AN INTRODUCTION TO METHODS & MODELS IN Ecology, Evolution, & Conservation Biology This page intentionally left blank AN INTRODUCTION TO METHODS & MODELS IN Ecology, Evolution, & Conservation Biology STANTON BRAUDE & BOBBI S. LOW Editors PRINCETON UNIVERSITY PRESS PRINCETON AND OXFORD Copyright 2010 © by Princeton University Press Published by Princeton University Press, 41 William Street, Princeton, New Jersey 08540 In the United Kingdom: Princeton University Press, 6 Oxford Street, Woodstock, Oxfordshire OX20 1TW All Rights Reserved Library of Congress Cataloging-in-Publication Data An introduction to methods and models in ecology, evolution, and conservation biology / Stanton Braude and Bobbi S. ISBN 978-0-691-12723-1 (hardcover : alk. paper) — ISBN 978-0-691-12724-8 (pbk.
Conservation biology— Research.072—dc22 2009012206 British Library Cataloging-in-Publication Data is available This book has been composed in Minion Printed on acid-free paper.edu Printed in the United States of America 1 3 5 7 9 10 8 6 4 2 Contents Figures vii Tables xi Preface xv Acknowledgments xvii Introduction xix Section I Evolutionary Biology 1 Evolution and Pesticide Resistance: Examining Quantitative Trends Visually Stanton Braude and John Gaskin 3 2 Lizard Ecomorphology: Generating and Testing Hypotheses of Adaptation Kenneth H. Kozak 12 3 Phylogenetic Inference: Examining Morphological and Molecular Datasets James Beck 22 4 Life History Tradeoffs in Avian Clutch Size: Interpreting Life History Data and Evaluating Alternative Hypotheses Jon Hess 36 5 Mimicry: Experimental Design and Scientific Logic James Robertson 51 Section II Demography and Population Ecology 6 Life Table Analysis Stanton Braude 63 7 Lotka-Volterra Competition Modeling Stanton Braude, Tara Scherer, and Rebecca McGaha 69 8 Explosive Population Growth and Invasive Exotic Species Jon Hess and James Robertson 79 9 Island Biogeography: Evaluating Correlational Data and Testing Alternative Hypotheses James Robertson 91 Section III Population Genetics 10 Hardy-Weinberg: Evaluating Disequilibrium Forces Jason J. Kolbe 107 11 Drift, Demographic Stochasticity, and Extinction in Woggles James Robertson, Anton Weisstein, and Stanton Braude 117 12 Conservation of Small Populations: Effective Population Sizes, Inbreeding, and the 50/500 Rule Luke J. Harmon and Stanton Braude 125 vi • CONTENTS 13 Dispersal and Metapopulation Structure James Robertson 139 Section IV Quantitative Ecological Tools 14 Understanding Descriptive Statistics Beth Sparks-Jackson and Emily Silverman 155 15 Understanding Statistical Inference Emily Silverman and Beth Sparks-Jackson 179 16 Sampling Wild Populations Stanton Braude and James Robertson 189 17 Quantifying Biodiversity Cawas Behram Engineer and Stanton Braude 198 18 Environmental Predictability and Life History Bobbi S.
Low and Stanton Braude 214 19 Modeling Optimal Foraging Stanton Braude and James Robertson 226 Section V Synthetic Exercises and Writing Assignments 20 Evaluating Competing Hypotheses of Regional Biodiversity Stanton Braude 235 21 Preparing and Evaluating Competitive Grant Proposals for Conservation Funding Stanton Braude 239 22 Tracing the History of Scientific Ideas: From Darwin, Connell, or Soule to the Present Bobbi S. Low 245 Glossary 251 Contributors 263 Index 265 Figures Figure 1. Growth in number of pesticide-resistant species with widespread use of insecticides and herbicides (Modified from National Research Council Report, 1986). Different methods of clinging to branches.
Evolutionary tree depicting the evolution of toepads in anoline lizards. Running speed of species of Anolis on dowels of different diameters (Irshick and Losos, 1999). Anolis habitat generalists tend to be less sensitive to branch diameter than specialists. Phylogenetic tree constructed using the data in table 3.
Polytomies result when no synapomorphies (shared derived traits) can be found for a part of the tree. Choice of the tree that is most likely to represent true evolutionary relationships. (a) Side view of a typical mammal skull, with major tooth types noted. (b) This mammal jaw exhibits a diastema, a prominent gap between teeth.
Eastern cottontail rabbit (Sylvilagus floridanus). 350 bases of DNA sequence for the enzyme RuBisCO in six species of carnivorous plants. Yellow-Shafted Flicker Colaptes auratus auratus. Black-Billed Magpie Pica pica.
Wood Duck Aix sponsa. Partridge Perdix perdix. Great Tit Parus major. Relationship between brood size and nestling size.
An insect of the order Phasmatoidea. Monarch butterflies. Red-spotted newt Notophthalmus viridescens and red-backed salamander Plethodon cinereus. Example of Verhulst-Pearl population growth.
Isoclines for species 1 and species 2 illustrating the population sizes under which each species will increase or decrease. 72 viii • LIST OF FIGURES Figure 7. Four different isoclines illustrate how carrying capacity and competition result in extinction of one or the other species, unstable equilibrium, or stable equilibrium and coexistence. aurelia (left) is much smaller than P.
Zebra mussel Dreissena polymorpha. Percent live and dead unionids collected at one site in offshore waters of western Lake Erie 1969–1991 (from Schloesser et al. Difference between cage and control area biomass (cage density – control density) of zebra mussels plotted against biomass found in duck gizzards (from Hamilton et al. Zebra mussel biomass per unit area (including mussels of all size classes) for cage and control treatments (from Hamilton et al.
Zebra mussel number per unit area for cage and control treatments, September 1991– May 1992 (from Hamilton et al. Species area curve: Amphibians and reptiles of the West Indies. Long-horned beetle. Species-area curve: Cerambycid beetle species of the Florida Keys (from Brown and Peck 1996).
Species-area curve: Land birds of the West Indies (from Terborgh 1973). Map of the Caribbean. Relationship between species number and both the rate of colonization and the rate of extinction for a particular island (adapted from MacArthur and Wilson, 1967). Both the rate of colonization and the rate of extinction vary for islands of different size and distance from the mainland or source population (adapted from MacArthur and Wilson, 1967).
Collared lizards Crotaphytus collaris. Maps of the Missouri Ozark and New Mexico populations of collared lizards Crotaphytus collaris. Two hypothetical populations illustrate subdivision and free exchange of migrants. The mythical woggle Treborus treborsonii.
European adder Vipera berus. The black rhinoceros has been in severe decline in recent years and is entering a population bottleneck (source data: Emslie and Brooks, 1999). The southern white rhinoceros has recovered from near extinction at the turn of the last century and the current growing population is descended from a bottleneck population of only 20 animals (source data: Emslie and Brooks, 1999). Distribution of the fictional California desert woggle Treborus treborsonii.
This simulation examines the effect of migration on loss of alleles and extinction of demes. Bar chart representing invasive species by taxon. Top five reasons for introduction of invasive species. Skewed and symmetric distributions illustrating the relative location of the mean (dashed line) and median (solid line).
Elements of a histogram. Histograms illustrate the frequency distribution of a sample. 163 LIST OF FIGURES • ix Figure 14. Examples of symmetric and skewed histograms and boxplots.
Variable X graphed in three ways illustrates how the shape of a histogram may change with the number of bins. Elements of a boxplot. Boxplots also illustrate the distribution of a sample. They highlight skew and very large and small values.
Variable X presented in a histogram and a boxplot. Examples of commonly observed distributions. Comparison of these two scatterplots shows that correlation does not accurately describe nonlinear relationships. Various scatterplots and associated r.
Illustration of least squares regression. Example scatterplots of y versus x and associated least squares regression lines and R2 values. Quayle’s old data on clutch size. Breeding tenure of Western Gulls on Alcatraz Island.
Percentage of fish in parents and offspring diets. Kenya is located in the eastern horn of Africa and Namibia is located in southwest Africa. Mouse-tailed bat Rhinopoma kardwickei. Springbok Antidorcas marsupialis.
African elephant Loxodonta africana. No matter what level of biodiversity you are concerned with, it is affected by biodiversity at the other levels. Missouri’s geologic history has resulted in a rich variety of habitat types. The Galapagos Archipelago lies 960 km west of the South American mainland.
Little trick-or-treaters like this will be the subject of your foraging model. This figure represents the point at which a trick-or-treater who is optimizing happiness units is predicted to switch neighborhoods. 230 This page intentionally left blank Tables Table 1. Farm pesticide use in the United States 1964–1990.
Types of pesticide, volume of use in the United States, and major crop uses. Tobacco budworm (Heliothis virescens) resistance to methyl parathion. Average number of two phytophagous (plant-eating) mites (Tetranychus mcdanieli [T. mc] and Panonychus ulmi [P.
ul]) and one predatory mite (Metaseiulus occidentalis [M.]) in an apple orchard. Body mass and clinging ability for fifteen species of anoline lizards. Morphology, perch diameter, and sprint performance of the six Anolis ecomorphs. Morphological and ecological measurements for seventeen species of Anolis.
Comparisons of sprint sensitivities for eight species of Anolis over the range of dowel diameters. Trait information for four plant species and their designated outgroup. Life history comparison of five avian species. Partridge and Wood Duck reproductive data.
Survival of young Great Tits (1945–1955). Reproductive output in the Magpie. Nestling weight and survival in the Great Tit. Predation on salamanders.
Percent survival of salamanders. Blank life table. Life table with age-specific survivorship. Life table with age-specific survivorship and fecundity.
Life table with age-specific survivorship, fecundity, and fertility. Killer mouse life table. Projected population growth for the released killer mice. Age distribution across generations.
Population sizes of P. caudatum when cultured separately. Population sizes of P. caudatum when cultured together.
Population growth data for P. bursaria cultured together. North American populations of the zebra mussel, Dreissena polymorpha. Species of Unionid mollusks infested by zebra mussels in the Great Lakes and connecting rivers.
Herpetological biodiversity in the West Indies. Worksheet for calculating coefficient of determination for West Indies amphibian and reptile species-area curve. Long-horned beetle species numbers in the Florida Keys. Comparison of MacArthur and Wilson’s model across taxa.
Channel Island bird species tallies. 102 xii • LIST OF TABLES Table 10. Assumptions, processes, and conservation issues relevant to H-W theory. Collared lizards in the New Mexico region.
Collared lizards in the Ozark region. Pairwise comparisons of FST and geographic distances for seven subpopulations from New Mexico and six subpopulations from the Missouri Ozarks. Sample simulation results. Simulation results for the small Mojave population.
Simulation results for the large Bakersfield population. Time until extinction for woggle populations of various sizes. Sceloporus population at Tyson from 1996 to 2000. Total number of adult adders at Smygehuk.
Adult male and female adders in each year. Northern white rhinos by country, 1960–1998. Population census and effective sizes of African rhinos. Sample calculation of RR statistic.
Worksheet for deme 1 simulation. Worksheet for deme 2 simulation. Worksheet for deme 3 simulation. Calculation of RR statistic.
RR values for metapopulations with different migration rates. Breeding time for young, middle-aged, and old Western Gulls. “Raw” stomach content data for 20 Western Gulls. Types of error.
Common statistical tests and their uses. Gull age and nest initiation. Random number table. Illustrative diversity data for two regions.
Recalculated diversity comparison. Costa Rica biodiversity. Estimated numbers of plant species in various taxonomic groups and their status in Missouri. Estimated numbers of animal species in various taxonomic groups and their status in Missouri.
Eastern Mongolian fauna. Herps of the Galapagos. Mammals of the Galapagos. Birds of the Galapagos.1 Occurrences of rainfall in different seasons over a period of nine years in (a) an Indonesian rainforest, (b) southern Ontario, and (c) central Australia.
Construction of Colwell’s predictability, constancy, and contingency matrix. Datura state matrices for six California locations. 220 LIST OF TABLES • xiii Table 18. Worksheet for calculating components of Datura predictability.
Further demographic data for six California woggle populations. Predictability (P), constancy (C), and contingency (M) of rainfall for 11 deserts around the world. Notes on CRUNCH grants reviewed. 241 This page intentionally left blank Preface Many biology courses are offered with laboratory sections that teach the techniques specific to that discipline as well as the broader tools of how we do science.
While this text cannot replace the hands-on experience of an ecology lab, it does introduce many of the theoretical and quantitative tools of ecology, conservation biology, and environmental sci- ence, and often shows how they intersect.