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Lower metazoans and lesser deuterosomes neil schlager

Grzimek’s
Animal Life Encyclopedia

Second Edition

● ● ● ●

Volume 16
Mammals V

Grzimek’s Animal Life Encyclopedia, Second Edition
Volume 16: Mammals V

Synapse, the Knowledge Link Corporation

Tracey Rowens, Jennifer Wahi

Margaret Chamberlain

Wendy Blurton, Dorothy Maki, Evi Seoud, Mary

Randy Bassett, Mary K. Grimes, Lezlie Light,

Christine O’Bryan, Barbara Yarrow, Robyn V.
Young

© 2004 by Gale. Gale is an imprint of The

not guarantee the accuracy of the data con-

Learning Inc.

http://www.gale-edit.com/permissions, or you
may download our Permissions Request form

cation of any organization, agency, institution,

are trademarks used herein under license.

ply endorsement of the editors and publisher.

The Gale Group, Inc.

MI, 48331-3535, Permissions hotline: 248-699- Errors brought to the attention of the pub-
lisher and verified to the satisfaction of the

publisher will be corrected in future editions.

Or you can visit our Internet site at

ISBN
cardinus avellanarius) by Stephen Dalton/Photo

ALL RIGHTS RESERVED

0-7876-5788-3 (vol. 12)

anemone by AP/Wide World Photos/University

No part of this work covered by the copy-

0-7876-5791-3 (vol. 15)

golden frog, and green python by JLM Visuals;
red-legged locust © 2001 Susan Sam; hornbill
by Margaret F. Kinnaird; and tiger by Jeff Lepore/

tronic, or mechanical, including photocopy-

Photo Researchers. All reproduced by permission.

ing, recording, taping, Web distribution, or

out the written permission of the publisher.

the reliability of the information presented

v. cm.

Includes bibliographical references.

QL7 .G7813 2004
590’.3—dc21 2002003351

Printed in Canada
10 9 8 7 6 5 4 3 2 1

Order MONOTREMATA
Monotremes ....................................................................... 227 Family: Echidnas......................................................... 235 Family: Duck-billed platypus ..................................... 243

Order DIDELPHIMORPHIA
New World opossums
Family: New World opossums................................... 249

Volume 13: Mammals II
Order PERAMELEMORPHIA
Bandicoots and bilbies ....................................................... 1 Family: Bandicoots...................................................... 9 Subfamily: Bilbies........................................................ 19

Order NOTORYCTEMORPHIA
Marsupial moles
Family: Marsupial moles............................................. 25

Order XENARTHRA
Sloths, anteaters, and armadillos....................................... 147 Family: West Indian sloths and two-toed tree
sloths............................................................................ 155 Family: Three-toed tree sloths................................... 161 Family: Anteaters ........................................................ 171 Family: Armadillos...................................................... 181

Order INSECTIVORA
Insectivores......................................................................... 193 Family: Gymnures and hedgehogs............................. 203 Family: Golden moles................................................. 215 Family: Tenrecs .......................................................... 225 Family: Solenodons..................................................... 237 Family: Extinct West Indian shrews.......................... 243 Family: Shrews
I: Red-toothed shrews............................................. 247 II: White-toothed shrews......................................... 265 Family: Moles, shrew moles, and desmans................ 279

Pteropus................................................................. 319 All other genera................................................... 333

Family: Mouse-tailed bats .......................................... 351 Family: Sac-winged bats, sheath-tailed bats,
and ghost bats ............................................................. 355 Family: Kitti’s hog-nosed bats ................................... 367 Family: Slit-faced bats ................................................ 371 Family: False vampire bats ......................................... 379 Family: Horseshoe bats .............................................. 387 Family: Old World leaf-nosed bats ........................... 401 Family: American leaf-nosed bats .............................. 413 Family: Moustached bats............................................ 435 Family: Bulldog bats ................................................... 443 Family: New Zealand short-tailed bats...................... 453 Family: Funnel-eared bats.......................................... 459 Family: Smoky bats..................................................... 467 Family: Disk-winged bats........................................... 473 Family: Old World sucker-footed bats...................... 479 Family: Free-tailed bats and mastiff bats................... 483 Family: Vespertilionid bats
I: Vespertilioninae................................................... 497

Glossary.............................................................................. 541 Mammals species list ......................................................... 546 Geologic time scale............................................................ 580 Index ................................................................................... 581

For further reading............................................................ 527 Order CETACEA

Organizations ..................................................................... 532 Whales, dolphins, and porpoises....................................... 1

The ungulates .................................................................... 131 Ungulate domestication..................................................... 145

Order TUBULIDENTATA
Aardvarks
Family: Aardvarks........................................................ 155

Order ARTIODACTYLA
Even-toed ungulates .......................................................... 263 Family: Pigs................................................................. 275 Family: Peccaries......................................................... 291 Family: Hippopotamuses............................................ 301 Family: Camels, guanacos, llamas, alpacas, and
vicuñas ......................................................................... 313 Family: Chevrotains.................................................... 325 Family: Deer
Subfamily: Musk deer ............................................ 335 Subfamily: Muntjacs............................................... 343 Subfamily: Old World deer................................... 357 Subfamily: Chinese water deer.............................. 373 Subfamily: New World deer ................................. 379 Family: Okapis and giraffes........................................ 399 Family: Pronghorn...................................................... 411

For further reading............................................................ 419 Organizations ..................................................................... 424 Contributors to the first edition ....................................... 426 Glossary.............................................................................. 433 Mammals species list ......................................................... 438

Grzimek’s Animal Life Encyclopedia vii

Contents

• • • • •
Foreword

Earth is teeming with life. No one knows exactly how many distinct organisms inhabit our planet, but more than 5 mil-lion different species of animals and plants could exist, rang-ing from microscopic algae and bacteria to gigantic elephants, redwood trees and blue whales. Yet, throughout this won-derful tapestry of living creatures, there runs a single thread: Deoxyribonucleic acid or DNA. The existence of DNA, an elegant, twisted organic molecule that is the building block of all life, is perhaps the best evidence that all living organ-isms on this planet share a common ancestry. Our ancient connection to the living world may drive our curiosity, and perhaps also explain our seemingly insatiable desire for in-formation about animals and nature. Noted zoologist, E. O. Wilson, recently coined the term “biophilia” to describe this phenomenon. The term is derived from the Greek bios mean-ing “life” and philos meaning “love.” Wilson argues that we are human because of our innate affinity to and interest in the other organisms with which we share our planet. They are, as he says, “the matrix in which the human mind originated and is permanently rooted.” To put it simply and metaphor-ically, our love for nature flows in our blood and is deeply en-grained in both our psyche and cultural traditions.

Grzimek’s Animal Life Encyclopedia ix

Foreword

While it is clear that modern conservation must take the needs of both wildlife and people into consideration, what will the quality of human life be if the collective impact of short-term economic decisions is allowed to drive wildlife popula-tions into irreversible extinction? Many rural populations living in areas of high biodiversity are dependent on wild an-imals as their major source of protein. In addition, wildlife tourism is the primary source of foreign currency in many de-veloping countries and is critical to their financial and social stability. When this source of protein and income is gone, what will become of the local people? The loss of species is not only a conservation disaster; it also has the potential to be a human tragedy of immense proportions. Protected ar-eas, such as national parks, and regulated hunting in areas out-side of parks are the only solutions. What critics do not realize is that the fate of wildlife and people in developing countries is closely intertwined. Forests and savannas emptied of wildlife will result in hungry, desperate people, and will, in the long-term lead to extreme poverty and social instability. Dr. Grz-imek’s early contributions to conservation should be recognized, not only as benefiting wildlife, but as benefiting local people as well.

Dr. Grzimek’s hope in publishing his Animal Life Encyclo-pedia was that it would “...disseminate knowledge of the ani-mals and love for them”, so that future generations would“...have an opportunity to live together with the great diver-sity of these magnificent creatures.” As stated above, our goals in producing this updated and revised edition are similar. However, our challenges in producing this encyclopedia were more formidable. The volume of knowledge to be summa-rized is certainly much greater in the twenty-first century than it was in the 1970’s and 80’s. Scientists, both professional and amateur, have learned and published a great deal about the animal kingdom in the past three decades, and our under-standing of biological and ecological theory has also pro-gressed. Perhaps our greatest hurdle in producing this revision was to include the new information, while at the same time retaining some of the characteristics that have made Grzimek’s Animal Life Encyclopedia so popular. We have therefore strived to retain the series’ narrative style, while giving the informa-tion more organizational structure. Unlike the original Grz-imek’s, this updated version organizes information under specific topic areas, such as reproduction, behavior, ecology and so forth. In addition, the basic organizational structure is generally consistent from one volume to the next, regardless of the animal groups covered. This should make it easier for users to locate information more quickly and efficiently. Like the original Grzimek’s, we have done our best to avoid any overly technical language that would make the work difficult to understand by non-biologists. When certain technical ex-pressions were necessary, we have included explanations or clarifications.

The overall structure of this reference work is based on the classification of animals into naturally related groups, a discipline known as taxonomy or biosystematics. Taxonomy is the science through which various organisms are discov-ered, identified, described, named, classified and catalogued. It should be noted that in preparing this volume we adopted what might be termed a conservative approach, relying pri-marily on traditional animal classification schemes. Taxon-omy has always been a volatile field, with frequent arguments over the naming of or evolutionary relationships between var-ious organisms. The advent of DNA fingerprinting and other advanced biochemical techniques has revolutionized the field and, not unexpectedly, has produced both advances and con-fusion. In producing these volumes, we have consulted with specialists to obtain the most up-to-date information possi-ble, but knowing that new findings may result in changes at any time. When scientific controversy over the classification of a particular animal or group of animals existed, we did our best to point this out in the text.

Readers should note that it was impossible to include as much detail on some animal groups as was provided on oth-ers. For example, the marine and freshwater fish, with vast

• • • • •
How to use this book

Gzimek’s Animal Life Encyclopedia is an internationally prominent scientific reference compilation, first published in German in the late 1960s, under the editorship of zoologist Bernhard Grzimek (1909-1987). In a cooperative effort be-tween Gale and the American Zoo and Aquarium Association, the series is being completely revised and updated for the first time in over 30 years. Gale is expanding the series from 13 to 17 volumes, commissioning new color images, and updat-ing the information while also making the set easier to use.

Organization of chapters within each volume reinforces the taxonomic hierarchy. In the case of the Mammals vol-umes, introductory chapters describe general characteristics of all organisms in these groups, followed by taxonomic chap-ters dedicated to Order, Family, or Subfamily. Species ac-counts appear at the end of the Family and Subfamily chapters To help the reader grasp the scientific arrangement, each type of chapter has a distinctive color and symbol:

● =Order Chapter (blue background)

Thumbnail introduction
Common name
Scientific name
Class
Order
Suborder
Family
Thumbnail description
Size
Number of genera, species
Habitat
Conservation status
Main essay
Evolution and systematics
Physical characteristics
Distribution
Habitat
Behavior
Feeding ecology and diet
Reproductive biology
Conservation status
Significance to humans
Species accounts
Common name
Scientific name
Subfamily
Taxonomy
Other common names
Physical characteristics
Distribution
Habitat
Behavior

xii Grzimek’s Animal Life Encyclopedia

All the color illustrations in Grzimek’s were created specif-ically for the project by Michigan Science Art. Expert con-tributors recommended the species to be illustrated and provided feedback to the artists, who supplemented this in-formation with authoritative references and animal skins from University of Michgan Zoological Museum library. In addi-tion to species illustrations, Grzimek’s features conceptual drawings that illustrate characteristic traits and behaviors.

About the contributors
The essays were written by scientists, professors, and other professionals. Grzimek’s subject advisors reviewed the com-pleted essays to insure consistency and accuracy.

Series advisor

Michael Hutchins, PhD
Director of Conservation and Science/William Conway Chair
American Zoo and Aquarium Association
Silver Spring, Maryland

Sean F. Craig, PhD
Assistant Professor, Department of Biological Sciences Humboldt State University

New York Aquarium
Brooklyn, New York

James B. Murphy, DSc
Smithsonian Research Associate
Department of Herpetology
National Zoological Park
Washington, DC

Arcata, California Volumes 8–11: Birds I–IV

Arthur V. Evans, DSc New York, New York

Research Associate, Department of Entomology Smithsonian Institution
Washington, DC

Volumes 4–5: Fishes I– II Paul V. Loiselle, PhD
Curator, Freshwater Fishes

Professor Emeritus of Environmental Science University of Calgary
Calgary, Alberta
Canada

Library advisors

Librarian
Oklahoma City Zoo
Oklahoma City, Oklahoma

Pittsburgh, Pennsylvania Richard Nagler

Linda L. Coates
Associate Director of Libraries
Zoological Society of San Diego Library San Diego, California

• • • • •
Contributing writers

Mammals I–V

Cynthia Berger, MS

Clarence L. Abercrombie, PhD
Wofford College

Jasmin Chua, MS

Spartanburg, South Carolina Durrell Institute of Conservation and

Freelance Writer

Cleber J. R. Alho, PhD
Departamento de Ecologia (retired)

Canterbury, Kent

Universidade de Brasília Far North Queensland Wildlife Res-

United Kingdom

Brasília, Brazil
Carlos Altuna, Lic
Sección Etología

Guillermo D’Elía, PhD

Facultad de Ciencias

Smithsonian Institution

Universidad de la República Oriental
del Uruguay

Justin S. Brashares, PhD

Montevideo, Uruguay

Montevideo, Uruguay

Anders Angerbjörn, PhD
Department of Zoology

University of Michigan Museum of

Zoology

Stockholm University
Stockholm, Sweden
William Arthur Atkins
Department of General Zoology Fac-
Atkins Research and Consulting

Andean Tapir Fund

ulty of Bio- and Geosciences

Normal, Illinois
Adrian A. Barnett, PhD
Centre for Research in Evolutionary

Susan Cachel, PhD

Department of Ecology and Evolution

Anthropology
School of Life Sciences
University of Surrey Roehampton

Stony Brook, New York

New Brunswick, New Jersey

West Will, London
United Kingdom
Leonid Baskin, PhD

University of Cambridge

National Museum Prague

Institute of Ecology and Evolution
Moscow, Russia
Paul J. J. Bates, PhD

Department of Biology

Harrison Institute

Institute of Vertebrate Biology

Sevenoaks, Kent
United Kingdom

Canada

Amy-Jane Beer, PhD

David J. Chivers, MA, PhD, ScD

Origin Natural Science
York, United Kingdom

South Windsor, Connecticut

xvi Grzimek’s Animal Life Encyclopedia

Contributing writers

Theodore H. Fleming, PhD
Department of Biology
University of Miami

Simon Fraser University Burnaby

Ecology

Coral Gables, Florida
Gabriel Francescoli, PhD

Australia

Sección Etología

Robin L. Hayes

Facultad de Ciencias
Universidad de la República Oriental
del Uruguay

ology

Montevideo, Uruguay

School of Earth and Environmental

Udo Gansloßer, PhD
Department of Zoology

Mike J. R. Jordan, PhD

Adelaide, Australia

Lehrstuhl I
University of Erlangen-Nürnberg
Fürth, Germany Department of Ethology and Ecology
Valerius Geist, PhD

German Primate Center

Upton, Chester

Professor Emeritus of Environmental
Science
University of Calgary

Science Journalist

Science Writer

Calgary, Alberta
Canada
Roger Gentry, PhD

Department of Biological Sciences

tensphysiologie

NOAA Fisheries
Marine Mammal Division
Silver Spring, Maryland

Serge Larivière, PhD

Kenneth C. Gold, PhD

Freelance Writer

Chicago, Illinois
Steve Goodman, PhD

Rodney L. Honeycutt, PhD

Adrian Lister

Field Museum of Natural History
Chicago, Illinois and Departments of Wildlife and Fisheries
WWF Madagascar
Programme Office

Genetics

Texas A&M University

Antananarivo, Madagascar
Nicole L. Gottdenker

Valdosta, Georgia

St. Louis Zoo

Geoff Lundie-Jenkins, PhD

University of Missouri
St. Louis, Missouri and The Charles
Darwin Research Station

Praha, Czech Republic

Galápagos Islands, Ecuador

Peter W. W. Lurz, PhD

Brian W. Grafton, PhD Centre for Life Sciences Modelling
Department of Biological Sciences
Kent State University

University of Newcastle

Canada

Kent, Ohio Newcastle upon Tyne, United King-
Joel H. Grossman
Freelance Writer

Facultad de Ciencias

Santa Monica, California School of Biological Sciences (Zool-
Universidad de la República Oriental

ogy)

Mark S. Hafner, PhD
Lowery Professor and Curator of
Mammals
Museum of Natural Science and De-

Jennifer U. M. Jarvis, PhD

James Malcolm, PhD

partment of Biological Sciences
Louisiana State University
Baton Rouge, Louisiana

Rondebosch, South Africa

Redlands, California

Grzimek’s Animal Life Encyclopedia xvii
David P. Mallon, PhD
Glossop

A.N. Severtsov Institute of Ecology

Derbyshire, United Kingdom

Birds

Robert D. Martin, BA (Hons), DPhil,

Randolph W. Rose, PhD

DSc
Provost and Vice President

Carsten Niemitz, PhD

Academic Affairs
The Field Museum

Hobart, Tasmania

Chicago, Illinois

Anthropology

Gary F. McCracken, PhD
Department of Ecology and Evolu-

Telemark University College

tionary Biology

Daniel K. Odell, PhD

University of Tennessee
Knoxville, Tennessee

Science and Environmental Writer

Colleen M. McDonough, PhD

Orlando, Florida

Department of Biology
Valdosta State University
University of Montana (adjunct retired
Valdosta, Georgia

professor)

William J. McShea, PhD

Georgia Southern University

Department of Conservation Biology
Conservation and Research Center John H. Seebeck, BSc, MSc, FAMS
Research Professor in African Ecology
Smithsonian National Zoological Park
School of Animal, Plant and Environ-
Washington, DC

mental Sciences

Melody Serena, PhD

Rodrigo A. Medellín, PhD
Instituto de Ecología
Universidad Nacional Autónoma de

Malcolm Pearch, PhD

Whittlesea, Australia

México
Mexico City, Mexico
Leslie Ann Mertz, PhD

United Kingdom

University of British Columbia

Fish Lake Biological Program
Wayne State University
Detroit, Michigan

Hiram, Ohio

Robert W. Shumaker, PhD

Gus Mills, PhD
SAN Parks/Head
Carnivore Conservation Group,

tute at George Mason University

EWT

Department of Conservation Biology

Skukuza, South Africa
Patricia D. Moehlman, PhD Smithsonian National Zoological Park

School of Life Sciences

IUCN Equid Specialist Group

Jan A. Randall, PhD

Paula Moreno, MS
Texas A&M University at Galveston

Karen B. Strier, PhD

Marine Mammal Research Program

San Francisco, California

Galveston, Texas
Virginia L. Naples, PhD
Department of Biological Sciences

Okapi Wildlife Associates

Hudson, Quebec

Northern Illinois University
DeKalb, Illinois
Ken B. Naugher, BS

Lehman College of The City Univer-

sity of New York

Conservation and Enrichment Pro-
grams Manager
Montgomery Zoo

Bettina Tassino, MSc

Montgomery, Alabama

Adelaide, Australia

xviii
Facultad de Ciencias

Randall S. Wells, PhD

Universidad de la República Oriental

PhD

del Uruguay
Montevideo, Uruguay

Sarasota, Florida

University of Pretoria

Barry Taylor, PhD
University of Natal
Pietermaritzburg, South Africa

Santa Barbara, California

Mammal Research Institute

Jeanette Thomas, PhD
Department of Biological Sciences
Western Illinois University-Quad

Christian C. Voigt, PhD

Brook

Cities
Moline, Illinois Leibniz-Institute for Zoo and Wildlife
Ann Toon

Berlin, Germany

Department of Anthropology and

Arnside, Cumbria
United Kingdom

Long Beach, California

Stephen B. Toon

Santa Rosa, California

Arnside, Cumbria
United Kingdom

Dalhousie University

Academy of Sciences of the Czech

Hernán Torres, PhD
Santiago, Chile
Grzimek’s Animal Life Encyclopedia xix

• • • • •
Contributing illustrators

Drawings by Michigan Science Art

Barbara Duperron, BFA, MFA Amanda Smith, BFA, MFA

Jarrod Erdody, BA, MFA Christina St.Clair, BFA

Thanks are due to the University of Michigan, Museum of Zoology, which provided specimens that served as mod-els for the images.

Paul Exner, Chief cartographer Laura Exner

Grzimek’s Animal Life Encyclopedia

XNR Productions, Madison, WI Andy Grosvold
Tanya Buckingham Cory Johnson
Jon Daugherity
Paula Robbins
xx

Antelopes, cattle, bison, buffaloes, goats, and sheep (Bovidae)