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Metabolic Ecology Cover image Photograph of a Laysan Albatross (Phoebastria immutabilis) by D. Costa. Drawings and photographs of organisms courtesy of R. Beckett, S. R. Jennings and J. H. Nichols. COMPANION WEBSITE This book is accompanied by a companion website: www.wiley.com/go/sibly/metabolicecology With figures and tables from the book for downloading, together with updates and additional resources M e ta b o l i c E c o l o g y A Scaling Approach Edited by Richard M. Sibly, James H. Brown, and Astrid Kodric-Brown A John Wiley & Sons, Ltd., Publication This edition fi rst published 2012 © 2012 by John Wiley & Sons, Ltd. Wiley-Blackwell is an imprint of John Wiley & Sons, formed by the merger of Wiley’s global Scientifi c, Technical and Medical business with Blackwell Publishing. Registered offi ce: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offi ces: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 111 River Street, Hoboken, NJ 07030-5774, USA For details of our global editorial offi ces, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/wiley-blackwell. The right of the author to be identifi ed as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. 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, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data applied for. Hardback: 9780470671535; paperback: 9780470671528. A catalogue record for this book is available from the British Library. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic books. Set in 9/11 pt PhotinaMT by Toppan Best-set Premedia Limited Contents Notes on contributors, vii 7 Population and community ecology, 77 NICK J.B. ISAAC, CHRIS CARBONE, Preface, xiv AND BRIAN MCGILL 8 Predator–prey relations and food webs, 86 Introduction: Metabolism as the basis for a theoretical OWEN L. PETCHEY AND JENNIFER A. DUNNE unifi cation of ecology, 1 JAMES H. BROWN, RICHARD M. SIBLY, 9 Ecosystems, 99 AND ASTRID KODRIC-BROWN KRISTINA J. ANDERSON-TEIXEIRA AND PETER M. VITOUSEK PART I FOUNDATIONS, 7 10 Rates of metabolism and evolution, 112 JOHN L. GITTLEMAN AND 1 Methodological tools, 9 PATRICK R. STEPHENS ETHAN P. WHITE, XIAO XIAO, NICK J. B. ISAAC, AND RICHARD M. SIBLY 11 Biodiversity and its energetic and thermal controls, 120 2 The metabolic theory of ecology and its central DAVID STORCH equation, 21 JAMES H. BROWN AND RICHARD M. SIBLY 3 Stoichiometry, 34 PART II SELECTED ORGANISMS AND MICHAEL KASPARI TOPICS, 133 4 Modeling metazoan growth and ontogeny, 48 12 Microorganisms, 135 ANDREW J. KERKHOFF JORDAN G. OKIE 5 Life history, 57 13 Phytoplankton, 154 RICHARD M. SIBLY ELENA LITCHMAN 6 Behavior, 67 14 Land plants: new theoretical directions and APRIL HAYWARD, JAMES F. GILLOOLY, empirical prospects, 164 AND ASTRID KODRIC-BROWN BRIAN J. ENQUIST AND LISA PATRICK BENTLEY v vi Contents 15 Marine invertebrates, 188 22 Conservation biology, 271 MARY I. O’CONNOR AND JOHN F. BRUNO ALISON G. BOYER AND WALTER JETZ 16 Insect metabolic rates, 198 23 Climate change, 280 JAMES S. WATERS AND JON F. HARRISON KRISTINA J. ANDERSON-TEIXEIRA, FELISA A. SMITH, AND S. K. MORGAN ERNEST 17 Terrestrial vertebrates, 212 WILLIAM KARASOV 24 Beyond biology, 293 MELANIE E. MOSES AND STEPHANIE FORREST 18 Seabirds and marine mammals, 225 DANIEL P. COSTA AND SCOTT A. SHAFFER 25 Synthesis and prospect, 302 JAMES H. BROWN, RICHARD M. SIBLY, 19 Parasites, 234 AND ASTRID KODRIC-BROWN RYAN F. HECHINGER, KEVIN D. LAFFERTY, AND ARMAND M. KURIS Glossary, 306 20 Human ecology, 248 MARCUS J. HAMILTON, OSKAR BURGER, References, 309 AND ROBERT S. WALKER Index, 361 PART III PRACTICAL APPLICATIONS, 259 21 Marine ecology and fi sheries, 261 SIMON JENNINGS, KEN H. ANDERSEN, AND JULIA L. BLANCHARD COMPANION WEBSITE This book is accompanied by a companion website: www.wiley.com/go/sibly/metabolicecology With figures and tables from the book for downloading, together with updates and additional resources Notes on contributors Ken H. Andersen is a pro fessor Lisa Patrick Bentley is in theoretical marine ecology currently an NSF post- at the National Institute of doctoral fellow at the Aquatic Resources at the University of Arizona. Technical University of She works on extending Denmark. He studies how the metabolic scaling theory marine ecosystem responds to to account for additional perturbations, in particular aspects of plant hydraulics, xylem function, and fi shing, using size - spectrum carbon fl ux. Her aim is to integrate key plant physio- models and metabolic argu- logical processes that affect ecosystem - level dynamics. ments. ttp://ken.haste.dk Her approach addresses an increasing need for predic- tive models that scale from the leaf to globe in order to inform future research and government policy Kristina J. Anderson - Teixeira decisions. received her PhD in Biology under James H. Brown and Julia Blanchard is a Lecturer also studied under Marcy E. in the Department of Animal Litvak at the University of and Plant Sciences at the New Mexico. She is currently University of Sheffi eld and an working as a postdoctoral Honorary Research Fellow of research associate with Evan Imperial College London. Julia H. DeLucia at the University of teaches and researches both Illinois. Her research focuses on understanding how the fundamental and applied climate shapes terrestrial ecosystems, quantifying the ecology of marine populations, communities, and climate regulation services of terrestrial ecosystems, ecosystems. Her current research involves linking mac ro- and applying knowledge of ecosystem – climate interac- ecology, food webs and fi sheries ecology to inform tions to inform land - use decisions in an era of climate marine ecosystem management. www.sheffi eld.ac.uk/ change. aps/staff-and-students/acadstaff/blanchard-julia vii viii Notes on contributors Alison Boyer is a Research Chris Carbone works at the Assistant Professor in the Institute of Zoology, Zoological Department of Ecology asd Society of London, Regent ’ s Evolutionary Biology at the Park, and is interested in University of Tennessee. She understanding drivers of uses ecological informatics and species distributions and the fossil record to examine abundance. His work focuses community ecology and extinction risk in island birds. particularly on the role of She is also engaged in research at broader scales to body size and consumer – resource relationships in examine processes governing biological diversity. shaping organism ecology and population processes, http://eeb.bio.utk.edu/boyer/index.html and he is developing a fondness for dinosaurs. James H. Brown is Distin- Daniel Costa is a Distinguished guished Professor of Biology Professor of Ecology and Evo- at the University of New lutionary Biology at the Uni- Mexico, Albuquerque. He led versity of California at Santa the development of the Meta- Cruz. He research focuses on bolic Theory of Ecology on the ecology and physiology of which this book is largely marine mammals and sea- based. He has a long history of birds. He has worked in almost research in biogeography and every habitat from the Galapa- macroecology, taking a large - scale statistical approach gos to the Antarctic with a broad range of animals to questions about abundance, distribution, and diver- including penguins, albatross, seals, sea lions, whales, sity. http://biology.unm.edu/jhbrown/index.shtml and dolphins. http://bio.research.ucsc.edu/people/ costa/ John Bruno is a marine ecologist in the Depart- Jennifer Dunne is a Professor ment of Biology at the at the Santa Fe Institute University of North Caro- ( www.santafe.edu ) and Co - lina at Chapel Hill. His Director of the Pacifi c Ecoin- research is focused on formatics and Computational marine biodiversity, coral Ecology Lab ( www.foodwebs. reef ecology and conservation and the impacts of org ). Her research seeks to climate change on marine ecosystems. John earned his identify fundamental patterns PhD in Ecology and Evolutionary Biology from Brown and principles of ecological University and was a postdoctoral fellow in disease network structure, robustness, ecology at Cornell University. www.brunolab.net and dynamics at multiple spatial and temporal scales. Oskar Burger is a Postdoctoral Fellow at the Max Planck Insti- tute for Demographic Research. He has many interests in both the social and natural sciences centered around understanding large - scale evolutionary con- straints on human/primate life history. His latest projects include the evolution of the post - reproductive lifespan, cross- species primate mortality patterns, life - history invariants across the tree of life, and the effects of variation in energy consumption on human demographics. Notes on contributors ix B rian Enquist is a Professor in Jamie Gillooly is an Assistant the Department of Ecology Professor of Biology at the and Evolutionary Biology at University of Florida. Jamie the University of Arizona played a primary role in devel- ( www.salvias.net/∼ brian ). He oping the Metabolic Theory uses physiological, theoretical, of Ecology, and is currently computational, and informat- working to extend these ics approaches in order to dis- energy - based models in new cover general principles that directions. Current projects shape the: (i) origin of biological scaling laws; (ii) include the study of animal communication, animal scaling of plant functional traits to ecology and evolu- migration, and aging and disease. In addition, Jamie tion; and (iii) origin and maintenance of functional has been working to foster collaboration between and phylogenetic diversity. He addresses these ques- artists and scientists as a “ scholar - in - residence ” in the tions in tropical forests and alpine ecosystems. School of Fine Arts and Art History at the University of Florida. S. K. Morgan Ernest is an Associate Professor in the John Gittleman is Dean and Department of Biology at Utah Professor of Ecology in the State University. She teaches Odum School of Ecology at the courses on Animal Community University of Georgia where Ecology, Macroecology, and he teaches Macroecology and non - majors courses in Biology. Conservation Ecology. His While she has been involved in research interests are large - research spanning from life - scale ecological and evolution- history theory to paleoecology, ary problems, specifi cally she is particularly interested in related to speciation, extinc- how energetic constraints impact the structure and tion, and global biodiversity. dynamics of communities. http://ernestlab.weecology. http://blackbear.ecology.uga.edu/gittleman/ org Marcus Hamilton is a Postdoc- Stephanie Forrest is Professor toral Fellow at the Santa Fe of Computer Science, Univer- Institute and an Adjunct Pro- sity of New Mexico, and Exter- fessor of Anthropology at the nal Professor of the Santa Fe University of New Mexico. His institute. Her research studies research focuses on general adaptive systems, including im- principles that drive the struc- munology, evolutionary com- ture, dynamics, and diversity putation, biological modeling, of human societies in the past, present, and future, and computer security. http:// integrating perspectives from across the social, biologi- www.cs.unm.edu/∼ forrest/ cal, and physical sciences. http://www.unm.edu/∼ marcusj/home.htm x Notes on contributors J on F. Harrison is a Professor Simon Jennings is a Principal in the School of Life Sciences Scientist at the Centre for at Arizona State University Environment, Fisheries and where he teaches a variety of Aquaculture Science, courses in biology and physiol- Lowestoft, and Professor of ogy. His research focuses on Environmental Science at the environmental and ecological University of East Anglia. His physiology of insects. http:// research focuses on assessing jharrison.faculty.asu.edu the sustainability of human and environmental impacts on marine populations, com- munities, and ecosystems as well as developing and April Hayward is a Postdoctoral applying tools to support marine environmental Associate at the University of management. Florida whose research ulti- mately focuses on understand- Walter Jetz is Associate Profes- ing how complex biological sor in the Ecology and Evolu- systems emerged from a prebi- tionary Biology Department otic soup. Current efforts at Yale University. Using toward this end center on mostly terrestrial vertebrates understanding the fl ow and and plants as study systems, retention of matter and energy his inter disciplinary research through different levels of biological organization. draws on elements of biogeog- raphy, community ecology, landscape ecology, macr- Ryan Hechinger is a research oecology, global change ecology, evolution, comparative scientist at the University of biology, biodiversity informatics, and conservation, California, Santa Barbara. He aiming to integrate across scales of geography and eco- has three overarching and logical organization – from local to global assemblages. related research goals. One is http://www.yale.edu/jetz/ to evaluate the importance of parasites in ecosystems. William Karasov is Professor in Another is to use parasites to test the Department of Forest and and refi ne general ecological Wildlife Ecology at University of and evolutionary theory. The Wisconsin - Madison, where he third is to “ keep it real.” http:// teaches Animal Physiological www.lifesci.ucsb.edu/∼ hechinge/ Ecology. He researches digestive physiology, nutritional ecology, Nick Isaac works at the animal energetics, and ecotoxi- Natural Environment Research cology of vertebrates. http:// Council’ s Centre for Ecology forestandwildlifeecology.wisc. and Hydrology in Wallingford, edu/facstaff/karasov.html Oxfordshire, on questions about species’ distribution and abun- dance over large scales, using data on mammals, insects, and birds. Of particular interest is the relative contribution of intrinsic biological traits and extrinsic environmental drivers in shaping biodi- versity, and how these patterns change at different spatial, temporal, and taxonomic scales.

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