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Rodent Societies Rodent Societies An Ecological & Evolutionary Perspective Edited by Jerry O Wolff and Paul W Sherman The University of Chicago Press Chicago and London paul w sherman is professor of[.]

Rodent Societies Rodent Societies: An Ecological & Evolutionary Perspective Edited by Jerry O Wolff and Paul W Sherman The University of Chicago Press Chicago and London paul w sherman is professor of animal behavior and a Weiss Presidential Fellow at Cornell University jerry o wolff was professor and chair of the biology department at the University of Memphis; he is now professor of biology at St Cloud State University The University of Chicago Press, Chicago 60637 The University of Chicago Press, Ltd., London © 2007 by The University of Chicago All rights reserved Published 2007 Printed in the United States of America 16 15 14 13 12 11 10 09 08 07 12345 isbn-10: 0-226-90536-5 (cloth) isbn-13: 978-0-226-90536-5 (cloth) isbn-10: 0-226-90537-3 (paper) isbn-13: 978-0-226-90537-2 (paper) Library of Congress Cataloging-in-Publication Data Rodent societies : an ecological & evolutionary perspective / edited by Jerry O Wolff and Paul W Sherman p cm Includes bibliographical references and index isbn-13: 978-0-226-90536-5 (cloth : alk paper) isbn-10: 0-226-90536-5 (cloth : alk paper) isbn-13: 978-0-226-90537-2 (pbk : alk paper) isbn-10: 0-226-90537-3 (pbk : alk paper) Rodents—Ecology Rodents—Evolution Social behavior in animals Animal societies I Wolff, Jerry II Sherman, Paul W., 1949 – ql737.r6r623 2007 599.35156 — dc22 2006021884  The paper used in this publication meets the minimum requirements of 䊊 the American National Standard for Information Sciences—Permanence of Paper for Printed Library Materials, ansi z39.48-1992 Contents Contributors ix Acknowledgments xiii Introduction Rodent Societies as Model Systems Jerry O Wolff and Paul W Sherman Rodent Evolution, Phylogenetics, and Biogeography Rodney L Honeycutt, Laurence J Frabotta, and Diane L Rowe Sexual Behavior Male Mating Strategies in Rodents Jane Waterman 27 Reproductive Strategies in Female Rodents Nancy G Solomon and Brian Keane 42 Sexual Selection: Using Social Ecology to Determine Fitness Differences Lara S Carroll and Wayne K Potts 57 A Phylogenetic Analysis of the Breeding Systems of Neotomine-Peromyscine Rodents Matina C Kalcounis-Rüppell and David O Ribble 68 Alternative Reproductive Tactics and Strategies of Tree Squirrels John L Koprowski 86 Life Histories and Behavior Fast and Slow Life Histories of Rodents F Stephen Dobson and Madan K Oli Acceleration and Delay of Reproduction in Rodents Lee C Drickamer 10 Sexual Size Dimorphism in Rodents Albrecht I Schulte-Hostedde 11 Facultative Sex Ratio Adjustment Robert S Sikes 129 12 The Role of the Stress Axis in Life-History Adaptations Rudy Boonstra, J M Barker, J Castillo, and Q E Fletcher 13 Dispersal and Philopatry Scott Nunes 115 139 150 14 Gene Dynamics and Social Behavior F Stephen Dobson 163 15 Social Behavior and Self-Regulation in Murid Rodents Charles J Krebs, Xavier Lambin and Jerry O Wolff 173 Behavioral Development 16 Neural Regulation of Social Behavior in Rodents J Thomas Curtis, Yan Liu, Brandon J Aragona, and Zuoxin Wang 17 Ontogeny of Adaptive Behaviors Jill M Mateo 18 Social Learning by Rodents Bennett G Galef Jr 195 207 185 106 99 vi Contents 19 Kin Recognition in Rodents: Issues and Evidence Warren G Holmes and Jill M Mateo 216 Social Behavior 20 Parental Care Betty McGuire and William E Bemis 231 21 The Ecology of Sociality in Rodents Eileen A Lacey and Paul W Sherman 22 Scent Marking S Craig Roberts 243 255 23 Nonparental Infanticide Luis A Ebensperger and Daniel T Blumstein 24 Social Organization and Monogamy in the Beaver Peter Busher 267 280 25 Evolution of Pacifism and Sociality in Blind Mole-Rats Eviatar Nevo 291 Antipredator Behavior 26 Social and Antipredator Systems: Intertwining Links in Multiple Time Frames Donald H Owings and Richard G Coss 305 27 The Evolution of Alarm Communication in Rodents: Structure, Function, and the Puzzle of Apparently Altruistic Calling Daniel T Blumstein 317 28 Fear and the Foraging, Breeding, and Sociality of Rodents Hannu Ylönen and Joel S Brown 328 Comparative Socioecology 29 Ecology, Kinship, and Ground Squirrel Sociality: Insights from Comparative Analyses James F Hare and Jan O Murie 345 30 Evolution of Sociality in Marmots: It Begins with Hibernation Kenneth B Armitage 356 31 Environmental Constraints and the Evolution of Sociality in Semifossorial Desert Rodents Jan A Randall 368 32 Comparative Social Organization and Life History of Rattus and Mus Manuel Berdoy and Lee C Drickamer 380 33 Social Organization and Resource Use in Capybaras and Maras David W Macdonald, Emilio A Herrera, Andrew B Taber, and José Roberto Moreira 393 34 Social Structure in Octodontid and Ctenomyid Rodents Eileen A Lacey and Luis A Ebensperger 403 35 Socioecology of Rock-Dwelling Rodents Karen J Nutt 416 36 African Mole-Rats: Social and Ecological Diversity Chris G Faulkes and Nigel C Bennett 427 37 Alarm Calling, Multiple Mating, and Infanticide among Black-Tailed, Gunnison’s, and Utah Prairie Dogs John L Hoogland 438 Conservation and Disease 38 Issues in Rodent Conservation William Z Lidicker Jr 453 39 Conservation of Ground Squirrels Beatrice Van Horne 463 40 Conservation of Prairie Dogs John L Hoogland 472 Contents vii 41 Social Behavior, Demography, and Rodent-Borne Pathogens Richard S Ostfeld and James N Mills 478 Conclusions 42 Conclusions and Future Directions Paul W Sherman and Jerry O Wolff References 491 Subject Index 599 Species Index 605 489 Contributors Brandon J Aragona Department of Psychology Florida State University Tallahassee, FL 32306 aragona@psy.fsu.edu Kenneth B Armitage Department of Ecology and Evolutionary Biology University of Kansas Lawrence, KS 66045 marmots@lark.cc.ukans.edu J M Barker Centre for the Neurobiology of Stress Department of Life Sciences University of Toronto at Scarborough Scarborough, Ontario, Canada, M1C 1A4 Rudy Boonstra Centre for the Neurobiology of Stress Department of Life Sciences University of Toronto at Scarborough Scarborough, Ontario, Canada, M1C 1A4 boonstra@utsc.utoronto.ca Joel Brown Department of Biological Sciences University of Illinois at Chicago Chicago, IL 60607 squirrel@uic.edu Peter E Busher Center for Ecology and Conservation Biology Boston University Boston, MA 02215 USA pbusher@bu.edu William E Bemis Department of Ecology and Evolutionary Biology Corson Hall Cornell University Ithaca, NY 14853 web24@cornell.edu Lara S Carroll Howard Hughes Medical Institute University of Utah Salt Lake City, UT 84112-5331 lara.carroll@utah.edu Nigel C Bennett Department of Zoology and Entomology University of Pretoria Pretoria, South Africa ncbennett@zoology.up.ac.za J Castillo Centre for the Neurobiology of Stress Department of Life Sciences University of Toronto at Scarborough Scarborough, Ontario, Canada, M1C 1A4 Manuel Berdoy Oxford University Veterinary Services Parks Rd, OX1 3PT Oxford, UK manuel.berdoy@vet.ox.ac.uk Richard C Coss Department of Psychology University of California One Shields Avenue Davis, CA 95616-8686 rgcoss@ucdavis.edu Daniel T Blumstein Department of Ecology and Evolutionary Biology University of California 621 Charles E Young Drive South University of California Los Angeles, CA 90095-1606 marmots@ucla.edu J Thomas Curtis, PhD Department of Psychology Florida State University Tallahassee, FL 32306 tcurtis@psy.fsu.edu x Contributors F Stephen Dobson Department of Biological Sciences Auburn University Auburn, AL 36849 fdobson@acesag.auburn.edu Emilio A Herrera Departamento de Estudios Ambientales Universidad Simón Bolívar Caracas 1080-A, Venezuela eherre@usb.ve Lee C Drickamer Department of Biological Sciences Northern Arizona University Flagstaff, AZ 86001 Lee.Drickamer@NAU.EDU Warren Holmes Department of Psychology and Center for Ecology and Evolutionary Biology University of Oregon Eugene, OR 97403-5289 wholmes@darkwing.uoregon.edu Luis A Ebensperger Departamento de Ecología P Universidad Católica de Chile Centro de Estudios Avanzados en Ecología & Biodiversidad Casilla 114-D, Santiago, Chile lebenspe@genes.bio.puc.cl Christopher G Faulkes Queen Mary & Westfield College School of Biological Sciences London E1 4NS UK C.G.Faulkes@qmul.ac.uk Quinn E Fletcher Centre for the Neurobiology of Stress Department of Life Sciences University of Toronto at Scarborough Scarborough, Ontario, Canada, M1C 1A4 Rodney L Honeycutt Texas A&M University Department of Wildlife & Fisheries Sciences College Station, Texas 77843-2258 rhoneycutt@neo.tamu.edu John L Hoogland The University of Maryland Appalachian Laboratory Frostburg, MD 21532 hoogland@al.umces.edu Matina Kalcounis-Rüppell Department of Biology The University of North Carolina at Greensboro Greensboro, North Carolina 27402-6170 matina_kalcounis@uncg.edu Laurence J Frabotta Department of Biology Texas A&M University College Station, Texas 77843-3258 Brian Keane Department of Zoology Miami University Hamilton, OH 45011 keaneb@muohio.edu Bennett G Galef Department of Psychology McMaster University Hamilton, Ontario L8S 4K1 Canada galef@mcmail.mcmaster.ca John L Koprowski Wildlife and Fisheries Science School of Renewable Natural Resources University of Arizona Tucson, AZ 85721 squirrel@ag.arizona.edu James F Hare Department of Zoology University of Manitoba Manitoba, Canada R3T 2N2 harejf@cc.umanitoba.ca Dr Charles J Krebs CSIRO Sustainable Ecosystems Canberra, A.C.T 2601 Australia Charles.Krebs@csiro.au Contributors xi Eileen Lacey Museum of Vertebrate Zoology University of California Berkeley, CA 94720 ealacey@socrates.berkeley.edu Xavier Lambin School of Biological Sciences Department of Biology University of Aberdeen Aberdeen, Scotland x.lambin@abdn.ac.uk William Z Lidicker, Jr Museum of Vertebrate Zoology University of California Berkeley, CA 94720 lidicker@socrates.Berkeley.edu Yan Liu Department of Psychology Florida State University Tallahassee, FL 32306 David W Macdonald Wildlife Conservation Research Unit Department of Zoology University of Oxford Tubney, Oxon OX13 5QL, UK david.macdonald@zoology.ox.ac.uk Jill M Mateo Department of Comparative Human Development Committee on Human Development Institute for Mind and Biology University of Chicago Chicago, IL 60637 jmateo@uchicago.edu Betty McGuire, PhD Department of Ecology and Evolutionary Biology Cornell University Ithaca NY 14853 bam65@cornell.edu James N Mills Special Pathogens Branch Centers for Disease Control and Prevention (MS G-14) Atlanta, GA, 30333 jum0@cdc.gov José Roberto Moreira Embrapa Recursos Genéticos e Biotecnologia Parque Estaỗóo Biolúgica 70770-900 Brasớlia DF Brazil jmoreira@cenargen.embrapa.br Jan O Murie Department of Biological Sciences University of Alberta Edmonton, AB T6G 2E9 Alberta, Canada jan.murie@ualberta.ca Eviatar Nevo Department of Environmental and Evolutionary Biology University of Haifa Mount Carmel, Haifa 31905 Israel nevo@research.haifa.ac.il Scott Nunes Department of Biology University of San Francisco San Francisco, CA 94117 nunes@ace.usfca.edu Karen J Nutt School of Biological Sciences University of Auckland Auckland, New Zealand knut006@ec.auckland.ac.nz Madan K Oli Department of Wildlife Ecology and Conservation University of Florida Gainesville, FL 32611 OliM@wec.ufl.edu Richard S Ostfeld Institute of Ecosystem Studies Millbrook, NY 12545 ROstfeld@ecostudies.org Donald H Owings Department of Psychology University of California One Shields Avenue Davis, CA 95616-8686 dhowings@ucdavis.edu xii Contributors Wayne K Potts Molecular Biology Program University of Utah Salt Lake City, UT 84112 potts@biology.utah.edu Nancy G Solomon Department of Zoology Miami University Oxford, Ohio 45056 solomong@muohio.edu Jan Randall Department of Biology University of San Francisco San Francisco, CA 94117 jrandall@sfsu.edu Andrew Taber Wildlife Conservation Society 2300 Southern Boulevard Bronx, NY 10460 ataber@wcs.org David O Ribble Biology Department Trinity University San Antonio, TX 78212 dribble@trinity.edu Beatrice Van Horne 1829 Elgin Dr Vienna, VA 22182 bvanhorne@fs.fed.us S Craig Roberts School of Biological Sciences University of Liverpool Liverpool UK L69 7ZB craig.roberts@liv.ac.uk Diane L Rowe Texas A&M University Department of Wildlife & Fisheries Sciences College Station, Texas 77843-2258 Albrecht I Schulte-Hostedde Department of Biology Laurentian University Sudbury, Ontario, Canada P3E 2C6 aschultehostedde@laurentian.ca Paul W Sherman Department of Neurobiology and Behavior Cornell University Ithaca, NY 14853 pws6@cornell.edu Robert S Sikes Biology Department University of Arkansas, Little Rock Little Rock, AR 72204 rssikes@ualr.edu Zuoxin Wang Department of Psychology Department of Biological Sciences Neuroscience Program Florida State University Tallahassee, FL 32306 zwang@darwin.psy.fsu.edu Jane M Waterman Department of Biology University of Central Florida Orlando, FL 32816-2368 waterman@mail.ucf.edu Jerry O Wolff Department of Biological Sciences St Cloud State University St Cloud, MN 56301 jowolff@stcloudstate.edu Hannu Ylönen Department of Biological and Environmental Science University of Jyväskyla FIN 40351 Jyväskyla, Finland hylonen@dodo.jyu.fi Acknowledgments Rodney L Honeycutt, Laurence J Frabotta, and Diane L Rowe: Research reported on hystricognath rodents was funded by National Science Foundation Grant DEB 9615163 to RLH Nancy G Solomon and Brian Keane: We thank the editors for the invitation to participate in this book Kristen Lucia, Gail Michener, Paul Sherman, and Jerry Wolff provided valuable feedback on a previous version of this manuscript We also are grateful to Michelle Edwards, Stephanie Kortering, Samantha Lowe, Lisa Walter, and Beth Widen for assistance with references NGS and BK were supported by NSF DEB-0316818 and NGS was supported by R15 6M069409-01 during the preparation of this book chapter Lara S Carroll and Wayne K Potts: We would like to thank the editors for substantive comments on an earlier version of this manuscript This manuscript was written while WKP was supported by NIH (GM39578) and NSF (IBN-9904609) grants Matina C Kalcounis-Rüppell and David O Ribble: We would like to acknowledge the assistance of Stacy Huff, Michelle Icenhower, and Adrian Sherman in compiling literature for our review and analysis Robert Bradley helped us to understand Neotomine-Peromyscine systematics Comments of Jack Millar, Maarten Vonhof, Eileen Lacey, and two anonymous reviewers improved earlier versions of this manuscript We thank the editors for their insight, suggestions, and editorial guidance F Stephen Dobson and Madan K Oli: We owe special thanks to the editors for suggesting that we review rodent life histories D R Broussard, P H Harvey, T J Karels, and S C Stearns provided excellent comments and suggestions for improvement of the manuscript FSD’s contribution to the chapter was supported by a National Science Foundation grant for research (DEB-0089473) Lee C Drickamer: I thank the many undergraduates, particularly at Williams College, graduate students and undergraduates at Southern Illinois University, and faculty colleagues at those two schools and at Northern Arizona University for their invaluable assistance, discussions, and enthusiasm Portions of the work in my laboratory and in field settings were supported by grants from the National Institutes of Health and National Science Foundation I thank the editors for their foresight 10 11 12 13 14 15 16 in organizing this volume and for their diligence in putting it all together Albrecht I Schulte-Hostedde: I thank the editors for the opportunity to contribute this chapter and for improving it with their insightful comments Thanks to all who responded to my queries, including T Best, S Boutin, J Hoogland, I Khokhlova, H Levenson, and R Sweitzer Climate data from weather stations were provided by J Pither A Oey provided valuable assistance Robert S Sikes: I am grateful for constructive comments on a previous version of this manuscript by T G Finley, S Krackow, P W Sherman, and J O Wolff This work was supported in part by NSF Grant 9975445 R Boonstra, J M Barker, J Castillo, and Q E Fletcher: The Natural Sciences and Engineering Research Council of Canada supported this research We thank Jim Kenagy for helpful comments on an earlier draft of this chapter Scott Nunes: I thank Jerry Wolff and Steve Dobson for constructive, incisive, and helpful comments on earlier versions of this chapter F Stephen Dobson: I owe special thanks to the editors for suggesting that I review the genetic properties of social breeding groups Ron Chesser provided patient instruction on the gene dynamics of social breeding groups during the summers of 1993–1996, and the Savannah River Ecology Laboratory provided support via Visiting Faculty Fellowships in 1993, 1994, and 1996 In 1995, I was an Oak Ridge Institute for Science and Education Research Fellow at SREL I especially appreciate the support of the director of SREL, M H Smith, during these years R K Chesser, N Perrin, and M F Winterrowd provided excellent comments and suggestions for improvement of the manuscript I also owe special thanks to John Hoogland, who generously made his incredible field data so freely available, and to Dave Foltz, for encouraging analyses on his and John’s allozyme data Preparation of the current manuscript was completed while I was supported by a National Science Foundation grant for research (DEB-0089473) Charles J Krebs, Xavier Lambin, and Jerry O Wolff: We thank Peter Brown, Grant Singleton, and Alice Kenney for their comments and suggestions on the manuscript J Thomas Curtis, Yan Liu, Brandon J Aragona, and Zuoxin Wang: We are grateful to Christie Fowler and xiv Acknowledgments 18 21 22 23 25 Mike Smeltzer for critical reading of this manuscript This work was partially supported by NIH grants HD48462 and HD40722 (JTC), MH67396 (BJA), MH58616 and MH67396 (ZXW) Bennett G Galef Jr.: I thank Paul Sherman for his thoughtful comments on earlier drafts of the manuscript The author’s research described here was supported for 35 years by the Natural Science and Engineering Research Council of Canada Eileen A Lacey and Paul W Sherman: The ideas presented in this paper reflect 25 years of collaborative interactions between the authors regarding sociality in rodents For insights and encouragement along the way we thank R D Alexander, J U M Jarvis, M L Morton, C K Sherman, J S Sherman, J R Wieczorek, J O Wolff, E Yensen, the Animal Behavior Lunch Bunch at Cornell University, and the many graduate and undergraduate students who have assisted with our field and laboratory studies of subterranean rodents Our work with social rodents has been supported by the National Science Foundation, the National Geographic Society, Sigma Xi, the Center for Latin American Studies at the University of California, Berkeley, and the College of Agriculture and Life Sciences at Cornell University S Craig Roberts: I thank Morris Gosling, Paul Sherman, Jerry Wolff, and Sarah Zala for their comments on the manuscript Luis A Ebensperger and Daniel T Blumstein: We thank the editors for inviting us to write this chapter, and for comprehensive suggestions, which improved the original manuscript considerably Comments by Bob Elwood and Stefano Parmigiani also are greatly appreciated During the writing of this article, LAE was supported by the Centro de Estudios Avanzados en Ecología & Biodiversidad (FONDAP 1501-001) and by a FONDECYT grant No 1020861 Eviatar Nevo: This chapter is based on the research program of speciation and adaptive radiation in Israeli subterranean mole-rats that started in the early 1950s The subset of aggression studies was initiated in 1975 The Spalax ehrenbergi superspecies research program incorporates the extensive Israeli and worldwide collaborations of active students, colleagues, and collaborators of subterranean mammals The Spalax research program has been documented since 1961 in some 280 multidisciplinary scientific publications Much of the evidence in this chapter stems from the aforementioned collaborations We acknowledge all participants’ contributions with much appreciation I wish to extend my deepest gratitude to my colleagues at the Institute of Evolution, whose generous and devoted help made this chapter possible: Mr Michael Margulis and Mrs Robin Permut 26 27 28 31 32 I also appreciate the continuous financial support of the University of Haifa, Israeli Discount Bank Chair of Evolutionary Biology, the Ancell-Teicher Research Foundation for Genetics and Molecular Evolution, the Israeli Ministries of Science and Absorption, and grants from the Israel Science Foundation, BSF, the Israel Academy of Sciences, and Guggenheim Foundation Donald H Owings and Richard G Coss: We are indebted to Eric Charles and the editors for their careful reviews of drafts of this manuscript Daniel T Blumstein: I thank Alexander Nikolskii for introducing me to the Russian literature on alarm communication and Luis Ebensberger for an introduction to South American rodents; neither is responsible for anticipated omissions I thank Janice Daniel, Kim Pollard, and my reviewers—Jan Randall and Ron Swaisgood— for constructive comments on previous versions, and Erin Shelly, who found reports of genera of alarm calling rodents that I originally missed I am extremely grateful to Paul Sherman, who went out of his way to improve the clarity of this chapter, and to remind me of the importance of the “limits of nepotism”—a hypothesis that is likely to explain interspecific variation in the adaptive utility of alarm communication and other nepotistic behaviors Hannu Ylönen and Joel S Brown: We thank the editors for inviting us to join the team of Rodent Societies It was a great honor For our experiences and journeys into the fascinating world of desert rodents, voles, and the ecology of fear, we owe much to the help, instruction, and good ideas of a number of colleagues and friends, too many to mention by name, but denizens of Fennoscandia, North America, the Middle East, and Australia We are grateful to them all Jan A Randall: I am grateful to my Russian collaborators on the gerbil project, especially Kostya Rogovin, who kindly provided references from the Russian literature and constructed table 31.1 I thank all the students who participated in my research on kangaroo rats and gerbils over the years, and Bruce MacEvoy for his support and editing skills He, Debra Shier, Kostya Rogovin, and Jerry Wolff provided helpful comments on the manuscript I thank the National Science Foundation, National Institute of Health, National Geographic Society, and the Research and Development Foundation for grant support This chapter is dedicated to John Eisenberg, who conducted the early studies on the behavior of desert rodents and encouraged me to study them Manuel Berdoy and Lee C Drickamer: The authors would like to thank J Galef, A Hanson, P Honess, G Singleton, A Voight, J Webster, and S Wolfensohn for helpful comments on this chapter Acknowledgments xv 33 David W Macdonald, Emilio A Herrera, Andrew B Taber, and José Roberto Moreira: We are grateful to Ruth Feber for considerable help in the preparation of this chapter, and to our many colleagues in the WildCRU who have commented on and encouraged our research 34 Eileen A Lacey and Luis A Ebensperger: We thank the editors for inviting us to write this chapter Figures 34.1– 34.4 were compiled by Karen Klitz During the preparation of this chapter, LAE was supported by the Centro de Estudios Avanzados en Ecología and Biodiversidad (FONDAP 1501-001) The authors’ ongoing studies of degus, cururos, and tuco-tucos are supported by funding from the National Science Foundation (DEB-0128857; EAL), the National Geographic Society (EAL and LAE), and by FONDECYT grant No 1020861 (LAE) 35 Karen J Nutt: I would like to thank the Marshall Commission and Roger Nutt for providing financial support for me while writing this chapter Thanks also to Paula White, Eileen Lacey, and the members of both LARG and Molecular Ecology at Cambridge University for their continued moral support I am also grateful to the editors for their kindness, patience, and constructive comments on this chapter Thanks also to the gundis for making the study of rock-dwelling rodents a truly fascinating and worthwhile experience 36 Chris G Faulkes and Nigel C Bennett: We are most grateful for financial support from the following: National Research Foundation (NCB and JUMJ), The University of Pretoria and the Mellon Foundation (NCB), The University of Cape Town and the National Geographic Society (JUMJ), and the Natural Environment Research Council (CGF) Thanks to Jenny Jarvis, Steve Le Comber, and Pippa Faulkes for proofreading and many helpful comments, and to the editors for further suggestions that greatly improved the manuscript 39 Beatrice Van Horne: NatureServe compiled and formatted a ground squirrel custom dataset that was useful in developing the cross-species summarizations Last, Jerry Wolff and Paul Sherman sincerely thank all of the authors for contributing their professional expertise to Rodent Societies and for their diligence, patience, and good humor in working with us to complete this anthology We also thank Janet Sherman for assistance with the index and Christie Henry, Monica Holliday, and Jennifer Howard of the University of Chicago Press and Susan Dodson of Graphic Composition for their support, encouragement, and editorial expertise in bringing this book to fruition Introduction Chapter Rodent Societies as Model Systems Jerry O Wolff and Paul W Sherman he Rodentia is the largest order of mammals, consisting of more than 2,000 species and comprising 44% of all mammals Rodents come in a variety of body shapes, from cylindrical to spherical, and sizes, from less than 10 g to more than 66 kg The characteristic that unites this order—and is its most conspicuous trait— is a single pair of razor-sharp incisors, which are used to gnaw food, excavate tunnels, and defend themselves The name “rodent” derives from the Latin word rodere, which means “to gnaw.” No matter where you live, a rodent is probably not far away Rodents inhabit all continents except Antarctica, and they occur in terrestrial, subterranean, arboreal, and aquatic habitats—from the high arctic tundra to equatorial rain forests, temperate bogs and swamps to hot, arid deserts, and rocky mountaintops to sandy canyon bottoms Many species live in close association with humans With such varied characteristics and expansive ecology, rodents provide a range of attributes that have captivated scientists and annoyed laypersons for hundreds of years The diversity of rodents, and the ease with which many species can be maintained in captivity, has led to their choice as model systems for observational and experimental studies in genetics, ecology, demography, physiology, and psychology The social and reproductive behaviors of rodents also are diverse and intriguing Although most rodents are nocturnal, a surprising amount of research has been conducted on rodent social biology As a result, large databases are available, and these can be used to test hypotheses about the ecological and evolutionary forces that mold mammalian social and reproductive behaviors Many T aspects of behavioral ecology are similar across species (e.g., the effects of resource distributions on mating systems [Slobodchikoff 1984; Ostfeld 1990]; the role of ecological factors in favoring group-living [Hoogland 1995; Ebensperger and Cofré 2001]; and the role of kinship in structuring social interactions [Sherman 1981a; Lacey and Wieczorek 2003]) Unifying theories developed from other taxa, such as primates, ungulates, or canids, can be experimentally tested in the field and laboratory with rodents These options are not so readily available for the larger, widerranging taxa Thus rodents are not only models for testing hypotheses developed from rodents, but they have become models for other taxa as well The relevant information, however, is widely scattered and sometimes conflicting Hence this volume Our goals were to synthesize and integrate the current state of knowledge about the social behavior of rodents, to provide ecological and evolutionary contexts for understanding rodent societies, and to highlight emerging conservation and management issues to preserve these societies Thus we attempted to emulate Primate Societies (1987) and Cetacean Societies (2000), the two outstanding preceding volumes in this series published by the University of Chicago Press In selecting topics and contributors for the present volume, we first chose areas of behavioral biology and model species that we considered essential for understanding the adaptive significance of rodent social behavior generally Then we invited contributions from researchers who have demonstrated their preeminence in illuminating these areas and in studying the focal taxa We urged authors to use the ... Sherman 22 Scent Marking S Craig Roberts 24 3 25 5 23 Nonparental Infanticide Luis A Ebensperger and Daniel T Blumstein 24 Social Organization and Monogamy in the Beaver Peter Busher 26 7 28 0 25 Evolution... 0 -22 6-90537-3 (pbk : alk paper) Rodents—Ecology Rodents—Evolution Social behavior in animals Animal societies I Wolff, Jerry II Sherman, Paul W., 1949 – ql737.r6r 623 20 07 599.35156 — dc 22 2006 021 884... 978-0 -22 6-90536-5 (cloth) isbn-10: 0 -22 6-90537-3 (paper) isbn-13: 978-0 -22 6-90537 -2 (paper) Library of Congress Cataloging-in-Publication Data Rodent societies : an ecological & evolutionary perspective

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