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American Journal of Botany 90(6): 960–964. 2003.
BOOK REVIEW
PLANT ECOLOGY TEXTBOOKS: A NEW CONTENDER1
RAGAN M. CALLAWAY,2,4 ANNA SALA,2 ELIZABETH CRONE,3
JOHN MARON2
2
Division of Biological Sciences, University of Montana, Missoula, Montana 59812 USA; and 3Wildlife Biology, University of
Montana, Missoula, Montana 59812 USA
In their new text, The Ecology of Plants, Gurevitch et al.
have produced a unique and valuable resource for plant ecol-
ogists. As a tool for teaching, this book is an improvement
over existing plant ecology texts, due to its comprehensive
scope, its rich illustration, and the level at which material is
presented. The Ecology of Plants covers ecophysiology, soils,
evolution, population biology, life history, community struc-
ture and dynamics, ecosystems, landscapes, climate, biomes,
paleoecology, and global change. As we present in detail be-
low, the primary weakness of the text is its organization. How-
ever, because of its strengths, this text is in use at the Uni-
versity of Montana as we write and will be used by some or
all of us in the future. Below we have organized our detailed
comments and opinion on The Ecology of Plants by the order
of the general sections used by the authors.
Gurevitch et al. begin with a short chapter on ecology as a
science. This section will be appreciated by those teaching at
the undergraduate level in particular. The presentation is clear,
the authors do not fall into the trap of Popperian dogmatism,
nor do they simply insist that ecology is more than environ-
mentalism. Instead, we are provided with a broad perspective
on scientific epistemology, the value of experimentation, and
a brief history of ecology. This is a nice opening to the section
that follows, although the leap from Darwin and Haeckel to
John Harper left some of us wondering where Warming,
Schimper, Clements, and Gleason had been placed. We found
Clements and Gleason in wonderful detail in Chapter 12.
The first meaty section is the unit titled ‘‘The Individual and
its Environment.’’ This section deals with aspects of plant
physiological ecology and is divided into chapters on light and
photosynthesis, water relations and energy balance, and soils,
mineral nutrition, and belowground interactions. Within each
chapter, the authors chose a traditional descriptive approach in
which a great deal of useful information is provided. However,
we thought that a more contemporary approach, perhaps em-
phasizing how plant physiological responses under different
environments relate to overall plant strategies within popula-
tion, community, and ecosystem contexts might be more ben-
eficial for students of plant ecology. Overall, there is a lack of
conceptual integration in this section of the book, and the in-
formation does not always follow the most logical structure.
There is also little emphasis on the integration of responses at
different time (from short-term to evolutionary responses) and
spatial scales (from cells to entire plant responses). The student
ends up with an excellent source of ecophysiological infor-
1
The ecology of plants. Jessica Gurevitch, Samuel M. Scheiner, and Gordon
A. Fox. Sinauer Associates, Sunderland, Massachusetts, USA. 2002. 523 pp.
ISBN 0-87893-291-7.
4
E-mail: callaway@selway.umt.edu
960
AND
mation, but little appreciation that plants deal with numerous
physiological and biophysical constraints that impose impor-
tant trade-offs that prevent plants from excelling in each of
their main functional outcomes (e.g., growth, stress tolerance,
defense against consumers, reproduction). There is little de-
velopment in this unit of the historical progression of the field,
and there is insufficient reference to classical work.
Given the traditional approach taken by the authors in Part
I, it is surprising that there is no stand-alone treatment of plant
responses to temperature (there’s a little in Chapter 22). After
all, temperature and water availability are the two most influ-
ential factors on productivity and distribution. While the au-
thors include an energy balance section within the water re-
lations chapter, there is no mention of responses to low and
high temperature. We thought that an initial section on energy
would have provided a stronger general framework for the
following sections on carbon, water, and nutrient relations.
This organization would also emphasize the role of plants as
primary drivers of energy flow within ecosystems and provide
a clear tie to Parts IV and V.
Chapter 2, ‘‘Photosynthesis and the Light Environment,’’
does not emphasize the concept that assimilated carbon is par-
titioned into many competing functions and that overall carbon
allocation strategies have important consequences on produc-
tivity, growth rates, reproductive potential, stress tolerance,
and competitive interactions. While not crucial, this omission
is likely to cause physiology-challenged ecologists a little re-
organizational heartburn. Without reorganization, students may
not fully appreciate that overall biomass accumulation may be
substantially limited by respiration costs, which in turn are
regulated by growth and allocation strategies as well as the
environment (e.g., temperature). A more comprehensive chap-
ter on carbon relations would provide this perspective. As it
is, the chapter quickly moves to explanations of the light and
dark reactions of photosynthesis without providing an overall
conceptual overview of the function of these two important
phases and their interdependence from the onset. While there
is substantial detail on light harvesting, the conceptual under-
standing of how solar energy is used to synthesize ATP in the
chloroplast is not clear (Fig. 2.2 may be misleading). The dark
reactions are described in one short paragraph with no mention
on how the products of the light reactions are used in the dark
reactions. There is no mention at this point that actual carbon
‘‘fixation’’ and reduction occur during this phase. This is un-
fortunate as it is this initial reduction by plants that allows
energy transfer through trophic levels within ecosystems. The
dual affinity of rubisco to CO2 and O2 and the consequent
photorespiratory cycle are not brought up until the discussion
on photosynthetic pathways. Adding to the organizational con-
June 2003] BOOK REVIEW
fusion, the chapter moves into a discussion of the effects of
light and carbon uptake limitations prior to a more in-depth
description of the dark reactions; these are included in an ex-
planation of the different photosynthetic pathways. The chap-
ter includes strong sections on evolutionary aspects of pho-
tosynthetic pathways, geographic distribution, habitat prefer-
ences, and adaptations to the light environment. However, the
last section should be titled ‘‘Plant Responses’’ because it in-
cludes both adaptation and acclimation. The information is ex-
cellent, but we believe the chapter would be improved if the
light and dark reactions in C3 plants were presented in suffi-
cient conceptual detail first, and afterwards the chapter pro-
ceeded to the ‘‘problem’’ of rubisco and photorespiration in
C3 plants and the evolution of alternative photosynthetic path-
ways with consequent costs and benefits. With this organiza-
tion, the progression to the responses of plants with different
pathways to light, water, and temperature, the relationship of
pathways to habitat preference and geographic distribution,
and general ecological strategies would flow naturally. Links
between this chapter and others could be made by emphasizing
the links between photosynthetic traits and plant life history
traits (e.g., fast- vs. slow-growing plants, early vs. late suc-
cessional species).
The chapter on water relations suffers from the same lack
of integration and cohesive structure. The abundant informa-
tion is often scattered (e.g., responses to flooding or drought)
or missing all together (e.g., there is no mention of the concept
of water-use efficiency or the importance of xylem structure
in water relations). Students and instructors without strong
ecophysiological backgrounds would benefit from a simple
discussion of why plants lose so much water, the main com-
partments that water moves through in the plant (introducing
here the concept of the relative role of these compartments to
maintain a functional homeostasis with respect to water trans-
port), and then a clear explanation of water potential as the
driving force for water movement. In the discussion of water
potential the authors make a minor but common mistake by
portraying matric potential as an independent component of
the total water potential. Matric potential is actually a com-
bination of osmotic and hydrostatic pressures and should be
omitted from the equation. A more linear explanation of the
water potential components along the overall plant–atmo-
sphere continuum could logically lead to an explanation of
why xylem water potential is such a good indicator of overall
plant water status and a variable commonly measured by plant
ecologists. (Fig. 4.10 in the soil chapter would be much more
relevant here.) The sections on strategies for coping with water
availability are quite unorganized. The student should be pro-
vided with a clearer understanding that plants exhibit a variety
of phenological, structural, anatomical, and physiological re-
sponses to variation in water availability and that these re-
sponses may operate at different times (short-term to evolu-
tionary time scales) and spatial scales (from cells to entire
plants). The ‘‘strategies’’ employed by plants to cope with wa-
ter stress are presented more as a list and could have been
divided into groups—for example, species that ‘‘avoid’’ any
activity during water stress (drought escape) vs. plants that
remain more or less active during drought (drought resistance).
Perhaps it is reasonable for a general text on plant ecology to
omit some important strategies for drought resistance, but in-
creased water transport and storage tissues in woody plants
and increased resistance to xylem cavitation are missing and
961
hydraulic lift (which should probably be generalized to ‘‘hy-
draulic redistribution’’) by roots comes later in the soil chapter.
Of the three chapters in Part I, the chapter on soil and min-
eral nutrition is the most deficient. The chapter starts with a
great deal of descriptive detail on soil composition and struc-
ture without an initial discussion of why these details are im-
portant to plants. There is no emphasis on the importance of
plant mineral nutrition or its effect on functional strategies and
ecosystem properties. While the chapter goes into great detail
on soil properties it does not explore in a logical order the
many strategies plants employ to deal with limiting nutrients
(mycorrhizae and nitrogen fixation are in independent sec-
tions) and the trade-offs associated with these strategies. Not
including these potential trade-offs is not crucial, but they
would offer a great opportunity to link this chapter to the car-
bon and water relations chapters and to subsequent chapters
on ecological strategies and plant–animal interactions. The
chapter omits fundamental concepts such as nutrient-use effi-
ciency, nutrient residence time (or life span), nutrient recy-
cling, and the relationship of these traits to growth forms (e.g.,
a distinction between evergreen and deciduous species, etc.).
There is little or no emphasis on how mineral nutrition feeds
back to soil properties (including microbes) and the implica-
tions for community and ecosystem dynamics. In much the
same way as we would like to see for the water relations
chapter, why not start with an exploration of why mineral nu-
trients are needed for plant growth and how soil properties
affect nutrient availability and end with a clear overview of
how plants respond to variation in soil fertility and physical
characteristics?
Like much of the rest of the text, the population biology
section (Chapters 7–9, or 5–9 if you include the two chapters
on evolution) is comprehensive, but the organization is some-
what obscure. The section begins with a general discussion of
evolutionary processes (Chapters 5 and 6), followed by pop-
ulation growth (Chapter 7), individual growth (Chapter 8), and
life history theory (Chapter 9). We agreed that for teaching
plant ecology, we would almost certainly present the material
in a different order than in the text—at a minimum, by putting
individual growth before population growth and after evolu-
tion, which has traditionally been approached by plant ecolo-
gists from an individual perspective (for example, estimating
fitness from individual survival and fecundity, rather than by
projecting genotype-specific lambdas). Yet Gurevitch et al.
should be commended for including chapters on evolutionary
processes in their book, as these important topics are missing
from many plant ecology texts. Indeed, with the growing in-
terest among ecologists in rapid evolution and the role of evo-
lutionary processes in plant invasions, there is ample scope for
even broader coverage and integration of ecology with evo-
lution in textbooks.
The material covered in the subsequent chapters on popu-
lation biology was also presented in a somewhat haphazard
manner. For example, the chapter on population growth only
briefly mentions dispersal and density dependence, and almost
entirely focuses on age- or stage-structured transition matrices,
with an extensive discussion of matrix modeling methods, in-
cluding calculation of sensitivity, elasticity, reproductive val-
ue, as well as methods for incorporating environmental sto-
chasticity. In our experience, undergraduates would benefit
from a more complete discussion of basic (i.e., unstructured)
population models before moving so quickly into a detailed
discussion of matrix projections. Gurevitch et al.’s emphasis
962 AMERICAN JOURNAL
may well reflect the state of the art in plant demography, but
our sense is that the groundwork for this material isn’t ade-
quate. For example, there is only a brief discussion of life
tables, but no real discussion of the elements of a life table or
different approaches to collecting data for life tables (i.e., co-
hort, partial cohort, or static analyses), or how one integrates
life table statistics into projection matrices. Although the au-
thors cite a few alternative methods to transition matrices, al-
ternative mechanisms of population growth are scattered
among other chapters. For example, clonal spread is discussed
in detail in Chapter 8, without reference to methods for infer-
ring population growth and spread from clonal tillering, which
is, in our opinion, an approach much more naturally suited to
rhizomatous herbaceous perennials than transition matrices.
Density-dependent population growth models are presented in
the context of r vs. K selection in Chapter 9, but only dis-
cussed qualitatively in the population growth chapter. Meta-
population models and source–sink dynamics appear in Chap-
ter 17 (landscape ecology), creating the impression that spatial
dynamics and population dynamics describe different process-
es, rather than acting as ends of a continuum of population
structures. Interspecific competition is presented in Chapter 10,
but the relationship to density-dependent population growth is
not clear from the text alone; it would need to be made by a
course instructor.
Similarly, Chapter 8 combines basic plant morphology; a
generalized life cycle; clonal growth, foraging, and reproduc-
tion (awkwardly split into two disjunct sections); a large sec-
tion on pollination biology; and a discussion of seed dispersal
and seed banks. There is no real synthesis of the different
methods; e.g., comparison of ‘‘dispersal’’ by branching, ra-
mets, pollen, or seeds. Neither is it clear why dispersal belongs
with individual growth, rather than population growth. In the
discussion of clonal growth, as in other places throughout the
book, those whom we perceive as movers and shakers in the
field (de Kroon and van Groenendael [1997], Cain [1994], and
Alpert [1991], for example) were not mentioned. This chapter
would have been better split into three; one dealing with mod-
ularity and clonal growth, the second discussing reproduction,
pollination ecology, and pollination syndromes, and the third
discussing frugivory, seed dispersal, and seed predation. We
found the section on the ecology of fruits and seeds in Chapter
8 particularly frustrating, as the coverage of frugivory, seed
dispersal and predation, and seed dormancy was rudimentary
at best, although granivory is covered in more detail in Chap-
ter 11.
The chapter on life history (Chapter 9) includes a general
discussion of life history strategies (annual vs. perennial; se-
melparous vs. iteroparous; seed dormancy and germination re-
quirements), seed size vs. seed number trade-offs, phenology,
and a brief section on using demographic models to quantify
life history trade-offs. The demography section is a bit repet-
itive with Chapter 7; for example, the justification for why
geometric mean population growth is the appropriate measure
of fitness and long-term population growth is given in both,
though in slightly different words. We believe that these sec-
tions would also be greatly improved by tighter links among
chapters; for example it is almost—but not quite—clear that
matrix models are central to life history theory because ge-
notype-specific population growth rates are estimates of fit-
ness. A brief appendix is presented to discuss simple statistics,
with description of simple statistics like mean and variance,
and verbal explanations of statistical tests and P values. The
OF BOTANY [Vol. 90
combination of the very basic and general appendix with the
detailed section on matrix calculations in Chapter 7 seems tar-
geted to different audiences. We were impressed by the effort
to explain to readers how to do calculations for population
models; it can be frustrating when texts mention that models
exist and provide nifty results, but do not explain relevant
calculations.
Criticisms stated, we agree that The Ecology of Plants is
the text we would choose if population biology was the em-
phasis in an ecology course. As for other sections, our opin-
ion was swayed by the impressive breadth of engaging ma-
terial.
Following Part III, ‘‘From Populations to Communities,’’
the text springs immediately into competition. The commu-
nity context comes in Chapter 12 in a rather traditional dis-
cussion of Clements and Gleason. The chapter on competi-
tion is strong, but is missing or only perfunctorily addresses
current advances in indirect interactions, allelopathy, and, to
the chagrin of at least one of our reviewing crew, facilitation.
Competitive hierarchies are mentioned, but the fascinating
implications of nontransitive competitive webs (indirect in-
teractions among competitors á la Thomas Miller [1994] and
Jonathan Levine [1999] are not explored). One of us thought
that this was the most serious oversight in the text. Positive
interactions among plants (Callaway, 1995) are mentioned in
a later chapter in the obligatory discussion of the mechanisms
driving succession, but are otherwise ignored. Allelopathy is
written off in the standard manner of the 1980s with the add-
ed twist of misinterpreting a recent paper written by one of
our review group (Callaway and Aschehoug, 2000). For a
general text, the sections on modeling competition are clearly
written and nicely comparative, and the section on competi-
tion along abiotic gradients is up to date and excellent. This
chapter may be improved moderately by incorporating classic
statements by John Harper (1961) on distinguishing compe-
tition for resources from other forms of interference.
Herbivory and pathogens are treated in the same chapter,
appropriately in our opinion, and this chapter includes a short
but highly effective consideration of biological control of ex-
otic plants by insects, but with no mention of pathogens as
biocontrols. The subsections in the chapter ‘‘Effects of Her-
bivory on the Community Level’’ begin with the consequences
of herbivore behavior, move smartly through subsections on
introduced and native herbivores and on to a too-small portion
of ‘‘generality.’’ One glaring hole in this section is the omis-
sion of a discussion on McNaughton’s (1983) classic work in
Africa and on Richard Root’s (1996) work on the community
impacts of herbivores feeding on goldenrod. However, the sec-
tion on plant defenses is top-notch and thorough, although a
clear treatment of the controversy on the costs of defense
would have rounded this section out and possibly provided a
tie to earlier chapters on carbon allocation. Following this are
two pages on ‘‘evolutionary consequences’’ of herbivory that
are not connected in any way to communities. Ecology teach-
ers beware; if your goal is a clear organizational picture of
plant ecology for relatively naive students, you will have to
reorganize. The pathogen section is a quick mention of some
cool ecological tidbits, but is not tied to the previous text on
herbivory at all.
Chapter 12, on community properties, comes without apol-
ogy or comment after the chapters on competition and con-
sumers. We think that this organizational strategy could have
worked, but it would have taken some clear explanation of
June 2003] BOOK REVIEW
how the stuff just digested on interactions leads in some way
to the properties of communities. One approach to bridging
chapters on competition and herbivory to the community prop-
erties material might have been to include a chapter explicitly
dealing with food webs. Rich topics such as trophic cascades,
keystone species (Power et al., 1996), subsidies, and subsi-
dized food webs are mostly ignored in The Ecology of Plants,
and inclusion of these topics would have made for engaging
reading. In this section the enormous body of work describing
plant communities is limited to species richness, diversity,
dominance, and basic sampling strategies—multivariate ordi-
nation and classification come four chapters later in an entirely
different section, a chapter titled ‘‘Communities in Land-
scapes’’ in Part IV, ‘‘From Ecosystems to Landscapes.’’
Chapter 13, on succession, is standard but strong. Students
will come away with a clear understanding of succession and
why it is important to plant communities. Missing however, is
any mention of cyclic or shifting mosaic dynamics, and the
subsection on primary section follows sub-subsections on vol-
canoes, fire, colonization, and others, but no parallel subsec-
tion on secondary succession. This is also one of the only
chapters that simply does not include enough current literature
on succession.
Chapter 14 is a peach, and here Gurevitch et al. have made
a significant stride in teaching ecology. By thoughtfully prob-
ing dominance, abundance, and rarity, and including theory
developed by Deborah Goldberg and colleagues, David Til-
man, MacArthurian distribution curves, and niche theory, this
chapter in The Ecology of Plants will provide advanced stu-
dents and most professors with stimulating thoughts and new
directions. These concepts are blended smoothly with a fol-
lowing section on invasives that will no doubt help make the
material more relevant to undergraduates. In our opinion, how-
ever, the treatment of invasives could have been bolstered by
both a clear explanation of the natural enemies hypothesis as
the primary hypothesis for invasive success and a broader in-
clusion of the processes driving invasion such as presented by
Richard Mack and colleagues (2000). More generally, inclu-
sion of neutral community theory such as that promoted by
Steven Hubbell (2001) would have fit well in the discussion
of abundance and dominance.
The Ecology of Plants leaves communities and proceeds to
ecosystems cleanly. ‘‘Processes’’ in Chapter 15 are nutrient
cycling, carbon flow, and productivity, but nowhere is the
movement of energy explicitly and clearly explained as a cru-
cial ecosystem process. This chapter includes two very nice
subsections on nitrogen and phosphorus in ecosystems, inter-
rupted oddly by an equivalent subsection titled ‘‘Ecosystem
Nutrient Cycling and Plant Diversity,’’ after which puzzled
readers will find processes described for other elements. A
simple move of the diversity section to the end would solve
the problem.
Chapter 16 probably comprises the most unusual large-scale
organizational choice of all. We could find no connection be-
tween ‘‘communities in landscapes’’ and ecosystems. On the
other hand, multivariate ordination is just skipped by many
instructors, and this may be easier to do when it is hiding in
an ecosystems unit. This said, The Ecology of Plants deals
with the messy issues of ordination and classification quite
nicely. Direct and indirect ordination are clearly contrasted,
several modern techniques are presented, and this is all fol-
lowed by a connection with remote sensing. This chapter is
followed by a nice chapter on landscape ecology, which com-
963
bines the explanation of remote techniques with island bio-
geography and metapopulations theory. The latter two bodies
of theory link perfectly, but these were not tied to remote
landscape analysis.
Part V, ‘‘Global Patterns and Processes,’’ is one of the best.
Beginning with a superb unit on basic global climate that
should be read just for fun, the transition to biome physiog-
nomy in the last sections and the following chapter (Chapter
19) is smooth. Chapter 20 on regional and global diversity is
a unique and brilliant follow-up to the biome chapters, the
ideas are stimulating, and the way that both old and new ma-
terial are integrated ought to change some syllabi. Although
none of the review team can claim paleological expertise, this
short but nicely placed chapter is almost as good a read as the
unit on global climate. Our one recommendation is that this
chapter could be better connected with the section on com-
munities.
Global change issues such as global warming, nitrogen de-
position, invasive species, and habitat fragmentation are pre-
sented in well-organized detail in the concluding chapter of
The Ecology of Plants. These topics are increasingly an im-
portant component of ecology that have traditionally been
given short shrift in texts. Global warming is clearly tied to
Chapter 2 on carbon uptake and Chapter 15 on the carbon
cycle and productivity. Increasing atmospheric CO2 is ex-
plained, and these effects on plants are separated clearly from
the cascading effects of CO2 on temperature and the effects
of temperature on plants. The fluxes in the carbon cycle fig-
ure correspond reasonably well with those in Houghton et al.
(1995); however, the section would be improved by a simple
layman’s explanation of how greenhouse molecules compris-
ing a tiny fraction of the atmosphere can force global warm-
ing. We were told to go to Chapter 2 for comments on the
effects of CO2 on temperature, but this didn’t help much. We
would have loved to have seen the Croll-Milancovič theory
for long-term climate change integrated with their presenta-
tion of the greenhouse-gas forcing theory for global warm-
ing. To one of us, Chapter 22 did not express reasonable (read
‘‘any’’) skepticism for the predictive global change scenarios
presented and took on a decidedly biased political tone when
discussing fossil fuel combustion. However, these opinions
should not be taken as reasons to dismiss a very good final
chapter in an excellent Part V.
Our complaints about the organization of The Ecology of
Plants should not be taken as a dismissal of the book as a
high-quality text. On the contrary, even with its organizational
shortcomings, this book provides the most comprehensive,
well-integrated, and accessible treatment of plant ecology
available and is perfectly suited for undergraduate teaching.
Gurevitch et al. use pictures, graphs, and diagrams liberally.
They use box insets frequently and effectively. A wonderful
aspect of the book is the sprinkling of names and pictures of
many ecologists that are cited in the text. All of these features
make for an engaging text that should provide stimulating
reading for undergraduates and a useful reference for more
advanced students.
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