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Evolutionary Ecology 7th Edition Eric R. Pianka Digital
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Author(s): Eric R. Pianka
ISBN(s): 9780321042880, 0321042883
Edition: 7
File Details: PDF, 14.40 MB
Year: 2011
Language: english
Evolutionary
Ecology
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Evolutionary
Ecology
Seventh Edition - eBook
Eric R. Pianka
Text Designer and Compositor: Eric R. Pianka
Copy Editor: Eric R. Pianka
Cover: Thorny devil lizard, Eric R. Pianka
Evolutionary Ecology, Seventh Edition – eBook
Copyright © 2011 by Eric R. Pianka
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, mechani
cal, photocopying, recording, or any other media embodiments now known or
hereafter to become known, without the prior written permission of the publisher.
Manufactured in the United States of America.
Previously published in earlier editions by Harper and Row, HarperCollins, and
Addison-Wesley Longman (Benjamin Cummings)
Library of Congress Cataloging-in-Publication Data
Pianka, Eric R.
Evolutionary Ecology / Eric R. Pianka. – 7th edition - eBook
Includes bibliographical references and index
1. Ecology. 2. Evolution (Biology) I. Title.
QH541 .P7 2011
577--dc21
v
| For this generation,
who must confront the
shortsightedness of their ancestors
vi
vii
| Contents
| Preface xii
1 | Background 1
Scaling in Biology 1
Scientific Methods and Human Knowledge 2
Domain of Ecology: Definitions and Groundwork 5
The Urgency of Basic Ecological Research 8
Natural Selection 10
Levels of Approach in Biology 12
Debates and Progress in Ecology 13
2 | History and Biogeography 16
Self-Replicating Molecular Assemblages 16
The Geological Past 17
Classical Biogeography 22
Continental Drift 24
3 | Meteorology 29
Earth’s Physical Environment 29
Major Determinants of Climate 30
Local Perturbations 35
Variations in Time and Space 37
Global Weather Modification 42
viii
4 | Climate and Vegetation 48
Plant Life Forms and Biomes 48
Microclimate 49
Primary Production and Evapotranspiration 53
Soil Formation and Primary Succession 58
Ecotones and Vegetational Continua 61
Classification of Natural Communities 63
Aquatic Ecosystems 65
5 | Resource Acquisition and Allocation 70
Limiting Factors and Tolerance Curves 70
Resource Budgets and the Principle of Allocation 72
Time, Matter, and Energy Budgets 73
Leaf Tactics 75
Foraging Tactics and Feeding Efficiency 78
Physiological Ecology 83
Physiological Optima and Tolerance Curves 84
Energetics of Metabolism and Movement 86
Adaptation and Deterioration of Environment 90
Heat Budgets and Thermal Ecology 93
Water Economy in Desert Organisms 97
Other Limiting Materials 98
Sensory Capacities and Environmental Cues 99
Adaptive Suites 100
Design Constraints 103
6 | Rules of Inheritance 110
Basic Mendelian Genetics 110
Nature versus Nurture 115
Selfish Genes 116
Population Genetics 117
Maintenance of Variability 118
ix
Units of Selection 120
Genetic Engineering 122
7 | Evolution and Natural Selection 124
Agents of Evolution 124
Types of Natural Selection 124
Ecological Genetics 126
Allopatric and Sympatric Speciation 127
Reproductive Isolating Mechanisms 128
Galápagos Finches 129
8 | Vital Statistics of Populations 134
Individuals Versus Populations 134
Life Tables and Tables of Reproduction 135
Net Reproductive Rate and Reproductive Value 142
Stable Age Distribution 147
Leslie Matrices 148
Intrinsic Rate of Natural Increase 150
Demographic and Environmental Stochasticity 153
Evolution of Reproductive Tactics 154
Reproductive Effort 155
Expenditure per Progeny 160
Patterns in Avian Clutch Sizes 163
Evolution of Death Rates and Old Age 169
Joint Evolution of Rates of Reproduction and Mortality 171
9 | Population Growth and Regulation 177
Verhulst–Pearl Logistic Equation 177
Derivation of the Logistic Equation 182
Density Dependence and Density Independence 182
Opportunistic versus Equilibrium Populations 184
Population Regulation 188
x
Population “Cycles”: Cause and Effect 193
10 | Sociality 200
Use of Space: Home Range and Territoriality 200
Sex 203
Sex Ratio 207
Sexual Selection and Mating Systems 211
Fitness and an Individual’s Status in Its Population 220
Social Behavior and Kin Selection 220
The Evolution of Self-Deceipt 225
11 | Interactions Between Populations 228
Direct Interactions 228
Complex Population Interactions 229
Mutualistic Interactions and Symbiotic Relationships 231
Indirect Interactions 236
12 | Competition 240
Mechanisms of Competition 240
Lotka–Volterra Competition Equations 241
Competitive Exclusion 248
Balance Between Intraspecific and Interspecific Competition 249
Evolutionary Consequences of Competition 252
Laboratory Experiments 252
Evidence from Nature 254
Other Prospects 264
13 | The Ecological Niche 267
History and Definitions 267
The Hypervolume Model 269
Niche Overlap and Competition 271
xi
Niche Dynamics 274
Niche Dimensionality 275
Niche Breadth 279
Evolution of Niches 289
14 | Experimental Ecology 294
Design of Experiments 294
Ecological Experiments 294
A Defaunation Experiment 299
15 | Predation and Parasitism 302
Predation 302
Predator–Prey Oscillations 304
“Prudent” Predation and Optimal Yield 312
Selected Experiments and Observations 313
Evolutionary Consequences: Prey Escape Tactics 315
Parasitism 322
Epidemiology 326
Darwinian Medicine 328
Coevolution 329
16 | Phylogenetics in Ecology 337
Phylogenetic Systematics 337
Vicariance Biogeography 338
Phylogeny and the Modern Comparative Method 338
Phylogenetically Independent Contrasts 340
Evolutionary Ecomorphology 341
17 | Community and Ecosystem Ecology 345
Systems and Macrodescriptors 345
Systems Ecology 347
xii
Compartmentation 348
The Community Matrix 352
Biogeochemical Cycles in Ecosystems 355
Principles of Thermodynamics 356
Pyramids of Energy, Numbers, and Biomass 358
Energy Flow and Ecological Energetics 359
Secondary Succession 363
Evolutionary Convergence and Ecological Equivalence 366
Community Evolution 368
Pseudocommunities 369
Landscape Ecology and Macroecology 379
18 | Biodiversity and Community Stability 388
Saturation with Individuals and with Species 388
Species Diversity 390
Latitudinal Gradients in Diversity 394
Types of Stability 403
Community Stability 406
19 | Island Biogeography and Conservation Biology 413
Species–Area Relationships 414
Equilibrium Theory of Island Biogeography 416
Islands as Ecological Experiments: Some Examples 420
The Taxon Cycle 422
Experimental Biogeography 424
Conservation Biology 424
| References 431
| Index 505
| Preface
During the past five decades, evolutionary ecology has blossomed into a
massive discipline that has assimilated and largely replaced other ecological
subdisciplines. Until this intellectual revolution, most biologists merely
accepted a broad range of biological phenomena as essentially immutable,
such as the fact that sex ratios are usually near equality (1:1), without con
sidering why such facts might be so or how they could have evolved. Rigor
ous application of the theory of natural selection in population biology has
greatly increased our understanding of numerous phenomena, including the
evolution of such things as genetic dominance, foraging modes, reproductive
tactics, senescence, sex ratios, a wide range of social behaviors, mate
choice, mating systems, predator escape tactics, parasite virulence, niche
breadth, resource partitioning, and so forth. The evolutionary approach has
opened up whole new areas of endeavor such as optimal foraging, life his
tory tactics, sexual selection, and coevolution, each of which became an
instant subdiscipline and then quickly exploded into a field in its own right.
When I began teaching ecology over thirty years ago, only two textbooks
existed — both totally lacked an evolutionary perspective. In 1970, I grew
tired of apologizing to my students for the inadequacies of those books and I
decided to write a more modern one based on my own lectures. My approach
to ecology had been strongly influenced by the work of Robert H. Mac
Arthur, an intellectual giant who held center stage in ecology until his pre
mature death in 1972. Ecology today would be very different if Robert had
not died. I like to think that my text is close to the book that MacArthur
would have written, had he lived. Evolutionary Ecology is a “Citation Clas
sic” (Current Contents, 20 June 1988) and has now been translated into
Greek, Japanese, Polish, Russian, and Spanish. Several other books and even
a journal have adopted the same title, and courses in evolutionary ecology
are now being offered around the world.
Some ecologists favor a more mechanistic approach than I do. Some people
perceive Evolutionary Ecology to be more “advanced” than other ecology
texts, perhaps because I do not shy away from simple mathematics. I use my
text in a freshman-level classes as well as in an advanced upper-division
class called Evolutionary Ecology. In the freshman class I downplay the
xiv Preface
math, but in the upper division course I emphasize that ecology is necessar
ily a quantitative subject, and I insist that students make an effort to under
stand all the math, which is really only high school algebra.
My primary goal in teaching has always been to encourage my students to
think clearly. Natural selection is frequently misunderstood and naive, group
selectionist explanations are often given for biological phenomena. Thinking
about the evolution of various ecological attributes must be done carefully
and correctly. Evolutionary Ecology is a strongly conceptual book. I have
found that the best way to use my book is to adorn its concepts to students in
lectures using real examples from my own personal field experience.
In previous editions, I have always tried to present evolutionary ecology as a
“pure” science. Humans now dominate ecosystems to such an extent that
pure ecology has all but vanished from the face of the earth! Hence, in this
edition, multitudinous anthropogenic effects are interwoven into every chap
ter: these include overpopulation, many different kinds of pollution of the
atmosphere, water, and land (and the manifold effects of such pollution on
the health and livelihood of plants and animals, including ourselves), habitat
destruction and fragmentation, endangered species, loss of genetic variabil
ity, extinction, disruption of natural ecosystems, human transportation of
organisms and the resultant homogenization of earth’s biota, evolution of
microbes that infect humans as hosts, and murder rates among humans. A
new chapter has been added on the role of phylogeny and modern compara
tive methods in deducing the probable actual course of evolution with exam
ples of the utility of this approach in modern ecology. Other new topics
include subjects such as conservation biology, demographic stochasticity,
Leslie matrices, the sunspot and time lags explanations for cyclic popula
tions, global weather modification, experimental biogeography, evolution
ary ecomorphology, the complexity-stability controversy, Jacobian matrices,
evolutionary epidemiology, and Darwinian medicine.
In this edition, island biogeography is presented late in the book and it is
integrated with metapopulations and conservation biology.
Language forces us to express ourselves in a one-dimensional stream of
words. Nature, however, is multidimensional. This is particularly true of
ecology because its subject matter includes many complexly interrelated
concepts and phenomena involving several different levels of organization.
There is no such thing as an “ideal” outline or a perfect sequential order for
presentation of the subject matter treated here. To obtain an overview of
modern ecology, a student needs to assimilate a great many ideas. Ideally, a
reader would know everything in this book even before beginning to read it!
Preface xv
Perhaps the only solution is to read it twice. To assist readers in following
different trains of thought than I have chosen to use, various chapters and
sections are cross-referenced.
Evolutionary Ecology is my own eclectic blend and distillation of what I
consider to be significant ideas and principles; they represent the residue
remaining after considerable sifting and sorting through many other facts
and ideas. My approach is abstract and conceptual, and I strive to provide a
reasonably crisp overview of a fairly vast subject matter. This edition of
Evolutionary Ecology represents an image of a part of my mind, a part that
should mirror an external reality common to all living systems anywhere in
the cosmos.
People have a tendency to stop citing (and quit reading) someone once he/
she dies, in favor of stroking living egos who pass judgment on grant pro
posals and publications. As a result, the older literature is seriously under
valued and well-meaning scientists are doomed to continually reinvent the
wheel. Wherever possible, I continue to give credit to originators of concepts
even though it requires citing material that some might consider “dated.”
Most scientific endeavors are quite pedestrian. Thus, the research projects in
which most scientists engage themselves are relatively trivial, constituting
mere building blocks for major advances. Such “normal” science is, of
course, absolutely essential in that it provides the raw material for progress
in understanding. Periodically an extraordinary event occurs that enables a
novel breakthrough. Occasionally, this may be just a serendipitous discovery
by a more or less “ordinary” scientist (provided, of course, that someone has
the wisdom to appreciate the true significance of the discovery and the cre
ativity to develop it). But more often than not, major new directions are
charted by rare individuals with incredible intellectual prowess. Population
biology has attracted a few of these people in the past, and ecology today
stands poised, awaiting its next such genius.
Eric R. Pianka, Austin, Texas, June 2011
xvi Preface
1 Background
Humans seem to delight in animal motifs — thus, we have automobiles, airplanes,
and athletic teams named after various animals: cougar, jaguar, lynx, mustang,
pinto, ram, eagle, falcon, nighthawk, roadrunner, and the list goes on. Zoos are a
popular form of entertainment, particularly for children. Yet, many people feel
threatened by a free-ranging wild creature, even by a tiny mouse or a harmless
snake. Indeed, urbanization is now so complete that, aside from cockroaches and
songbirds (and perhaps while on vacation), most of us seldom encounter wild ani
mals.
What is the essential difference between a wild animal versus one in a cage?
Clearly, a rattlesnake behind glass does not pose nearly as much physical danger to
a human observer as does a wild rattlesnake. For the study of many kinds of bio
logical phenomena, there is in fact no difference between a caged specimen, so long
as it remains alive, and its wild cousin. The constrained one still has intact cells,
molecules, physiological processes, and, to some extent, behavior. But the caged
animal, removed from its habitat, is out of context — it has been stripped of its nat
ural history and it no longer interfaces with the environment to which it is adapted
and in which it evolved.
The biological discipline of ecology deals with a myriad of ways in which organ
isms (plants, animals, and other heterotrophs such as bacteria and fungi) interact
with, influence, and are in turn influenced by their natural surroundings. Wild plants
and animals in their natural communities constitute the subject matter of ecologists.
To these scientists, caged organisms might as well be dead for they have no ecol
ogy. Ecology differs from other sorts of biology in that its perspective is directed
upward and outward from the individual organism to its environment. Other kinds
of biology focus on organismal and suborganismal processes and thus involve a
reductionistic viewpoint. Ecology has a more holistic perspective.
Scaling in Biology
Biology is a vast discipline ranging from molecules to cells to tissues to whole
organisms to kin groups to populations to communities (and clades) to entire
ecosystems, or even the entire global biosphere. Across this broad range of scale,
factors vary by many orders of magnitude (Figure 1.1).
2 Scientific Methods and Human Knowledge
Figure 1.1. Diagrammatic representation of the time-space scaling of various
biological phenomena. Community and ecosystem phenomena occur over longer
time spans and more vast areas than suborganismal- and organismal-level processes
and entities. [Redrawn from Osmund et al. (1981)]
Molecular biology can be done quickly in small spaces, but community ecology
requires decades and square kilometers. Biogeographic and historic events take
place over many millennia. Continental plates have moved across thousands of
kilometers over geological time. Until recently, ecologists have been preoccupied
with local phenomena and events occurring on relatively short time scales. How
ever, emerging new subdisciplines of landscape ecology and macroecology (pp.
379–385) offer promising new regional and global perspectives.
As an example of the effects of scale, consider movements of organisms across
landscapes (Brooks 1988). At the shortest temporal and spatial scale, individual
organisms move during their lifetimes and disperse; over an intermediate scale in
space and time, immigration and emigration occur between and among populations
(see also section on metapopulations, pp. 426-427); over a much greater spatial
extent and at a much longer time scale, geographical ranges shift in response to cli
matic changes and to vicariant events such as geotectonic movements, leading ulti
mately to the formation of geographical patterns in species diversity.
Scientific Methods and Human Knowledge
Scientists are motivated by curiosity about their surroundings; they go to great
lengths to satisfy their desire to understand natural events and phenomena. All sci
Chapter 1Background 3
entists assume that an organized reality exists in nature and that objective principles
can be formulated that will adequately reflect this natural order. A fundamental and
important way in which biological phenomena can be ordered is by simple and
direct enumeration, as in the classification of organisms or biotic communities.
Thus, ecologists recognize different ecological systems such as tundra, desert, prai
rie, savanna, deciduous forest, coniferous forest, and rain forest. Early ecology was
primarily descriptive; the originators of the science spent most of their time
describing, itemizing, and classifying various ecological elements. This process
was absolutely essential before a more process-oriented ecology could be
developed.
Founded and firmly based on this older body of descriptive information, modern
ecology seeks to develop general theories with predictive powers that can be com
pared against the real world. Ecologists want to understand and to explain, in gen
eral terms, the origin and mechanisms of interactions of organisms with one
another and with the nonliving world. To build such general theories of nature,
ecologists construct hypotheses, hypothetical “models” of reality. All models must
make simplifying assumptions — some sacrifice precision for generality, whereas
others sacrifice generality for precision. Some models actually sacrifice certain
aspects of realism itself! Models have been described as “mere caricatures of nature
designed to convey the essence of nature with great economy of detail.” No model
is “correct” or “true” — any given model merely represents one particular attempt
to mimic reality. All models are to some extent incorrect. To be most useful, models
are usually designed to generate testable predictions. Most models can therefore be
confronted with reality and can be falsified. But not all models are refutable; some
conceptual models have proven to be useful in an abstract way even though they do
not suggest direct tests.
When a model’s predictive powers fail, it is either discarded or revised (Popper
1959). Models and hypotheses that do not conform adequately to reality are gradu
ally replaced by those that better reflect the real world (Lakotos 1970). The scien
tific method is thus self-regulating; as time progresses, knowledge expands and is
continually refined and improved to reflect external reality better and better. Well
substantiated hypotheses become theories. We are very fortunate indeed to be able
to benefit from past genius and research effort; in a few hours of careful reading,
you can now learn material that required many lifetimes to acquire.
Due to the multiple meanings of words, verbal models are usually somewhat
ambiguous and imprecise and therefore of limited utility. The great complexity of
ecological systems necessitates the use of graphical and mathematical models, so
much so that ecologists often employ nearly as much mathematics as biology. How
Other documents randomly have
different content
Plate 272
GATE OF THE CONVENT OF SANTO TOMÁS
AVILA
Plate 273
DOOR OF SANTO TOMÁS
AVILA
Plate 274
INTERIOR OF SANTO TOMÁS
AVILA
Plate 275
THE COURT OF SILENCE, IN THE CONVENT OF SANTO TOMÁS
AVILA
Plate 276
CONVENT OF SANTO TOMÁS. THE COURT OF SILENCE
AVILA
Plate 277
CONVENT OF SANTO TOMÁS. COURT OF THE KINGS
AVILA
Plate 278
CONVENT OF SANTO TOMÁS. COURTYARD OF THE INFIRMARY
AVILA
Plate 279
CLOISTERS OF SANTO TOMÁS
AVILA
Plate 280
CLOISTERS OF THE CONVENT OF SANTO TOMÁS
AVILA
Plate 281
CLOISTERS OF THE CONVENT OF SANTO TOMÁS
AVILA
Plate 282
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