of many species that might otherwise have
survived. As humans have come to dominate the surface of the planet,
and our impact on ecological systems has grown exponentially. The rate of
species loss today is estimated to be far greater than the rate of extinction
that occurred in prehistoric times. Some estimates place the current losses
at about 1 species per hour.
As shown with tadpoles and their predators, the identity of a predator
has consequences for diversity. In the same manner, the loss of a species
will alter interactions of the remaining species. This could impact biodi-
versity in a negative manner. Even the loss of a single species could have far
ranging effects on the remaining species in terms of their interactions and
abundance. Whether or not we should prevent the extinction of species
whose populations are in decline because of human activities is the subject
of much debate by conservation biologists, philosophers, and ethicists.
To consider our ethical obligations to preserve species that we have
endangered through our activities, people should familiarize themselves
with several arguments. One argument is based on the fact that humans
are part of naturemerely one species among many competing for lim-
ited resources. Humans are subject to the same biological processes as
other species, including competition, disease, and evolution. If humans
do not use available resources to flourish, other species will. Our current
population size of over seven billion people and our global distribution
on Earth are testament to the fact that we can exploit many resources to
increase our population size. Is it morally wrong to optimize our own
survival as other species do? When framed this way, it seems logical to
conclude that we have no moral obligation to protect any species other
than our own.
However, as humans drive other species to extinction, we negatively

impact our own ability to survive. Ecological systems with low biodiver-
sity are less able to withstand further pressure from humans and provide
fewer resources for our continued existence. Fish, for instance, will be-
come less available if we continue to overexploit the oceans. Unlike other
species, humans have more information about ecological systems. Per-
haps our strategy should be for long-term rather than immediate survival,
and we should alter our current behavior.
Humans exploit ecological systems to gain resources such as food, fiber,
and medicines. A counter to the humans as part of nature argument is
that biodiversity provides unique benefits to humans and ecological sys-
tems. If we destroy the habitats that provide us with resources in order to
obtain lumber from forests or to clear land for shopping malls, we damage
the diversity that provides us with resources. These resources could be used
to improve the human condition by providing us with new knowledge
and medical treatments. This argument stipulates that biodiversity offers
potential benefits and preserving biodiversity is in our own best interest.
Because preservation of biodiversity would benefit subsequent generations
more than the current one, many people consider this longevity argument
to be a moral imperative rather than an ecological argument.
There are many benefits to biodiversity that support this longevity argu-
ment. Ecological systems and human populations suffer when biodiversity
is diminished by human activities. Humans are dependent upon animal
pollination, where animals spread pollen among flowers to produce a large
proportion of our food. Plants benefit from pollination, as do animals that
eat the fruits and vegetables produced. Loss of pollinating insects will af-
fect our food production. Benefits from biodiversity can be both biological
and economic. Pollinating insects provide an estimated $8billion annual
benefit to the agricultural industry in North America alone.
Humans also benefit from the useful chemicals derived from plants.
Aspirin, antibiotics, caffeine, cocoa, digitoxin for heart arrhythmia, qui-
nine to prevent malaria, and taxol, which has cancer fighting properties,
are all obtained from biological sources. The loss of organisms with me-
dicinal properties from ecological systems represents a reduction of biodi-
versity and information, as well as a loss of the human benefits that accrue
from biodiversity.
A third argument in the debate over our obligation to preserve species

is the ethics-based inherent value argument, which states that the mere
existence of a species gives it value. Apart from the benefits that we or
other species gain from any species, all species have an inherent right to
exist. There may be species from which humans derive no benefit, yet
proponents of this view argue that humans have an obligation to pre-
serve all creatures. The origins of this inherent value argument spring from
consideration of the unique evolutionary history and unique information
and interactions of each species. To destroy a species habitat for human
gain effectively denies the right of the species to exist. If one accepts the
premise that every species has the right to exist, then they must conclude
that it is unethical to destroy that habitat and cause the extinction of any
organism. The loss of any species and its unique evolutionary lineage is
considered unacceptable, and proponents have concluded that humans
should protect all species from extinction regardless of their utility to
humans.
Bibliography
Chalcraft DR, Resetarits WJ: Predator identity and ecological impacts:
functional redundancy or functional diversity, Ecology 84(9):2407
2418, 2003.
Lloyd M, Zar JH, Karr JR: On the calculation of information-theoretical
measures of diversity, Am Midl Nat 79(2):257272, 1968.
Picard K: Biodiversity and ethics: do we have a responsibility to preserve?
J Sci Health at U Alabama 4:4446, 2006.
Conclusion
Information in an environment comes in a variety of forms. Sounds that
can be heard, chemicals that can be detected, and shapes that can be
recognized are all sources of biologically significant information about
the environment. Different modes of communication, from the chemical
to the visual to the acoustic, are interpreted by species when interacting
with other species. Organisms collect information from other species and
use it to exploit those species as resources, and they collect information
from the abiotic environment to help locate resources. Although the focus
was primarily on non-heritable information in this chapter, a substantial
portion of biological information arises from the heritable information
encoded in DNA, which provides continuity of life. Finally, ecological
systems contain information that is related to the amount of biodiversity
in the system. The number and types of species that are present in an
ecological system can be a source of information about that ecological
system for all organisms living there, including humans. Changes in the
abundance and types of species change the information content of eco-
logical systems.
Glossary
ammonium. An ion of NH41 derived from ammonia by combination with a
hydrogen ion.
asexually. To reproduce without sex or union of gametes.
biodiversity. All the species, their abundances, and their interactions with the
environment.
biomass. The total amount of organic matter in an organism, population, or
other ecological system after water is removed.
choice experiment. An experiment used to determine the choices organisms
make under certain conditions, usually between two alternatives.
communication. Transmission of information so that it is satisfactorily received
or understood.
competition. An interaction defined by the demand by two or more organisms
for limited environmental resources.
composition. The variety, or types, of species existing at a particular point in
time.
coral settlement. The landing and attachment of coral larvae that precedes matu-
ration into adults.
density. The quantity of a substance or population per unit volume, unit area,
or unit length.
diminishing returns. A result of a yield rate that declines because a constant pro-
portion is removed from a finite resource over time.
ecological systems. An ecological community together with the abiotic environ-
ment, usually considered to function as a unit.
global extinction. The permanent loss of a species from the planet when no indi-
viduals of that species are left living.
host. The organism upon which parasites feed, or the organism that is used as a
resource by a parasite.
information. Information is stored, communicated, and implemented or
interpreted.
interactions. Positive or negative associations between species that favor or in-
hibit growth and evolution of populations. It may take the form of competition,
predation, parasitism, commensalism or mutualism.
larvae. The immature, wingless, and often wormlike form of an insect.
marginal value theorem. Predicts that optimal foragers leave a food patch when
the rate of capture at that patch falls below the overall habitat rate.
natural enemies. Organisms that use other organisms as a resource.
48 GLOSSARY
niche. Where a species lives, how the environment affects it, and how it affects
the environment.
nitrate. An ion of NO3 and has a negative one charge.
nutrients. Food or chemicals that organisms obtain from their environment and
need to live and grow.
optimal foraging theory. The theory that investigates evolution of animals to
forage in a manner that maximizes their rate of energy intake per unit time spent
feeding.
parasites. Organisms that live on or in another organism.
phosphate. An ion of PO43 derived from phosphoric acid H3PO4.
predators. Organisms that kill and feed on other organisms.
pupation. The process in many insects of transforming from a larva to a pupa.
resource. A natural feature, substance, or organism used as a supply of energy or
nutrients to other organisms.
spawning. The process of releasing gametes, eggs or sperm, into the environment,
usually synchronized within a species.
Index
Acropora millepora, 31 Competition, 26
Acropora tenuis, 31 Composition, 3940
Ambystoma opacum. See Marbled Coralline algae
salamander coral settlement on, 3132
Ammonium, 19, 20 correlations between ability for
Amphiuma means. See Two-toed extract, 33
amphiuma from great barrier reef, 32
Anderson, P., 2729 Coral reefs, 31
ANR1, 22, 23 Corals
Arabidopsis thaliana, 22 larva. See Larva
Asexual reproduction, 31 settlement, 3133
spawning, 31
Banded sunfish, 3840 using information during
Biodiversity settlement, 3035
affected by species, 39 Corbet, Sarah, 26
benefits to, 42 Crickets. See also Mole crickets
in ecological system, 37 male, using sound, 4
obligation to preserve, 4142 songs exploited by natural
Biological diversity. See Biodiversity enemies, 26
Biomass, 20, 21
Bufo marinus. See Cane toads Decaying animal carcass, 19
Bufo terrestris. See Southern toad Diminishing returns, 1415
Cane toads, 1011 Eastern mud minnow, 3840

Caterpillars, 26 Ecological niche, 37
Chalcraft, David, 3839 Ecological system
Changa, 56 biodiversity in, 37
Chemical communication as discrete patches, 3738
corals, using information during humans and, 4142
settlement, 3035 interactions between species
egg-laying moths, detecting larvae affecting, 3840
of other moths, 2530 loss of organisms with
Choice experiment, 27, 28 medicinal properties
Choruses, by frogs, 612 from, 42
Communication with low biodiversity, 42
of chemical information. See Enneacanthus obesus. See Banded
Chemical communication sunfish
between individuals of other species Environmental assessment
cricket songs exploited by natural lizards, optimal foraging behavior
enemies, 26 of, 1318
frog choruses attract predators, by plants in search of nutrients,
612 1823
50 INDEX
Ephestia kuehniella. See Mediterranean Lizards

flour moth average capture rate, 17
average consumption rate of ants,
Fabricius, Katharina, 3133 1617
Farley, Rebecca. 1923 feed on ants, 14
Finger coral, 31 habitat rate, 16, 17
Fireflies, risk to, 1 marginal value theorem of,
Fitter, Alastair, 1923 1516
Foraging process, of lizard, 1318 optimal foraging theory, 1314
Forde, B. G., 2324 reasons for leaving nest, 18
Fowler, Harold, 56 travel between nests, 1718
Fringe-lipped bats, 7 Lofqvist, J., 2729
hunting, 9
preying on frogs, 8 Marbled salamander, 3840
vocalization, 912 Marginal value theorem, 1516
Frog-eating bats. See Fringe-lipped Mating vocalization
bats crickets, 2, 4
Frogs frogs, 7
mating vocalization, 7 Mediterranean flour moth, 2630
predators of, 712 Messenger RNA (mRNA), 22, 23
Milli staghorn coral, 31
Gambusia holbrooki. See Mole crickets, 2
Mosquitofish, 38 experimental design for testing, 3
Gene expression, 22 southern, 3
Global extinction, 41 tachinid fly and, 6
Grasshoppers, 2, 4 tawny, 3
Great Barrier Reef, 3135 trapping at speakers, 4
Montipora verrucosa, 31
Habitat, selection of, 32 Mosquitofish, 3840
Host, 1 Munger, James, 1418
Imitator mole cricket, 5 Natural enemies, 1

Indian meal moths, 2730 cricket songs exploited by, 26
Information Neocurtilla hexadactyla, 6
communication. See Niche, 37
Communication Nitrates, 19, 20
interception of, 12 Northern mole cricket, 6
Inherent value, 43 Notophthalmus viridescens. See
Interactions Red-spotted newt
between species, 40 Nutrient-enriched patches, 2021
for transfer of information, 37 Nutrients, plant assessing
environment
Katydid, 2 location of, 18
in soil, 1920
Larva, 4, 31
settlement and survival of, 3334 Optimal foraging theory, 1314
settlement patterns of, 32 Ormia deplete, 5
INDEX
51
Parasites, 1 Scapteriscus vicinus. See Tawny mole

flies, 46 cricket
Philander opossum, 7 Shannon diversity index, 39
Phosphates, 19, 20, 22 Southern leopard frog, 3840
Phrynosoma cornutum. See Texas Southern mole cricket, 3
horned lizard Southern toad, 3840
Phrynosoma modestum. See Spawning, 31
Round-tailed horned lizard Spring peeper, 3840
Physalaemus pustulosus. See Tungara
frogs Tachinid fly, mole cricket and, 6
Plants, environmental assessment Tawny mole cricket, 3
in search of nutrients, 1823 Texas horned lizard, 14
Plodia interpunctella. See Indian meal Trachops cirrhosis. See Fringe-
moths lipped bat
Potassium, 22 Tubulin, 22
Predators, 1 Tungara frogs, 7
-prey interactions, 7 Two-toed amphiuma, 3840
Pseudacris crucifer. See Spring peeper
Pupation, 26 Ulagaraj, S., 2
Umbra pygmaea. See Eastern mud
Rana sphenocephala. See Southern minnow
leopard frog
Red-spotted newt, 3840 Vocalization
Resetarits, William, 3839 cane toad, 1011
Resource, 1 of crickets, 26
Root length density, 20, 21 benefit of making, 4
Round-tailed horned lizard, 14 experimental design for testing, 3
Ryan, Merlin, 712 mating, 2, 4
by predators of frogs, 712
Scapteriscus borellii. See Southern mole tungara frogs, 7
cricket
Scapteriscus didactylus. See Changa Walker, Thomas, 2
Scapteriscus imitates. See Imitator mole
cricket Zhang, H., 2324
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FOR THE Christopher J. Paradise A. Malcolm Campbell

Christopher J. Paradise, PhD

All rights reserved. No part of this publication may be reproduced, stored

First published in 2016 by

ISBN-13: 978-1-60650-969-2 (print)

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Cover and interior design by S4Carlisle Publishing Services Private Ltd.,

Printed in the United States of America.

Organisms Have Evolved

The mechanisms and purposes of communication between individuals

Cricket Songs Are Exploited by Natural Enemies

Figure 1 A katydid (left) and a mole cricket (right). Note the

Figure 2 Experimental design for testing mole cricket response to

Figure 3 Responses of southern and tawny mole crickets to broadcast

Male crickets use sound to attract mates, and this communication is

Figure 4 Monthly captures of female parasitic flies at speakers

Table 1 Numbers of flies captured at traps playing vocalizations from

Frog Choruses Attract Predators

investigators led by Merlin Ryan studied frogs in Panama. Male frogs,

leading to capture of a frog 50

Figure 5 Results of experiments on fringe-lipped bats preying on

Ryan and his colleagues set up an experiment to test the reactions of a

choice of frogs of different sizes, bats preferred the vocalizations of frogs

Table 2 Number of trials needed for fringe-lipped bats to learn to

Organisms Assess Their

Optimal Foraging Behavior of Lizards

diminishing returns. As the number of ants remaining decreases, it gets

Source: From Munger, 1984, Figures 3 and 5.

relatively quickly. There was no point in a lizard staying at a low-quality

Plants Assess their Environment In Search of Nutrients

the scientists extracted mRNA to determine presence and abundance of

Foraging organisms assess their environment and take in information

Egg-Laying Moths Detect Larvae of Other Moths

in a patch. Females of some species have evolved the ability to locate

100 Med moth laying in 100 Indian moth laying in

100 Med moth laying in 100 Indian moth laying in

Figure 6 Results of choice experiments by female Mediterranean

Table 4 Mean mass of adult female and male Mediterranean flour

Source: From Anderson and Lofqvist, 1996, Figures 4 and 5.

by their avoidance of cups that previously held high densities of either

or 20 larvae of the same species, or 10 or 20 larvae total, with half of them

Information Is Used by Corals

maximum survival and growth of coral

Figure 8 Extract concentrations that produced maximum survival and

Algal species D and E, on which settlement and survival of corals were

The data suggest that information is transmitted by species to sig-

A Change in the Number

Ecological systems are made up of many species and environmental fac-

by David Chalcraft and William Resetarits who were interested in how

With newts and salamanders as predators, southern toads nearly or

types, of species existing at a particular point in time of an ecological

Ethical, Legal, Social Implications: We Have an

However, as humans drive other species to extinction, we negatively

A third argument in the debate over our obligation to preserve species

Cane toads, 1011 Eastern mud minnow, 3840

Ephestia kuehniella. See Mediterranean Lizards

Imitator mole cricket, 5 Natural enemies, 1

Parasites, 1 Scapteriscus vicinus. See Tawny mole

Behavior and Information Exchangeby Christopher J. Paradise and A. Malcolm Campbell

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