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HARVARD UNIVERSITY 
e 
Library of the 


Museum of 


Comparative Zoology 


t 


tent 


4 


SEVENTY-FIVE YEARS OF 
MAMMALOGY 
(1919-1994) 


ELMER C. BIRNEY 
AND 
JERRY R. CHOATE 
Editors 


SPECIAL PUBLICATIONS 


This series, published by the American Society of Mammalogists, has been established 
for papers of monographic scope concerned with some aspect of the biology of mam- 
mals. 

Correspondence concerning manuscripts to be submitted for publication in the series 
should be addressed to the Editor for Special Publications, Michael A. Mares (address 
below). 

Copies of Special Publications of the Society may be ordered from the Secretary- 
Treasurer, H. Duane Smith, 501 Widtsoe Bldg., Dept. Zoology, Brigham Young Uni- 
versity, Provo, UT 84602. 


COMMITTEE ON SPECIAL PUBLICATIONS 


MIcHAEL A. Mares, Editor 
Oklahoma Museum of Natural History 
University of Oklahoma 
Norman, Oklahoma 73019 


JosepH F. Merritt, Managing Editor 
Powdermill Biological Station 
Carnegie Museum of Natural History 
Rector, Pennsylvania 15677 


ll 


SEVENTY-FIVE YEARS OF 
MAMMALOGY 
(1919-1994) 


EDITED By 


ELMER C. BIRNEY 
Bell Museum of Natural History 
100 Ecology Building 
University of Minnesota 
St. Paul, Minnesota 55108 


JERRY R. CHOATE 
Sternberg Museum of Natural History 
Fort Hays State University 
Hays, Kansas 67601 


SPECIAL PUBLICATION NO. 11 
THE AMERICAN SOCIETY OF MAMMALOGISTS 
PUBLISHED 27 May 1994 


il 


MCZ 
LIBRARY 
OCT 19 1994 


HARVARD 
UNIVERSITY 


Library of Congress Catalog Card No. 97-71464 
© 1994 
ISBN No. 0-935868-73-9 


lv 


LIST OF AUTHORS 


Sydney Anderson 
American Museum of Natural History 
Central Park West at 79th Street 
New York, NY 10024 


David M. Armstrong 
Department of Evolutionary, Population, 
and Organismic Biology 
University of Colorado 
Boulder, CO 80309 


Robert J. Baker 
Department of Biological Sciences and the 
Museum 
Texas Tech University 
Lubbock, TX 79409 


Elmer C. Birney 
Bell Museum of Natural History 
100 Ecology Building 
University of Minnesota 
St. Paul, MN 55108 


James H. Brown 
Department of Biology 
University of New Mexico 
Albuquerque, NM 87131 


Guy N. Cameron 
Department of Biology 
University of Houston 
Houston, TX 77004 


Jerry R. Choate 
Sternberg Museum of Natural History 
Fort Hays State University 
Hays, KS 67601 


Mark D. Engstrom 
Department of Mammalogy 
Royal Ontario Museum 
100 Queen’s Park 
Toronto, Ontario, CANADA MSS 2C6 


John F. Eisenberg 
Florida Museum of Natural History 
Department of Natural Sciences and 
School of Forest Resources and 
Conservation 
University of Florida, Gainesville, FL 
32611 


Gregory L. Florant 
Department of Biology 
Temple University 
Philadelphia, PA 19122 


Hugh H. Genoways 
University of Nebraska State Museum 
University of Nebraska-Lincoln 
Lincoln, NE 68588 


Ayesha E. Gill 
Institute of Health Policy Studies 
University of California 
1388 Sutter Street, 11th Floor 
San Francisco, CA 94109 
Present Address 
2308 Jefferson Avenue 
Berkeley, CA 94703 


Mark S. Hafner 
Museum of Natural Science and 
Department of Zoology and Physiology 
Louisiana State University 
Baton Rouge, LA 70803 


Robert S. Hoffmann 
Smithsonian Institution 
Washington, D.C. 20560 


Donald F. Hoffmeister 
Museum of Natural History 
University of Illinois 
Urbana, IL 61801 


Rodney L. Honeycutt 
Department of Wildlife and 
Fisheries Sciences and 
The Faculty of Genetics 
210 Nagle Hall 
Texas A&M University 
College Station, TX 77843 


Murray L. Johnson 
501 N. Tacoma Avenue 
Tacoma, WA 98403 


G. J. Kenagy 
Department of Zoology 
University of Washington 
Seattle, WA 98195 


Gordon L. Kirkland, Jr. 
The Vertebrate Museum 
Shippensburg University 
Shippensburg, PA 17257 


James N. Layne 
Archbold Biological Station 
P.O. Box 2057 
Lake Placid, FL 33852 


William Z. Lidicker, Jr. 
Museum of Vertebrate Zoology 
University of California 
Berkeley, CA 94720 


vl 


Jason A. Lillegraven 
Departments of Geology/Geophysics and 
Zoology/Physiology 
The University of Wyoming 
Laramie, WY 82071 


Michael A. Mares 
Oklahoma Museum of Natural History 
University of Oklahoma 
Norman, OK 73019 


Bruce D. Patterson 
Field Museum of Natural History 
Roosevelt Road at Lake Shore Drive 
Chicago, IL 60605 


Oliver P. Pearson 
Museum of Vertebrate Zoology 
University of California 
Berkeley, CA 94720 


Randolph L. Peterson (Deceased) 
Royal Ontario Museum 
100 Queen’s Park 
Toronto, Ontario, CANADA MS5S 2C6 


Carleton J. Phillips 
Department of Biological Sciences 
Illinois State University 
Normal, IL 61790 


Duane A. Schlitter 
Edward O’Neil Research Center 
Carnegie Museum of Natural History 
5800 Baum Blvd 
Pittsburgh, PA 15206 


David J. Schmidly 
Texas A&M University 
P.O. Box 1675 
Galveston, TX 77553 


James H. Shaw 
Department of Zoology 
Oklahoma State University 
Stillwater, OK 74078 


H. Duane Smith 
Department of Zoology 
Brigham Young University 
Provo, UT 84602 


Keir Sterling 
324 Webster Street 
Bel Air, MD 21014 


J. Mary Taylor 
Cleveland Museum of Natural History 
Wade Oval, University Circle 
Cleveland, OH 44106 


B. J. Verts 
Department of Fisheries and Wildlife 
Oregon State University 
Corvallis, OR 97331 


John O. Whitaker, Jr. 
Department of Life Science 
Indiana State University 
Terre Haute, IN 47809 


Don E. Wilson 
National Museum of Natural History 
Smithsonian Institution 
Washington, D.C. 20560 


Vil 


Jerry O. Wolff 
Department of Fisheries and Wildlife 
Oregon State University 
Corvallis, OR 97331 


W. Chris Wozencraft 
Division of Natural Sciences 
Lewis-Clark State College 
Lewiston, ID 83501 


Bruce A. Wunder 
Department of Biology 
Colorado State University 
Fort Collins, CO 80523 


Terry L. Yates 
Department of Biology and 
Museum of Southwestern Biology 
University of New Mexico 
Albuquerque, NM 87131 


Richard J. Zakrzewski 
Department of Geosciences and 
Sternberg Museum of Natural History 
Fort Hays State University 
Hays, KS 67601 


PREFACE 


he ad hoc committee to plan the 75th 
anniversary of the American Society 
of Mammalogists (abbreviated ASM here 
and throughout this book), was established 
by President Hugh Genoways, seemingly 
only a short time after we celebrated our 
50th anniversary. The first meeting of that 
committee that we recall was at the annual 
gathering of ASM in Madison, Wisconsin, 
in 1986, and was chaired by Craig Hood. 
The coeditors of this book volunteered early 
in that meeting to oversee the preparation 
of a book covering the 75 years that ASM 
had been in existence, 1919-1994, admit- 
ting at the time that we had no specific plan 
but that we thought we could get something 
of this nature done in the 8 years avail- 
able. 

We are not the first, nor will we be the 
last, to learn that every job expands to con- 
sume all available time. Certainly, this book 
was no exception. It was remarkably easy 
for authors to agree to participate and for 
editors to develop ‘“‘firm’’ deadlines when 
the target dates were such a long time in the 
future. Today, 10 February 1994, as we are 
drafting this final note to accompany the 21 
chapters in this book, we still lack and shall 
never see one important chapter, and Allen 
Press is bending over backwards to get page 
proofs to the authors. Managing Editor Joe 
Merritt and Special Publications Editor Mi- 
chael Mares are probably the only authors 
and editors who did not have time to pro- 
crastinate! Nevertheless, production pres- 


ipa fe 


ently is on schedule to have this book in the 
hands of ASM members at the anniversary 
meeting in Washington, D.C., as promised 
so long ago. 

As we employ the latest in word-process- 
ing software to draft this document, send 
e-mail messages between Kansas and Min- 
nesota in seconds, fax manuscripts between 
authors and editors, and generally make the 
most of this electronic era and its infor- 
mation superhighways, we contemplate 
1919, the year ASM was founded. The First 
World War had just ended, Woodrow Wil- 
son was President of the United States, and 
Mexico was experiencing a revolution. What 
was the state of the discipline of mammal- 
ogy, and what has ASM done in its 75-year 
existence to promote and facilitate the sci- 
ence? That is the topic of this book, which 
was conceived without much forethought in 
an otherwise forgotten committee meeting, 
and which underwent early embryogenesis 
along the banks of Lake Mendota on the 
beautiful University of Wisconsin campus, 
then survived a lengthy period of delayed 
development following some rapid growth 
that took place while the list of chapters was 
finalized and authors were recruited. A few 
individual cells underwent mitosis now and 
then, but real gestation began in the spring 
and summer of 1992. Subsequently, all 
chapters were subjected to two reviews by 
peers, mostly during the spring of 1993, and 
final revisions of most chapters were com- 
pleted that summer. 


The book is in two parts, one on the so- 
ciety and its members (the first eight chap- 
ters) and the other on the intellectual growth 
and development of the discipline of mam- 
malogy during the past 75 years. The charge 
to authors of the two parts was different. 
Those writing chapters for Part I were asked 
simply to treat the topic, and in all cases the 
emphasis was on ASM, its members, its 
growth, and its activities to promote mam- 
malogy. Thus, those chapters address his- 
tory, and their topics are less about science 
than its facilitation. 

Authors writing chapters for Part II were 
given the following, much more specific, 
guidelines: ““We envision that chapters in 
this section will briefly review the pre-1919 
state of knowledge of the assigned subdis- 
cipline, if appropriate, then trace the intel- 
lectual development of the field through the 
75-year period that ASM will have been in 
existence. Chapters in Part II are expected 
to take a global perspective of the history, 
with no special emphasis on either ASM or 
its members, of the field’s development.” 
We judge that all authors have more than 
adequately fulfilled the charge. 

Authors originally were selected in pairs 
with an eye toward diversity. In some pairs 
our strategy was to select collaborators rep- 
resenting different eras, in others different 
schools of thought, and in still others we 
sought authors whose expertise encom- 
passed the extremes of a broad or complex 
subdiscipline. A few prospective authors re- 
signed for one reason or another, one died, 
one pair decided their chapter was not nec- 
essary, and fora host of other reasons author 
lines changed. We attempted to maintain 
the two-author-per-chapter philosophy 
throughout in order to get the best ideas of 
at least two individuals into every chapter, 
but in three instances that was not possible 
and in one a third author was recruited. 
Historical details of author selection pale in 
comparison to the heartfelt thanks we ex- 
tend to all authors—it was our very real 
pleasure to work with each of them. 

We are equally appreciative of the con- 
siderable effort donated by Jane Waterman, 
who drew the vignettes used on the first 


pages of chapters. We like each one very 
much, Jane. Our thanks go also to a long 
list of reviewers, some of whom dropped 
everything in order to help us meet our 
deadline, then employed fax and e-mail as 
necessary to provide nearly instantaneous 
turn-around of excellent, insightful reviews. 
We greatly appreciate the time and efforts 
of all reviewers, several of whom reviewed 
more than a single chapter: Sydney Ander- 
son, David M. Armstrong, Robert J. Baker, 
Patricia J. Berger, James H. Brown, William 
A. Clemens, Mark D. Engstrom, James S. 
Findley, G. Lawrence Forman, Enk K. 
Fritzell, Hugh H. Genoways, Sarah B. 
George, Donald W. Kaufman, Gordon L. 
Kirkland, Jr., Thomas H. Kunz, Norman 
C. Negus, Bruce D. Patterson, Anne E. Pu- 
sey, O. J. Reichman, Eric A. Rickart, Duke 
S. Rogers, Robert K. Rose, William D. 
Schmid, Robert S. Sikes, Donald B. Siniff, 
Norman A. Slade, H. Duane Smith, Robert 
H. Tamarin, Robert M. Timm, Michael R. 
Voorhies, Jane M. Waterman, Michael R. 
Willig, Don E. Wilson, and Robert M. Zink. 

Finally, we thank three people who made 
our jobs easy, and without whose untiring 
energies at crucial times this book would 
not have been completed in time for the 
anniversary celebration: Joseph F. Merritt, 
Managing Editor for Mammalian Species 
and Special Publications, put a prodigious 
amount of time and energy (with occasional 
lapses into jocularity) into making certain 
that no important detail of production was 
slighted; Michael A. Mares, Editor for Spe- 
cial Publications, processed manuscripts as 
fast as the two of us could send them to 
him; and Ken Blair at Allen Press adopted 
this project and simply made it happen on 
time no matter what the obstacle. 

The proof of the pudding, as always, is in 
the eating. We hope that you, the reader, 
like our idea of pudding. 


ELMER C. BIRNEY 
St. Paul, Minnesota 


JERRY R. CHOATE 
Hays, Kansas 
February 1994 


CONTENTS 


PART I. History of ASM and Its Most Prominent Members 


ORIGIN Donaid F ehojmeisterand Keir B. Stine, tse pec culo. 14 ee De eee ese 
RIO O CHR GUO Mie tk vc ee Ree rhea hee haan coy an aanias versie ita a pee 4 ke eee MRS net A 
The Development of Scientific Societies in Europe in the 18th and 19th Centuries ...... 
North American Mammatogy Before the 20th Century «2.2... d22c2 2 nocecee cies ce sen: 
The Early History of the American Society of Mammalogists ......................... 
EN CLCEUCI OT ASIN CACTI ES pert te asec ai aubecie ana ep i taed Gad tte he an es ara neh ate oak Oa ee catia aad 
tc rare Site dh 0 Aachen eee oe ood eGo Oe ok 8 tae eek tee hg ee a 2 


PRESIDENTS James N. Layne and Robert S: HOPMOnNN 2.2 oss edhe thea eed sesS 
EHROGUCHION Wace Mist 22 dui, eq: pain ich aes Bu anes yeah cates mat msad Sate Slates 
Ee SIC ential p PEO lier Sets errant tat aot ham eee Rett tean dae Sa ea ale hg aes ee orale 
BIOS rap MICS KEICNES a man oa ba aS od ott aaa doe hs ee ee espana oem Mee cat eden eee 
NClMOWICGRINEDS Sn eG Oar eaiee cages se ee he alae ee ete Ae be 3.65 bau ee oot aes 
Tre neitnpres CGM tes aN tig, Rag sree ey tan pe teh ta ne al oh ants a _ateee ca 


AWARDEES J. Mary Taylorand Duane A, SCHUMCl 2) nce oo ee ow eee Pees goa deeeees 
TALEO GU GLIOM pee Mets e5.2 Mire ee re ae ale eee a tt tare EM Ree nis SG Ae ete 
ILONORARY IVICINDErS ite a ate eG SRO AT FR erates a Neg Se ae 
Caan MERaInek WaldCeS da.0e. hoe ae Naoto en CAA a hae PER Uae ee Bee nee eee 
Brantley Elli FAGKSOMUAWATOUCES Daren: gals a.com ae eatin nnn cde a areas eds is eed occas 
CEQ CIU SIO tS ree ee aaa OR ple tee ca etm care Sete ie 2 laa, Mat cols lhe, LI eS a O 
FACRMOWICGO MENUS cde, Sidi sete ack ee eahe 2 Bon acar meat Thon anal ke ier tee ee eae? We eal ate oe 


OTHER PROMINENT MEMBERS David M. Armstrong, Murray L. Johnson, and 
RONG OWNER RELErSOMs fans tte eee ae ee ee eG eae ees, anaes 
IU OGUCIOler eee haere eae en eat eet oie rancor Ake Ne tune ee Rae a 
BU gl 22 0h oe cee eee eae eee vee EL ag Se ey eC ee 2 eee gn OES ny 
VEL pYoroal 108183 0 ILO nen om teed ea OF tegen En ES) OCT SO rg ARE reg ates OE errs sc eee 
BURLY 1090) ae ere cree nce eee en a ee pC Sn an ek Cy nS A. 
Ua Ge lbs) Serene rete, 0, PR eee TR oe eee, ante eI Et A as Allee Wi ea cha 
BING sD. Sareea wees erates wer ee ee ee eee ee cate ke aes ee ne ees 
GE 0 an) AS 27 (San aS cS  ea e Y e e a ae e 


ACADEMIC PROPINQUITY SONMLO AVUILGKCI gy ere ee este =, Ae, ee 
MVE THO CUNG ENO TIC ce enero ae eee ee ay eee ek eric nape ears ern 
Ihe: NViemiam: Group) 4, ...242. 055. eo ee ee ces een ee 
Me the Agassiz/Glover Allen Group (Harvard) {is.i..2..0e07-8 eee ose eee ee 
III. The Joseph Grinnell/E. A. Hall Group (Berkeley and Kansas) ..................... 
IV Phe Walliam. Hamulton, Jr. (Group (Cornell) 2). os eo oe i ee 
LALO 18 Fes col S100) 6 oo ae ae era en ne ne a Dy no Og Oe oP 
Acknowledgments 
ite rapinkc: Cie creamy eee yee ee ce ee a ee me ere nan weve ee 


PUBLICATIONS Bio). Verts and bner GC. BUney wen ones a6 cask on tes es een eae ee 
Ja eC (0 117) 50) 1 eee eee nme Oem RY 0> SPEER Te SETA PORE tS TR RE RE TY Ec eee ee ee 
lieed ournal Of MIGWIMNGIOCY .24. -chsacs gem bats eae nk eo ne ee ee BR eg ok 
VA GITIIGUI GANS DECICS ie awe ay esa rues ee ee ek ee ES ces HN og ae 
Monographs and Special Publications 3.25 24. 2c. eens an cesta ces eee eene Seana ee: 
Cumulative Indices and Miscellaneous Publications ................0.0 0000 ce cece 
ACKHOWICCOTMCIUS: = ote. i ees ee See ee, Uae Be Geter ee Redeem eee 
Piteratune riled, a422 aaa ane ee ee ee ee ce Ae ie ee eee eee ema 


N 


NO — 
NO S&S CO he Kee 


N 


COMMITTEES AND ANNUAL MEETINGS Ayesha E. Gill and W. Chris Wozencraft .. 
INMtFOGUCUON 4.252.025 fas 358826 bitte ere ete ere aa Oe eee oe ese a ts ae eee 
Histor or che Comumittees of ASM 1c... 222/502 ites nc alacuat ee een eeteeee ee aa 
ihe Hlustory of ASMcAnnial Mecunes: . 2 sic 2 6.50, ee gon sha ion a eng See ee oe 
Biterature: Cited % 202 2 ie certs ec oa hike oe AS Sea ee on es a een ee een eee 

MEMBERSHIP AND FINANCE Gordon L. Kirkland, Jr. and H. Duane Smith ......... 
RTEPOCUCHON:” 4, 358 ba a arnien se Qe oe nS any on hae oan eee ras ee See ee 
Mero bership Glassesir oucctesy sce xs tau as ea ater ado cots atl a ra ee 
Membership History... Wee 25 2 dar0 ind. iut aud baat als Cate ae eet 
Intemational Micmibersiiip 2, gaint sa ere soars acs orate eae ne ire ee ee 
Corresponding Secretary, Treasurer, and’ Secretary- lireasurer ...22...2.. sae. 2 eee eee 
RESET V Gai TUINC AS errata eaten cc icici eer Ng Bars Nes LeU en coe eens a 
AAS MAV BUC BCES Au ecte cestia se dens) eR asics wo ous waa 6 aaah tact n eee: ee a ae ne 
po] 10060 800) @) OMe sae ieee tena Cee eek eG Rrra hme © Gelea eeebed terran ede ne Nee Meee ee nerarerinnes tn ree a 
AACKMNOWICASINGIIS. 2 (orem. anaes dene wee eee tee en aed nn ae ee 
Heiterature | Wed oo. ed ee ee eae ee ne es Le ck en ee rae 


PART II. Intellectual Development of the Science of Mammalogy 


TAXONOMY Mark D. Engstrom, Jerry R. Choate, and Hugh H. Genoways ........... 
| Goi ieova Ub Tey oY) ote memrebmeaer eearaee Wl ultra Ur patty rae ural verge esir ream we arse nr erie) Met yw ery es eanny rae eee ee ck A 
Histoncall Perspective qe ee ne ns ee eee ee eee ee 
BiglopicaleSpecies Concept... ei ie ets se ee ee te ea eee 
SUIOSWECICS CC OMCs eer ae em eg ye Seen eee 
Figher evel axOnOmiy ger. eee oe ee aa ee eee ease ee 
auinal@Survicy Sse icc ae ere eer eis eee rae cir ene a een cee mice see eg 
PREKTOW ECO IT TALS tent eee ee eee eg eee ee 
Wikerature:Citedi ee a. eee ete eee ea Stee ete ae eee er eee 


PALEOMAMMALOGY Richard J. Zakrzewski and Jason A. Lillegraven .............. 
DtEO CIC TIO ae eros oes eee eee apres yore ne Sa Ee pe ee 
Compartmentalization of Mammalopgy +22... 1. -.o2-.225..-24s 224560105 ss asntee nee 
General Advancements...) 4-e oe et ee e e e e eee 
Geologically Directed Paleomammialogy ...::....2-. 2.246.462.5454. 004-6eu140sse08 
Biologically Directed Paleomammmalogy ...... 2.2.4 ..-202h26..605 ne sndnendw toned sees 
The Blending of Geologically and Biologically Directed Paleomammalogy ............. 
EDilOSUC se ce et oe aoe te So tae Meta AME Ate te ooh 2 ieee Ue Se ct 
ACKNOWICAUSIMENIS 66 okct ean ened xP Fo Ed Pods Ga £ LA Sod we antl dis 4 atin ho ee eee 
MiteratureCed: 25.456 ea ok eh oe eee adeee ee tee the. hey hele Peele eee 


BIOGEOGRAPHY Sydney Anderson and Bruce D. Patterson ......:4:.. 5202002 ee 
IntrodUCtION( «54 = eccidelenc eaddlonandtee Echadoe garde cates aoa ganaus nee s eee 
Histoncal Trends. 23 422644 28 4 Soden Pad aod toned oar ood nhae eninge ce te Oe pe eee 
Species Over Ecological Time Penods.2.222...4 424.2022 oo sss eon es soos cae eee 
Biotas Over Ecolopical Time Periods. .¢2.).%).4.4.-6.4%40060% mtn der sake eaeesaes seems 
Species Over Evolutionary Time Permods:,¢. 52.42. 4.40452552e02se4nsee eee eee 
Bietas Over Evolutionary Time PerodS<22...2224c2. 222. 340805e ne se a eee eee 
iterature Cited) =) 3.5.4.ceddi6 8a Crd. Sasha cba eh ee See Reheat ea eee eee 


ANATOMY Carleton TS GPITS: 4 Peres S6 Ge ok Bo CaS Oyo SA ee ee 
Introduction... ...«. odes See 28 Wi ih eao Det ea ea aa aula th @ a dtd bE dee Gale Se Ng eee 
Paradiems.and Conceptual Frameworks 2.4; auc n¢ani+ ac. 2 6ee eal ee eee eee 
he BarivieHlistOny: £5 «4 e\sck 2, fam canhealeracidon tate: ind 4 ae ghs panes ee Gen Oe ee ee 
ihelnivence:of Laxonomy 2.4.5.6 Ss cheno fnew eae ee eee ee oe 
‘Phe Iniluence of Natutal History: <..2..4 428.4 icn Sa eRe ee ee es ee 
The Future-of Mammalogical Anatomy: 3442.6 2005 ine das fa aoe ce oe ee 
ACkKnOWIEdSINEGHtS, 24.4 uaete © Site in mode CE Oe Reh SP oe Clone Lue waren cee ae ane 
HeTte TAGE CNC! Fs 216.0 Bho arcicgia ec beare ae wid Eee PASE Ge ee eee ee 


PHYSIOLOGY Bruce A. Wunder and Gregory L. Florant .......0. 00000000 ccc eee. 
INET EINE UT Oe a ree res tes he eaters weer iat eae ty Le ne SE tee a a anon eas toe a 
Physiology Backoroundhy 63.6 2526.05 5s Soe thee 1 hae nates dhe vetoed we baued andes 
RG Views MICUMOOUS, 24 xc. nenaet yet an tear eA tone eee eae heute eee sehen feo 


Ep MUG Ue ine Ne gc NN a ec 2 1 abs dae tee eM Rae a ee SO be tain yah daha teh 
PERS OUI Se OILEO Mat oa etn Lh etl te ete ees een ears akarmeeet ns ae not eee ec ola ra 


REPRODUCTION Oliver’? Pearsonand Go J. ReEnGoy 2) ces, bite oa oe is St he 
ITSO CEC UTOTE ee cis 5 Pie cal tacts Mie ate ete lls Sati tans iecasea cea nent sto, Ren A Ste bak Sok eee Fae 
Panly22 OUR Cent uiys ato oa os SO coe tan on wate hones Ones ewes aeuacen 
the Cambria se keg ag re elena ck aied edeee Nee cannes eit te nd btn 
ne Vols 1OpKInS* beCAaCy me. 3 cokes OO he Be es ae all ae cee Gece th aoe: 
ONT EC SACI CS erat ee Mt PANN eich senses ans 2 S0E tsa Eeas ets SOs ein aeeR ald elie ee 
PuntheriNotablesPUblicatlons @ saserees os ey oes Sea nik we Beek 6 ns ke ec KG 
Bate 20 thi Gemtunny ae... crs een aaa Oe old rate a hae ake Pens e a ts tag oe ano 
Reproduction, Neuroendocrinology, and Molecular Biology .......................... 
Environmental ‘Physiology and Regulatory Processes i... . 0421054 5:-5.605.-¢c08es uns 
Reproduciuycienerey Ie xpencivures sete ky manok cw niche cet ens any Gees eee oe nd go tna 9 ak 
Oltiction and Regulation of Reproduction. .¥24.0ye soe tase See te eee eee oe 
Behavior and Neuroendocrinology «22: 225.2 eee s en od eh eo es dns See ae eta eae eee 
INTARS UPI alSeerre mete et ee te cM so a Cc ed i 7 acer cs ae anes <A Oe 
Reproductive Mechnologies sa. aaa. ae. or ase Menace Aimee tote Kae en a 
INaturalsrnstory-and the Utne a. oye een cease es ka ek oa a eee hoe es key eeees 
PXCKMO WILE GOIN CTL Sa ene ties ape onan areas SN aa reas Cl Oe My Ui, mes ne a 
PEt trea tame Gc Leen Sree es ee a ae Se uen eect eet ee as 


MOLECULAR SYSTEMATICS Rodney LoHoneyoutl and Terry Lk. Yares, 4. acest. 
Meraitiecy ne ERO Tne eee eee, weenie os ee a Rel Hg cee onc ees oe ee Os, Sar el Yee pe 
Molecular Techniques in Mammalian Systematics ................ 00.0. .0 cece ences 
Molecular Glock Concept we is... cue one ey ee ee eee eee Se ees 
Emereineissucs-and Future DirectionS..04..0- sor sete eee ee ae ee 
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X1V 


PART I 


HISTORY OF ASM AND ITS MOST 
PROMINENT MEMBERS 


ORIGIN 


DONALD F. HOFFMEISTER AND KEIR B. STERLING 


Introduction 


BY, in the 20th century, groups of sci- 
entists with common interests were 
banding together to share their mutual re- 
search and activities. Some of these groups 
organized themselves to form scientific so- 
cieties. Among vertebrate zoologists this was 
especially true of the ornithologists, but 
much field work and research on mammals 
also was being conducted during this period. 
For example, the United States Bureau of 
Biological Survey had field workers study- 
ing and collecting mammals in many parts 
of North America, and by 1919 this group 
had published 44 monographs on mammals 
in the North American Fauna series. Uni- 
versities and museums in the United States 
also were becoming increasingly more ac- 
tive in mammalian research. Thus, it is not 
surprising that in 1919 a group at the United 
States Bureau of Biological Survey would 
consider forming a scientific society for 
mammalogists (Hoffmeister, 1969). 

This chapter summarizes the historical 
details of the formation of this society, which 
became known as The American Society of 
Mammalogists (ASM). To accomplish this, 
it is necessary first to review briefly some 
of the earlier developments of mammalian 


research, collections, and societies in the Old 
World, as well as the work conducted with 
mammals in North America prior to the 
formation of the ASM. 


The Development of Scientific 
Societies in Europe in the 18th 
and 19th Centuries 


In the Middle Ages, interest in mammals 
was directed more to their economic im- 
portance—as a source of food and furs, as 
beasts of burden, as animals that could be 
domesticated. People were most familiar 
and concerned with those mammals they 
could readily observe, especially those that 
were diurnal or of large size. As interest in 
identifying and describing all of the mam- 
mals developed, it was advantageous and 
even necessary to collect and preserve them. 
With more parts of the world being explored 
and colonized and new and unusual mam- 
mals being encountered, these needed to be 
collected and described. The naturalists in- 
volved in research on these new discoveries 


2 HOFFMEISTER AND STERLING 


were slow in joining together to form sci- 
entific societies. However, as their numbers 
increased and printing became possible and 
affordable, they formed groups with com- 
mon interests which were the forerunners 
of our scientific societies. 

Already in the late 1600s, collections of 
animals and plants increased so much that 
museums, usually called cabinets, were be- 
ing developed, especially by wealthy indi- 
viduals. In Paris alone there reportedly were 
over 200 private cabinets and a visitor’s 
guide to Paris, published in 1787, listed 45 
notable cabinets of natural history for that 
year. It was said that every member of the 
leisured class felt it necessary to have a cab- 
inet of natural curiosities. The owners of 
such cabinets and collections usually were 
not scientists or naturalists. Some spon- 
sored collecting expeditions, both locally and 
to distant places, and hired persons to do 
the collecting for them. Among the Euro- 
pean collections in the 1700s were those of 
the English East India Company, the cabi- 
nets of Becoeur and Mauduyt de la Varenne, 
Sir Hans Sloane, Lady Margaret Cavendish 
Bentinck, Comte de Reaumur, Coenraad 
Jacob Temminck, William Yarrel, Lord 
Stanley, Rene-Antoine Ferchault de Reau- 
mur, and the Cabinet du Roi. Most of these 
were private cabinets or collections, al- 
though some were opened to the public for 
a fee. 

In the 1700s the preservation of mammal 
and bird specimens was haphazard and poor 
(Farber, 1982). Many specimens were with- 
out adequate data. Specimens obtained from 
foreign travelers were often of even poorer 
quality. Attempts were made to improve 
the quality of the materials. Some of the 
larger private collections hired curators. 
Many of these were trained in universities 
or apprenticed under medical doctors. Of- 
ten those persons working with the collec- 
tions published catalogues of the specimens 
in their cabinets. For example, Mathurin- 
Jacques Brisson’s Ornithologie of 1760 has 
been referred to as a collection-catalogue of 
natural history, and Etienne Geoffroy Saint- 


Hilaire’s work of 1803 was a Catalogue des 
Mammiferes due Museum. Some of the 
wealthy cabinet owners sought to advance 
their prestige by producing lavishly illus- 
trated books, based in part upon their col- 
lections. Because of the great expense, such 
works were not widely distributed or avail- 
able. There were exceptions, such as the vol- 
umes of Histoire Naturelle by Buffon 
(Georges Louis Leclerc, Comte de Buffon), 
Starting in 1749. 

In the early 1700s, natural history soci- 
eties were practically non-existent. By the 
latter half of that century, groups of literary 
and scientific men frequently met for infor- 
mal discussions. Natural history was further 
stimulated by the publicity Carl Linnaeus 
was receiving. In England, James Edward 
Smith and a few friends formed a “‘Society 
for the Investigation of Natural History” 
and a group that was the forerunner of the 
Linnean Society of London was started in 
1782 by William Forsyth and nurtured by 
James Smith. In France, several local or 
provincial societies started, as in Caen, 
Cherbourg, Lyons, Nancy, and others, but 
most lasted only for a short time and not 
beyond the French Revolution. 

At the end of the 18th century, natural 
history became increasingly noteworthy be- 
cause many turned to improving upon Lin- 
naeus’ Sytema Naturae. Lamarck’s ideas 
stirred interest, as did the debates of Georges 
Cuvier and Etienne Geoffroy Saint-Hilaire 
(Appel, 1987). This was the period when 
18th-century natural history was changing 
to 19th-century biology. With such interest, 
local and national governments became 
more involved with making collections for 
the scientific communities. Government 
ships were sent to chart distant seas and 
coasts and to survey overseas holdings, as 
well as to trade with the new colonies. These 
were outfitted with naturalists whose intents 
were to bring back specimens. Examples of 
such voyages are those of Maximilan Prinz 
von Wied-Neuwied in 1815 to the New 
World, John Natterer to Brazil in 1817, 
sponsored by the Imperial Natural History 


ORIGIN | 


Museum of Vienna, Robert Herman Schom- 
burgh to British Guiana in 1835, sponsored 
by the Royal Geographical Society of Lon- 
don, Johann Jacob von Tschudi in 1838 to 
Peru, the voyages of the Beagle, and the 
explorations of Captain Cook. 

Museums and collections maintained by 
government agencies grew in size and im- 
portance, became available to the public, 
and began to incorporate some of the pri- 
vate collections (McClellan, 1985). The 
British Museum in its infancy was located 
in the Montagu House and was not open to 
the public. Sir Hans Sloane’s large collection 
was turned over to the museum and in 1830 
it moved to a new building, was recognized 
as the national museum, and soon addi- 
tional private collections were acquired. For 
example, James Edward Smith bought Carl 
Linnaeus’ library, manuscripts, herbarium, 
and specimens in 1788. Shortly thereafter 
the Linnean Society was formed, with Smith 
as president. Collections of the Society went 
to the British Museum, as did those of some 
other British societies. In the ensuing parts 
of the 19th century, the British Museum was 
curated by such mammalogists as George 
Robert Waterhouse, Richard Lydekker, and 
William Henry Flower. 

The Jardin du Roi became the French 
Museum National d’Histoire Naturelle in 
1793. Buffon had built the Cabinet at the 
Jardin into an outstanding collection during 
the mid-1700s. Etienne Geoffroy Saint-Hi- 
laire continued this endeavor when, at age 
21, he was placed in charge of the newly 
established national museum. Other per- 
sons with mammalogical interests associ- 
ated with the Muséum d’Histoire Naturelle 
were Georges Cuvier, Georges Duvernoy, 
Henri Milne Edwards, and Etienne’s son, 
Isidore Geoffroy St-Hilaire. Anselme-Gaé- 
tan Desmarest served part-time as a pre- 
parator for Cuvier. 

Germany had fewer private collections in 
the 1700s, but a museum was included with 
the establishment of the Universitat zu Ber- 
lin in 1810. Johann Carl Wilhelm Illiger was 
the first curator. Incorporated into this mu- 


seum was the Pallas Collection and the Cab- 
inet of Count Johann Centurius von Hoff- 
mannsegg. 

In Holland, there were many collectors 
(reportedly more than in all the rest of Eu- 
rope) and many private collections in the 
late 1700s. In 1820, the Rijksmuseum van 
Natuurlijke Histoire was started in Leiden 
and many of these private collections were 
incorporated. This included the private col- 
lection of Coenraad Jacob Temminck, who 
became the first director of the new muse- 
um. The Dutch government provided spec- 
imens from its possessions. Max Wilhelm 
Carl Weber, known for numerous studies 
on mammals including Die Saugetiere, was 
associated with the University of Amster- 
dam. 

The beginning of the 19th century saw 
the birth of many new scientific societies 
and the beginning of many new scientific 
journals. As collections and museums grew, 
so did the international community of 
scholars. At about the same time, it became 
possible to publish one’s findings more 
readily. For example, before 1802 there were 
few permanent journals in Europe for pub- 
lishing research in natural history. How- 
ever, the development of steam-driven 
printing presses in the early 1800s made it 
possible to produce up to 20 times as many 
impressions in a given time span. The price 
of production of periodicals and books de- 
clined. Also, the wars that had ravaged parts 
of Europe from about 1790 greatly subsided 
after 1815. 

As the number of scholars interested in 
natural history and biology grew, they began 
to associate into mutually beneficial groups 
or societies. There are many examples of 
societies and journals started in the late 
1700s and early 1800s. The Zoological So- 
ciety of London was founded in 1826, in- 
corporated in 1829, and shortly thereafter 
started publishing the Proceedings. The Lin- 
nean Society of London began in 1788 and 
published its Transactions in 1789 and its 
Zoological Journal in 1824. The Scottish 
counterpart of the Linnean Society, the 


4 HOFFMEISTER AND STERLING 


Wernerian Society of Edinburgh, was 
founded in 1808. Although the Royal So- 
ciety of London began at an earlier time, it 
was in 1820 that it reportedly threw off its 
social club aura and became a society of 
professional scientists. In 1823, the Plinian 
Society of Edinburgh, and in 1836 the Bo- 
tanical Society of Edinburgh, were founded, 
and soon they started the Magazine of Nat- 
ural History. The British Association for the 
Advancement of Science was founded in 
1831 and the British Ornithological Union 
in 1858. 

In France, the numerous provincial so- 
cieties that had been established began to 
wane; during the French Revolution, a de- 
cree was issued in 1793 that eliminated all 
societies patented or endowed by the na- 
tion. The Paris Academy of Sciences was 
revived in 1795; in 1822, the Societe d’His- 
toire Naturelle de Paris was founded and in 
1824 began publishing the Annals des Sci- 
ences Naturelles, and the French journal 
Magasin de Zoologie was founded in 1831. 

In Germany and northern Europe, soci- 
eties and academies were only beginning to 
emerge in the late 1700s. Most were asso- 
ciated with universities. One of the early 
societies was the Berlin Gessellschaft Na- 
turforschender Freunde, first established as 
a private society in 1742. In 1812, the Aka- 
demie der Wissenschaften became aligned 
with the University of Berlin. In 1822, the 
Deutsche Naturforscher Versammlung was 
organized; in 1831, the Gesellshaft Deutsch- 
er Naturforscher und Artze. The Archiv fur 
Naturgeschichte began publication in 1835. 
The Society for Finnish Zoology and Bot- 
any, later called the Societas pro Fauna et 
Flora Fennica, was founded in 1821, and 
published Fauna Fennica. The Society of 
Naturalists of the Imperial University of 
Moscow initiated publishing in 1811 the 
Memoires Moskovskoe Olshchestov Ispyta- 
telei Prierody, which was republished in Paris 
as Memoires de la Societe Imperiale des Na- 
turalistes de Moscou. 

During the 18th and 19th centuries, nu- 
merous factors contributed to the evolving 


study of mammals. The scientific explora- 
tion of many parts of the world and the 
growth of collections were early factors. The 
development of private collections that lat- 
er became public or university museums was 
significant. The growth of numerous sci- 
entific societies and the proliferation of sci- 
entific journals encouraged the study of fau- 
na and flora. Numerous persons during this 
time made an imprint on scientific thought 
and research, especially Carl Linnaeus, 
Georges Cuvier, Geoffroy Saint-Hillaire, 
Jean Baptiste de Lamarck, Alfred Russel 
Wallace, and Charles Darwin. 


North American Mammalogy Before 
the 20th Century 


The fauna and flora of North America 
were of considerable interest to the natu- 
ralists, travellers, colonists, and other visi- 
tors who arrived there beginning in the late 
15th century. Mammals they encountered 
were either eaten or had their fur or hides 
utilized for clothing, decoration, and other 
purposes. Thomas Hariot’s Briefe and True 
Report of the New Found Land of Virginia, 
published in London in 1588, was the first 
scientific effort to describe the natural re- 
sources of any part of what is now the Unit- 
ed States. Hariot accompanied Sir Walter 
Raleigh’s 1585 expedition to North Caro- 
lina. His 44-page account mentioned deer, 
rabbits, opossum, raccoons, squirrels, bears, 
“lyon,” wolves, and ‘‘Wolfish Dogges,”’ al- 
though he did not personally observe all of 
these. Hariot’s book underwent 17 editions 
before the 1620s. 

Spanish observers, notably Gonzalo Fer- 
nandez de Oviedo y Valdez, whose Historia 
general y natural de las Indias, Islas y Ti- 
erra-Firme del Mar Oceano (a natural his- 
tory of the West Indies) was published in 
1526, had preceded Hariot in reporting on 
the mammals of the New World, but these 
earlier writers were active in the Spanish 
colonies of the Caribbean and in Central 
and South America from the beginning of 


ORIGIN 5 


the 16th century. The first person to give 
concentrated attention to the natural history 
of Canada was Samuel de Champlain. How- 
ever, his studies at the end of the 16th cen- 
tury were principally concerned with plants. 

Other notable travelers and observers who 
mentioned the mammals of the English col- 
onies in their writings before the 18th cen- 
tury included Ralph Hamor, author of A 
True Discourse of the Present State of Vir- 
ginia (1615); Captain John Smith, in his 
General Historie of Virginia, New England, 
and Summer Isles (1624); William Wood, 
author of New Englands Prospect (1634), 
with a listing of New England mammals in 
verse; Thomas Morton’s New English Ca- 
naan (1637); and John Josselyn’s New En- 
gland’s Rarities Discovered (1672) and An 
Account of Two Voyages to New England 
(1674). 

John Lawson, surveyor general of the 
North Carolina Colony from 1708 until his 
death at the hands of Indians in 1711, pro- 
vided a full and detailed account of the 
mammals of that region. Pehr (or Peter) 
Kalm (1715-1779), a protegé of Linnaeus, 
traveled in the American colonies between 
1748 and 1751 and was a principal con- 
tributor to his mentor’s understanding of 
North American species for successive edi- 
tions of the Systema Naturae. His En Resa 
Til Norra America, published in Stockholm 
between 1753 and 1761, was translated by 
John Reinhold Forster in 1770-1771. The 
narrative provided ethological information 
for some species, and he mentioned fossil 
elephants found in the Ohio country. 

Undoubtedly the single most outstanding 
work to appear before the American Rev- 
olution was Mark Catesby’s The Natural 
History of Carolina, Florida, and the Ba- 
hama Islands, which first appeared between 
1729 and 1747. A later revision was com- 
pleted by Catesby’s friend George Edwards 
in 1754. Although a popular as opposed to 
a scientific account, Catesby’s was the first 
attempt at a detailed description of the 
mammals he observed. His two volumes 
contained illustrations of only nine mam- 


mals, as compared with 113 birds, 33 am- 
phibians, 46 fish, and 31 insects, most of 
them set against a background of plant life, 
but these combinations introduced for the 
first time many American ecological asso- 
ciations. Not until the early 19th century 
would there be any further notable advances 
in general descriptive mammalogy. 

The Revolutionary and post-Revolution- 
ary period offered some useful details about 
North American mammals in the works of 
such men as the Marquis Francois de Chas- 
tellux, a French army officer whose Travels 
in North America (1786) included a detailed 
account of opossum gestation written by a 
friend, and William Bartram’s Travels 
Through North and South Carolina, Geor- 
gia, ... (1791), which included a short de- 
scriptive narrative of the mammals en- 
countered on his travels. When Buffon 
published his account of New World fauna 
in 1769, detailing principally mammals, he 
was clearly unimpressed by his subject (Pe- 
den, 1955). He implied that American spe- 
cies were “‘shrivelled and diminished”’ both 
in size and variety because of excessive 
moisture and less heat than was to be found 
in Europe. In Thomas Jefferson’s Notes on 
the State of Virginia, written during the 
American Revolution and later revised, the 
future president went to considerable pains 
to amass statistical data with which he ef- 
fectively demolished the French savant’s 
views. Buffon appeared to be convinced by 
the weight of Jefferson’s evidence, and 
promised that suitable corrections would be 
published in the next volume of his Histoire 
Naturelle. However, he died before this 
could be accomplished. 

Philadelphia was the first important cen- 
ter of research in natural history in the Unit- 
ed States, and it maintained its dominance 
in the field from the late 1790s until the late 
1830s. There were a number of reasons for 
this. Curious naturalists in the Quaker city 
had closer ties with their English and French 
colleagues than did naturalists in any other 
part of the country. Here, the American 
Philosophical Society, the oldest scholarly 


6 HOFFMEISTER AND STERLING 


= s ———— .. 


Charles Willson Peale* John D. Godman Richard Harlan 
(1741-1826) (1794-1830) (1796-1843) 
Courtesy of the Pennsylvania Academy of Courtesy of the Library, College of Courtesy of the Library, College of 
Fine Arts, Philadelphia. Gift of Mrs. Sarah Physicians of Philadelphia Physicians of Philadelphia 
Harrison (The Joseph Harmison, Jr. 
Collection) 


Elliott Coues, M.D. Spencer F. Baird Rev. John Bachman 
(1842-1899) (1823-1887) (1790-1874) 
Courtesy of the Smithsonian Institution, Courtesy of the Charleston Museum, 
Washington, D.C. Charleston, N.C. 


Fic. 1.—Eminent early North American mammalogists. 


* The self-portrait of 1822, showing his museum on the second floor of Independence Hall, Philadelphia. A mastodon skeleton 
exhumed and mounted by Peale stands to the right, partially obscured by the curtain. 


ORIGIN y 


organization in America, had been founded 
in 1743. Here too, other organizations, in- 
cluding Peale’s Museum, begun in 1784, the 
Academy of Natural Sciences, founded in 
1812, and several medical colleges were in 
operation. Philadelphia also led the rest of 
the nation both in the number of libraries 
and in the numbers of books they contained. 
At least 40% ofall scientific periodicals pub- 
lished in the United States were published 
in Philadelphia by 1832, at a time when 
none was being published in New York City. 

Art and medicine were the two major av- 
enues through which Americans ap- 
proached the study of mammals in the 19th 
century. The Maryland-born Charles Will- 
son Peale (1741-1826) was a largely self- 
taught artist whose interest in natural his- 
tory began to manifest itself when he was 
in his mid-40s (Fig. 1). The museum he 
founded in his Philadelphia home in 1784 
was not the first in the country, but was the 
first successful one to be started north of the 
Mason and Dixon Line. It survived for more 
than 60 years (Sellers, 1980). The Charles- 
ton Museum in South Carolina had begun 
operations in 1773, and still operates today, 
but was slower to develop its natural history 
collections. Peale’s was the focal point for 
those working on mammals in Philadel- 
phia. Most books on the subject published 
from about 1815 until the early 1840s were 
largely based on specimens examined there. 
Peale’s Museum operated for many years 
on the top floor of Independence Hall, and 
he was probably the first to supply painted 
backgrounds suggestive of habitat for the 
cases in which many of his specimens were 
mounted. The natural history specimens in 
Peale’s Museum were exhibited as a unit by 
Charles’ sons and grandsons until forced to 
sell, with most going to P. T. Barnum and 
some to the Boston Museum. Peale also at- 
tempted a series of public lectures on what 
was then (1799-1800) known concerning the 
mammals and birds of the world. Peale’s 
Museum housed specimens brought back 
by the leaders of the Lewis and Clark Ex- 
pedition of 1805-1807, the first federally- 


sponsored scientific expedition. It cost the 
government about $2,500, and produced 39 
new species and subspecies of mammals. 
Most of the specimens were later lost in a 
fire, but a few survive to this day. 

The first attempt at a comprehensive 
compilation of mammals by an American 
was George Ord’s “North American Zool- 
ogy,” which appeared anonymously in the 
third edition of William Guthries’ 4 New 
Geographical, Historical, and Commercial 
Grammar, and Present State of the Several 
Kingdoms of the World (1815). Of the 167 
species listed, Samuel Rhoads determined 
in 1894 that “fifteen are undeterminable, 
twenty-four are Mexican and South Amer- 
ican species, eighteen are synonyms of other 
names on the list and ten are old world forms 
having no specific affinities with those of 
America” (Baird, 1859). Nevertheless, Ord’s 
24-page contribution was the first effort by 
an American to place American species in 
some scientific arrangement. 

Two Philadelphia physician-naturalists 
(Fig. 1), Richard Harlan (1796-1843) and 
John Godman (1794-1830), produced no- 
table works focusing on American mam- 
mals in the 1820s. Harlan’s Fauna Ameri- 
cana (1825) was largely a compilation, 
although he added 10 new American species 
and discussed the role of tooth structure in 
speciation. Godman’s American Natural 
History: Part I: Mastology, appeared in three 
volumes (1826-1828), and was the first es- 
sentially original work on mammals com- 
pleted by an American. The illustrations 
were based on mounted specimens in Peale’s 
Museum. The first part of the English ex- 
plorer-naturalist Sir John Richardson’s 
Fauna Boreali Americana dealing with 
mammals was published in London in 1829. 
Richardson focused on Canadian forms, 
some of them native to the United States, 
and his descriptions were still considered 
authoritative at the end of the 19th century. 

With the creation of the various state geo- 
logical and natural history surveys in the 
1830s, and a rapid increase in new infor- 
mation, a greater degree of specialization 


8 HOFFMEISTER AND STERLING 


entered into American natural science. A 
number of studies centered on particular 
states were published, such as Ebenezer Em- 
mons’ Report on the Quadrupeds of Mas- 
sachusetts (1840) and James De Kay’s five- 
volume Zoology of New York (1842-1844), 
which included a volume on mammals. Such 
publications helped expand the horizons of 
Americans interested in their native mam- 
malian fauna. 

The famous collaboration of John James 
Audubon (1785-1851) and his colleague, the 
New York-born Charleston-Lutheran cler- 
gyman John Bachman (1790-1874), result- 
ed in the brilliant three-volume Quadrupeds 
of North America (1845-1854) (Fig. 1). 
Bachman supplied much of the scientific 
information in this work, while Audubon 
(until his mind and eyes failed him in 1846) 
and his sons Victor Gifford (1809-1860) and 
John Woodhouse Audubon (1812-1862) 
completed the excellent mammal paintings. 
Audubon and Bachman tried to deal with 
all known species from the Tropic of Cancer 
north to Canada and Alaska. The work was 
intended for the widest appeal, necessitated 
in some measure by the costs of producing 
this very expensive set of books. As a con- 
sequence, there was no particular sequence 
of orders, families, and genera, although this 
weakened the finished product from a sci- 
entific standpoint. Today, the 155 forms de- 
scribed in the Imperial Folios of 1845-1848 
have been reduced to about 118. 

In June 1840, the 17-year-old Spencer 
Fullerton Baird (1823-1887) (Fig. 1), on the 
point of graduating from Dickinson College, 
Pennsylvania, diffidently wrote Audubon for 
help in identifying a flycatcher, which proved 
to be a new species. Audubon was kind, 
agreed that the bird was probably unde- 
scribed, and asked Baird’s help in capturing 
small mammals. Baird gave up the study of 
medicine, taught natural history at Dick- 
inson, and in 1850, was named the first As- 
sistant Secretary of the new (1846) Smith- 
sonian Institution at the comparatively 
young age of 27. Baird, a seminal figure in 
American zoology, brilliantly orchestrated 


the collecting talents of eager but unpaid 
civilian naturalists who accompanied the 
field parties exploring railroad surveys sent 
out by Secretary of War Jefferson Davis in 
the mid 1850s. From the materials thus de- 
rived, Baird wrote and published his fa- 
mous report on the mammals of the expe- 
ditions in 1857, which was commercially 
reprinted 2 years later as Mammals of North 
America. This substantial volume listed 52 
new species and 18 previously described 
forms not mentioned by Audubon and 
Bachman. Baird also listed 37 other species 
and varieties he had not personally seen or 
identified, together with 16 species of squir- 
rels and skunks, which he thought might be 
located in the United States. This totaled 
273 forms, although Baird was careful to 
state that some might prove invalid. Baird’s 
work was a model of accuracy for its time, 
with emphasis placed upon morphological 
detail and geographical range. 

As Baird had been encouraged by Au- 
dubon and Bachman, so he in turn provided 
all possible support to his contemporaries 
and to the next generation of individuals 
just beginning their professional careers. A 
wise and patient official, he doled out the 
limited practical assistance at his command 
and carefully gathered many of the speci- 
mens and field observations that form the 
basis of the excellent government collec- 
tions of today (Lindsay, 1993; Rivinus and 
Youssef, 1992). Among his protegés may be 
mentioned Elliott Coues (Fig. 1), Joseph 
Leidy, Robert Kennicott, Robert Ridgway, 
and C. Hart Merriam. 

The last several decades of the 19th cen- 
tury coincided with a period of ferment in 
American intellectual life and in American 
natural science. A few American and Ca- 
nadian colleges and universities began to 
offer modern biological training after the 
American Civil War. At the same time, the 
federal government became increasingly 
concerned with scientific research and ex- 
ploration. The creation of a federal De- 
partment of Agriculture in 1862 (which 
achieved cabinet status in 1889) provided 


ORIGIN 9 


the needed home for a number of research 
components. These included an Entomo- 
logical Commission, established in 1877, 
and the antecedents of work in animal in- 
dustry, begun in 1879. These and several 
other agencies separately organized, includ- 
ing the Fish Commission in 1871 (placed 
in the Commerce Department in 1903) and 
the U.S. Geological Survey in 1878 (placed 
in Interior), all helped to create a large group 
of professions interested in various kinds of 
scientific activity in Washington (Dupree, 
1957). The capitol city rapidly became an 
important center of scientific inquiry. In- 
deed, Congress gave some consideration to 
the establishment of a Department of Sci- 
ence in the early 1880s. One authority has 
pointed out that the 1,812 members of the 
Agriculture Department employed in sci- 
entific research in the year 1913 was larger 
than the number of American scientists 
known to be active in the first 5 decades of 
the 19th century. 

A number of professional organizations 
in zoology began making their appearance 
in the 1880s. The American Society of Nat- 
uralists and the American Ornithologists’ 
Union were founded in 1883, the Ento- 
mological Society of America in 1889, and 
the American Morphological Society, and 
later the American Society of Zoologists, in 
1890. These organizations and others that 
followed helped bring about a rise in sci- 
entific standards. The articles and reviews 
appearing in their journals made possible 
the dissemination of modern scientific in- 
formation. 

Mammalian paleontology, in which Har- 
lan and Godman had been early American 
pioneers, prospered with the work of Joseph 
Leidy (1823-1891), the first of whose two 
most famous works was published under the 
aegis of the Smithsonian. This was his An- 
cient Fauna of Nebraska (1854). The other, 
On the Extinct Mammalia of Dakota and 
Nebraska, was published by the Academy 
of Natural Sciences of Philadelphia in 1869. 
Other leading paleontologists who made 
contributions to the study of fossil mam- 


mals included Othniel Charles Marsh (183 1- 
1899), including his studies on fossil horses; 
Edward Drinker Cope (1840-1897), includ- 
ing work with mammals of the Paleocene, 
and Henry Fairfield Osborn (1857-1935), 
whose Age of Mammals in Europe, Asia, 
and North America (1910) and later works 
on the Equidae, on titanotheres, and on the 
Proboscidea were important additions to the 
literature. 

A good number of late 19th century lead- 
ers in ornithology were simultaneously ac- 
tive in studying mammals, both in the field 
and in the laboratory. They included Baird 
at the Smithsonian and Joel Asaph Allen 
(1838-1921) at Harvard, who moved in 
1885 to the American Museum of Natural 
History in New York City as its first curator 
of ornithology and mammalogy. Two pro- 
tegés of Baird also began making substantial 
contributions to mammalogy in the 1860s 
and 1880s, respectively. One was Elliott 
Coues (1842-1899), an Army physician in- 
volved with several of the federal geograph- 
ical and geological surveys of the west, and 
later a free-lance naturalist. The other was 
Clinton Hart Merriam (1855-1942), trained 
as a medical doctor (Columbia University, 
1879), who wrote Mammals of the Adiron- 
dacks (1882 and 1884) at an early age. 

In 1888, the Federal government estab- 
lished an agency in the Department of Ag- 
riculture called the ‘Division of Economic 
Ornithology and Mammalogy,” under the 
direction of C. Hart Merriam. This division, 
later to evolve into the Bureau of Biological 
Survey, developed in a most indirect way. 
The American Ornithologists’ Union in 
1883 created a committee concerned with 
the migration and geographical distribution 
of birds. Merriam was the chairman, and 
his group was so successful at gathering data 
that they soon had more on their hands than 
they could handle. Into this emergency 
stepped Spencer F. Baird and Senator War- 
ner Miller of New York, an old Merriam 
family friend. In 1884, they pushed a bill 
through Congress calling for a $5,000 sub- 
vention for the establishment of an Office 


10 HOFFMEISTER AND STERLING 


of Economic Ornithology to be placed with- 
in the Entomological Commission at the 
Agriculture Department. As head of this di- 
vision, Merriam invited Albert Kenrick 
Fisher (1856-1948), a friend and fellow 
alumnus of the College of Physicians and 
Surgeons in New York, to join the fledgling 
agency as his assistant. They began opera- 
tions in July, 1885 (Cameron, 1929; Ster- 
ling, 1977, 1989). Their task was to research 
“the interrelation of birds and agriculture 
[and] an investigation of the food, habits 
and migration of birds in relation to plants, 
and publishing report[s] thereon....” The 
intent of this operation was to benefit Amer- 
ican agriculture by collecting data and de- 
veloping information that farmers could use 
in fending off the depredations of harmful 
species. The relationship between birds and 
insects was to be an important element in 
this work. Within a year, Merriam’s re- 
sponsibilities had been expanded to include 
mammals and birds as they related to ag- 
riculture, horticulture, and forestry. 

By 1886, Merriam’s agency had achieved 
emancipation from the parent Entomolog- 
ical Commission; within 10 years, it had 
been redesignated the Division of Biological 
Survey, and by 1906, it had become the 
Bureau of Biological Survey, the name it 
would retain until 1940. In that year, it was 
combined with the old Fish Commission, 
then in the Commerce Department, to form 
the U.S. Fish and Wildlife Service, which 
was placed in the Interior Department. The 
United States National Museum was estab- 
lished in 1879 as an adjunct of the Smith- 
sonian. It is reported that the National Mu- 
seum started back-handedly when an 
unauthorized sign “‘National Museum of the 
United States” appeared in the hall with the 
collections of the Smithsonian. 


The Early History of the 
American Society of Mammalogists 


The United States Biological Survey 
flourished under the direction of C. Hart 
Merriam (Fig. 2). Merriam’s personal agen- 


da involved nothing less than a continent- 
wide biogeographical reconnaissance, and 
Congress officially incorporated this in its 
authorization of his agency’s expenditures 
in 1894. Merriam assembled an impressive 
cadre of young workers in Washington, D.C. 
Some of these came college-trained, but 
many had only a high school education. 
Merriam was critical of the educational phi- 
losophies of some American universities that 
stressed laboratory work to the exclusion of 
field work. He preferred to give his men on- 
the-job training, using field methods he had 
developed (Cameron, 1929; Sterling, 1977, 
1989). Included in those associated in the 
early history of the Biological survey were 
Vernon Bailey, Clarence Birdseye, A. K. 
Fisher, Frederick Funston, Edward A. 
Goldman, Ned Hollister, Arthur H. Howell, 
Hartley H. T. Jackson, John Alden Loring, 
Marcus Ward Lyon, Jr., Waldo Lee Mc- 
Atee, Gerrit S. Miller, Jr., Edward M. Nel- 
son, Wilfred H. Osgood, T. S. Palmer, Ed- 
ward Preble, Walter P. Taylor, W. E. Clyde 
Todd, and Stanley P. Young. 

Another event that gave great impetus to 
the study of mammals at this time was the 
invention and adoption of the cyclone mouse 
trap in the late 1880s. This trap and its var- 
ious modifications, including the Museum 
Special and live traps, opened up new vistas 
in the study of mammals. 

In the early 1900s an increasing number 
of persons outside of the Washington, D.C., 
area were publishing or lecturing about 
mammals. Some of these were associated 
with universities and others with large mu- 
seums. Included in this group were Joel A. 
Allen, Glover M. Allen, Rudolph Ander- 
son, Joseph Grinnell, W. D. Matthew, Er- 
nest Thompson Seton, and Alfred H. Wright. 

Many important events mentioned above 
led to the formation of the American So- 
ciety of Mammalogists: 1) the establish- 
ment and objectives of the U.S. Bureau of 
Biological Survey with its cadre of enthu- 
siastic, eager mammalogists; 2) the forma- 
tion and growth of the U.S. National Mu- 
seum with curators in mammalogy, 
including Elliott Coues, Gerrit S. Miller, Jr., 


ORIGIN 11 


oe 


Fic. 2.—Bureau of Biological Survey members working at the U.S. National Museum at the turn 
of the century. From left to right: Vernon O. Bailey, Wilfred H. Osgood, Edward W. Nelson, Albert 
K. Fisher. Photograph from the files of the U.S. Fish and Wildlife Service. 


and Frederick True; 3) the formation and 
growth of successful scientific societies for 
the other “‘ologies’’; 4) the use of the mu- 
seum-special trap and the associated in- 
crease in numbers of specimens of mam- 
mals in collections with uniform, standard 
data; and 5) the increased interest in teach- 
ing mammalogy at the college level. 

One young mammalogist working at the 
Bureau of Biological Survey had often 
thought about a society of persons interest- 
ed in mammals. This was Hartley H. T. 
Jackson (Fig. 3). In 1902, when a junior at 
Milton College, Wisconsin, young Jackson 
discussed such a society with his admired 
mentor, Professor Ludwig Kumlien, and his 
boyhood friend, Ned Hollister. Although the 
others were somewhat skeptical, Jackson 
was not. In 1910, Jackson who by then was 
working for the Bureau of Biological Survey 
in Washington, D.C., attended the annual 
meeting of the American Ornithologists’ 


Union held in that city. This meeting en- 
forced his earlier views and “‘I became more 
thoroughly convinced that we could make 
a success of a mammal society” (H. H. T. 
Jackson letter, 1968, in archives of ASM). 
For the next few years, Jackson ““muddled 
along with ideas, worked on a possible con- 
stitution or bylaws, figured on possible 
sources of members” (ibid). He discussed 
such an organization with Edward Gold- 
man when in the field on Horseshoe Cie- 
nega, Arizona, in 1915, and again with 
Goldman and Walter Taylor on the Nantan 
Plateau, Arizona, in 1916. Walter Taylor 
was enthusiastic about such an organiza- 
tion. 

The Bureau of Biological Survey was un- 
der the leadership of Edward W. Nelson in 
1918, Merriam having stepped down in 
1910. The Survey, by custom, held staff 
meetings periodically, but they gradually had 
become disorganized, according to Jackson. 


12 HOFFMEISTER AND STERLING 


Fic. 3.—Hartley Harrad Thompson Jackson, 
the one person whose dream, dedication, and 
perseverance contributed the most to the suc- 
cessful origin of the American Society of Mam- 
malogists. 


To rectify this situation, a committee of 
three—A. K. Fisher, Vernon Bailey, and 
Walter Taylor—was appointed to plan such 
meetings. Among other things, this com- 
mittee recommended that the scientific staff 
hold evening meetings monthly at the home 
of different staff members. 

At the third such meeting, held at Vernon 
Bailey’s home on 5 December 1918, Jack- 
son wrote (ibid) that he ‘“‘thought there might 
not be too much to talk about at the De- 
cember meeting, and suggested to Mr. Bai- 
ley [who would preside] ahead of the meet- 
ing that it might be a good time to bring up 
the matter of a mammal society.”’ Dr. Jack- 
son continued to write that ‘“‘near the close 
of a busy session the question as to the ad- 
visability of launching a new organization 
for the promotion of mammal study was 
brought up for discussion by Chairman Bai- 
ley. I had already appraised Bailey of some 
of my ideas such as that [A. H.] Howell, 
[Ned] Hollister, [E. A.] Preble, and [W. P.] 


Taylor should be on the committee, and 
that it would be advisable to have five other 
representatives, one from each of five other 
institutions outside of Washington. I had 
already done considerable work such as out- 
lining a constitution or by-laws, searching 
lists for possible members, etc.”’ At this 
meeting it was moved that a committee be 
appointed to canvas the situation, and this 
committee consisted of Dr. Jackson as 
chairman and the other recommended 
members. It was further suggested that a 
report be made at the next meeting on work- 
ing plans for the proposed organization. 
Eight days after the committee was ap- 
pointed, on 13 December 1918, the five 
Washington members met, discussed a con- 
stitution for the proposed society, suggested 
a first regular meeting in the spring of 1919, 
and added these non- Washingtonians to the 
committee: G. M. Allen, J. A. Allen, J. 
Grinnell, W. H. Osgood, and later, Witmer 
Stone. On 21 December the Washington- 
members of the committee met again and, 
quoting from Walter Taylor’s notes (in ASM 
archives), “decided upon the following rec- 
ommendations: (1) That there be organized 
a society for mammal study to be known as 
the American Society of Mammalogists. (2) 
That the constitution attached hereto be 
proposed as a basis for further considera- 
tion. (3) That the report of the Committee 
on Organization appointed by the Chair- 
man of the meeting of the Scientific Staff of 
the Biological Survey on Dec. 5, 1918, be 
received and the Committee discharged, it 
being understood that the Committee would 
be immediately reorganized as a permanent 
Committee on Organization independent of 
the Survey. (4) That plans be made for hold- 
ing a formal organization meeting of the 
new Society if possible in March, 1919.” 
At the next staff meeting at the home of 
Walter Henderson, 9 January 1919, the re- 
port of the committee was approved, the 
committee was discharged, and Jackson as 
chairman appointed a new committee con- 
sisting of the same ten persons. Also, the 
original notice of ‘““A proposed American 


ORIGIN i 


A PROPOSED AMERICAN SOCIETY OF MAMMALOGISTS 


You are cordially invited to join in a movement to organize a 
society for the promotion of the interests and study of mammalogy. 
It is intended that the society shall devote itself to the subject in a 
broad way, including studies of habits, life histories, evolution, ecology, 
and other phases. Plans call for the publication of a journal in which 
both popular and technical matter shall be presented, for holding meet- 
ings, both general and sectional, aiding research, and engaging in 
such other activities as may be deemed expedient. It is hoped that 
you will actively participate, and, if possible, attend the organization 
meeting which will be held in the New National Museum, Washing- 
ton, D. C., April 3 and 4, 1919, sessions commencing at 10.00 a.m. 
and 2.00 ep. m. No program of papers has been planned for this 
meeting. 

Prevalent opinion indicates that annual dues for members will be 
about three dollars. 

Kindly bring this notice to the attention of others who may be 
interested in the movement. 

Respectfully submitted, 


( HARTLEY H. T. JACKSON, Chairman, 
U. S. Biological Survey. 
WALTER P. TAYLOR, Secretary, 
U. S. Biological Survey. 
GLOVER M. ALLEN, 
Boston Society of Natural History. 
| J. A. ALLEN, 
American Museum of Natural History. 
; JOSEPH GRINNELL, 
Committee 2 University of California. 
on N. HOLLISTER, 
Organization National Zoological Park. 
ARTHUR H. HOWELL, 
H U. S. Biological Survey. 
| WILFRED H. OSGOOD, 
Field Museum of Natural History. 
EDWARD A. PREBLE, 
U. S. Biological Survey. 
WITMER STONE, 
Academy of Natural Sciences of Philadelphia. 


The following blank properly filled and sent to the Chairman or 
Secretary of the Committee, Biological Survey, U.S. Department of 
Agriculture, Washington, D. C., will constitute application for charter 
membership. 


TATA ATA ATA ALATALALATATALALALALATALA LA TALATALALAUATALAVATATACAVAUL YA UA VAVACLULUL YA VALAUL CALL TANI 2 a VAUA UD VAVAVALYAVAYA YALA A VAYAULVIYIVAVIVIVAVILYIVLY 


I desire to become a charter member of the American Society of 
Mammalogists. I shall attend the organization meeting. 


Fic. 4.—This announcement of the proposed Society was sent to prospective members in the 
United States and other countries. 


14 HOFFMEISTER AND STERLING 


A PROPOSED AMERICAN SOCIETY OF 
MAMMALOGISTS 


A COMMITTEE of representative American 
mammalogists, including men from different 
parts of the country in its membership, has 
recently been at work on plans to organize a 
society for the promotion of interest in the 
study of mammalogy. It is intended that the 
society shall devote itself to the subject in a 
broad way, including investigations of habits, 
life histories, evolution and ecology. The 
plans call for the publication of a journal in 
which both popular and technical matter will 
be presented, for holding meetings both gen- 
eral and sectional, aiding research, and en- 
gaging in such other activities as may be 
deemed expedient. It is hoped to secure the 
active participation of all interested. The 
organization meeting will be held at the New 
National Museum, Washington, D. C., April 


3 and 4, 1919, sessions commencing at 10:00 
AM. and 2:00 p.m. 


No program of papers 
has been planned for this meeting. The or- 
ganization committee includes the following: 
Hartley H. T. Jackson, Chairman, U. S. Bio- 
logical Survey; Walter P. Taylor, Secretary, 
U. S. Biological Survey; Glover M. Allen, 
Boston Society of Natural History; J. A. 
Allen, American Museum of Natural History; 
Joseph Grinnell, University of California; 
N. Hollister, National Zoological Park; 
Arthur H. Howell, U. S. Biological Survey: 
Wilfred H. Osgood, Field Museum of Natural 
History; Edward A. Preble, U. S. Biological 
Survey; Witmer Stone, Academy of Natural 
Sciences of Philadelphia. Further informa- 
tion will be furnished by either the chairman 
or the secretary, to whom applications for 
charter membership should be transmitted. 


Fic. 5.—Account of the proposed American 
Society of Mammalogists as it appeared six weeks 
before the first meeting in Science, n.s., XLIX, 
21 February 1919. 


Society of Mammalogists” was printed and 
mailed in early February, 1919, to prospec- 
tive members (Fig. 4). A notice of the forth- 
coming organizational meeting was pub- 
lished in Science, n.s., 49:189, 21 February 
1919 (Fig. 5). 

With an official committee set up for the 
organization of a society of mammalogists, 


five meetings were held in late January to 
March, 1919. The out-of-town members 
were usually unable to attend. Jackson was 
busily drawing up a list of prospective mem- 
bers, gathering funds to start such an or- 
ganization, and drafting the by-laws. These, 
Jackson said, were modeled after the con- 
stitution and by-laws of the A. O. U., Amer- 
ican Society of Naturalists, Wisconsin Nat- 
ural History Society, Wisconsin Academy 
of Sciences, and the Biological Society of 
Washington. Jackson learned that under the 
laws of the District of Columbia, where the 
Society was to be incorporated, the phrase- 
ology of “‘bylaws and rules, had to be used, 
not constitution.” On 23 January 1919, a 
most important meeting of the ““Committee 
on the Organization of Mammal Society” 
was held in Room 61 of the “New Muse- 
um,” Washington, D.C. Four typescript 
pages of this meeting are in the ASM ar- 
chives. Jackson chaired the meeting with 
other committee members consisting of A. 
H. Howell, Ned Hollister, and Walter Tay- 
lor. Other “resident mammalogists’” who 
were present included J. W. Gidley, E. W. 
Nelson, H. H. Sheldon, Charles Sheldon, C. 
Birdseye, William Palmer, T. S. Palmer, 
Vernon Bailey, C. Hart Merriam, George 
Field, W. C. Henderson, W. D. Bell, and 
M. W. Lyon, Jr. Most of the meeting was 
devoted to a discussion of the by-laws. Mer- 
riam ‘‘advocated simplicity in the consti- 
tution as the best way to promote effective 
business administration and permanence.” 
He also opposed ‘“‘the division of the mem- 
bership into different classes and favoring 
one general class of members, with possibly 
an honorary class composed of foreign 
members.” Thereby a section on “Fellows” 
was deleted by committee action, but a sec- 
tion on Honorary Members was included. 
The suggestion of meeting with other soci- 
eties was discussed but remained undecid- 
ed. Persons in the Washington area were 
encouraged to make voluntary contribu- 
tions of $2.00 for preliminary operations, 
and Jackson said the response was good. A 
total of $52 had been collected by the time 


ORIGIN ibs: 


of the first meeting, and of this $47.31 had 
been spent for 1,000 circulars and stamped 
envelopes, 500 printed membership cards, 
and 100 printed programs. The first annual 
meeting started with a balance of $4.69. 

The organizational meeting was held on 
3 and 4 April 1919 at the U.S. National 
Museum (Fig. 6). Eight members of the orig- 
inal organizing committee were elected to 
top positions in the new society. Although 
many persons were anxious for J. A. Allen 
to be the first president, he declined because 
of failing health. C. Hart Merriam was elect- 
ed President, E. W. Nelson and Wilfred H. 
Osgood Vice-presidents, H. H. T. Jackson 
Corresponding Secretary, W. P. Taylor 
Treasurer, H. H. Lane Recording Secretary, 
and Joel A. Allen Honorary Member. Ten 
members were elected to the Council (now 
Board of Directors), five for a 1-year term — 
R. M. Anderson, M. W. Lyon, Jr., W. D. 
Matthew, T. S. Palmer, E. A. Preble—and 
five for a 2-year term—G. M. Allen, J. Grin- 
nell, J. C. Merriam, G. S. Miller, Jr., and 
W. Stone. Every person who joined before 
or during the first meeting or the first year 
was regarded as a charter member and re- 
ceived a card signed by Jackson and Mer- 
riam. About 60 persons attended the meet- 
ing. 

At the first meeting, often referred to as 
the Organizational Meeting, there were three 
sessions of the Council (Fig. 7). These were 
held at 8 p.m., April 3; 9 a.m., April 4; and 
11:15 a.m., April 4. The original By-laws 
and Rules had a “Council or Board of Man- 
agers” (Journal of Mammalogy, 1:50, 1919). 
Before the society was incorporated in the 
District of Columbia, this was changed to 
Directors (Journal of Mammalogy, |, inside 
cover of No. 4, 1920). 

At the first session of the organizational 
meeting, Marcus W. Lyon was elected tem- 
porary chairman, H. H. Lane, temporary 
secretary. Two hundred and forty persons 
were accepted as charter members. The 
original list is on file in the ASM archives. 
At the afternoon session the officers and 
“councillors” were elected. Wilfred H. Os- 


good gave an “illustrated lecture on North 
American Mammals” at the evening session 
(1919 minutes, ASM archives). 

The business that transpired at the third 
(Friday morning) session can be summa- 
rized thus: 1) J. A. Allen unanimously elect- 
ed Honorary Member; 2) persons qualified 
for charter membership if they enroll before 
the next annual meeting; 3) incorporation 
of the Society under the laws of the District 
of Columbia; 4) plans to issue a quarterly 
publication known as the Journal of Mam- 
malogy, 5) appointed a Committee on 
Membership; 6) J. C. Merriam was elected 
‘““Councillor” to replace Ned Hollister, who 
was appointed Editor of the Journal; 7) es- 
tablished a Committee on the Study of Game 
Mammals; 8) next annual meeting in New 
York City. 

There was “‘quite a difference of opinion 
regarding the name of the Journal. Some 
favor a short name, like ‘Bison’, ‘Puma’, or 
something of that sort. Others like “Bairdia’, 
but I think that most of us here, at least, 
agree with you that ‘American Journal of 
Mammalogy’ is the most appropriate name 
suggested to date. Or, more simply ‘Journal 
of Mammalogy’ [letter from Walter P. Tay- 
lor to Glover Allen on 11 March 1919].” 
On 11 July 1919, Williams and Wilkins 
Company of Baltimore, Maryland, solicited 
the new society through President C. Hart 
Merriam and Glover Allen to print the 
Journal of Mammalogy. The report of the 
“Committee on Publications,” chaired by 
Ned Hollister, pointed out that Williams 
and Wilkins was the only company to sub- 
mit a bid. 

By January 1920, there were 11 life mem- 
bers with their membership fees invested in 
United States Liberty and Victory bonds. 
By the end of the second annual meeting, 
there were 441 members (Fig. 8), of which 
25 resided outside of the United States and 
Canada. Income for this period amounted 
to $3,003.58; expenses for printing and 
mailing the Journal and all other expenses 
were $748.44: monies invested in bonds and 
in the bank, $2,255.14. A memorandum of 


16 HOFFMEISTER AND STERLING 


APrey sg Pe eg nt tee 


Fic. 6.—This is the only known photograph of the first (organizational) meeting of the American 
Society of Mammalogists, 4 April 1919, taken at the Administration Building, National Zoological 
Park, Washington, D.C. 1. C. H. M. Barrett; 2. Walter P. Taylor; 3. Charles M. Hoy; 4. Arthur J. 
Poole; 5. Vernon Bailey; 6. Ned Hollister; 7. Marcus W. Lyon, Jr.; 8. George A. Lawyer; 9. Frank 
M. Jarvis; 10. H. H. T. Jackson; 11. A. K. Fisher; 12. Leo D. Miner; 13. W. B. Bell; 14. Witmer 
Stone; 15. Wilfred H. Osgood; 16. C. Hart Merriam; 17. J. W. Gidley; 18. W. H. Cheesman; 19. 
James S. Gutsell; 20. C. C. Adams; 21. G. W. Field; 22. Ned Dearborn; 23. T. S. Palmer; 24. Charles 
Batchelder; 25. Charles Sheldon; 26. E. A. Preble; 27. Rudolph M. Anderson; 28. Mrs. Witmer Stone; 


ORIGIN i, 


29. Mrs. T. S. Palmer; 30. Mrs. E. A. Preble; 31. A. B. Baker; 32. Mrs. C. H. Merriam; 33. Harry 
Oberholser; 34. Mrs. F. M. Bailey; 35. B. H. Swales; 36. Waldo L. Schmitt; 37. Alexander Wetmore; 
38. Mrs. Leo D. Miner; 39. Mrs. Waldo Schmitt; 40. Miss Catherine Baird; 41. Miss May T. Cooke; 
42. Mrs. G. W. Gidley; 43. J. W. Scollick; 44. Jonathan Dwight; 45. Mrs. Ned Hollister; 46. Mrs. 
Jane Elliott; 47. John P. Buwalda; 48. Leland C. Wyman; 49. H. W. Henshaw; 50. Warren Craven; 
51. Mrs. Marcus W. Lyon; 52. Remington Kellogg; 53. Viola S. Schantz; 54. Mrs. Anna Jackson; 55. 


E. W. Nelson; 56. H. H. Lane; 57. W. C. Henderson. 


18 HOFFMEISTER AND STERLING 


Ameriran Soriety of Mammalogists 


ORGANIZATION MEETING 
NEW NATIONAL MUSEUM 
WASHINGTON, D. C. 


APRIL 3 AND 4, 1919 


ALL BUSINESS SESSIONS WILL BE HELD IN ROOMS 42 AND 43 


Program 
April 3. Business session : : , 10:00 A. M. 
Luncheon for members. 1:00 P. M. 


Members are asked to assemble at 12.45 P. M. at B Street or 
North entrance of the Museum and proceed in a body to May- 
nard Cafe, formerly Tea Cup Inn, 611 12th Street, N. W. 


Business session : : . : 2:00 P. M. 


Informal program and conversazione . 7:30 P. M. 


Auditorium, New National Museum. 


April 4. Business session : . ; 10:00 A. M. 


Luncheon for members and their wives 12:30 P. M. 


National Zoological Park, Administration Building. | Members 
are asked to assemble at the B Street entrance of the Museum at 
12:00 o'clock sharp. Following the luncheon there will be a tour 
of National Zoological Park, under direction of N. Hollister, 
Superintendent. 


Fic. 7.—Program of the first meeting of the American Society of Mammalogists. Note that this 
was called the “‘organization”’ meeting. 


ORIGIN 1) 


AMERICAN SOCIETY OF MAMMALOGISTS 


EO Lay ioe Daahao 


I have the honor to inform you that you were elected a Sore ee 


at the meeting peld Opes 2-ve CGT : 
ine Ue neers ere 


~~ 


President. 


Fic. 8.—Original membership card of Hartley H. T. Jackson. Cards so dated represented charter 
membership. The first four lines are in the hand-writing of Anna Marcia Jackson. 


H. H. T. Jackson’s (ASM archives) of 3 May 
1920, states that “After many distressing 
circumstances the Journal of Mammalogy 
is started. .. . A written agreement was made 
with Williams and Wilkins Company, Bal- 
timore, to print the first volume. . . . It seems 
to the corresponding secretary [Jackson] that 
an endowment—a publication fund—is es- 
sential if the Society is to live up to stan- 
dards worthy of its membership. The in- 
come from such a fund would nourish the 
Journal through its precarious infancy, and 
could later be utilized for publishing mono- 
graphs or for whatever worthy cause the So- 
ciety might deem desirable.” 

On 3 May 1921, Jackson re-emphasized 
this request in the Second Annual Report 
of the Corresponding Secretary. He wrote: 
“The Society can take pride in having es- 
tablished a creditable magazine without a 
single financial donation toward its publi- 
cation or general expenses. This has been 
done at a critical period in industrial history 
and at a time when printing costs were al- 
most prohibitive. It has been possible, how- 
ever, largely through the Charter Members, 
who willingly paid membership dues for the 


year 1919, yet received only one number of 
the Journal during that year. With a normal 
increase in the number of members and sub- 
scribers we can hope to continue to publish 
under present conditions between 200 and 
250 pages, and 10 halftones a year. The ac- 
tual costs of printing and distributing our 
present edition averages a trifle less than 
$8.00 per page. Indications are that we shall 
soon be receiving first class manuscript in 
quantity sufficient to publish 400 pages a 
year. Is the Editor to be placed in a position 
where it will be necessary for him to refuse 
valuable contributions? It would seem that 
the Society could ill afford to sanction such 
a predicament. Diffusion of knowledge 1s as 
essential as its creation. Immense endow- 
ments are given to be devoted to research, 
investigations, and explorations. Compar- 
atively small sums set aside as permanent 
publication funds would make available 
some of the results now buried in manu- 
scripts. It is, therefore, essential to the best 
interests of the Society, the Journal, and 
everybody concerned, that definite and pos- 
itive action be immediately taken to raise a 
Permanent Publication Fund. Any amount 


20 HOFFMEISTER AND STERLING 


raised would actually be worth double the 
amount to the Journal because of the as- 
sured increase in the number of subscrip- 
tions which would follow the improvement 
in the Journal.” 

The Board of Directors, on 17 May 1922, 
heard and approved the report of the Com- 
mittee on the Allen Memorial, chaired by 
Harold E. Anthony. This report recom- 
mended that: 1) a permanent fund be cre- 
ated to be known as the J. A. Allen Me- 
morial Fund; 2) this fund be invested and 
the income used for the publication of such 
memorial numbers of the Journal of Mam- 
malogy or other publications dedicated to 
the memory of Dr. J. A. Allen; 3) a com- 
mittee be appointed to raise such funds; 4) 
a minimum of $10,000 be raised in 2 years. 
By the end of 1924, the Fund had acquired 
$7,606. At the 1925 meeting, John Rowley, 
noted taxidermist, offered to apply all roy- 
alties from his book towards this Fund until 
$10,000 had been secured from all sources. 
By 10 July 1928, the goal had been reached. 
Also at the meeting in 1922, it was proposed 
that the by-laws be amended to provide for 
a board of three trustees. These trustees con- 
tinue to manage the society’s reserve fund. 

The new society received considerable 
early publicity. Science in its 21 February 
1919 issue carried a report of ““A Proposed 
American Society of Mammalogists” and a 
follow-up account on 18 April 1919, of the 
organizational meeting, with elected officers 
and “councilors,” committees, and refer- 
ence to a forthcoming Journal of Mam- 
malogy. 

Concerning the Fourth Annual Meeting, 
Science reported in its 16 May 1922 issue 
that “among the many interesting papers 
that were given before the mammalogists 
was the ‘Symposium on the Anatomy and 
Relationships of the Gorilla.’ At this session 
the attendance was probably greater than at 
any of the others, and representatives of the 
press were present to make the most of a 
subject in which the public is at present so 
keenly interested [the infamous Scopes tri- 
al].” 


Of the Sixth Annual Meeting of the So- 
ciety, the Boston Evening Transcript had an 
interesting story. It began: ““Mammalogists 
take their electioneering seriously. Twenty- 
five of them, all members of the American 
Society of Mammalogists, spent an hour and 
a half at the Harvard Museum in Cambridge 
this morning, making up a slate of six of- 
ficers and as many directors. The hitch came 
in choosing the directors. On the first ballot 
the names of twenty-four candidates ap- 
peared, one fewer than the number of men 
in the room. Eight ballots were taken before 
the choice was made.” 

At the beginning of the 20th century, there 
was a marked increase in the study of mam- 
mals in the United States. Museums and 
universities were training young people in 
mammalogy, both in the laboratory and 
field. Sooner or later there surely would be 
an organization of such scientists. However, 
this would not have come about as rapidly, 
effectively, and successfully without the 
dreams and determination of Hartley Jack- 
son and a group of dedicated fellow workers 
in Washington, D.C. Their work toward the 
formation of a new society is attested to in 
a small part by the fact that between 5 De- 
cember 1918 and 13 March 1919, Jackson 
and his colleagues held a recorded nine or- 
ganizational meetings, and undoubtedly 
many other private discussions. Once the 
ASM was started, many persons continued 
unselfishly to devote much time to the op- 
erations of the society. For the first 14 years, 
Henry H. Lane of the University of Kansas 
served as Recording Secretary. For 23 years, 
Viola S. Schantz served as Treasurer. Anna 
M. Jackson, Hartley’s wife, did most of the 
record-keeping and typing during the for- 
mative period and during the years that 
Hartley served as Corresponding Secretary. 
The work of these and many others provid- 
ed a sound basis for the rapidly growing 
society. 

The foregoing paragraphs have briefly re- 
viewed the events and circumstances that 
led to the formation of a scientific society 
of mammalogists in the Americas in the 


ORIGIN Al 


early 1900s. A group of energetic and far- 
sighted mammalogists working in the Unit- 
ed States National Museum seized the mo- 
ment to spearhead the organization of the 
ASM. As stated in Article 1, Section 2, of 
their by-laws, it was their intention that: 
“The object of the Society shall be the pro- 
motion of the interests of mammalogy by 
holding meetings, issuing a serial or other 
publications, aiding research, and engaging 
in such other activities as may be deemed 
expedient.” The following chapters review 
how these aims and goals of 1919 have been 
accomplished during the ensuing 75 years, 
both through activities of the ASM and the 
growth and intellectual development of the 
discipline of mammalogy. 


Additional Readings 


One volume of an international history 
of mammalogy has been published (Ster- 
ling, 1987) and another is in progress. His- 
torical accounts of mammalogy in the USA 
include contributions by Hamilton (1955) 
and Gunderson (1976); Allen (1916) pro- 
vided insights into the career of a major 
American mammalogist and founder of the 
ASM. 


Literature Cited 


ALLEN, J. A. 1916. Autobiographical notes and a bib- 
liography of the scientific publications of Joel Asaph 
Allen. American Museum of Natural History, New 
York, 215 pp. 


ApPEL, T. A. 1987. The Cuvier-Geoffroy debate: 
French biology in the decades before Darwin. Oxford 
University Press, New York, 305 pp. 

Bairp, S. F. 1859. Mammals of North America. J. 
B. Lippincott, Philadelphia, 764 pp. 

CAMERON, J. 1929. The Bureau of Biological Survey. 
Johns Hopkins University Press, Baltimore, 339 pp. 

Dupree, A. H. 1957. Science in the federal govern- 
ment; a history of policies and activities to 1940. 
Harvard University Press, Cambridge, 460 pp. 

FARBER, P. L. 1982. The emergence of ornithology 
as a scientific discipline, 1760-1850. Dordrecht, 
Holland, 191 pp. 

GUNDERSON, H.L. 1976. Mammalogy. McGraw-Hill 
Book Company, New York, 483 pp. 

HAmILTon, W. J., JR. 1955. Mammalogy in North 
America. Pp. 661-688, in A century of progress in 
the natural sciences, 1853-1953 (E. L. Kessel, ed.). 
California Academy of Sciences, San Francisco, 807 
pp. 

HOFFMEISTER, D. F. 1969. The first fifty years of the 
American Society of Mammalogists. Journal of 
Mammalogy, 50:794-802. 

LinpsAy, D. 1993. Science in the subarctic: trappers, 
traders, and the Smithsonian Institution. Smithson- 
ian Institution Press, Washington, D.C., 176 pp. 

McCLELLAN, J. R. 1985. Science reorganized: sci- 
entific societies in the Eighteenth Century. Columbia 
University Press, New York, 413 pp. 

PEDEN, W. (ED.) 1955. Thomas Jefferson’s notes on 
the state of Virginia. University of North Carolina 
Press, Chapel Hill, 315 pp. 

Ruoaps, S. N. (Ep.) 1894. A reprint of the North 
American zoology, by George Ord. . . . George Stokely 
Printer, Haddonfield, New Jersey, 290-361; 1-90 


SELLERS, C. C. 1980. Mr. Peale’s museum: Charles 
Willson Peale and the first popular museum of sci- 
ence and art. W. W. Norton, New York, 370 pp. 

STERLING, K. B. 1977. Last of the naturalists: the 
career of C. Hart Merriam (revised edition). Arno 
Press, New York, 478 pp. 

.(ED.) 1987. An international history of mam- 

malogy. One World Press, Bel Air, Maryland, Vol. 

1, 198 pp. 

. 1989. Builders of the Biological Survey, 1885- 

1930. Journal of Forest History, 30:180-187. 


PRESIDENTS 


JAMES N. LAYNE AND ROBERT S. HOFFMANN 


Introduction 


he President is one of four elective of- 
ficers of the ASM, the others being the 
First and Second Vice-presidents and the 
Recording Secretary. The President is the 
official representative of the Society. His or 
her duties include presiding over the meet- 
ings of the Board of Directors and the gen- 
eral business meeting, appointment of chairs 
and members of standing committees, es- 
tablishing ad hoc committees to carry out 
specific tasks, designating representatives to 
other organizations, and preparation of an 
annual budget proposal with the help of the 
Secretary-Treasurer. Past-presidents are au- 
tomatically members of the Board of Di- 
rectors. 

The term of office of the President and 
other elective and appointed officers of the 
society extends from the end of the annual 
meeting at which elected or appointed to 
the end of the following annual meeting, 
normally from June of one year to June of 
the next. Prior to 1973, the President was 
elected for a 1-year term and was eligible 
for reelection. In 1974, the By-laws and 
Rules were revised to extend the term of 
office to 2 years, with no provision for re- 
election. 

Unlike many scientific societies in which 


22 


Se 9. the President is empowered to speak 
for the Society... 


election of officers is by mail ballot, the ASM 
has followed the practice of holding elec- 
tions at the annual general business meet- 
ing. Nominations are made from the floor 
and voting is by written ballot. The pros 
and cons of this policy have been debated 
over the years, but it has survived succes- 
sive revisions of the By-laws and Rules. The 
prevailing view has been that members who 
regularly attend annual meetings and take 
an active part in the affairs of the society 
are best qualified to judge the qualifications 
of candidates. The long succession of pres- 
idents who have ably served the society at- 
tests to the effectiveness of this system. 

The 38 presidents of the society during 
its 75-year history and their terms of office 
are as follows (living individuals indicated 
with an asterisk): 


1. C. Hart Merriam (1919-1921) 

2. Edward W. Nelson (1921-1924) 
3. Wilfred H. Osgood (1924-1926) 
4. William D. Matthew (1926-1927) 
5. Glover M. Allen (1927-1929) 

6. Witmer Stone (1929-1931) 

7. Marcus W. Lyon, Jr. (1931-1933) 
8. Vernon Bailey (1933-1935) 

9. Harold E. Anthony (1935-1937) 


PRESIDENTS 25 


10. Joseph Grinnell (1937-1938) 

11. Hartley H. T. Jackson (1938-1940) 
12. Walter P. Taylor (1940-1942) 

13. A. Brazier Howell (1942-1944) 

14. E. Raymond Hall (1944-1946) 

15. Edward A. Goldman (1946-1947) 

16. Remington Kellogg (1947-1949) 

17. Tracy I. Storer (1949-1951) 

18. William J. Hamilton, Jr. (1951-1953) 
19. William H. Burt (1953-1955) 

20. William B. Davis* (1955-1958) 

21. Robert T. Orr* (1958-1960) 

22. Stephen D. Durrant (1960-1962) 

23. Emmet T. Hooper, Jr. (1962-1964) 
24. Donald F. Hoffmeister* (1964-1966) 
25. Randolph L. Peterson (1966-1968) 
26. Richard G. Van Gelder* (1968-1970) 
27. James N. Layne* (1970-1972) 

28. J. Knox Jones, Jr. (1972-1974) 

29. Sydney Anderson* (1974-1976) 

30. William Z. Lidicker, Jr.* (1976-1978) 
31. Robert S. Hoffmann* (1978-1980) 
32. James S. Findley* (1980-1982) 

33. J. Mary Taylor* (1982-1984) 

34. Hugh H. Genoways* (1984-1986) 

35. Don E. Wilson* (1986-1988) 

36. Elmer C. Birney* (1988-1990) 

37. James H. Brown* (1990-1992) 

38. James L. Patton* (1992-1994) 


Presidential Profile 


Several of the early presidents played a 
key role in the prehistory of the ASM. Grin- 
nell was one of the founders, in 1903, of the 
short-lived Pacific Coast Mammalogical 
Club, apparently the first attempt to form a 
professional mammalogy society in North 
America (Jackson, 1948). The major figure 
in the establishment of the ASM was Jack- 
son. As early as 1902, he discussed with Ned 
Hollister the formation of a mammal so- 
ciety (Hoffmeister, 1969). More serious 
consideration of the idea took place while 
Jackson and Goldman were collecting in the 
White Mountains of Arizona in the summer 
of 1915 and when Jackson, Goldman, and 
Taylor were working on the Natanes Plateau 


in Arizona in 1916 (Hoffmeister, 1969). 
Jackson, together with three others (Bailey, 
Nelson, and W. Taylor) destined to become 
ASM presidents, was a member in 1918 and 
1919 of the informal group from the Wash- 
ington area known as the Biological Survey 
Association that formally proposed the for- 
mation of the ASM; and he served as chair- 
man, with W. Taylor as secretary, at the first 
meeting of the society in April 1919. Four 
presidents (Bailey, Jackson, Merriam, Nel- 
son) were signatories to the articles of in- 
corporation of the society in April 1920 
(Anon., 1923). With the exception of Hall, 
who became a member of the society in 
1923, all of the first 17 presidents, from 
Merriam to Storer, were charter members. 
Nelson and Osgood, the second and third 
presidents, were the first vice-presidents, 
serving in that capacity from 1919 to 1921 
and 1924, respectively. 

Typical of other scientific organizations, 
the sex ratio of the elective officers of ASM 
has been strongly male-biased; and it was 
not until 1982 that the first woman, J. Mary 
Taylor, was elected president. Prior to that 
time, Viola S. Schantz and Caroline A. Hep- 
penstall were the only women to hold office, 
that of treasurer, which together with sec- 
retary, was the traditional post of women 
in scientific and other organizations in ear- 
lier days. 

Most presidents (excluding charter mem- 
bers) joined the society in their early 20s 
(average age 23), with Findley and Jones the 
youngest (18) and Davis and Durrant the 
oldest (32). Presidents who were charter 
members averaged 46 years of age at the 
time ASM was formed. Storer (27) and An- 
thony (29) were the youngest and Merriam 
and Nelson the oldest (64). Van Gelder was 
the youngest president (40) at the time of 
election, followed by Wilson (42), Jones (43), 
and Genoways, Layne, and Lidicker (44). 
Goldman (73) was the oldest, followed by 
Bailey (69), Nelson (66), and Merriam (64). 
Considering only non-charter members, 
Durrant (58) was the oldest president at the 
time of election. As a group, ASM presi- 


24 LAYNE AND HOFFMANN 


dents have been relatively long-lived, with 
an average life span of 76 years, with Jack- 
son holding the record for longevity (95) 
and Matthew being the youngest at time of 
death (59). The average age of living pres- 
idents (as of June 1993) was 64, ranging 
from 49 (Wilson) to 91 (Davis). With six 
exceptions, all presidents have served for 2 
years. Goldman died within a few months 
of election, Matthew and Grinnell served 
only | year, and Nelson and Davis were 
elected for 3 years. Davis’s extended tenure 
was the result of a desire of the membership 
to maintain administrative continuity dur- 
ing a period of reorganization of the socie- 
ty’s finances. 

The usual path to the presidency of the 
society has been through membership in 
standing committees, service as a director, 
and election to the vice-presidency. Mat- 
thew, Allen, Stone, and Lyon were members 
of the original Council. With the exception 
of Merriam, Bailey, and Anderson, presi- 
dents have served from 1 (Nelson, Jackson, 
Kellogg, Van Gelder) to 9 (Patton) terms as 
vice-president, with a mean of 3 years. Ten 
presidents have held other elective offices 
in the society besides those of Director and 
Vice-president. Jackson, Howell, Burt, 
Hooper, and Hoffmeister served as Corre- 
sponding Secretary and Orr, Peterson, Van 
Gelder, and Anderson as Recording Secre- 
tary. W. Taylor was Treasurer. Anthony, 
Davis, and Anderson served as Trustees of 
the Reserve Fund. Thirteen presidents held 
editorial posts. These included Jackson 
(Journal of Mammalogy), Howell (Journal 
of Mammalogy), Burt (Journal of Mam- 
malogy, Special Publications), Davis (Jour- 
nal of Mammalogy), Van Gelder (Recent 
Literature), Layne (Special Publications), 
Jones (Managing Editor, Review Editor, 
Journal of Mammalogy), Anderson (Mam- 
malian Species), Hoffmann (Review Edi- 
tor), Genoways (Journal of Mammalogy, 
Special Publications), Wilson (Mammalian 
Species, Special Publications), Birney (Man- 
aging Editor, Journal of Mammalogy, Spe- 


cial Publications), and Patton (Review Ed- 
itor). 

With the exception of Matthew, who was 
born in New Brunswick, Canada, all ASM 
presidents have been born in the United 
States. Nine were born in the Northeast 
(Maryland [1], New Hampshire [3], New 
York [4], Pennsylvania [1]), 18 in the Mid- 
dle West (Illinois [4], Iowa [1], Kansas [4], 
Michigan [1], Missouri [1], Nebraska [2], 
Ohio [1], Oklahoma [2], Wisconsin [2]), and 
8 in the West (Arizona [1], California [2], 
Idaho [1], Oregon [2], Texas [1], Utah [1]). 
There is a historical trend in the geographic 
origins of the presidents, with the Northeast 
and Middle West predominating in the pe- 
riod up to the 1940s and increasing repre- 
sentation of western states in subsequent 
years. Interestingly enough, the Southeast 
has produced no presidents thus far in the 
history of the society. 

Slightly more than half (55%) of the pres- 
idents were born and spent at least their 
early childhood in a rural setting, while the 
remainder, with the exception of Lyon, who 
spent his youth on different army posts 
around the country, were born in larger cit- 
ies. The proportion of presidents born and 
raised in cities increases after the late 1940s. 
Regardless of the environment of their 
youth, almost all of the presidents devel- 
oped a consuming interest in natural history 
at an early age, sometimes through an in- 
terest in collecting objects or in hunting or 
other outdoor activity such as falconry 
(Layne). Birney and J. Taylor divided their 
interests between natural history and sports, 
football and tennis, respectively; and Ham- 
ilton was a champion boxer during his un- 
dergraduate years. 

Almost all presidents were strongly influ- 
enced in their pursuits of natural history by 
their mothers or fathers; particular friends; 
high school, college, and, in the case of 
Hamilton, Sunday school teachers; or mu- 
seum curators or keepers in zoological parks. 
The majority of presidents focused on 
mammalogy as a career during their college 


PRESIDENTS 25 


years as a result of the influence of an un- 
dergraduate or graduate professor or, in 
some cases, a fellow student. Seven presi- 
dents have had students who themselves be- 
came president. These include (students in 
parentheses) Grinnell (Burt, Davis, Hall, 
Hooper, Orr), Hall (Hoffmeister, Durrant, 
Jones, Anderson, Findley), Jones (Geno- 
ways, Birney), Hooper (Brown), Hoffmeis- 
ter (Lidicker, Van Gelder), Hamilton 
(Layne), and Findley (Wilson). A more-de- 
tailed “family tree’ of ASM presidents and 
other North American mammalogists is 
given in the chapter by Whitaker (1994) in 
this volume. Major influences on the careers 
of the earliest presidents were Spencer Ful- 
lerton Baird, second Secretary of the Smith- 
sonian Institution, who encouraged Merri- 
am as a youth and supported Nelson at an 
early stage in his career, and the famous 
ichthyologist and president of Stanford 
University, David Starr Jordan, who ad- 
vised Osgood to take a position under Mer- 
riam in the Bureau of Biological Survey 
while he was still an undergraduate. Osgood 
was not only one of ““Merriam’s Men” in 
the Survey but also lived in Merriam’s home. 
Merriam also played an important role in 
the career of Bailey, purchasing specimens 
from him when he was a youth and later 
bringing him into the Bureau of Biological 
Survey. The famous team of Nelson and 
Goldman was born when Nelson, who 
needed a field assistant for a survey of the 
southern San Joaquin Valley of California, 
happened to stop at the Goldman ranch to 
have his wagon repaired. Goldman’s father 
told him of his son’s interest in natural his- 
tory and suggested that Nelson might like 
to hire him, which he did. 

Nelson and Osgood were bachelors. Of 
the presidents who were married, six (Stone, 
Jackson, Storer, Burt, W. Taylor, Patton) 
had no children. The remainder had from 
one to five children, with an average of 2.7. 

Except for Matthew, a geologist and pa- 
leontologist, all ASM presidents have been 
neomammalogists, although some, such as 


Anthony and Kellogg, also published on 
fossil mammals. Other than Howell, who 
was primarily a mammalian anatomist, the 
major research fields of the remainder of 
the presidents can be broadly defined as ei- 
ther taxonomy or ecology. This categori- 
zation is, however, rather arbitrary, as one 
of the hallmarks of the work of many pres- 
idents has been the wide scope of their in- 
terests. Thus, persons who might be classed 
as taxonomists on the basis of the major 
body of their research may well have pub- 
lished significant papers in the area of life 
history, ecology, behavior, morphology, or 
physiology; and workers whose major re- 
search has been in ecology and life history 
have often done taxonomic or distribution- 
al studies as well. Given this qualification, 
the presidency of ASM has been dominated 
by taxonomists (67%). The early presidents 
were exclusively taxonomists, W. Taylor 
being the first president whose interests were 
in areas of ecology and life history, which 
in the present day would probably be de- 
fined as “wildlife biology.”’ Although begin- 
ning with Storer and Hamilton, ecology and 
life history interests have been more strong- 
ly represented in the ASM presidency, tax- 
onomy still prevails as the major field. 

In addition to the wide recognition of the 
research of ASM presidents among mam- 
malogists at the national and international 
levels, the work of several of the presidents 
has had an impact beyond the field of mam- 
malogy in the broader areas of evolution, 
ecology, and education. Examples include 
Merriam’s life zone concept, Matthew’s 
volume Climate and Evolution, Storer’s 
classic text General Zoology, Burt’s work on 
territoriality and home range, and Brown’s 
research on desert ecology. 

In addition to the diversity of their mam- 
malian research, most presidents have pub- 
lished on other taxonomic groups or in oth- 
er fields. Merriam and Nelson, for example, 
conducted ethnographic research and Mat- 
thew published many papers on geology. Of 
the other taxonomic groups of interest to 


26 LAYNE AND HOFFMANN 


ASM presidents, birds predominate. Allen, 
Grinnell, and Stone are as well known as 
ornithologists as they are mammalogists, 
and at least 17 other presidents have pub- 
lished one or more papers on birds. Also 
appearing in the bibliographies of presi- 
dents are publications on fishes, amphibians 
and reptiles, insects and other invertebrate 
groups, botany, plant ecology, conserva- 
tion, and a wide range of other topics. One 
of the most versatile researchers among 
ASM presidents was Hamilton, who, be- 
sides work on a broad range of mammalian 
subjects, published extensively on the ecol- 
ogy and life histories of other vertebrates. 
In addition to their service to ASM in 
many capacities, presidents have played an 
active role in over 20 other scientific soci- 
eties as president or other elective officer. 
These include the Ecological Society of 
America (W. Taylor, Hamilton, Brown), 
American Society of Naturalists (Brown), 
Wildlife Society (Storer, W. Taylor), Pale- 
ontological Society (Matthew), Biological 
Society of Washington (Osgood, Bailey, 
Jackson, Wilson), Texas Academy of Sci- 
ence (W. Taylor), Florida Academy of Sci- 
ences (Layne), Society of Systematic Zool- 
ogy (Durrant, Hoffmann, Peterson), 
Midwest Museums Conference (Hoffmeis- 
ter), Texas Mammal Society (Jones), Or- 
ganization of Biological Field Stations 
(Layne), Southwest Association of Natural- 
ists (Genoways), Nebraska Museum Asso- 
ciation (Genoways), Nuttall Ornithological 
Club (Allen), American Ornithologists’ 
Union (Merriam, Grinnell), Cooper Orni- 
thological Society (Osgood, Storer), New 
York Academy of Sciences (Anthony), New 
York Explorers Club (Anthony), Organi- 
zation of Tropical Studies (Jones), Ameri- 
can Society of Ichthyologists and Herpetol- 
ogists (Storer), and Association of Science 
Museum Directors (J. Taylor). ASM pres- 
idents have also served as editors of journals 
of other organizations, including the Auk 
(Allen, Stone), Condor (Grinnell), Ecologi- 
cal Monographs (Hamilton), The American 
Midland Naturalist (Hoffmeister, Birney), 


Evolution (Jones), The Journal of Wildlife 
Management (Storer), and The Texas Jour- 
nal of Science (Jones). In addition to these 
activities, presidents have served as board 
members of numerous conservation, aca- 
demic, and museum organizations, as well 
as scientific consultants or advisors to var- 
ious local, state, federal, and international 
agencies. 

ASM presidents have frequently received 
recognition from the society for their re- 
search, service to the society, and other con- 
tributions to the field of mammalogy. Mer- 
riam and Jackson have been memorialized 
through the creation of the C. Hart Merriam 
and the H. H. T. Jackson awards. Seven 
(Layne, Jones, Lidicker, Findley, Geno- 
ways, Brown, Patton) of the 12 presidents 
since the establishment of the Merriam 
Award have been recipients; and the Jack- 
son Award has gone to Jones and Anderson. 
Honorary Membership has been bestowed 
on Merriam, Nelson, Lyon, Anthony, Jack- 
son, W. Taylor, Howell, Hall, Storer, Ham- 
ilton, Burt, Davis, Orr, Durrant, Hooper, 
Hoffmeister, Peterson, Layne, Jones, and 
Anderson. Early in their careers, Anderson 
and Layne received ASM Graduate Student 
Honoraria. 

ASM presidents also have been the recip- 
ients of numerous honors and awards from 
other professional organizations as well as 
from academic institutions, governmental 
bodies, and environmental groups. Merri- 
am is the only president to have been elected 
to the National Academy of Sciences. 

Ten of the 38 presidents have served with 
distinction in the armed forces of the United 
States. These include Lyon, Anthony, Gold- 
man, Kellogg, and Storer who served in var- 
ious branches of the army in World War I; 
Hamilton and Findley (army) and Peterson, 
Hooper, and Layne (air force) during or just 
after World War IJ; Jones (army) in the Ko- 
rean War; and Birney (navy) in the 1960s. 

The educational backgrounds of the ear- 
lier ASM presidents were more diverse than 
those of later years. Merriam and Lyon were 
MDs, and Stone had an honorary D.Sc. Nel- 


PRESIDENTS 2d 


son, Bailey, Howell, and Goldman were 
largely self-trained scientists, and were 
known by some of their contemporaries as 
“range-raised naturalists and biologists” 
(Young, 1947). With these exceptions, pres- 
idents have invariably had bachelor’s and 
Ph.D.s, and a large percentage has also re- 
ceived master’s degrees. Presidents have at- 
tended 25 different undergraduate institu- 
tions, with the University of California at 
Berkeley, University of Kansas, and Cornell 
each having been attended by four future 
presidents; the University of Arizona, Yale, 
and Stanford by two; and the remaining 17 
colleges or universities by a single president. 
The list of institutions from which presi- 
dents have received their doctorates is much 
shorter (12), with over half (58%) of the 
degrees having been awarded by the Uni- 
versity of California (11) and University of 
Kansas (6) and a maximum of two by other 
institutions. 

The careers of ASM presidents have cov- 
ered a broad spectrum of employment, and 
summarization of their professional posts is 
complicated by the fact that in many cases 
persons have held a number of appoint- 
ments, either concurrently or successively, 
during the course of their careers. Thus, the 
following breakdown, based upon the pre- 
dominant, if not exclusive, type of positions 
held by ASM presidents during their careers 
is of neccessity somewhat arbitrary. Seven 
presidents have been employed in various 
agencies or organizations of the federal gov- 
ernment, including the original Biological 
Survey (Merriam, Nelson, Bailey, Jackson, 
Goldman, Kellogg), U.S. Fish and Wildlife 
Service (W. Taylor, Wilson), and the Smith- 
sonian Institution (Kellogg, Hoffmann). 
Over half (55%) of the presidents are iden- 
tified primarily with museums. Nine of these 
have been associated with public museums, 
including the American Museum of Natural 
History (Anthony, Van Gelder, Anderson), 
Field Museum of Natural History (Osgood), 
Academy of Natural Sciences of Philadel- 
phia (Stone), California Academy of Sci- 
ences (Orr), Cleveland Museum of Natural 


History (J. M. Taylor), and the Royal On- 
tario Museum (Peterson). Twelve more have 
been members of the curatorial staffs, and 
with professorial appointments in academic 
departments as well, of museums afhliated 
with universities, including the Museum of 
Comparative Zoology at Harvard (Allen), 
Museum of Vertebrate Zoology at the Uni- 
versity of California, Berkeley (Grinnell, 
Lidicker, Patton), Museum of Natural His- 
tory at the University of Kansas (Hall, Jones, 
Hoffmann), Museum of Natural History at 
the University of Illinois (Hoffmeister), 
Museum of Zoology at the University of 
Michigan (Burt, Hooper), Museum of 
Southwestern Biology at the University of 
New Mexico (Findley), The Museum of 
Texas Tech University (Jones), University 
of Nebraska State Museum (Genoways), and 
the Bell Museum of the University of Min- 
nesota (Birney). 

Five presidents have been teachers and 
researchers in academic departments at the 
University of California at Davis (Storer), 
Cornell University (Hamilton), Texas A&M 
University (Davis), University of Utah 
(Durrant), and University of New Mexico 
(Brown). One president (Layne) left acade- 
mia (Cornell) to spend a major portion of 
his career as a research biologist at the Arch- 
bold Biological Station, one (Howell) was a 
professor in a medical school (Johns Hop- 
kins), and one (Lyon) did much of his re- 
search while a practicing physician in In- 
diana. 

In addition to their research, teaching, and 
other professional activities, many (74%) 
ASM presidents have held administrative 
posts during the course of their careers. 
Merriam, Nelson, Bailey, Jackson, W. Tay- 
lor, and Wilson served as heads of sections 
or programs of federal agencies, including 
the Biological Survey and U.S. Fish and 
Wildlife Service Cooperative Research 
Units. Anthony, Peterson, Van Gelder, and 
Anderson were chairmen of museum mam- 
mal departments; Osgood, Matthew, and 
Allen were chief curators at museums; Orr 
and Patton were associate directors of mu- 


28 LAYNE AND HOFFMANN 


seums; and Stone, Grinnell, Hall, Kellogg, 
Hoffmeister, Jones, Findley, M. J. Taylor, 
Birney, and Genoways served as museum 
directors. Hall, Davis, Hoffmann, and Fin- 
dley served stints as university department 
chairman. Layne was director of research 
and executive director of the Archbold Bi- 
ological Station. Kellogg and Hoffmann held 
the post of assistant secretary for science at 
the Smithsonian, and Jones served as grad- 
uate school Dean and Vice-President for re- 
search at Texas Tech University. 


Biographic Sketches 


Following are brief biographies, arranged 
chronologically by term of office, of the 38 
persons who have served as presidents of 
the ASM during the 75 years of the society’s 
history. Published source materials used in 
preparation of the accounts of deceased 
presidents are given at the end of the ac- 
counts. 


Clinton Hart Merriam: 1919-1921 


C. Hart Merriam (Fig. 1) was a founding 
member and the first president of the ASM. 
His selection as the founding president of 
the new society was a logical choice, given 
the preeminence he had attained in the field 
of mammalogy by age 64 when he assumed 
the presidency. His career spanned the for- 
mative period of the science of mammal- 
ogy. He was born on 5 December 1855 at 
Locust Grove, New York, and at age 16 
joined the Hayden Survey of the American 
West. Throughout a long and extremely 
productive career that ended with his death 
in 1942, he helped shape the modern sci- 
ence of mammalogy. His parents lived in 
comfortable circumstances, in a “rural 
mansion surrounded by ample acres and 
shadowed by the Adirondack Mountains,” 
(Osgood, 1943). His early schooling appears 
to have been routine, and it is likely that he 
was much influenced by his natural sur- 


roundings. In his teens he began to collect 
birds and eggs and early came under the 
influence of Spencer Fullerton Baird, the 
second Secretary of the Smithsonian Insti- 
tution. At age 17, he was sent to a day pre- 
paratory school, Pingry Military, in Eliza- 
beth, New Jersey. After 2 years, he enrolled 
at Yale University to study medicine. How- 
ever, his interest in natural history contin- 
ued unabated, and he had already accu- 
mulated a significant series of publications 
when he enrolled at age 24 in medical school 
at Columbia University. While still a med- 
ical student, he was involved in organizing 
the Linnaean Society of New York and cho- 
sen its first president, having previously been 
involved in the organization of the Nuttall 
Ornithological Club. Graduating from med- 
ical school in 1879, he returned home to 
Locust Grove to practice, but continued to 
pursue his natural history avocation; at this 
time his increasing interest in mammals be- 
came evident. Through the early 1880s, most 
of his publications were devoted to mam- 
mals, and this early phase culminated with 
publication of Mammals of the Adirondacks 
in 1884. Nevertheless, his interest in birds 
had not flagged, and he was also active in 
the formation of the American Ornitholo- 
gists’ Union, becoming the first secretary of 
that organization. 

By 1885 Merriam was ready to give up 
his medical practice and accepted the po- 
sition of ornithologist in the Division of En- 
tomology of the Department of Agriculture. 
His position soon became a division and in 
1888 was expanded to include mammalogy, 
at the same time separating itself from en- 
tomology. This new scientific bureau of the 
government provided the vehicle for his 
principal life work; Merriam’s name is syn- 
onymous with the Bureau of Biological Sur- 
vey, and with the “life zone’ concept he 
pioneered. He inaugurated the North Amer- 
ican Fauna series and in the first four num- 
bers (1889-1890) described 71 new species 
and several new genera of mammals. He 
developed an ambitious program of field 
collecting throughout North America, aided 


PRESIDENTS 29 


, ° 


RS 


N 


C. Hart Merriam Edward W. Nelson Wilfred H. Osgood a 
(1919-1921) ‘anions ees 


William D. Matthew Glover M. Allen Witmer Stone 
(1926-1927) (1927-1929) (1929-1931) 


hy 


x Ne 
N 


r 


Vernon O. Bailey Harold E. Anthony 
(1931-1933) (1933-1935) (1935-1937) 


Fic. 1.—Presidents of the ASM from 1919 to 1937. 


30 LAYNE AND HOFFMANN 


by people such as Vernon Bailey, A. K. Fi- 
scher, T. S. Palmer (Fig. 5), and the incom- 
parable duo of Nelson and Goldman. 
Equally important was the recent invention 
of a cheap portable “‘mouse trap,” the Cy- 
clone. 

The avidity with which Merriam named 
new species ultimately led him to write a 
revision of the brown and grizzly bears of 
North America in which he described a total 
of 84 species, including one of separate ge- 
neric rank. However, from about 1900, at 
the age of 55, he began to devote most of 
his time to the ethnology of California In- 
dians, having become the beneficiary of the 
Harriman Trust. His work on bears was thus 
published when he no longer was devoting 
himself primarily to mammalogy. Paradox- 
ically, the nearly universal rejection of his 
systematic concept was balanced by his rep- 
utation, which resulted in his systematic ar- 
rangement nevertheless being employed 
even after his death. 

Among his many honors was election to 
the National Academy of Sciences in 1902. 
He married Elizabeth Gosnell in 1886 and 
they had two daughters (Sources: Grinnell, 
1943; Osgood, 1943). 


Edward William Nelson: 1921-1923 


The second president of the ASM and an 
Honorary Member, E. W. Nelson (Fig. 1) 
was born near Manchester, New Hamp- 
shire, on 8 May 1855 and, like his prede- 
cessor, Merriam, is said to have been in- 
terested in the out-of-doors as a child. 
During the Civil War he lived with his 
grandparents on a farm in the northern Ad- 
irondacks while his father served in the 
Union Army and his mother nursed in a 
hospital in Baltimore. He attended a one- 
room rural school until 1886 when his 
mother, now widowed, moved to Chicago 
and enrolled him in schools there. His for- 
mal education appears to have been some- 
what spotty, but continued until 1875. 


However, even by 1872 he had participated 
in a field collecting expedition to the western 
United States and after assuming a teaching 
position in Dalton, Illinois, in 1875 began 
to publish on birds. Like Merriam, he also 
went to Washington, met Spencer Baird at 
the Smithsonian, and was sent on a govern- 
ment expedition to Alaska. During this trip 
he carried out a variety of observing and 
collecting activities, including geography, 
ethnography, and zoology. Other expedi- 
tions to Alaska followed, resulting in a series 
of ethnographic and biological publications; 
although not trained as a scientist, Nelson 
was obviously an excellent self-trained nat- 
ural historian. 

By 1890 he was working for the Bureau 
of Biological Survey, as a special agent on 
the Death Valley Expedition. Thereafter, he 
and Edward A. Goldman began a series of 
field studies in Mexico, which continued al- 
most unabated until 1929 when he retired. 
During his later years, he became increas- 
ingly involved in administration of the Sur- 
vey, being named Assistant Chief in 1914 
and Chief in 1916, and serving until 1927. 
For the next 4 years, he continued his re- 
search as a Principal Biologist for the Sur- 
vey. Subsequent to his retirement, he spent 
some time in California, but died in Wash- 
ington, D.C. on 19 May 1934 (Sources: 
Goldman, 1935; Lantis, 1954). 


Wilfred Hudson Osgood: 1924-1926 


W. H. Osgood (Fig. 1) was born 8 De- 
cember 1875 in Rochester, New Hamp- 
shire; he was the first of five children. When 
the family moved to California in 1888, they 
settled in the Santa Clara Valley in a rural 
area at the south end of San Francisco Bay. 
Osgood’s primary schooling was in Roch- 
ester, and he attended three years of high 
school in Santa Clara, but the family then 
moved into the city of San Jose. Osgood 
had become interested in birds and egg col- 
lecting and was involved in the organization 
of the Cooper Ornithological Club in San 


PRESIDENTS 51 


Jose, which has subsequently become a ma- 
jor professional organization. 

After graduating from high school, Os- 
good accepted a teaching position in a small 
school in Wilcox, Arizona, for a year and 
then entered Stanford University shortly af- 
ter its founding. Here he came within the 
orbit of the eminent zoologist David Starr 
Jordan, then president of the university. It 
was Jordan’s suggestion that he leave Stan- 
ford before completing his BA degree in or- 
der to take a position in C. Hart Merriam’s 
Bureau of Biological Survey, but he was 
eventually awarded his degree in 1899. He 
spent over a decade with the Survey, pub- 
lishing a number of papers in the North 
American Fauna series, culminating in his 
monographic revision of the genus Pero- 
myscus in 1909. In that year he joined the 
staff of the Field Museum of Natural His- 
tory in Chicago, the second of his two posts. 
He was Assistant Curator of Mammals and 
Birds, receiving his Ph.D. from University 
of Chicago in 1918 for a dissertation enti- 
tled ““A Monographic Study of the Ameri- 
can Marsupial, Caenolestes,” which was 
published a few years later by the Field Mu- 
seum. He served as Chief Curator of Zo- 
ology for 20 years, until his retirement in 
1941. During his career at the Field Mu- 
seum, he alternated between studying col- 
lections, both at the Field and in museums 
in other parts of the world, and conducting 
field expeditions. He participated in about 
20 expeditions, 8 of which were major for- 
eign ventures. Asa result, the Field Museum 
mammal collections grew greatly in size and 
importance during his tenure. From his re- 
tirement until his death 6 years later on 20 
June 1947, he remained fully engaged in 
publishing scientific papers. He was active 
not only in scientific societies, including the 
Biological Society of Washington, the Chi- 
cago Zoological Society, the American Or- 
nithologists’ Union, and the British Orni- 
thologists’ Union, but also in a number of 
other clubs such as the Explorers Club. 

Like Nelson, his predecessor, he re- 
mained a bachelor (Source: Sanborn, 1948). 


William Diller Matthew: 
1926-1927 


William D. Matthew (Fig. 1) was born on 
19 February 1871 in St. John, New Bruns- 
wick. He acquired his interest in the natural 
sciences from his father, Dr. George F. Mat- 
thew, who was a well-known and highly 
skilled amateur paleontologist and an au- 
thority on the geology, paleobotany, and 
fossil amphibian tracks of New Brunswick. 

In graduate work at Columbia Univer- 
sity, he studied geology, mineralogy, and 
metallurgy, which provided a solid back- 
ground for his subsequent research in pa- 
leontology. He received the doctorate in 
1895 and the same year joined the staff of 
The American Museum of Natural History 
as an assistant in the Department of Ver- 
tebrate Paleontology. He rose to Assistant 
Curator and then Curator in the department 
and Curator-in-chief of the Division of Ge- 
ology, Mineralogy, and Paleontology. In 
1927, after 32 years service with the Mu- 
seum, he left to become Professor of Pale- 
ontology and Curator of the Paleontological 
Museum of the University of California at 
Berkeley. His courses in paleontology, de- 
spite their reputation as difficult, were taken 
by hundreds of students, many of whom 
went on to distinguished careers in the field. 

Although his early publications were in 
the field of geology, for example, crystal- 
lography and the structure of rocks in New 
Brunswick, the main body of Matthew’s re- 
search dealt with mammalian paleontology. 
His first major project after coming to the 
American Museum was to catalog, pack, and 
ship to the Museum the extensive collec- 
tions of E. D. Cope. This task introduced 
him to the mammal fauna of the Basal Eo- 
cene of New Mexico, which he later desig- 
nated as the Paleocene. In the course of his 
career he was to work on fossils of nearly 
every major group of mammals, including 
carnivores, insectivores, primates, marsu- 
pials, rodents, edentates, and ungulates. He 
played a leading role in fossil collecting ex- 


a2 LAYNE AND HOFFMANN 


peditions to many localities in the western 
states and Florida, as well as Mongolia, Chi- 
na, and Java. In addition to his basic studies 
on the phylogeny of various groups, he also 
contributed to general theories concerning 
the arboreal origin of mammals, the mode 
of formation of the mammal fossil-bearing 
strata in the western United States, and the 
major patterns of the origin and dispersal 
of the mammalian fauna of the world. It 
was the latter subject, treated in his book 
Climate and Evolution, published in 1915, 
for which he was most widely known out- 
side the field of paleontology. The book was 
a healthy antidote to the tendency at the 
time of erecting hypothetical land bridges 
to explain the distribution of related groups 
separated by ocean barriers. Although some 
of the major conclusions have not stood the 
test of time, the book remains one of the 
classic works in biogeography. 

In addition to his technical writing, Mat- 
thew contributed many articles to Natural 
History magazine and authored handbooks 
and guide leaflets on various fossil exhibits 
at the museum. He was active in prepara- 
tion of public exhibits. He was especially 
concerned with mounting fossils in a life- 
like posture and was a pioneer in the use of 
comparative myology and osteology for this 
purpose. 

His scholarship and solid contributions 
to paleontology brought him numerous 
honors from scientific societies during the 
course of his career, including election as a 
Fellow of the Royal Society of England. He 
was a Charter Member of ASM and, in ad- 
dition to his term as president, also was a 
member of the original Council and Vice- 
president. He also served as President of the 
Paleontological Society in 1929. 

He was married and had two daughters 
and a son. He died on 24 September 1930, 
following an illness of several months 
(Source: Gregory, 1930a, 19306, 1931). 


Glover Morrill Allen: 1927-1929 


Glover M. Allen (Fig. 1), son of Reverend 
Nathaniel Allen and Harriet Ann (Schouler) 


Allen, was born on 8 February 1879, in 
Walpole, New Hampshire. He developed a 
keen interest in natural history at an early 
age and by the time he was in high school 
had become an expert in bird identification 
and an authority on local mammals. He at- 
tended Harvard College on a John Harvard 
Scholarship, was elected to Phi Beta Kappa 
in his junior year, and graduated magna cum 
laude in 1901. He remained at Harvard for 
graduate studies, receiving an A.M. in 1903 
and a Ph.D. in 1904. His doctoral thesis 
was on the heredity of pelage color in mice. 
In addition to scientific subjects, he studied 
several foreign languages and acquired broad 
knowledge of classical European and Rus- 
sian literature. He was married in 1911 to 
Sarah Moody Cushing, and they had one 
daughter, Elizabeth Cushing Allen (Mrs. 
Arthur Gilman). 

Upon receiving his doctorate, he was ap- 
pointed Secretary, Librarian, and Editor of 
the Boston Society of Natural History. He 
returned to the Harvard Graduate School 
in 1906 and 1907 and in the latter year be- 
gan work on the mammal collections of the 
Museum of Comparative Zoology. In 1924, 
he became Lecturer in Zoology at Harvard 
and Curator of Mammals in the Museum 
of Comparative Zoology, where he re- 
mained for the remainder of his career. 

His research involved both mammals and 
birds. He had a keen interest in the fauna 
of New England and also conducted re- 
search in the Bahamas, Labrador, Africa, 
West Indies, Brazil, and Australia. AI- 
though small and slight of build, he had 
unusual stamina and capacity for work when 
in the field. His mammal research was pri- 
marily concerned with taxonomy and dis- 
tribution, and he also published a number 
of papers on fossil sirenians, cetaceans, and 
bats. Among his major contributions were 
the books Bats, Checklist of African Mam- 
mals, Mammals of China and Mongolia, 
and Extinct and Vanishing Mammals of the 
Western Hemisphere with Marine Species 
of all Oceans. His ornithological work in- 
cluded The Birds of Massachusetts coau- 
thored with R. H. Howe, Jr., and Birds and 
Their Attributes. He also published numer- 


PRESIDENTS 33 


ous distributional records and regional 
checklists of birds and was a prolific re- 
viewer of ornithological works. 

He was a charter and life member of the 
ASM and, in addition to the presidency, was 
Vice-president, a Director, and member of 
a number of standing committees. The Life 
Histories and Ecology, Conservation of Land 
Mammals, and Nomenclature committees 
were established during his presidency. He 
was a Fellow of the American Ornitholo- 
gists’ Union and Editor of the Auk and also 
served as Editor of the American Naturalist 
and Secretary and President of the Nuttall 
Ornithological Club. 

Glover Allen was known for his modest 
nature, kindly presence, diplomacy, and ac- 
cessibility to all who wished his advice or 
help. Although not given to “hearty ca- 
maraderie,” as one of his friends put it, when 
encouraged he would greatly entertain lis- 
teners with whimsical and humorous tales 
of his travels, often enhancing his accounts 
with appropriate quotes drawn from his vast 
knowledge of literature. W. M. Tyler (1943) 
cited an example of Allen’s ability to come 
up with a quote from the classics to fit the 
occasion. After they had gone to bed in a 
hotel on Cape Cod after a day in the field, 
someone in the room above tramped heavi- 
ly across the floor. Allen, nearly asleep, mut- 
tered: ‘““The Wild Ass stamps o’er his Head, 
but cannot break his Sleep.”’ Glover Allen 
died on 14 February 1942 in Cambridge, 
Massachusetts (Sources: Barbour et al., 

1943; Tyler, 1943). 


Witmer Stone: 1929-1931 


Witmer Stone (Fig. 1) was born in Phil- 
adelphia, Pennsylvania, on 22 September 
1866. His parents were Frederick D. Stone 
and Anne E. Witmer. He developed an in- 
terest in natural history at an early age and 
as a small boy was a regular visitor to the 
Academy of Natural Sciences of Philadel- 
phia, where he was later to spend his entire 
career. While a student at the Germantown 
Academy in 1877, he and several school- 
mates founded the Wilson Natural Science 


Association. Regular meetings were held at 
which formal papers were presented, and 
scientific collections were maintained. In- 
cluded among the mammals were speci- 
mens he collected during summers spent at 
his uncle’s home in Chester County, Penn- 
sylvania. He was married to Lillie May Laf- 
ferty in 1904. 

He received A.B. and A.M. degrees from 
the University of Pennsylvania in 1887 and 
1891, respectively. His first position follow- 
ing graduation was that of assistant in the 
library of the Historical Society of Penn- 
sylvania, where his father was librarian. In 
1888, he became affliated with the Acad- 
emy of Natural Sciences of Philadelphia 
where he served in many capacities until his 
death on 23 May 1939. He was Conservator 
of the Ornithological Section (1891-1918); 
Assistant Curator (1893-1908) and Curator 
(1908-1918) of the Museum; Executive Cu- 
rator (1918-1925); Director (1925-1929); 
Emeritus Director (1929-1939); Curator of 
Vertebrates (1918-1936); Honorary Cura- 
tor of Birds (1938-1939); and Vice-presi- 
dent of the Academy (1927-1939). 

Although Stone authored 19 publications 
on mammals, he was primarily an orni- 
thologist. Reflecting his broad interest in 
natural history, he also conducted research 
on plants, reptiles, amphibians, insects, and 
land molluscs. He published two books on 
mammals: American Animals coauthored 
with W. E. Cram and The mammals of New 
Jersey. His other mammal publications in- 
cluded descriptions of several new taxa; re- 
ports on collections from Alaska, Sumatra, 
western United States, and Ecuador; and 
studies of the Hawaiian rat and pumas in 
western United States. One of his best known 
ornithological works is Bird Studies of Old 
Cape May, which earned him comparison 
with Thoreau and Burroughs as a writer. A 
major botanical contribution was The Plants 
of Southern New Jersey with Especial Ref- 
erence to the Flora of the Pine Barrens. 

One of his major accomplishments as cu- 
rator of the bird and mammal collections 
at the Philadelphia Academy was rescuing 
many valuable historic specimens that had 
been exposed to moisture, mold, and insects 


34 LAYNE AND HOFFMANN 


and the dust and grime of the city while on 
exhibit. He also performed the monumental 
task of salvaging and rehabilitating E. D. 
Cope’s large collection of reptiles, which 
came to the Academy after Cope’s death. 
The state of preservation of many of the 
valuable specimens was questionable and 
the alcohol had to be poured off carefully 
before the condition of the specimens could 
be determined. J. A. Rhen, who assisted 
him in the task, wrote that “‘the tedium of 
this work was greatly enlivened by Stone’s 
vivid classification and nomenclature of the 
various color shades and consistencies re- 
ferred to as ‘gorum,’ ‘gee,’ and ‘goo,’ to be 
found in the five-gallon glass jars used to 
receive the discarded solution.” 

Witmer Stone was a Charter Member of 
the ASM. He was a member of the original 
Council and served as Vice-president prior 
to assuming the presidency. Two important 
standing committees established during his 
tenure as President were the Editorial and 
Membership committees. He also was an 
active member of the American Ornithol- 
ogists’ Union, serving as Editor of the Auk 
from 1912 to 1937. Among honors he re- 
ceived was an Honorary Sc.D. and the 
Alumni Award of Merit from the Univer- 
sity of Pennsylvania (Source: Huber, 1940). 


Marcus Ward Lyon, Jr.: 1931-1932 


Marcus Ward Lyon, Jr. (Fig. 1), was born 
at Rock Island Arsenal, in Illinois, to Cap- 
tain Lyon and his wife on 5 February 1875. 
Little appears to be known of his early life, 
which was spent at Army posts in various 
parts of the country. One of these was Wa- 
tertown Arsenal near Boston, Massachu- 
setts. His scientific interests apparently stem 
from his childhood days there when he be- 
gan to make collections of insects and other 
animals. Later, his father apparently was 
again posted to Rock Island, because Lyon 
graduated from high school there in 1893 
and entered Brown University that same 
year, receiving his bachelor’s degree in 1897. 
His college training in biology led to his 


being offered an instructorship in bacteri- 
ology at North Carolina Medical College in 
1897. After serving in that post for a year, 
he moved to Washington, D.C., where he 
was appointed an Aid in the Division of 
Mammals, U.S. National Museum (USNM), 
Smithsonian Institution. Concurrent with 
this part-time position, he began graduate 
studies at George Washington University, 
obtaining his M.S. degree in 1900 and his 
M.D. in 1902. In that same year he married 
Martha Maria Brewer of Lanham, Mary- 
land. Lyon continued to work in the Na- 
tional Museum, but embarked upon a par- 
allel teaching career in the Howard 
University Medical School in Washington. 
He taught physiology, bacteriology, and pa- 
thology there until 1917. With the outbreak 
of World War I, he joined the U.S. Army 
and served as pathologist in Walter Reed 
Army Hospital from 1917 to 1919, attain- 
ing the rank of Major. At the same time, he 
taught veterinary zoology and parasitology 
at the Medical School of George Washing- 
ton University. During that 18-year stretch 
of medical teaching and practice, his wife 
also obtained an M.D. from Howard Uni- 
versity, and in 1919 they jointly accepted 
an invitation to join the staff of the South 
Bend Clinic in Indiana. This decision re- 
sulted in a major change of direction for 
Lyon. Previously while associated with the 
Division of Mammals at USNM, he had 
published a series of significant papers on 
the morphology, systematics, and zooge- 
ography of wild mammals. Most notable 
among these are his paper on the classifi- 
cation of the hares and their allies (1904) 
and an account of the mammalian family 
Tupaitidae (1913), for which he was awarded 
a doctorate by George Washington Univer- 
sity. Although his formal relationship with 
the USNM ended in 1912, he continued to 
publish broadly in mammalogy until his 
move to Indiana. In addition, he published 
a number of basic medical studies during 
that period. 

After he and his wife set up their medical 
practice in South Bend, Indiana, his scien- 
tific contributions were almost all devoted 


PRESIDENTS 35 


to Indiana subjects, focusing particularly on 
the region around South Bend. His medical 
publications also drew from his practice 
more frequently than during his time in 
Washington. Perhaps the most significant 
publication from this period is his book, 
Mammals of Indiana, published in 1936. In 
this last period of his life, he became an 
ardent conservationist and spokesman for 
wildlife protection. His last paper was in 
press when he died on 19 May 1942; it de- 
scribed the changes, mostly negative, that 
had occurred in the Kankakee Region along 
the Indiana border near his home as a result 
of human activities (Source: Anon., 1942). 


Vernon Orlando Bailey: 1933-1934 


Vernon Bailey (Fig. 1) was born of pio- 
neer parents, the fourth child of Hiram and 
Emily Bailey, on 21 June 1864 in Man- 
chester, Michigan. His father had learned 
the mason’s trade, but was by preference a 
woodsman and hunter, and when Vernon 
was about 6 years old the family moved west 
to Elk River, Minnesota, on the western 
frontier. This move was accomplished in a 
horse-drawn wagon and must have taken 
some months to cover the 700 miles. The 
only opportunity for schooling in a frontier 
homestead such as his parents established 
was at home, but late in 1873 the families 
of the adjacent homesteads built a school- 
house and formal coursework began. Like 
most early mammalogists, Bailey began by 
collecting the organisms in his surround- 
ings. Self-taught in taxidermy, he began to 
prepare museum specimens, which he sold 
to firms in Ontario, Canada, and in Halle, 
Germany. Some of these specimens were in 
turn purchased by C. Hart Merriam, leading 
him to contact Bailey who was then 19. This 
was prior to Merriam’s being named to his 
government position, eventually in the Bu- 
reau of Biological Survey, and their lifelong 
association gained Bailey entreé into the Bu- 
reau. In 1887 Bailey was appointed asa field 
naturalist and sent to the northern Great 
Plains and Rocky Mountains. For virtually 


every year thereafter, until his final trip to 
Nevada in 1937, he collected for the Bureau 
and for the U.S. National Museum. How- 
ever, he found time to take course work at 
the University of Michigan in 1893 and at 
George Washington University in 1894- 
1895. 

He retired from the Biological Survey in 
1933, having gained the rank of Chief Field 
Naturalist, but continued to work until his 
death in Washington on 20 April 1942. He 
was survived by his wife, Florence Merriam 
Bailey, herselfa biologist, whom he married 
in 1904. In addition to the presidency of the 
American Society of Mammalogists, he 
served as President of the Biological Society 
of Washington (Sources: Smithsonian In- 
stitution Archives, Record Unit 7098; 
Zahniser, 1942). 


Harold Elmer Anthony: 1935-1937 


Harold E. Anthony (Fig. 1) was born in 
Beaverton, Oregon, on 5 April 1890. His 
father was a well-known Pacific Coast or- 
nithologist and collector. From an early age, 
he hunted and trapped and loved the out- 
doors and, although his primary field came 
to be mammalogy, he retained a broad in- 
terest in natural history throughout his life. 
He was married in 1916 to Edith Demerell, 
who died shortly after their son, Alfred 
Webster Anthony, was born. Four years lat- 
er he married Margaret Feldt, and they had 
a daughter, Margery Stuart, and a son, Gil- 
bert Chase. He was an officer (1st Lieuten- 
ant and Captain) in the field artillery during 
World War I (1917-1919) and saw action 
in France. 

He attended Pacific University for 2 years 
(1910-1911) and received B.S. and M.A. 
degrees from Columbia University in 1915 
and 1920, respectively. 

He began his career as a field collector for 
the Biological Survey in 1910 and in the 
same year was employed by The American 
Museum of Natural History as naturalist on 
the Albatross Expedition to Lower Califor- 
nia. The following year he joined the Mu- 


36 LAYNE AND HOFFMANN 


Joseph Grinnell Hartley H. T. Jackson Walter P. Taylor 
(1937-1938) (1938-1940) (1940-1942) 


oe 


{ 


A. Brazier Howell E. Raymond Hall Edward A. Goldman 
(1942-1944) (1944-1946) (1946-1947) 


Remington Kellogg Tracy I. Storer William J. Hamilton, Jr. 
(1947-1949) (1949-1951) (1951-1953) 


Fic. 2.—Presidents of the ASM from 1937 to 1953. 


PRESIDENTS oF 


seum staff full-time as a cataloger and gen- 
eral handyman in the Department of 
Mammals and Ornithology. He was ap- 
pointed Associate Curator in the Depart- 
ment of Mammalogy in 1919, Curator in 
1926, and Emeritus Curator upon his re- 
tirement in 1958. In addition to serving as 
Chairman of the Department of Mammal- 
ogy from 1942 to 1958, he held the posts 
of Dean of the Scientific Staff (1942-1948) 
and Deputy Director (1952-1957) of the 
Museum. After retirement, he was Appoint- 
ed Curator of the Frick Laboratory, a pa- 
leontological research laboratory at the Mu- 
seum supported by the Charles Frick 
Foundation, and served in that capacity un- 
til 1966. 

Anthony’s research involved both Recent 
and fossil mammals, with an emphasis on 
the Caribbean and Central and South Amer- 
ican regions. In addition to his work in the 
Neotropics, he participated in expeditions 
to various regions of western United States, 
Alaska and the Arctic Ocean, Canada, Af- 
rica, and Burma. Among his major contri- 
butions were the two volume Mammals of 
Puerto Rico, Living and Extinct and Field 
Book of North American Mammals, which 
for many years was the major guide to mam- 
mals of the region. He was active in the 
Museum’s exhibition program, playing a key 
role in the creation of the Hall of North 
American Mammals, the Akeley Hall of Af- 
rican Mammals, and the Hall of South Asi- 
atic Mammals. An ardent conservationist, 
he served as Chairman of the Committee 
on Preservation of Natural Conditions of 
the National Research Council’s Division 
of Biology and Agriculture. 

Anthony was a Charter Member of the 
ASM. Besides the presidency, he served as 
a Councillor, Trustee, and Vice-president. 
He also was a director of both the New York 
Explorers Club and National Audubon So- 
ciety, Treasurer of the New York Academy 
of Sciences, and an Honorary Life Member 
of the Sociedad Colombiana de Ciencias 
Naturales. 

In addition to his scientific interests, An- 
thony was a financial expert. As was once 


stated in an article in an American Museum 
employee newsletter “he knew that a bear 
market wasn’t always a place where grizzlies 
and kodiaks are sold, and that there are two 
kinds of bulls.” His financial expertise made 
him a particularly valuable member of the 
Museum’s Pension Board and Welfare 
Committee. 

As a youth, he discovered the pleasure 
and satisfaction of growing plants and this 
became a lifetime avocation. Orchids were 
his specialty. He served as President of the 
Greater New York Orchid Society and 
Treasurer of the American Orchid Society, 
from which he received a gold medal in rec- 
ognition of his contributions. Cooking was 
another of his long-time interests, and his 
culinary skills were attested to by his in- 
duction into the Society of Amateur Chefs. 

He died of a heart attack on 29 March 
1970, while on a family outing in Paradise, 
California (Sources: Anon., 1958a, 19585, 
1970). 


Joseph Grinnell: 1937-1938 


Joseph Grinnell (Fig. 2) was born on 27 
February 1877, at Ft. Sill (then Indian Ter- 
ritory) in what is now Oklahoma. His family 
was of New England origin, but his father, 
a physician, moved the family to California 
when Grinnell was still young. Joseph’s 
schooling through college was in Pasadena. 
He attended Pasadena High School and then 
enrolled in what was known as Throop 
Polytechnic Institute (now the California 
Institute of Technology) where he received 
a bachelor’s degree in 1897. He began his 
graduate studies at Stanford University 
shortly thereafter, receiving his M.A. degree 
in 1901. Even as a high school student he 
had displayed an interest in natural history 
and had begun to amass a collection of ver- 
tebrates. In 1896, while only 19 years old, 
he made his first visit to Alaska, where he 
collected around Sitka. Two years later he 
returned to Kotzebue and the Bering Sea 
Region where he not only collected verte- 
brates but also apparently prospected for 


38 LAYNE AND HOFFMANN 


gold. An apocryphal tale suggests that he 
found a rich claim but was robbed of it by 
claim jumpers; however, this cannot be sub- 
stantiated. Between these early expeditions, 
he served as instructor at Throop Polytech, 
teaching assistant at Stanford, and instruc- 
tor in the Palo Alto High School. He re- 
ceived an appointment at the University of 
California, Berkeley, in 1905, and almost 
all of his subsequent field collecting was car- 
ried out within the state of California. 
Shortly after joining the Berkeley faculty, 
however, he returned to coastal Alaska in 
1907 on an expedition headed by Annie M. 
Alexander, who became his life-long bene- 
factor. In 1908 she founded the California 
Museum of Vertebrate Zoology at the Uni- 
versity of California, Berkeley, of which 
Grinnell was named Director. Together with 
Louise Kellogg, Alexander supported the 
Museum and Grinnell until his death at age 
63 on 29 May 1939. During those 31 years 
as Director of the Museum of Vertebrate 
Zoology, Grinnell developed a highly or- 
ganized approach to field collecting, which 
has had an influence far beyond the state of 
California, to which he restricted not only 
his own efforts, but if possible, those of his 
students. Most of his many publications 
were devoted to birds, but 76 treat wholly 
or in part of mammals. 

In addition to his systematic and ecolog- 
ical work, he played a significant role in the 
developing field of conservation. His im- 
pact on teaching biology at Berkeley was 
profound, as is suggested by the fact that 15 
years after his death his principal course 
“Zoology 113°. and his graduate seminar 
“Vertebrate Review” were still essentially 
Grinnellian (Source: Hall, 1939). 


Hartley Harrad Thompson Jackson: 
1938-1939 


Although Hartley H. T. Jackson (Fig. 2) 
was only the eleventh president of the ASM, 
he was one of those Biological Survey sci- 
entists who first developed the idea of such 
a society, and he chaired the first Organizing 


Committee. He served first as Correspond- 
ing Secretary (1919-1925), was elected Vice 
President in 1937, and in addition held a 
number of committee posts. Hartley Jack- 
son was born in Milton, Wisconsin, on 19 
May 1881, the son of English immigrants 
to the United States. He was the last of their 
eight children and the only one born in this 
country. Like so many other field biologists, 
he began when still young to collect birds, 
and his first scientific paper on screech owls 
appeared when he was 16 years old. Jackson 
attended primary and secondary schools in 
Milton, and then enrolled in Milton College, 
where he received his bachelor’s degree in 
1904. Upon graduating, he taught at Car- 
thage Collegiate Institute in Missouri, where 
he met Anna Marcia Adams who he mar- 
ried in 1910, having already entered the 
University of Wisconsin 2 years earlier for 
graduate work. His master’s degree was 
awarded in 1909, and the following year he 
joined the Bureau of Biological Survey in 
Washington. He also enrolled in George 
Washington University, and attained a doc- 
toral degree in Zoology in 1914. 

In 1917, E. W. Nelson, Chief of the Sur- 
vey, arranged with the State of Wisconsin 
for a cooperative study of the fauna. Jackson 
was designated principal investigator from 
the Biological Survey, with the state sup- 
plying a field assistant and other support. 
Jackson had, even prior to this formal agree- 
ment, carried out field work in Wisconsin, 
but thereafter field work was conducted reg- 
ularly each summer by a team directed by 
Jackson until 1922 when the agreement be- 
came inactive. It was, however, reactivated 
in 1940, and eventually led to one of Jack- 
son’s most important works, The Mammals 
of Wisconsin, published in 1954. 

Increasing administrative duties cur- 
tailed Jackson’s field research, and he be- 
came more involved in wildlife manage- 
ment as Chief of the Division of Wildlife 
Research, later renamed Wildlife Surveys. 
This unit sponsored a great many important 
studies of game birds and mammals in the 
period just prior to World War II, during 
which Jackson served on several War Pro- 


PRESIDENTS Sy) 


duction Board committees. After a 41-year 
period of government service, Hartley Jack- 
son retired in 1951. He continued to utilize 
his office in the National Museum of Nat- 
ural History after retirement, but worked 
primarily on a history of the Bureau of Bi- 
ological Survey, which apparently was nev- 
er published. His wife Anna died in 1968, 
but 2 years later he married Mrs. Stephanie 
Hall of Durham, North Carolina, whose fa- 
ther was the former president of Milton Col- 
lege in Wisconsin. He died at age 95 in Dur- 
ham (Source: Aldrich, 1977). 


Walter Penn Taylor: 1940-1942 


Walter P. Taylor (Fig. 2) was born 31 Oc- 
tober 1888 near Elkhorn, Wisconsin, to 
Benton Ben and Helen West Taylor. No in- 
formation could be found concerning the 
family or Taylor’s childhood and early ed- 
ucation. That the family had moved by the 
time he reached his teens can be inferred 
from the fact that he received his secondary 
education from Throop Polytechnic Insti- 
tute in Pasadena, California, between 1902 
and 1908. This was the same school at- 
tended by Joseph Grinnell a few years pre- 
viously. He then spent one semester at Stan- 
ford University before transferring to the 
University of California at Berkeley, where 
he received a bachelor’s degree in 1911. He 
continued on at Berkeley in graduate school, 
marrying Mary E. Fairchild in 1912, and 
completing his doctorate in zoology in 1914. 
Both at Throop and at the University of 
California he was employed while a student. 

His first post-doctoral appointment was 
as Assistant Curator and then Curator of 
Mammals at the University of California 
Museum of Vertebrate Zoology under its 
director, Joseph Grinnell. In 1916, as so 
many of his colleagues had, he joined the 
U.S. Biological Survey first as Assistant and 
subsequently Senior Biologist. He remained 
full time with the Survey until 1932, when 
he joined the faculty of the University of 
Arizona under a cooperative arrangement 
with the Survey. From 1935 to 1947 he oc- 


cupied a comparable position at Texas A&M 
College; during this time, the Biological Sur- 
vey was transformed to the U.S. Fish and 
Wildlife Service, and he headed one of the 
first Cooperative Wildlife Research Units 
within the Service. He then transferred to 
Oklahoma State University in Stillwater 
(then the Agricultural and Mechanical Col- 
lege) where he served as Wildlife Research 
Unit Leader until 1951, when he retired from 
federal service. In 1954 he was appointed 
Professor of Conservation Education and 
Biology at the Claremont Graduate School 
of the Claremont Colleges group in southern 
California. During this time he also taught 
at LaVerne College, Murray State College, 
and Southern Illinois University. He retired 
from this position in 1962, but entered on 
a second career in politics, serving on the 
City Council and as Vice-mayor of Clare- 
mont. 

He was the recipient of many honors, in- 
cluding the Distinguished Service Medal of 
the Department of the Interior, and the Le- 
opold Award of The Wildlife Society. He 
was President of The Wildlife Society, Eco- 
logical Society of America, and Texas Acad- 
emy of Sciences, as well as of the ASM. 

Although Taylor held a number of dif- 
ferent appointments in the course of a long 
career, his principal focus after he joined 
the Biological Survey in 1916 was on what 
would now be called wildlife biology. He 
was a prolific writer, authoring about 300 
scientific and technical papers and pam- 
phlets, and was co-author or editor of sev- 
eral books, including The Birds of the State 
of Washington (1953) and Deer of North 
America (1956). 

He died on 29 March 1972, and was sur- 
vived by his wife, Clara, two sons, and two 
daughters, one of whom, Elizabeth, married 
Randolph Peterson (Sources: Cottam, n.d.; 
Lehmann, 1972; E. Peterson, pers. comm.). 


Alfred Brazier Howell: 1942-1944 


A. Brazier Howell (Fig. 2) was born on 
28 July 1886 in Catonsville, Maryland. His 


40 LAYNE AND HOFFMANN 


parents were Darius Carpenter and Kath- 
erine Hyatt Howell. As a youngster, Howell 
became interested in birds and egg collect- 
ing. At age 13 his mother gave him a bird 
book by William E. D. Scott inscribed “A. 
Brazier Howell from Mother,” which may 
have been the reason he later dropped his 
first name from most of his publications. 
He married Margaret Gray Sherk in 1914, 
and they had three daughters and a son. His 
wife enjoyed the out-of-doors and frequent- 
ly accompanied him in the field. He died 
on 23 December 1961 at his home in Ban- 
gor, Maine. 

Howell’s formal college education was 
limited to a year at Yale after graduation 
from the Hill Preparatory Boys School in 
1905. In 1908 he and his mother moved to 
Pasadena, California. There he developed a 
serious interest in research, which began with 
a study of the birds of the Channel Islands 
off the southern California coast. In 1911, 
having sufficient financial means, he pur- 
chased a home and small orange grove in 
Covina, California, where he housed his ex- 
panding collections and library. He could 
afford to spend considerable time in the field, 
and from time to time he employed collec- 
tors, among whom were A. J. Van Rossem, 
Chester Lamb, and Laurence Huey. In 1918, 
under the direction of E. W. Nelson, he and 
Luther Little conducted a collecting expe- 
dition in southern Arizona. They were kept 
out of one area by an uprising of Yaqui 
Indians. From 1922 to 1928, the Howells 
lived in Washington, D.C., and Brazier 
worked as a “‘dollar-a-year-man” in the Di- 
vision of Biological Survey with the title of 
Scientific Assistant. In 1928, he accepted a 
position in the Department of Anatomy of 
Johns Hopkins Medical School. He taught 
gross human anatomy, in which he had nev- 
er had a formal course, until his retirement 
in 1943. 

Although Howell is best known for his 
work on mammalian anatomy, his early re- 
search was primarily on the distribution, 
taxonomy, and life histories of birds and 
mammals. His first anatomical paper, ““On 
the alimentary tracts of squirrels with di- 


verse food habits,’ appeared in 1925. His 
best known contribution to mammalian 
anatomy was the volume Anatomy of the 
Woodrat, which appeared as the first mono- 
graph of the ASM in 1926 and remains one 
of the classics in the field. Among his other 
important contributions to mammalogy 
were a revision of the genus Phenacomys 
and the life history of the red tree mouse 
and a revision of the genus Synaptomys 
published in the North American Fauna se- 
ries. 

Howell was a Charter Member of the 
ASM, and, in addition to the presidency, 
was a Director, Corresponding Secretary, 
and member of various committees. He also 
served on the Council for the Conservation 
of Whales and other Marine Mammals or- 
ganized in 1929 under the ASM. A few years 
before his death he provided an endowment 
to the ASM for a graduate student award, 
now designated the A. Brazier Howell 
Graduate Student Honorarium. He also was 
active in the Cooper Ornithological Society, 
serving for some time as an aid to the Busi- 
ness Manager and in managing the endow- 
ment fund. 

Brazier Howell was a talented artist, as 
reflected in his anatomical illustrations, a 
gifted musician, and an accomplished wood 
worker. Among his other interests were re- 
furbishing old cars, stamp collecting, raising 
tropical fish, and collecting antiques. He was 
a quiet, friendly man, but as a result of an 
inherited hard-of-hearing condition tended 
to avoid meetings and large groups of peo- 
ple. One of the Bill Hamilton anecdotes 
concerns A. Brazier Howell. As a graduate 
student, the well-known Cornell anatomist 
and shark expert, Perry Gilbert, was greatly 
impressed by the work of Howell. Thus he 
was delighted when he came to Hamilton’s 
office one day and found him with a man 
Bill introduced as his old friend Brazier 
Howell. After going to great lengths to dis- 
play his knowledge of anatomy and How- 
ell’s research, Gilbert was disappointed that 
Howell remained silent and seemingly un- 
impressed. It was not until later that he 
learned that “Brazier Howell” was a local 


PRESIDENTS 41 


farmer who had come to ask Hamilton how 
to get rid of some mammal pest (Source: 
Little, 1968). 


Eugene Raymond Hall: 1944-1946 


E. Raymond Hall (Fig. 2) (students and 
colleagues never called him Eugene) was 
born on 11 May 1902, in the small town of 
Imes, in eastern Kansas, a town that no lon- 
ger appears on most maps. He grew up on 
the family farm in nearby Le Loup and spent 
his boyhood helping in farming activities 
and in fur trapping. After an initial educa- 
tion in rural schools, he spent his final year 
of high school in Lawrence, Kansas, and 
then enrolled in the University of Kansas. 
His first scientific publication, ““The First 
Record of a Golden-Winged Warbler from 
Kansas,” was published in 1921 while he 
was still an undergraduate majoring in zo- 
ology. During his KU years, he was influ- 
enced by Remington Kellogg, who, 10 years 
his senior, had graduated from the univer- 
sity and was then working in the Bureau of 
Biological Survey in Washington. Kellogg 
urged him to enroll in graduate studies at 
the University of California at Berkeley, as 
Kellogg had. Hall did so, first marrying Mary 
Harkey, also a University of Kansas un- 
dergraduate. At Berkeley, he worked under 
the direction of Joseph Grinnell, who was 
to become president of ASM and was Di- 
rector of the Museum of Vertebrate Zool- 
ogy. In 1927, still a year short of earning 
his Ph.D., he became Curator of Mammals 
in the museum. During the next decade, 
expanding beyond Grinnell’s preoccupation 
with California, Hall carried out intensive 
field work on mammals in Nevada. This led 
to what many regard as his most notable 
publication, The Mammals of Nevada, in 
1946. In 1938 he became Acting Director 
of the Museum of Vertebrate Zoology upon 
the resignation of the founding director, 
Grinnell, who died the next year. Hall served 
as acting director until 1944; in that year he 
abruptly left Berkeley to return to the Uni- 


versity of Kansas as Chairman of the De- 
partment of Zoology and Director of the 
Museum of Natural History, holding the 
latter position until he retired in 1967. Many 
have speculated that his sudden departure 
from Berkeley was occasioned by the failure 
of the university to name him as Director 
of the Museum of Vertebrate Zoology dur- 
ing the 6 years he served there in an acting 
capacity. 

At Kansas, he took a museum with a 
strong tradition and built it into one of the 
leading research and graduate education 
museums of natural history in the country. 
His own productivity was prodigious, re- 
sulting in an output of 350 publications be- 
fore his death at age 84 in 1986. In addition 
to his other contributions to mammalogy, 
his major work was The Mammals of North 
America, first published in 1959 and revised 
in 1981. 

He attracted a large number of students 
to Kansas, many of whom have gone on to 
make major contributions to the ASM. Hall 
was respected by many, disliked by some, 
and feared by a few. He had an exceptionally 
strong personality, through which he in- 
spired respect and loyalty among his grad- 
uate students. Few ever saw Hall’s human 
side, but for those who did, he was a proud 
father and husband, and loyal friend. “E. 
Raymond Hall was a farmer, trapper, and 
naturalist at heart, and a prodigiously suc- 
cessful scientist by profession. He was a 
uniquely prominent and tremendously in- 
fluential figure in twentieth century mam- 
malogy” (Findley and Jones, 1989) (Sources: 
Findley and Jones, 1989; Jones, 1990). 


Edward Alphonso Goldman: 1946 


Edward A. Goldman (Fig. 2) was born to 
Jacob and Laura Goltman in Mount Car- 
roll, Illinois, in 1873. His parents were 
farmers, and little is known about his child- 
hood, although he was presumably educat- 
ed in rural schools. When he was around 10 
years old, his parents left Illinois for Falls 


42 LAYNE AND HOFFMANN 


City, in eastern Nebraska, driving 300 head 
of cattle seeking “‘greener pastures.”” Two 
signal events marked their short residence 
in Nebraska; Jacob Goltman changed the 
family name to Goldman and a grasshopper 
plague resulted in the family losing most of 
its livestock to starvation. In 1888, the fam- 
ily again resettled, this time in Tulare Coun- 
ty, California. No details concerning his 
schooling in either Nebraska or California 
could be found, but as was the case with 
some other early presidents of the society, 
he was thoroughly self-tutored as a natu- 
ralist. His interest in natural history appears 
to have come from his father, who was him- 
self an amateur student of nature. Goldman 
was taught to shoot a shotgun while on the 
Nebraska ranch and began then to collect 
specimens of birds and mammals, a hobby 
he continued after the move to California. 
At age 17 Goldman left home to accept a 
job as vineyard foreman near Fresno, about 
120 km north of the family ranch at Earli- 
mart in the southern San Joaquin Valley. In 
that same year, there came his fateful meet- 
ing with E. W. Nelson, who had been in 
California participating in the famous Death 
Valley Expedition of the Bureau of Biolog- 
ical Survey. Nelson had been asked by the 
survey director, C. Hart Merriam, to con- 
duct a survey of the southern San Joaquin 
Valley and needed an assistant. He stopped 
at the Goldman ranch for help in repairing 
his wagon, learned of Jacob Goldman’s in- 
terest in natural history, and received the 
suggestion that son Edward might serve as 
a field assistant. From this fortuitous meet- 
ing came the famous collecting team of Nel- 
son and Goldman. 

The first joint expedition was a short one, 
of about 3 months duration, but it was fol- 
lowed by Merriam’s order to collect in west- 
ern Mexico. What was planned as a 3-month 
stay in Mexico lengthened to 4 years, during 
which time Goldman worked his way up 
from the status of temporary field assistant 
to a permanent position in the Biological 
Survey. Together Nelson and Goldman col- 
lected in every state and territory in Mexico, 


obtaining a combined total of nearly 23,000 
mammal specimens by the time of Nelson’s 
death in 1934. In addition to Mexico, Gold- 
man worked in many parts of the United 
States, as well as in Panama, where his re- 
sults were published as Mammals of Pan- 
ama by the Smithsonian in 1920. 

During World War I he entered the U.S. 
Army, attaining the rank of Major in the 
Sanitary Corps in France. After the war, he 
retained his rank in the Sanitary Reserve 
Corps of the U.S. Army Medical Depart- 
ment until 1937. Although the war had in- 
terrupted his work with Nelson, this was 
resumed until Nelson’s retirement in 1929 
terminated the active collaboration. The 
previous year, however, Goldman had been 
relieved of all administrative duties so that 
he could carry on the Mexican work, which 
he did very productively until his retire- 
ment at the end of 1944. He continued to 
work on the ““Mammals of Mexico”? manu- 
script until his untimely death from a heart 
attack on 2 September 1946, which cut short 
his service as President of the American So- 
ciety of Mammalogists. He was survived by 
his widow, Emma May Chase, and three 
sons, Nelson, Orville, and Luther. The latter 
followed his father’s ir terest in natural his- 
tory. 

Edward Goldman was one of the small 
group who had the vision to organize an 
American Society of Mammalogists during 
the years immediately following World War 
I. He published over 200 scientific papers, 
among them classic volumes on the puma 
and gray wolf (Sources: Jackson, 1947; Tay- 
lor, 1947; Young, 1947). 


Arthur Remington Kellogg: 
1946-1949 


Another midwesterner, Remington Kel- 
logg (Fig. 2), was born in Davenport, Iowa, 
on 5 October 1892, the son of Claire and 
Rolla Remington Kellogg. His father was a 
printer by profession and his mother taught 
school. When young Remington was 6 years 


PRESIDENTS 43 


old, his parents moved to Kansas City, Mis- 
souri, where, after grammar school, he com- 
pleted Westport High School, and then en- 
rolled at the University of Kansas in 1910. 
As an undergraduate at the university, he 
was strongly influenced by two men, Charles 
Dean Bunker, who was then Curator of Birds 
and Mammals in the Museum of Natural 
History, and Alexander Wetmore, a Kansan 
who was an upper division student in zo- 
ology and who was to become an eminent 
American ornithologist. While initially in- 
terested in insects, Kellogg shifted his focus 
to marine mammals and paleontology dur- 
ing the course of his undergraduate work. 
After graduation, he enrolled at the Uni- 
versity of California, Berkeley, in 1916. It 
had taken him 6 years to graduate from 
Kansas because of the necessity of working 
to support his college career, but in Cali- 
fornia he was awarded a teaching fellowship 
under Dr. John C. Merriam, who was to be 
another important influence in Kellogg’s life. 
His graduate work was interrupted by World 
War I, and he enlisted in late 1917. Several 
months later he was promoted to sergeant 
and transferred to the Central Medical De- 
partment Laboratory, whose commander 
was Major Edward Goldman and whom he 
succeeded as President of the American So- 
ciety of Mammalogists, serving not only 
Goldman’s unexpired term but a regular 
2-year term subsequently. Receiving a dis- 
charge from the Army in 1919, he returned 
to the University of California to complete 
his residence requirements for the doctoral 
degree and at the end of the fall semester 
was appointed Assistant Biologist in the Bi- 
ological Survey. Later that year he married 
fellow student Marguerite Henrich, and they 
spent their entire married life in Washing- 
ton, D.C., until his death from a heart attack 
at age 77, in 1969. 

Around the same time that Kellogg joined 
the Biological Survey, his former mentor 
John C. Merriam accepted appointment as 
President of the Carnegie Institution in 
Washington. Merriam arranged for Kellogg 
to be made a Research Associate of the In- 


stitution, a position he held from 1921 until 
1943. This arrangement allowed Kellogg to 
receive funding from Carnegie to pursue his 
research on marine mammals at the same 
time that he carried out assigned projects 
for the Biological Survey. This in turn al- 
lowed Kellogg to complete the research nec- 
essary to write his dissertation, which com- 
pleted the requirements for his Ph.D.from 
the University of California in 1928. That 
same year, Kellogg left the Biological Sur- 
vey to fill a position of Assistant Curator of 
Mammals at the U.S. National Museum. 
Under Gerritt S. Miller’s supervision, Kel- 
logg was able to devote more time to marine 
mammals, and he became recognized as the 
American authority. As a result he found 
himselfin 1937 with an appointment by the 
Department of State as U.S. Delegate to the 
International Conference on Whaling, the 
forerunner of the International Whaling 
Commission (IWC). Further appointments 
followed in 1944-1946, and he served as 
Commissioner of the IWC from 1949 until 
1967, being Chairman from 1952 to 1964. 

In 1948, Kellogg was appointed Director 
of the U.S. National Museum, and 10 years 
later, Assistant Secretary for Science of the 
Smithsonian Institution. His heavy admin- 
istrative burdens deprived him of the time 
he was used to spending on research, but he 
still attempted to spend several hours a day 
on his own research projects. As an admin- 
istrator, his tenure in both the museum and 
the Office of the Assistant Secretary were 
characterized by an innate negativism that 
led him to be referred to at times as the 
‘abominable no man’’ (Source: Setzer, 
POT): 


Tracy Irvin Storer: 1949-1951 


Like many presidents of the ASM, Tracy 
I. Storer (Fig. 2) had a wide array of interests 
beyond the study of mammals. He was also 
well known as an ornithologist and herpe- 
tologist, published in the field of wildlife 


44 LAYNE AND HOFFMANN 


management and animal control, and au- 
thored the most successful general zoology 
text of its time. He was born in San Fran- 
cisco, California, on 17 August 1889 and 
grew up in Elmhurst, south of Oakland. He 
and his brother were raised by their father, 
his mother having died when he was 9 years 
old. Tracy went to local public schools, and 
in 1908 at the age of 18 entered the Uni- 
versity of California at Berkeley. He re- 
ceived his bachelor’s degree in zoology with 
honors in 1912 and his master’s degree the 
next year. He worked his way through col- 
lege as a printer, primarily of handbills and 
cards, and his well known frugality was un- 
doubtedly instilled by the often straitened 
financial circumstances of his motherless 
boyhood. Upon completion of his master’s 
degree he was hired as an assistant by Charles 
Koford in Berkeley’s Department of Zool- 
ogy but transferred the next year to the staff 
of the Museum of Vertebrate Zoology, es- 
tablished five years previously and directed 
by Joseph Grinnell. With the financial sup- 
port of Annie M. Alexander, Grinnell had 
mapped out several ambitious projects. 
Storer’s first assignment was to work with 
Grinnell and Harold Bryant on a project 
that resulted in the publication of The Game 
Birds of California in 1918. While this was 
going on, Storer, along with Walter P. Tay- 
lor and a number of others, began field work 
on an ambitious transect survey across the 
Sierra Nevada in the region of Yosemite 
National Park. This field work was inter- 
rupted by World War I, during which Storer 
served in the U.S. Army Sanitary Corps in 
Texas; his commanding officer was his for- 
mer employer, Koford. Upon his return 
from the Army he resumed the Yosemite 
survey, completing field work in 1920 and 
then drafting the report, which he and Grin- 
nell completed as Animal Life in the Yo- 
semite in 1924. 

Storer had married Ruth Risdon just be- 
fore his Army tour. His wife, one of the first 
women graduates of the University of Cal- 
ifornia Medical School, suffered from tu- 
berculosis during the first years of their mar- 


riage, which further strengthened Storer’s 
habits of frugality. In order to fund field 
work for his doctoral dissertation, Storer 
accumulated vacation time for 6 years, and 
his doctoral dissertation on Amphibia of 
California earned him a Ph.D. in 1924, just 
as he and Grinnell completed the Yosemite 
work. 

With these two milestones achieved, 
Storer accepted a position at the University 
of California at Davis as the first member 
of its new Division of Zoology. The ap- 
pointment was not only as Assistant Pro- 
fessor but also as Assistant Zoologist in the 
Experiment Station, and his research hence- 
forth focused on control and manipulation 
of vertebrate population densities— wildlife 
management and pest control. After a de- 
cade, undergraduate enrollment in zoology 
at Davis had increased so much that addi- 
tional faculty could be added to his one- 
person division. When a program in wildlife 
management that he had sought for the ex- 
panding division was instead awarded to the 
Berkeley campus, Storer turned his atten- 
tion to undergraduate teaching and wrote a 
text first published in 1943 with the title 
General Zoology. The book, with its sys- 
tematic organization, profuse illustration, 
and large information content, quickly came 
to dominate the freshman zoology market, 
and made Storer relatively wealthy. At the 
same time, the book and its many subse- 
quent editions and associated teaching aids 
came to usurp much of his time. He nev- 
ertheless continued to produce a steady flow 
of short papers and reviews, as well as sev- 
eral major monographs, most notably those 
on the California grizzly in 1955 and on 
Pacific Island rat ecology in 1963. 

The Storers were childless. While he was 
frugal, Tracy Storer was generous. In ad- 
dition to gifts to U.C. Davis, he also re- 
membered the ASM in his will with a large 
bequest, which has become a major portion 
of the trust funds administered by the so- 
ciety. He died from a heart attack on 25 
June 1973 at age 84 (Source: Salt and Rudd, 
L975); 


PRESIDENTS 45 


William John Hamilton, Jr.: 
1951-1953 


William J. Hamilton, Jr. (Fig. 2), was born 
on 11 December 1902 in Corona, New York, 
the son of William J. Hamilton and Char- 
lotte Richardson Hamilton. Bill, as he was 
known to all, was a quintessential naturalist, 
whose interest in natural history stemmed 
from the experience of caring for a plant he 
received as a gift when he was 7 years old. 
Gardening and horticulture remained a ma- 
jor avocation throughout his life. Bill met 
his future wife, Nellie Rightmyer, when she 
took a class in which he was an instructor. 
They were married in 1928 and had three 
children, Ruth, June, and William J. III, 
who also is a well-known zoologist. Bill was 
commissioned a captain in the U.S. Army 
Medical Corps in 1942 and worked on ro- 
dent and typhus control problems and as 
one of the military governors of Manheim, 
Germany, after the war. He was discharged 
in 1945 with the rank of Major. 

He received all of his degrees from Cor- 
nell University, including a B.S. in 1926; 
M.S. in entomology in 1928; and a Ph.D. 
in vertebrate zoology in 1930. His major 
professor for the doctorate was the herpe- 
tologist Albert H. Wright, and his doctoral 
thesis was on the life history of the star- 
nosed mole. 

In 1930, he was appointed Instructor in 
Zoology in the New York State College of 
Agriculture at Cornell, where he remained 
for his entire career. He became Assistant 
Professor in 1937, Associate Professor in 
1942, Professor in 1947, and Professor 
Emeritus upon his retirement in 1963. At 
various times he was a member of the de- 
partments of Entomology, Zoology, and 
Conservation (now Natural Resources). He 
taught vertebrate zoology, mammalogy, 
herpetology, literature of vertebrate zoolo- 
gy, economic zoology, and conservation and 
served as major professor of over 60 grad- 
uate students in mammalogy and herpetol- 
ogy. 


Bill’s principal research interests were the 
ecology and life history of mammals. Much 
of his work was done in New York, reflect- 
ing his belief that one did not have to go to 
far off places to find interesting and signif- 
icant problems to study. Among his major 
contributions were studies on microtine cy- 
cles, food habits of mammals and other ver- 
tebrates, and various aspects of reproduc- 
tive biology. He also published life history 
accounts for a substantial proportion of 
eastern United States mammals. In addi- 
tion to over 200 papers, he authored Amer- 
ican Mammals, the first textbook in mam- 
malogy, and Mammals of Eastern United 
States and was coauthor of Conservation in 
the United States. 

His professional honors included election 
as an Honorary Member of the ASM and 
as a Fellow of the American Association for 
the Advancement of Science, New York 
Academy of Sciences, and the Royal Hor- 
ticultural Society of England. He also was 
the recipient of the LePiniec Award from 
the American Rock Garden Society and the 
Outstanding Alumni Award from Cornell 
University. 

He joined the American Society of Mam- 
malogists in 1924. In addition to the pres- 
idency, he was Vice-president, a Director, 
and a member of numerous committees. He 
was also Secretary and President of the Eco- 
logical Society of America and Zoological 
Editor of Ecological Monographs. He served 
on the Environmental Biology Panel of the 
National Science Foundation and as Chair- 
man of the Scientific Advisory Committee 
of the E. N. Huyck Preserve and for many 
years was a Research Associate in the De- 
partment of Mammalogy of the American 
Museum of Natural History. 

Bill Hamilton was one of the most col- 
orful of the mammal society presidents. His 
sense of humor, which earned him the title 
“Wild Bill,’ was legendary. He was able to 
weave the most outlandish tall tales into a 
conversation with such apparent sincerity 
that the listener often did not realize he was 
joking. He died at his home in Ithaca, New 


46 LAYNE AND HOFFMANN 


York, on 27 July 1990 (Source: Layne and 
Whitaker, 1992). 


William Henry Burt: 1953-1955 


Like several other presidents of the ASM, 
William H. Burt (Fig. 3) was born and raised 
in Kansas. He was born on 22 January 1903 
in Haddam, near the border with eastern 
Nebraska. His parents were Frank and Hat- 
tie Burt, and no references to siblings have 
been found. He grew up on the Burt farm, 
but was reticent to talk about his early years. 
He did, however, comment once that his 
observations of prairie dogs on the family 
farm were the basis of his later thoughts on 
territoriality and home range in mammals, 
the field in which he made a singular con- 
tribution to biology. He attended the Uni- 
versity of Kansas, graduating in 1926; he 
was thus an undergraduate together with E. 
Raymond Hall. He completed a master’s 
degree at Kansas in its Museum of Natural 
History in 1927, after which, like Hall, he 
enrolled in graduate school at the Univer- 
sity of California, Berkeley. He began grad- 
uate work in paleontology, even though his 
earlier interest had been in ornithology and 
mammalogy. In 1928 and 1929, he was 
awarded a research fellowship at the Cali- 
fornia Institute of Technology, which al- 
lowed him to complete his doctoral disser- 
tation on the morphology and evolution of 
woodpeckers, and he was awarded a Ph.D. 
by the University of California in 1930. 
While in graduate school, he married Leona 
Suzan Galutia. 

His doctorate was awarded at the begin- 
ning of the Great Depression, and Burt re- 
mained at Cal Tech as a research fellow for 
6 years working on a variety of projects. In 
1935, he was awarded a tenure-track posi- 
tion at the University of Michigan, where 
he remained for the rest of his career. He 
also held a joint appointment as Curator of 
Mammals in the Museum of Zoology there. 
His career at Michigan was marked by the 
mentoring of over 20 graduate students, 


many of whom have become important 
mammalogists in their own right. He suc- 
cessfully guided the growth of the mammal 
collections of the Museum of Zoology to 
their current level of excellence. He also 
published pioneering studies on territorial 
behavior and home range in mammals. In 
addition to his scientific publications, he au- 
thored A Field Guide to the Mammals, which 
with its illustrations by Richard Grossen- 
heider, became a best-selling classic. 

Burt retired in 1969 and took up resi- 
dence in Boulder, Colorado, where he con- 
tinued his scientific studies as Honorary Cu- 
rator and Lecturer at the University of 
Colorado Museum. He and his wife also 
were able to indulge in the foreign travel 
they both loved until her death in 1973. He 
died in 1987 at the age of 84. His alma 
mater, the University of Kansas, was be- 
queathed the royalties from his field guide 
(Source: Muul, 1990). 


William B. Davis: 1955-1958 


William B. Davis (Fig. 3) was born to 
Bennoni Washington Davis and Mary Ann 
Matilda (Owens) Davis on 14 March 1902 
in Rexburg, Idaho, a small agricultural and 
lumber community on the Snake River 
about 50 miles southwest of Yellowstone 
National Park. His father and grandfather 
operated a small sawmill east of Rexburg. 
When Bill was 3 years old his father was 
killed in an accident at the sawmill. This 
tragedy left Buill’s mother with two small 
children and no visible means of support. 
Fortunately she was a competent cook so 
she spent the next 2 years cooking for min- 
ing crews in northern Utah. In 1907 she 
found employment as a cook in a new 
boarding house and hotel in Rupert, Idaho, 
a small community in an irrigation project 
on the north side of the Snake River. Bill 
received all of his elementary and high 
school education there and graduated in 
February 1920. 

At that time Idaho law permitted high 


PRESIDENTS 47 


William H. Burt William B. Davis Robert T. Orr 
(1953-1955) (1955-1958) (1958-1960) 


Stephen D. Durrant Emmet T. Hooper Donald F. Hoffmeister 
(1960-1962) (1962-1964) (1964-1966) 


Randolph L. Peterson Richard G. Van Gelder James N. Layne 
(1966-1968) (1968-1970) (1970-1972) 


Fic. 3.—Presidents of the ASM from 1953 to 1972. 


48 LAYNE AND HOFFMANN 


school graduates to qualify for a teaching 
certificate upon completion of two summer 
school courses at a normal college. Bill was 
not enthused with the labor involved in 
farming, so he followed the suggestion of 
his fiancée and qualified for a grade three 
teaching certificate. That autumn he began 
his teaching career in a rural school near St. 
Anthony, Idaho. During the next 13 years 
he alternated going to summer school and 
teaching in elementary schools in Idaho, 
Washington, and California, ranging from 
a single-room school with seven students in 
six grades to a three-room school where he 
was principal and teacher of the sixth to 
eighth grades. On 21 April 1923 he married 
Pearl Kathryn Tansey, and they have two 
children, a daughter, LaNell, and a son, 
Robert Lee. 

In 1932, Bill matriculated at Chico State 
College in California, where he received a 
B.A. in Education in 1933. However, even 
before entering college, he had developed a 
professional interest in ornithology. His first 
paper is dated 1923, and by the time he had 
finished at Chico State he had published 10 
papers, all but one on birds, based on ob- 
servations made during the course of his 
teaching career. 

His association with the University of 
California at Berkeley began the summer 
after he completed his B.A., when he served 
as a field assistant to E. R. Hall in Nevada, 
a position also held by Bob Orr. Dr. Joseph 
Grinnell agreed to chair Bill’s graduate 
committee if he switched his research to the 
field of mammalogy. This appears to have 
been the stimulus that turned him from birds 
to mammals, and in the following four sum- 
mers he conducted his graduate field work 
in Idaho, collecting mammals throughout 
the state, while supporting himself by work- 
ing as a graduate assistant in the Depart- 
ment of Zoology. His dissertation, The Re- 
cent Mammals of Idaho, was published in 
1939, 2 years after he received his Ph.D. By 
that time he had also published an addi- 
tional 26 papers, mostly based on work done 
while a graduate student. It is interesting to 


see the increasing emphasis on mammals in 
his scholarly output during this period. 

Upon completing his doctorate, Bill ac- 
cepted a professorship in the Department 
of Wildlife Science at Texas A&M Univer- 
sity. The following year (1938) he became 
Curator of the Texas Cooperative Wildlife 
Collections. He also served as Head of the 
Department from 1947 to 1965. During his 
academic career, he supervised the theses 
and dissertations of many well-known 
mammalogists. Upon his retirement from 
administration, he received the Governor’s 
Award for Outstanding Service in Conser- 
vation Education. 

He first became active as an officer of the 
ASM in 1937, as Corresponding Secretary, 
which he held for 3 years. He was elected 
President in 1955, and re-elected in 1956 
and 1957. He was appointed Chairman of 
the Board of Trustees, strong evidence of 
his colleagues’ confidence in his judgment 
and financial acumen. Bill remained active 
in research following his retirement in 1967, 
and to date has published a total of 188 
scholarly contributions. Failing eyesight fi- 
nally forced him to curtail his scholarly ac- 
tivities, but his interests remain strong. He 
now lives quietly with his second wife of 8 
years, Leola, in Bryan, Texas. She has two 
children by a former marriage. 


Robert Thomas Orr: 1958-1960 


Robert T. Orr (Fig. 3) recently told a friend 
and colleague that he has always considered 
himself ‘‘a real naturalist, not a specialist.” 
He attributed his initial interests in the out- 
of-doors to his physician father who took 
the whole family camping and encouraged 
him to hunt and fish. Bob was born on 17 
August 1908 in San Francisco, California, 
to Robert H. and Agnes K. Orr; he was one 
of three children. His grandfather had a 
ranch in Tehama County, and Bob spent 
many vacations while growing up collecting 
vertebrates on the ranch, although it is not 
clear where those specimens were deposit- 


PRESIDENTS 49 


ed, if they still survive. After grammar and 
high school in San Francisco, he enrolled in 
the University of San Francisco, receiving 
a Bachelor of Science in 1929. One of his 
teachers there, George Haley, was a person- 
al friend of Joseph Grinnell at the Univer- 
sity of California, Berkeley, to whom he in- 
troduced Orr. It was natural then that Bob 
should enroll in graduate school at Berkeley, 
receiving a master’s degree in 1931. At this 
time, E. Raymond Hall was also in the Mu- 
seum of Vertebrate Zoology, and employed 
him in field studies on the mammals of Ne- 
vada, which Hall later published through 
the U.C. Press. He also was befriended there 
by Alden Miller, the highly respected or- 
nithologist who was to become director of 
MVZ after Grinnell’s death. Bob accom- 
panied Miller on collecting trips, and cred- 
ited Miller with teaching him the funda- 
mentals of field ornithology, whose study 
he pursued throughout his career. 

His doctoral research was on the rabbits 
of California and was supervised by Grin- 
nell. He received the Ph.D. in 1937, 2 years 
after he had accepted a position as Wildlife 
Biologist with the National Park Service with 
assignments at various places in central Cal- 
ifornia. In 1936 he began a lifetime asso- 
ciation with the California Academy of Sci- 
ences when he was appointed Assistant 
Curator in the Department of Ornithology 
and Mammalogy, ultimately being awarded 
its Fellow’s Medal in 1973. 

Although his work prior to his doctoral 
dissertation was primarily on terrestrial 
mammals, his research interests at the 
Academy began to focus on marine mam- 
mals, although he continued to publish 
widely in both ornithology and mammal- 
ogy. His advancement at the California 
Academy of Sciences was steady, and be was 
named Full Curator in 1945, a rank he held 
for 30 years until his retirement. He also 
assumed the additional administrative duty 
of Associate Director in 1964 at the request 
of George Lindsay, whom he had supported 
for the directorship. Those additional duties 
finally forced him to terminate his courtesy 


teaching appointment at the University of 
San Francisco, which he had begun as an 
Assistant Professor of Biology in 1942, again 
rising through the ranks to Full Professor in 
1955. Upon his retirement in 1975, he was 
named Senior Scientist and Curator Emer- 
itus at the Academy. He has continued to 
publish, and his total bibliography now 
amounts to 267 titles. Only about one in 
ten were co-authored, one with his wife, 
Margaret C. Orr. They have one daughter. 

Bob has been honored as a Fellow and 
Honorary Member by a number of scientific 
societies and conservation organizations, 
including the American Association for the 
Advancement of Science, American Orni- 
thologists’ Union, and Explorers Club of 
New York. 


Stephen David Durrant: 1960-1962 


Stephen D. Durrant (Fig. 3) was born 1 1 
October 1902 in Salt Lake City, Utah, the 
son of Stephen Thomas and Martha Har- 
man Durrant. Following graduation from 
high school he spent several years (1922- 
1925) in Europe, mainly in Switzerland, on 
a mission for the Church of Jesus Christ of 
Latter-day Saints. During the summer of 
1933 while taking a course at the University 
of California at Berkeley, he met Sylvia Jane 
Burt, who was vacationing there from Salt 
Lake. They were married that December. 
They had two children, a daughter, Sue 
Marilyn, and a son, Stephen Carl. 

Steve began his undergraduate work at 
Weber Junior College, then transferred to 
the University of Utah, where he received 
the A.B. degree, with a major in Modern 
Languages (French), in 1929. As a result of 
courses taken with William W. Newby, who 
also taught him to prepare mammal skins, 
he decided to major in zoology for his Mas- 
ter’s degree, which he received in 1931. He 
began doctoral work at the University of 
Minnesota but after a year (1931-1932) ac- 
cepted an offer to return to the University 
of Utah as an instructor in comparative 


50 LAYNE AND HOFFMANN 


anatomy. While a full-time faculty member, 
he began doctoral work in mammalogy with 
E. R. Hall, first at the University of Cali- 
fornia at Berkeley (1938-1939) then at the 
University of Kansas when Hall moved 
there. He received his doctorate in 1950 and 
remained at the University of Utah for his 
entire career, rising from Assistant Profes- 
sor to Professor. 

Steve’s research dealt primarily with the 
distribution and systematics of Utah mam- 
mals. The genus 7homomys was a favorite 
subject. Of 37 new subspecies named by 
him and collaborators, 15 were pocket go- 
phers. He spent most of his summers in the 
field, often traveling by horseback. He par- 
ticipated in the Upper Colorado River Ba- 
sin Surveys from 1958 to 1962, serving as 
Field Director and mammalogist. His years 
of field work, during which he and graduate 
students amassed some 27,000 specimens, 
and his intimate knowledge of the mam- 
malian fauna of Utah were reflected in his 
book Mammals of Utah, Taxonomy and 
Distribution. 

Steve excelled as a teacher and, although 
a tough taskmaster, was revered by his stu- 
dents. His comparative anatomy course had 
the reputation of being both one of the best 
and hardest courses on campus. Mammal- 
ogy was offered once a year and was such a 
popular course that enrollment had to be 
limited. He had 36 graduate students, a 
number of which earned both master’s and 
doctorates under his direction. 

Steve joined the ASM in 1934. In addi- 
tion to the presidency, he was a Director 
and Vice-president. The International Re- 
lations Committee, one of the most pro- 
ductive in the Society, was formed during 
his tenure as president. He also served as a 
member, often chairman, of six standing 
committees. He participated in other sci- 
entific societies, including serving as Pres- 
ident of the Pacific Division of the Society 
of Systematic Zoology in 1956. Among hon- 
ors received during his career were election 
as Honorary Member of the ASM; the Dis- 
tinguished Teaching Award from the Uni- 
versity of Utah; the Distinguished Service 


Award from the Utah Academy of Sciences, 
Arts and Letters; and the establishment of 
the Stephen D. Durrant Memorial Schol- 
arship at the University of Utah. Perhaps 
his most cherished honor was a bronze cast- 
ing of a pocket gopher presented to him by 
graduate students and members of his last 
classes in comparative anatomy and mam- 
malogy. 

Steve was a warm and jovial person and 
a superb raconteur—a skill honed during 
many hours around a campfire with stu- 
dents and colleagues. He was an ardent duck 
hunter and a crack shot. He died on 11 No- 
vember 1975, and his remains after cre- 
mation were deposited near his favorite 
blind on Salt Lake where he had hunted for 
many years (Source: Behle, 1977). 


Emmet Thurman Hooper, Jr.: 
1962-1964 


Emmet T. Hooper (Fig. 3) was another of 
the many mammalogists inspired by Joseph 
Grinnell to pursue a professional career in 
biology. He was born 19 August 1911 in 
Phoenix, Arizona, the eldest of two chil- 
dren, to Emmet Thurman and Frances Jew- 
ell (McDonald) Hooper. His elementary 
schooling was in Phoenix, but after he had 
finished a year of high school, the family 
moved to San Diego, California. Complet- 
ing high school in that city, Emmet then 
enrolled in San Diego State University at 
the somewhat precocious age of 17. For his 
senior year, however, he transferred to the 
University of California at Berkeley where 
he came within the sphere of Joseph Grin- 
nell and the Museum of Vertebrate Zoology. 
Completing his bachelor’s degree in 1933, 
he then continued his graduate studies, re- 
ceiving a master’s degree in 1936 and his 
Ph.D. in 1939. His doctoral dissertation fo- 
cused on geographic variation in woodrats 
of the San Francisco Bay region. While at 
Berkeley, he also worked as a part-time as- 
sistant in the U.S. Bureau of Fisheries. 

He married Helen Bacon while a graduate 


PRESIDENTS Bil 


student, and they had two sons, Alan and 
Kim. Shortly before completing his doctor- 
ate, Emmet accepted what appears to have 
been a non-tenured position at the Univer- 
sity of Michigan Museum of Zoology, where 
he began a long professional association with 
fellow U.C. Berkeley graduates Bill Burt and 
Lee Dice. His tenure at the University of 
Michigan was interrupted by World War II, 
and he spent 4 years in the U.S. Army Air 
Corps, attaining the rank of Captain by the 
time of his discharge in 1946. He returned 
to the University of Michigan in that year 
and remained at the Museum of Zoology as 
Professor and Curator until his retirement 
in 1978. During those 3 decades, he served 
as major professor for many graduate stu- 
dents who have gone on to distinguished 
careers in mammalogy. His own research, 
published in 85 papers and monographs, 
was principally on the muroid rodents, es- 
pecially their morphology and systematics. 
Having lost his wife of 40 years in 1976, 
Emmet made the decision to relocate fol- 
lowing his retirement from the University 
of Michigan and accepted a position as lead- 
er of the sea otter research program of the 
U.S. Fish and Wildlife Service at the Center 
for Marine Studies, University of Califor- 
nia, Santa Cruz. He was very active in re- 
tirement, not only scientifically but in pub- 
lic service as well, serving as Commissioner 
of the Santa Cruz Museum and member of 
the Citizens Advisory Committee for Ni- 
senemarks State Park. In 1983 he remar- 
ried, to Leanore Theriot, and they resided 
in Aptos, California, until his death on 28 
June 1992 (Sources: Anon., 1988, 1992). 


Donald Frederick Hoffmeister: 
1964-1966 


Donald F. Hoffmeister (Fig. 3) was born 
in San Bernardino, California, on 21 March 
1916, and spent his youth in southern Cal- 
ifornia. Although his parents moved to Cal- 
ifornia from Iowa in 1906, his paternal 
grandfather had gone to California in the 


gold rush of 1849. Don and his wife, the 
former Helen Kaatz, were married in 1938 
and have two sons, Robert and Ronald. 

Don took his first 2 years of undergrad- 
uate work at San Bernardino Junior College 
and received his A.B. from the University 
of California, Berkeley, in 1938. Following 
in the footsteps of his grandfather and en- 
couraged by his parents, Don originally in- 
tended to become a medical doctor. How- 
ever, as the result of the influence of Dr. 
Elton R. Edge, one of his instructors in ju- 
nior college who had taken a field trip in 
Nevada with E. R. Hall, and a course in 
vertebrate zoology taught by Joseph Grin- 
nell and Hall, which he took in his senior 
year, Don decided to switch from medicine 
to mammalogy even though he had already 
been accepted to medical school. He re- 
mained at Berkeley for graduate study with 
E. R. Hall, who had a profound influence 
on his scientific career. He received the M.A. 
in 1940 and the Ph.D. in 1944. During his 
graduate work, he held a Teaching Assis- 
tantship and also served as Technical As- 
sistant and Research Assistant in the Mu- 
seum of Vertebrate Zoology. 

In 1944 he was appointed Assistant Pro- 
fessor and Assistant Curator of Modern 
Vertebrates at the University of Kansas, and 
in 1946 went to the University of Illinois 
as Assistant Professor and Assistant Cura- 
tor in the Museum of Natural History, where 
he remained for the remainder of his career. 
He became Associate Professor in 1956, and 
Professor in 1959. He was promoted to Cu- 
rator, with responsibility as director, in the 
Museum of Natural History in 1948 and 
was given the official title of Director of the 
museum in 1964. Upon his retirement in 
1984, he was appointed Emeritus Director 
and Professor. In addition to his research, 
administration, and teaching, he served as 
chairman of 14 Ph.D. and 18 master’s stu- 
dents. Two of his students are themselves 
past-presidents of ASM. 

Don’s research has dealt primarily with 
the distribution and taxonomy of mam- 
mals, with emphasis on Arizona and Illi- 
nois. However, his publications include a 


52 LAYNE AND HOFFMANN 


distributional note and study of growth and 
development of birds and papers on life his- 
tory and ecology, pelage coloration, and 
various aspects of anatomy of mammals. 
He has also described a number of mam- 
malian taxa. He is the author or coauthor 
of a number of semipopular and technical 
books on mammals, including Mammals/ 
A Guide to Familar American Species with 
H. S. Zim, Handbook of Illinois Mammals 
with C. O. Mohr, Fieldbook of Illinois 
Mammals with C. O. Mohr, Mammals of 
the Grand Canyon, Mammals of Illinois, 
and the monumental Mammals of Arizona. 
Sources of support for his work include the 
National Science Foundation, National In- 
stitutes of Health, Illinois Department of 
Conservation, Arizona Fish and Game De- 
partment, and the Max McGraw Wildlife 
Foundation. 

Don became a member of the ASM in 
1938. Other elective offices he held in ad- 
dition to the presidency include Director, 
Corresponding Secretary, and Vice-presi- 
dent. He was appointed in 1966 as the so- 
ciety’s first Historian and continues to serve 
in that capacity. He was a member of five, 
and chairman of three, standing committees 
and also chaired special committees on Sub- 
scriptions to the Journal of Mammalogy and 
Reprinting of the Journal of Mammalogy. 
Offices he has held in other professional or- 
ganizations include President of the Mid- 
west Museums Conference, Chairman of the 
Zoology Section and Councillor of the Ili- 
nois State Academy of Science, Councillor 
of the American Association of Museums, 
and Associate Editor of The American Mid- 
land Naturalist. 

In recognition of his outstanding service 
to the ASM, Don was awarded Honorary 
Membership in 1982 and the Hartley H. T. 
Jackson Award in 1986. Among his other 
professional honors are Honorary Mem- 
bership, Midwest Museums Conference; 
appointment to the Governor’s Board of the 
Illinois State Museum; and appointment as 
Research Associate of both the Museum of 
Northern Arizona and the Northern Ari- 
zona Society of Science and Art. 


The second meeting of the society outside 
the United States was held in Winnipeg, 
Canada, during Don’s presidency, and in 
what was probably a first for an ASM pres- 
ident he was made an Honorary Citizen of 
Winnipeg by Royal proclamation. 


Randolph Lee Peterson: 1966-1968 


Randolph L. Peterson (Fig. 3) was born 
on 16 February 1920 in Roanoke, Texas, 
one of five children of Omas and Margaret 
Francisco Peterson. Pete, as he was known 
to his friends and colleagues, spent his youth 
on the family farm, where he developed an 
interest in natural history, particularly 
mammals, plants, and ecology. In 1942 he 
married Elizabeth Fairchild Taylor, the 
daughter of the well-known mammalogist 
Walter P. Taylor, who was Pete’s mentor. 
They had one daughter, Penny Elizabeth. In 
addition to her role as wife and mother, 
Elizabeth participated with Pete in running 
a biological supply business and helped as 
his research assistant. During World War 
II, Pete served in the U.S. Air Force as pilot 
and instructor and Operations Officer with 
the Mediterranean Allied Air Force. 

He obtained his B.Sc. in 1941 in the De- 
partment of Fish and Game at Texas A&M 
University. During his undergraduate years 
he served as Assistant Curator of the Texas 
Cooperative Wildlife Research Collection 
under William B. Davis. He began graduate 
studies at Texas A&M, but went into service 
before completing his degree. After the war, 
he entered the graduate program of the Uni- 
versity of Toronto under J. R. Dymond and 
received the Ph.D. in 1950. 

While at the University of Toronto, Pete 
served as Acting Curator in Charge of the 
Mammal Division of the Royal Ontario 
Museum and upon receiving his degree was 
appointed Curator-in-Charge of the De- 
partment of Mammalogy of the Museum, a 
position he held until retirement in 1985. 
He was also on the faculty of the University 
of Toronto, as Special Lecturer in the De- 
partment of Zoology (1949-1962), Asso- 


PRESIDENTS 58, 


ciate Professor (1962-1968), and Professor 
(1968-1985). Upon retirement he was ap- 
pointed Curator Emeritus in the museum 
and Professor Emeritus in the university. 
He died on 29 October 1989. 

Pete’s doctoral research was on the bi- 
ology of the moose and was published as 
the book North American Moose in 1955. 
This was one of the most definitive studies 
of the species and has been reprinted several 
times. He also directed a survey of the 
mammals of Ontario and Quebec, which 
culminated in his second book, Mammals 
of Eastern Canada, in 1966. Bats became a 
consuming research interest later in his ca- 
reer and over a third of his publications deal 
with the taxonomy, distribution, habitats, 
and habits of bats, including descriptions of 
five new species. He led expeditions to many 
areas in North America and Mexico and 
abroad and built one of the largest and most 
complete collections of bats in the world at 
the Royal Ontario Museum. 

As Curator-in-Charge of mammalogy at 
the museum, he supervised the renovation 
and expansion of the department’s office and 
collection space. He also served as Editor 
and Chairman of the Life Sciences Publi- 
cations and was a member of the Promotion 
and Tenure Committees. As Professor of 
Zoology he taught mammalogy and directed 
the work of eight doctoral and eight master’s 
students, in addition to serving on the grad- 
uate committees of many others. 

Pete joined the ASM in 1940 and attend- 
ed 50 consecutive meetings. Besides the 
presidency, he was a Director, Recording 
Secretary, and Vice-president, in addition 
to serving as chairman and member of nu- 
merous committees. He played a key role 
in the establishment of the Future Mam- 
malogists Fund and in 1986 was awarded 
Honorary Membership. His participation 
in other scientific organizations included 
serving on the boards of the Metropolitan 
Toronto Zoological Society and the Met- 
ropolitan Toronto Zoo and as Councillor of 
the Society of Systematic Zoology. 

In addition to his active and productive 
professional life, Pete pursued interests in 


gardening, farming, wood-working, and oe- 
nology. In the 1950s, he and Elizabeth start- 
ed a thriving biological supply company and 
operated it until 1974. He also invented such 
items of equipment as automated calipers 
for measuring specimens, a cider press, and 
a large skeleton cleaning apparatus. 

Pete was a man of contrasts. When nec- 
essary, he was rough and ready, as might be 
expected given his Texas origins, but on 
other occasions he was the perfect country 
gentleman (Source: Eger and Mitchell, 1990). 


Richard George Van Gelder: 
1968-1970 


Richard G. Van Gelder (Fig. 3) was born 
in New York City on 17 December 1928, 
the son of Joseph and Clara DeHirsch Van 
Gelder. Despite growing up in an urban en- 
vironment, he developed an avid interest in 
natural history at an early age. Upon grad- 
uation from the Horace Mann School in 
New York with honors in biology and Span- 
ish, he entered Colorado A&M College. He 
received a B.S. with honors in 1950, then 
attended graduate school at the University 
of Illinois at Urbana, where he worked with 
Donald F. Hoffmeister. Dick received the 
M.S. in 1952 and Ph.D. in 1958. During 
graduate school, he spent one summer at 
the Marine Biological Laboratory at Woods 
Hole. He was married in 1962 to Rosalind 
Rudnick, and they have three children: Rus- 
sell Neil, Gordon Mark, and Leslie Gail. His 
son Russell probably holds the record of 
being the youngest person ever to join ASM, 
as Dick took out a life membership for him 
when he was a baby. 

Dick served as Curator in the Natural 
History Museum of Colorado A&M College 
in 1948-1949 and was an Assistant in the 
Mammal Department of the American Mu- 
seum of Natural History in 1952. He was a 
Research Assistant in the Museum of Nat- 
ural History of the University of Kansas 
from 1954 to 1956 and an Assistant Pro- 
fessor in 1955-1956. In 1956, he was ap- 
pointed Assistant Curator in the Depart- 


54 LAYNE AND HOFFMANN 


ment of Mammals of the American Museum 
of Natural History and was promoted to 
Associate Curator in 1961 and Curator in 
1969. He also served as Acting Chairman 
of the Mammal Department in 1958-1959 
and as Chairman from 1959 to 1974. He 
retired in 1986. During his tenure at the 
American Museum, Dick also held appoint- 
ments as Instructor and Assistant Professor 
at Columbia University; Adjunct Graduate 
Advisor at Albert Einstein Medical College, 
Columbia University, New York Univer- 
sity, and City College of New York; and 
Professorial Lecturer at Downstate Medical 
Center of the State University of New York. 

Dick’s areas of research reflect his broad 
interests in mammals, including mammal 
populations and physiology; taxonomy of 
carnivores, marine mammals, bats, artio- 
dactyls; behavior; hybridization and spe- 
ciation; color patterns, and mammals of New 
Jersey. He also has published on other ver- 
tebrates, including amphibians, reptiles, and 
birds. He has conducted field work in many 
parts of North America, as well as Mexico, 
Uruguay, Bolivia, Bahamas, Mozambique, 
Botswana, and South West Africa. Skunks 
were one of his favorite groups, and he pub- 
lished a definitive revision of the spotted 
skunks in 1959, as well as a number of other 
papers and semipopular articles on the tax- 
onomy, morphology, behavior, and habits 
of skunks. In later years, he worked on the 
behavioral ecology of African ungulates and 
directed a cooperative study of the status of 
mammals of New Jersey. He was author of 
the books Biology of Mammals, Mammals 
of the National Parks, and Animals and Man, 
Past, Present, Future, coauthor of Animals 
in Winter, and coeditor of Physiological 
Mammalogy volumes I and II. In addition 
to research, he directed graduate students 
in areas of mammalian anatomy, behavior, 
and history and was active in the Museum’s 
exhibits program, playing a lead role in the 
design, construction, and installation of the 
blue whale model that dominates the Hall 
of Fishes. He recounted his experience with 
the latter project in the humorous article 
‘*“Whale on my back” published in Curator. 


Dick also taught adult education courses at 
the museum for many years. 

Dick joined the ASM in 1948. Besides 
the presidency, he served as Director, Vice- 
president, and Recording Secretary. As 
chairman of the Committee on Recent Lit- 
erature, he edited the Recent Literature sec- 
tion of the Journal of Mammalogy from 
1965 to 1968. He also was a member of 
many other committees. The office of His- 
torian was established during his presiden- 
cy. 
Among other appointments, Dick was a 
member of the Board of Directors of Arch- 
bold Expeditions; a Director of the Quincy 
Bog Natural Area in New Hampshire; a 
member of the Board of Education, Har- 
rington Park, New Jersey; and a member of 
the Technical and Editorial Advisory Board 
of the Population Reference Bureau. 


James Nathaniel Layne: 1970-1972 


James N. Layne (Fig. 3) was born on 16 
May 1926 in Chicago, Illinois, to Harriet 
(Hausman) and Leslie J. Layne. He grew up 
in what Chicagoans call the “near north 
side,” Irving Park and Rogers Park. When 
he was 6 years old, his father left the family, 
and he and his younger brother were raised 
by his mother through the difficult days of 
the Great Depression. Despite the hard 
times, his mother encouraged his growing 
interest in natural history. By age 12 he had 
become an enthusiastic falconer, and his 
high school years were spent capturing and 
training hawks, and spending many hours 
observing raptors in the Cook County For- 
est Preserves. His high school biology teach- 
ers Doris Plapp and Susan Arenberg also 
encouraged his passion for raptors, taking 
him on field trips and, together with his 
English teacher Fred Thompson, encour- 
aging him to write. His first scientific paper, 
published in 1943 in the J/linois Audubon 
Society Bulletin, was completed while he 
was still in high school. 

Upon his graduation in 1944, Jim enlist- 
ed in the Army Air Force and served until 


PRESIDENTS 5D 


after World War II, being discharged in 
1946. While stationed in the southeastern 
U.S., he met Philip S. Humphrey, the well- 
known ornithologist who now directs the 
University of Kansas Museum of Natural 
History. Through his influence, Jim devel- 
oped a broad interest in birds and was com- 
mitted to ornithology when he enrolled in 
Cornell University in 1947 after a freshman 
year at Chicago City Junior College. How- 
ever, during his sophomore year he took the 
vertebrate zoology course taught by Ed Ra- 
ney and Bill Hamilton, and henceforth fish- 
es and mammals also competed for his in- 
terest. He completed his B.A. degree in 1950 
still uncommitted to a particular vertebrate 
group, until Hamilton offered him an assis- 
tantship to work on mammals. In that year, 
he not only acquired a mentor, but also a 
wife when he was married to Lois Linder- 
oth; they have five children, all daughters: 
Linda, Kimberly, Jamie, Susan, and Rachel. 

Jim continued to publish during his Air 
Force and undergraduate as well as graduate 
careers, producing 15 more papers by the 
time he completed his Ph.D. in 1954. His 
dissertation on the biology of the red squir- 
rel was published in that year, and he ac- 
cepted an assistant professorship in the De- 
partment of Zoology and the Cooperative 
Wildlife Research Laboratory at Southern 
Illinois University in Carbondale. The next 
year he moved to the University of Florida 
in Gainesville where he held both an aca- 
demic and a curatorial appointment (Assis- 
tant and Associate Professor of Biology and 
Assistant and Associate Curator in Charge 
of Mammals in the Florida State Museum) 
until 1963. This was a productive period 
for him, with papers in a number of different 
disciplines of mammalogy, and supervision 
of graduate students. Nonetheless, in 1963 
he heeded the call from his alma mater and 
returned to Cornell for 4 years as Associate 
Professor of Zoology, only to reverse his 
course in 1967 and return to Florida as Di- 
rector of Research at the Archbold Biolog- 
ical Station in Lake Placid, Florida, with a 
concurrent appointment as Archbold Cu- 
rator of Mammals in the American Muse- 


um of Natural History. From 1976 to 1985 
he served as Executive Director of the sta- 
tion, and now continues as Senior Research 
Biologist there. His research has continued 
unabated, not only over a broad field of 
mammalian topics, but extended to all as- 
pects of the natural history of Florida. 

His contributions to the ASM, beginning 
with his service on the Committee on Ma- 
rine Mammals in 1959, are many and var- 
ied, including membership and chairman- 
ship of many other committees, long service 
on the Board of Directors, and Editor of 
Special Publications. In 1976, he received 
the C. Hart Merriam Award and in 1993 
was elected an Honorary Member. His ac- 
tivities in other professional organizations 
include the presidency of the Organization 
of Biological Field Stations and Florida 
Academy of Sciences. He has also served 
on a number of boards of environmental 
organizations and advisory committees of 
the Florida Game and Fresh Water Fish 
Commission, U.S. Fish and Wildlife Ser- 
vice, and other governmental agencies. 


J. Knox Jones, Jr.: 1972-1974 


J. Knox Jones, Jr. (Fig. 4) was born in 
Lincoln, Nebraska, on 16 March 1929. He 
was married to Maryane Rountree Davis in 
1989. Knox had three daughters, Amy, Sar- 
ah, and Laura, from an earlier marriage to 
Janet Glock. He was an officer, with ter- 
minal rank of Captain, in the U.S. Army 
and served on active duty in Korea and Ja- 
pan from 1953 to 1955 and in the reserve 
from 1956 to 1965. 

Knox received his B.S. in 1951 from the 
University of Nebraska and both his M.A. 
(1953) and Ph.D. (1962) degrees under E. 
R. Hall from the University of Kansas. 

In 1962 he was appointed Assisiant Pro- 
fessor of Zoology and Assistant Curator of 
Mammals in the Museum of Natural His- 
tory, University of Kansas, and was pro- 
moted to Associate Professor and Associate 
Curator in 1965 and Professor and Curator 
in 1968. While at Kansas, he also served as 


56 LAYNE AND HOFFMANN 


J. Knox Jones, Jr. Sydney Anderson William Z. Lidicker, Jr. 
(1972-1974) (1974-1976) (1976-1978) 


Robert S. Hoffmann James S. Findley J. Mary Taylor 
(1978-1980) (1980-1982) (1982-1984) 


Hugh H. Genoways Don E. Wilson Elmer C. Birney 
(1984-1986) (1986-1988) (1988-1990) 


Fic. 4.—Presidents of the ASM from 1972 to 1990. 


PRESIDENTS 57 


Assistant (1965-1967) and Associate (1967— 
1971) Director of the Museum. In 1971 he 
became Professor of Biological Sciences at 
Texas Tech University and in 1986 was ap- 
pointed Paul Whitfield Horn Professor of 
Biological Sciences and Museum Science. 
He also served as Dean of the Graduate 
School (1971-1984), Associate Vice Presi- 
dent for Research (1972-1974), and Vice 
President for Research and Graduate Stud- 
ies (1974-1984). Additional appointments 
he held at Texas Tech included Adjunct 
Professor of Veterinary and Zoological 
Medicine and Director of Academic Pub- 
lications. He was Acting Director of the 
Texas Tech Museum in 1971-1972, a Re- 
search Associate in 1971-1984, and was 
Curator and Editor of Museum Publications 
at the time of his death. 

Knox taught a wide range of courses at 
both the University of Kansas and Texas 
Tech and served as major professor to 16 
doctoral and 15 master’s students in mam- 
malogy. His primary areas of research were 
mammalian systematics, evolution, bioge- 
ography, and natural history, with a focus 
on the Great Plains and the Neotropics. His 
more than 300 publications include original 
descriptions of over 30 mammalian taxa, as 
well as three invertebrate species. He was 
the author or editor and contributing author 
of 13 books, including Distribution and 
Taxonomy of Mammals of Nebraska, Re- 
cent Mammals of the World—a Synopsis of 
Families, Pleistocene and Recent Environ- 
ments of the Central Great Plains, Mam- 
mals of the Northern Great Plains, and 
Handbook of Mammals of the North-central 
States. 

Knox became a member of the ASM in 
1947 and played an active role in the affairs 
of the society. Other offices he held in ad- 
dition to the presidency include Director, 
Vice-president, Managing Editor, and Edi- 
tor for Reviews. He served on many of the 
standing committees of the society. Major 
initiatives he undertook as president in- 
cluded organizing, together with Bob Hoff- 


mann, the transportation for, and leading 
the ASM contingent to, the I st International 
Theriological Congress in Moscow in 1974; 
establishing the geographic rotation plan for 
annual meeting sites; initiating the estab- 
lishment of the Merriam Award; and ob- 
taining National Science Foundation sup- 
port for a committee to evaluate systematic 
resources in mammalogy. The Information 
Retrieval, Index, and Systematic Collec- 
tions committees were established during 
his presidency. 

Offices he held in other scientific organ- 
izations included Director and Treasurer, 
Organization of Tropical Studies; Managing 
Editor, Society for the Study of Evolution; 
Councillor, Society of Systematic Zoology; 
Editor, The Texas Journal of Science of the 
Texas Academy of Science; and President, 
Texas Society of Mammalogists. 

Knox received the three highest honors 
bestowed by the ASM: the C. Hart Merriam 
Award (1977), the H. H. T. Jackson Award 
(1983), and Honorary Membership (1992). 
Among the other honors that came to him 
during the course of his career were the Out- 
standing Research Award from the College 
of Arts and Sciences, Texas Tech Univer- 
sity, the Barnie E. Rushing Award for out- 
standing research, and election as Fellow of 
the Texas Academy of Science. Knox died 
at his home in Lubbock, Texas, on 15 No- 
vember 1992. His outstanding contribu- 
tions to science and education have been 
recognized by Texas Tech University 
through the creation of the J. Knox Jones 
Memorial Scholarship. 


Sydney Anderson: 1974-1976 


Sydney Anderson (Fig. 4) was born in To- 
peka, Kansas, on 11 January 1927. His early 
years were spent in Kansas. As a child, he 
was fascinated with collecting things, fore- 
shadowing his later career as a museum cu- 
rator, and at age five he began to include 
natural history objects in his collections. His 


58 LAYNE AND HOFFMANN 


decision to become a mammalogist came 
while completing his undergraduate work at 
the University of Kansas and was strongly 
influenced by the environment of the Mu- 
seum of Natural History under E. Raymond 
Hall and summer field work directed by 
Rollin H. Baker. Syd met Justine Klusmire, 
also a native Kansan, while both were stu- 
dents at the University of Kansas, and they 
were married in 1951. Justine has shared 
Syd’s professional interests, working with 
him at the museum and in the field. She is 
a familiar figure at the annual meetings. They 
have three children, Evelyn Lee, Charles 
Sydney, an ichthyologist, and Laura Lyn- 
nette. 

Syd attended Baker University for 3 years 
and completed his A.B. degree at the Uni- 
versity of Kansas in 1950. He remained at 
Kansas for graduate work, receiving the 
M.A. in 1952 and the Ph.D. in 1959. He 
spent the summer of 1952 at the Friday 
Harbor Oceanographic Laboratory of the 
University of Washington. 

He joined the staff of The American Mu- 
seum of Natural History as Assistant Cu- 
rator of Mammals in 1961 and was pro- 
moted to Associate Curator in 1965 and 
Curator in 1969. He was appointed Emer- 
itus Curator upon retirement in 1992. He 
also served as Chairman of the Mammalogy 
Department from 1975 to 1981 and as Ad- 
junct Professor at the City University of New 
York from 1968 to 1988. He became a Re- 
search Associate of the University of New 
Mexico in 1988. 

Syd’s research has dealt broadly with nat- 
ural history, ecology, distribution, evolu- 
tion, and systematics of mammals and has 
also included work on such diverse topics 
as areography of North American fishes, 
amphibians, and reptiles; food habits of 
owls; and patterns of geographic distribu- 
tion of birds. He also was one of the pioneers 
in the development of information retrieval 
systems for natural history museums and 
has pursued interests in history of science 
and the literature of natural history. He has 


conducted field work in a number of regions 
in the eastern and western United States, as 
well as Mexico, Uruguay, and, most re- 
cently, Bolivia. In addition to his other pub- 
lications, Syd is author or coauthor of nu- 
merous book chapters and symposium 
papers as well as coeditor and coauthor of 
four books: Recent Mammals of the World, 
a Synopsis of Families; Readings in Mam- 
malogy, Selected Readings in Mammalogy, 
and Orders and Families of Recent Mam- 
mals of the World. 

He was awarded a National Science 
Foundation Graduate Fellowship in 1952- 
1954 and received an ASM Graduate Stu- 
dent Honorarium in 1954. He is a Fellow 
of the American Association for the Ad- 
vancement of Science. For his contributions 
to the ASM and mammalogy in general, he 
received the H. H. T. Jackson Award in 
1986 and was elected to Honorary Mem- 
bership in 1992. 

Syd has been a member of ASM since 
1952 and has served the society in a number 
of capacities in addition to the presidency. 
These include Vice-president, Recording 
Secretary, Director, Trustee (Chairman), 
Editor of Mammalian Species, and mem- 
bership in many standing and ad hoc com- 
mittees. The Merriam Award was formally 
established during Syd’s presidency. He was 
a prime mover in the creation of the Mam- 
malian Species series and played a key part 
in the establishment of the society’s stand- 
ing committees on systematic collections 
and information retrieval and in clarifying 
the role of the Trustees and management of 
the Reserve Fund. As a member of the com- 
mittee on revision of the Rules and By-laws 
in 1974, he proposed the expansion of the 
elected directors from 10 to 15 and the re- 
tention of past presidents on the Board. He 
also has served as the society’s unofficial 
parliamentarian. Syd has been active in a 
number of other scientific, educational, and 
conservation organizations as well, includ- 
ing serving as trustee and officer of the Clos- 
ter Nature Center Association and the Ber- 


PRESIDENTS Do 


gen Museum of Art and Science in New 
Jersey. 


William Zander Lidicker, Jr.: 
1976-1978 


William Z. Lidicker, Jr. (Fig. 4) was born 
in Evanston, Illinois, on 19 August 1932, 
to William Z. Lidicker and Frida Schroeter 
Lidicker; his father is a civil engineer. Bill 
is the eldest of three sisters and one brother. 
As a child, he lived successively in Iowa, 
Missouri, Minnesota, New Mexico, Min- 
nesota, The Republic of Panama, Texas, and 
finally New York. Bill attended Forest Hills 
High School in the Borough of Queens, New 
York City, by which time he had already 
developed a serious interest in natural his- 
tory. He recalls that this was probably a 
result of his experiences beginning with his 
residence in Panama. The family lived close 
to swamps along the Panama Canal and he 
had opportunities to visit tropical rain for- 
ests, including Barro Colorado Island, al- 
ready a famous tropical research center, as 
well as cloud forests in northern Panama 
near the Costa Rican border. When the fam- 
ily returned to the United States to live in 
Galveston, Texas, he was an active member 
of a Boy Scout troop that regularly hiked 
and camped along the coast. By the time he 
was a sophomore in high school in New 
York, he had become an active bird watch- 
er, and his developing professionalism was 
encouraged by biologists such as H. M. Van 
Deusen, then at the American Museum of 
Natural History. 

He enrolled at Cornell University with 
the intention of studying ornithology under 
Arthur A. Allen, but quickly was swayed by 
the powerful personality of William J. 
Hamilton, Jr., to broaden his horizons to 
mammals. As an undergraduate, he had a 
series of summer jobs in biology, including 
work with Jess Low in Utah, Paul S. Martin 
in northeastern Mexico, and finally as a bi- 
ologist on an oceanographic expedition along 


the coast of Newfoundland and Labrador. 
He received his bachelor’s degree from Cor- 
nell in 1953, and immediately enrolled in 
the graduate program at the University of 
Illinois, having been accepted as a student 
by Donald F. Hoffmeister. He progressed 
through graduate school in near-record time, 
being awarded a master’s in 1954 and his 
Ph.D. in 1957 from the University of Cal- 
ifornia, Berkeley. He progressed regularly 
through the professorial/curatoral ranks at 
Berkeley, where he has remained. His career 
appears to depict a linear trajectory without 
interruption or deviation; it also reveals a 
fundamental and continuing expansion of 
his scientific interests. His publication rec- 
ord shows him to be focused in his student 
years on taxonomy and distribution of 
mammals. In his first years at Berkeley he 
was influenced by a number of colleagues, 
particularly Paul K. Anderson and Frank A. 
Pitelka, to expand his horizons to ecology, 
and particularly population ecology and ge- 
netics, where he became a leader in estab- 
lishing the inter-disciplinary field of genetic 
ecology. This shift was clearly recognizable 
in 1962, when he published two papers in 
this field that continue to be cited. In that 
same year, he accompanied a field expedi- 
tion to New Guinea, which led him to ex- 
pand his interests beyond North America. 
All of his later publications reflect a pro- 
gressive broadening of his research interests 
(social behavior, landscape ecology, con- 
servation biology) as does the work of most 
of the 21 doctoral and 10 master’s students 
who have received their degrees under his 
supervision. 

In addition to mammalogy, his passion 
is international folk dancing, which he shares 
with his wife, Louise. He has two sons, Jef- 
frey and Kenneth, by his former wife Naomi 
Ishino. 


Robert Shaw Hoffmann: 1978-1980 


Robert S. Hoffmann (Fig. 4) was born in 
Evanston, Illinois, a northern suburb of 


60 LAYNE AND HOFFMANN 


Chicago, on 2 March 1929. By the time he 
was in grade school his family had moved 
to a more rural town, and he was spending 
much of his time in the woods and fields 
around his home and in a nearby forest pre- 
serve. He frequently journeyed by streetcar 
to the Field Museum in downtown Chicago 
and haunted the Brookfield Zoo. At age 11 
he got a summer job at the zoo selling pea- 
nuts, which gave him the opportunity of 
getting to know all the keepers and animals. 
A fifth grade teacher, Nell Hashagen, also 
strongly encouraged his interest in natural 
history, and by the time he reached high 
school he had decided on a career in some 
field of biology. Phil Wright, his advisor 
during a year he spent at the University of 
Montana as an undergraduate, was a major 
influence in his selection of mammalogy as 
his major field of specialization. Bob and 
his wife, the former Sally Ann Monson, have 
four children: Karl, John, David, and Bren- 
na. 

As an undergraduate, Bob attended the 
University of Illinois Extension in Moline 
(1946-1947), University of Montana (1947- 
1948), and Utah State University (1948- 
1950), from which he received the B.S. He 
did his graduate work (M.A. in 1954, Ph.D. 
in 1955) at the University of California, 
Berkeley, where he was awarded two Na- 
tional Science Foundation Predoctoral Fel- 
lowships and the Alexander Museum of 
Vertebrate Zoology Scholarship. His major 
professor was A. Starker Leopold, and he 
was also strongly influenced by Frank A. 
Pitelka and O. P. Pearson. 

In 1955, Bob was appointed Instructor in 
the Department of Zoology, University of 
Montana. He was promoted to Assistant 
Professor in 1957, Associate Professor in 
1961, and Professor in 1965. He also served 
as Curator of the Zoological Museum of the 
University of Montana. He joined the fac- 
ulty of the University of Kansas in 1968 as 
Curator of Mammals in the Museum of 
Natural History and Professor in the De- 
partment of Zoology. For varying periods 
during his tenure at the University of Kan- 


sas, he also served as Chairman of the De- 
partment of Systematics and Ecology, Act- 
ing Chairman of the Division of Biological 
Sciences, and Associate Dean and Acting 
Dean of the College of Liberal Arts and Sci- 
ences. In 1986, he became Director of the 
National Museum of Natural History and 
since 1988 has served as Assistant Secretary 
for Science at the Smithsonian Institution. 

Bob’s research has dealt with birds and 
wildlife management, as well as both fossil 
and Recent mammals. A major focus in his 
work has been on various groups of Hol- 
arctic mammals and the Pleistocene history 
of Beringia. He is coauthor of Selected 
Readings in Mammalogy. 

He became a member of the ASM in 1955. 
In addition to the presidency, he has served 
as a Director, Vice-president, Review Edi- 
tor of the Journal of Mammalogy, and 
member or chairman of a number of com- 
mittees. He has been especially active in the 
Committee on International Relations, 
which he chaired in 1964-1968 and 1972- 
1978. Because of his research in Russia, fa- 
miliarity with Russian scientists, and 
knowledge of the language, he played an im- 
portant role in establishing liaison between 
the ASM and Russian mammalogists and 
laying the groundwork for the Ist Therio- 
logical Congress. The Education and Grad- 
uate Students Committee was formed dur- 
ing his presidency. 

Bob is a member of a number of other 
professional societies and has served as a 
consultant to, or member of, many national 
and international scientific bodies, includ- 
ing the U.S.-U.S.S.R. Joint Commission of 
Science Policies of the U.S. National Acad- 
emy of Sciences and the Board of Editors 
of Acta Zoologica Sinica (Beijing). He is a 
member of Phi Kappa Phi, Sigma Xi, and 
Phi Sigma and a Fellow of the American 
Association for the Advancement of Sci- 
ence. Included among other honors he has 
received are an Honorary Doctor of Science 
from Utah State University, the Summer- 
field Distinguished Professorship from the 
University of Kansas, and Honorary Mem- 


PRESIDENTS 61 


bership in the All-Union (U.S.S.R.) Therio- 
logical Society. 


James Smith Findley: 1980-1982 


James S. Findley (Fig. 4) was born in 
Cleveland, Ohio, on 28 December 1926. He 
grew up in a well-forested suburb of the city, 
an environment that fostered his early in- 
terest in birds, which was further nurtured 
by his membership in the Kirtland Bird Club 
at the Cleveland Museum of Natural His- 
tory. When in seventh grade, he met Nor- 
man Negus, who shared his interest in nat- 
ural history and fishing. Together they began 
to collect mammals, birds, herps, and in- 
sects and discovered Hamilton’s Mammals 
of Eastern United States, which greatly in- 
fluenced Jim’s later decision to become a 
mammalogist. When 13 years old, he be- 
came acquainted with Phil Moulthrop and 
B. P. Bole, Jr., mammalogists at the Cleve- 
land Museum of Natural History, and they 
and other museum staff members intro- 
duced him to the possibilities of a career in 
natural history. Jim’s father wrote to Harold 
E. Anthony at the American Museum of 
Natural History asking advice on what to 
do with a son with such interests. Anthony 
replied that the financial rewards of a career 
in biology would be modest and empha- 
sized the importance of attending a good 
university with broad offerings in biology 
and a rich local flora and fauna, recom- 
mending Berkeley as such a place. 

Jim served in the U.S. Army in 1945 and 
1946, and saw duty in Japan. He and and 
wife, Helen Muriel Thomson (‘““Tommie’’), 
were married in 1949 and spent their hon- 
eymoon surveying mammals in Jackson 
Hole, Wyoming. They have four children, 
sons Stuart and Douglas and daughters Hei- 
di and Joan. 

Jim took undergraduate courses at Kobe 
Central School during the year he was sta- 
tioned in Japan and spent the summer of 
1947 at the Rocky Mountain Biological 
Laboratory. He received the B.A. cum laude 


from Western Reserve University in 1950. 
He attended the University of California, 
Pacific Grove, in 1951, and completed his 
graduate work (Ph.D., 1955) at the Uni- 
versity of Kansas, with E. R. Hall as his 
major professor. 

From 1954 to 1955, he was an Instructor 
in the Zoology Department of the Univer- 
sity of South Dakota. In 1955 he joined the 
staff of the Biology Department of the Uni- 
versity of New Mexico as Assistant Profes- 
sor. He became Associate Professor in 1961 
and Professor in 1970. He served as Chair- 
man of the Biology Department from 1978 
to 1982 and was Director of the Museum 
of Southwestern Biology from 1983 to 1992. 
He retired in July 1992. 

The main focus of Jim’s research has been 
on zoogeography, distribution, and taxon- 
omy of mammals of southwestern United 
States, with emphasis on bats and shrews. 
He pioneered in the study of ecological cor- 
relates of morphology of bats and other 
mammals and has made important contri- 
butions to knowledge of the patterns and 
processes of small mammal community for- 
mation. Besides journal papers and other 
publications, he is an author of chapters in 
Recent Mammals of the World, Contribu- 
tions to Mammalogy, Ecology of Bats, Pat- 
terns in the Structure of Mammalian Com- 
munities; and The Butterflyfishes: Success 
on the Coral Reef (with Tommie Findley). 
He coauthored Mammals of New Mexico 
and is the author of Natural History of New 
Mexican Mammals and Bats: a Community 
Perspective. In addition to his research with 
mammals, Jim has published on distribu- 
tion of such diverse taxa as birds, amphib- 
ians, and river crabs and in recent years has 
been investigating patterns of community 
organization in reef fishes around the world. 

He has directed the work of 33 master’s 
and 24 doctoral students, and under his di- 
rection the Museum of Southwestern Biol- 
ogy grew in the size and significance of its 
collections and production of scholarly re- 
search. Together with Terry L. Yates, he 
was instrumental in obtaining National Sci- 


62 LAYNE AND HOFFMANN 


ence Foundation funding to upgrade the 
Museum’s mammal collection. 

Jim became a member of the ASM in 
1944 and besides the presidency has served 
as a Director and Vice-president. During his 
tenure as president, the decision was made 
for the society to participate in producing 
the first edition of Mammal Species of the 
World. In 1978, Jim was presented the C. 
Hart Merriam Award from the Society in 
recognition of his outstanding contributions 
to mammalogy. Among other honors he has 
received is the Leopold Conservation Award 
of the Nature Conservancy. Two species and 
one subspecies of mammals also have been 
named in his honor. 


Jocelyn Mary Taylor: 1982-1984 


J. Mary Taylor (Fig. 4) (she decided be- 
fore entering kindergarten not to use her 
first given name) was born in Portland, Or- 
egon, on May 30, 1931, to Kathleen and 
Arnold L. Taylor. She has an older brother, 
John Stewart Taylor. Growing up in that 
area, she had early and regular contacts with 
the out-of-doors through daily walks with 
her mother, which instilled in her a love of 
nature. However, being tall and athletic, she 
also developed a love of tennis, and toured 
the junior circuit, playing in a prestigious 
British tournament at age 17. She also played 
the piano and became devoted to chamber 
music. 

However, instead of pursuing the sport, 
she went East to Smith College, intending 
to study music. A biology course in her se- 
nior year of high school led her to take fur- 
ther biology courses in college and she 
switched her major to zoology, with an hon- 
ors thesis in protozoology. Mary received 
her bachelor’s degree from Smith in 1952 
and enrolled the same year at the University 
of California at Berkeley. She completed her 
master’s degree the next year, again spe- 
cializing in protozoology, and accepted a 
position in the Department of Zoology at 
Connecticut College. In 1954, upon receiv- 


ing a Fulbright Fellowship, she spent a year 
in Australia, which formed the basis for her 
life-long interest in Australasian mammals. 
Returning from her pre-doctoral fellowship, 
she enrolled again at Berkeley, completing 
her doctorate in mammalogy in 1959 and 
then accepting a position at Wellesley, not 
too far from her alma mater. She remained 
at Wellesley until 1965, collaborating with 
Betty Horner on research on Australian ro- 
dents and small marsupials. While at 
Wellesley she also began a long-term col- 
laboration with Dr. Helen Padykula of Har- 
vard Medical School on placentation in 
marsupials. 

In 1965 she heeded the call of the great 
Northwest, and returned across the conti- 
nent to the University of British Columbia 
in Vancouver, where she was the first wom- 
an to hold a professorial appointment in the 
Department of Zoology, and a curatorial one 
as Director of the Cowan Vertebrate Mu- 
seum. This was scientifically a particularly 
productive period of her career with many 
published papers and 10 graduate students. 
However, changing priorities at the univer- 
sity resulted in decreasing support for ver- 
tebrate zoology, and in 1982 Mary resigned 
her appointments and moved back to Port- 
land, this time with her husband, Dr. J. Wil- 
liam Kamp, an entomologist whom she met 
in British Columbia. 

She became a scientist at the Oregon Re- 
gional Primate Research Center in Beaver- 
ton, Oregon, and in addition held an hon- 
orary professorship at Oregon State 
University in Corvallis. Her move to Ore- 
gon also coincided with her election to the 
presidency of the ASM, the first woman to 
be so honored. 

In 1987, she became the first woman to 
become Director of the Cleveland Museum 
of Natural History. In 1989 she became 
Chairman of the Rodent Specialist Group, 
SSG/IUCN, and the following year was 
elected Vice-president of the Association of 
Science Museum Directors. A few years ago, 
she shepherded to completion a major ad- 
dition to the Museum that included ex- 


PRESIDENTS 63 


panded housing for the mammal collections 
(Source: Snow, 1987). 


Hugh Howard Genoways: 1984-1986 


Hugh H. Genoways (Fig. 4) was born on 
24 December 1940 at Scottsbluff, Nebraska, 
and grew up in the town of Bayard, Ne- 
braska, and on farms in the vicinity. As a 
youth he was active in the Boy Scouts of 
America with his father, and the scouting 
experience was a strong influence on his 
eventual decision to pursue a career in the 
field of biology. As an undergraduate at 
Hastings College in Nebraska, he was intro- 
duced to the science of biology by Professor 
Wendell Showalter. His interest in mam- 
mals was first awakened by another under- 
graduate professor, Gilbert Adrian, and it 
later matured under the guidance of Knox 
Jones, his major professor in graduate 
school. Hugh and Joyce Elaine Cox were 
married in 1963 and have a daughter, Mar- 
garet Louise, and son, Theodore Howard. 

Hugh received his A.B. in 1963 from Has- 
tings College and his Ph.D. from the Uni- 
versity of Kansas in 1971. While in graduate 
school, he was a part-time Instructor in the 
Department of Systematics and Ecology and 
a Research Assistant in the Museum of Nat- 
ural History. He also spent a year at the 
University of Western Australia. 

From 1971 to 1976 he held various ap- 
pointments on the faculty of Texas Tech 
University. These included Research As- 
sociate, Lecturer, Acting Coordinator of Re- 
search, and Curator of Mammals in the Mu- 
seum and Adjunct Assistant Professor of 
Veterinary and Zoological Medicine and 
Pathology in the School of Medicine. He 
was Curator of Mammals at the Carnegie 
Museum of Natural History from 1976 to 
1986. He assumed his present post as Di- 
rector of the University of Nebraska State 
Museum in 1986. He also holds appoint- 
ments as Professor of the Museum and Mu- 
seum Studies, Chair of the Museum Studies 
Program, Courtesy Professor in the School 


of Biological Sciences, and Faculty Fellow 
of the Graduate College. 

Hugh’s research has centered on the sys- 
tematics, biogeography, and ecology of New 
World mammals, with emphasis on ro- 
dents, particularly heteromyids and geo- 
myids, and bats. He has conducted field 
work in most regions of the United States 
and throughout the Caribbean, as well as 
Suriname, Mexico, Nicaragua, Venezuela, 
India, and Australia. In addition to numer- 
ous journal papers and other publications, 
he is author, coauthor, or editor of eight 
volumes, including Systematics and Evo- 
lutionary Relationships of the Spiny Pocket 
Mice of the Genus Liomys; Biological In- 
vestigations in the Guadalupe Mountains 
National park, Texas; Mammalian Biology 
in South America; and Contributions in 
Quaternary Vetebrate Paleontology. He also 
has been active in the development of tech- 
niques for use of computers in data analysis 
and retrieval and in museum collections 
management. 

Hugh became a member of the ASM in 
1963. In addition to the presidency, he has 
served as a Director, Vice-president, Man- 
aging Editor of the Journal of Mammalogy, 
and Editor of Special Publications, as well 
as chair/member of seven standing com- 
mittees. One of his major actions as presi- 
dent was the establishment of the Future 
Mammalogists Fund. He also focused on 
strengthening the committees and broad- 
ening their membership. The decision for 
ASM participation and the planning for an 
international meeting with the Mexican 
Mammal Society also took place during his 
term. 

Offices he has held in other scientific or- 
ganizations include the presidency of both 
the Southwestern Association of Naturalists 
and the Nebraska Museums Association. He 
also served as Publications Editor of the 
Carnegie Museum of Natural History and 
Editor of Museology published by Texas 
Tech University and is presently Editor-in- 
chief of Current Mammalogy. 

Hugh is a recipient of the C. Hart Mer- 


64 LAYNE AND HOFFMANN 


riam Award from the society and other hon- 
ors include a Fulbright Grant while a grad- 
uate student, selection by the United States 
Jaycees as one of the Outstanding Young 
Men in America, and election as a Fellow 
of the Center for Great Plains Studies of the 
University of Nebraska. 


Don Ellis Wilson: 1986-1988 


Don E. Wilson (Fig. 4) was born 30 April 
1944 in Davis, Oklahoma, and during his 
youth lived in Nebraska, Texas, Oregon, 
Washington, and Arizona. He developed an 
interest in natural history at an early age, 
and by the time he entered college had de- 
cided on a career in biology. While an un- 
dergraduate he worked for the National Park 
Service in Grand Canyon National Park and 
made his first trip to the tropics. He also 
spent a summer as a naturalist for the U.S. 
Forest Service in the Sandia Mountains of 
New Mexico. In 1962 he married Kathleen 
Hayes, and they have two daughters, Wendy 
and Kristy. 

Don received a B.S. in Wildlife Manage- 
ment from the University of Arizona in 1965 
and did his graduate work (M.S., 1967; 
Ph.D., 1970) at the University of New Mex- 
ico under the direction of James Findley. 
He held a Postdoctoral Fellowship in Ecol- 
ogy from the University of Chicago in 1970— 
1971 and also obtained advanced training 
in statistics, computer systems, automatic 
data processing, editing, and personnel and 
financial management through programs in 
the U.S. Fish and Wildlife Service, Smith- 
sonian Institution, and the U.S. Depart- 
ment of Agriculture Graduate School. 

In 1971 he was appointed Zoologist in 
the Bird and Mammal Laboratories of the 
U.S. Fish and Wildlife Service at the Na- 
tional Museum of Natural History and in 
1973 became Chief of the Mammal Section 
of the National Fish and Wildlife Labora- 
tory. In 1978, he became Chief of the Mu- 
seum Section of the Denver Wildlife Re- 
search Center and in 1984 was promoted to 


Chief of the Biological Survey. He was ap- 
pointed to his present post, Director of the 
Biodiversity Programs of the Smithsonian 
Institution, in 1990. He also has appoint- 
ments as Visiting Professor at the Univer- 
sity of Maryland, George Mason Univer- 
sity, and the Organization for Tropical 
Studies— Universidad de Costa Rica. 
Don’s principal research interests have 
been the biology of neotropical bats and 
tropical ecology in general. His studies of 
bats have ranged over a broad spectrum of 
topics, including taxonomy, distribution, 
community ecology, physiology, and repro- 
ductive biology. He has also worked on a 
variety of other mammals, including ro- 
dents, lagomorphs, carnivores, ungulates, 
and marsupials. In addition to mammal re- 
search, he has conducted studies on birds, 
amphibians, and reptiles, as well as seed 
predation, tropical strand plants, and in- 
sects. He is a coauthor of Mammals of New 
Mexico and author or coauthor of chapters 
in a number of volumes, including Biology 
of Bats of the New World Family Phyllos- 
tomatidae, Wild Mammals of North Amer- 
ica, Advances in the Study of Mammalian 
Behavior, Costa Rican Natural History, Bi- 
ology and Management of the Cervidae, 
Ecological and Behavioral Methods for the 
Study of Bats, and Tropical Rain Forest 
Ecosystems. He also coedited the second 
edition of the ASM-sponsored Mammal 
Species of the World published in 1993. 
Don became a member of the ASM in 
1966. Besides the presidency, he has served 
the society as a Director, Vice-president, 
Journal Editor, and Managing Editor of 
Mammalian Species and Special Publica- 
tions. Offices held in other professional or- 
ganizations include President of the Asso- 
ciation for Tropical Biology; Councilman 
and Treasurer of the Biological Society of 
Washington; Board of Managers, Vice-pres- 
ident, and President of the Washington Field 
Biologists Club; Board of Scientific Direc- 
tors of Bat Conservation International; and 
Board of Directors of Integrated Conser- 
vation Research. He also serves on the Chi- 


PRESIDENTS 65 


roptera Specialists Group of the IUCN Spe- 
cies Survival Commission and is Associate 
Editor of Revista de Mastozoologia Mexi- 
cana and a member of the editorial boards 
of Acta Zoologica Mexicana, Current Mam- 
malogy, and Anales del Instituto de Biolo- 
gia. In addition, he has served as a consul- 
tant to many U.S. and foreign governmental 
agencies and nongovernmental organiza- 
tions. 

Among his professional honors and 
awards are membership in Phi Sigma; Na- 
tional Science Foundation Predoctoral and 
Postdoctoral Fellowships; the Smithsonian 
Institution Award for Excellence in Tropi- 
cal Biology; Outstanding Publication Award 
from the Denver Wildlife Research Center; 
Reconocimiento, Asociacion Mexicana de 
Mastozoologia, and the Centennial Distin- 
guished Alumni Award from the University 
of New Mexico. 


Elmer Clea Birney: 1988-1990 


Elmer C. Birney (Fig. 4) was born on 26 
March 1940 at Satanta, Kansas, and grew 
up on a wheat farm and cattle ranch in a 
staunchly conservative family with a strong 
work ethic and belief in education as a key 
to one’s future. Sports were his major in- 
terest as a youth, and the closest he came 
to a biological interest was crossbreeding 
rare breeds of chickens to determine what 
combination of phenotypic traits were pro- 
duced in the hybrids. The possibility of ob- 
taining a football scholarship got him think- 
ing seriously about attending college. After 
his first year as an agriculture major, he en- 
listed in the Naval Reserve and served 2 
years on active duty on a destroyer in the 
Pacific. While on leave in 1960, he met 
Marcia Fayla McVey, and they were mar- 
ried in 1961, a day after his release from 
active duty. They have a daughter, Amelia 
Joleen, and son, Clayton Eugene. 

Elmer received his B.S. in 1963 and MLS. 
in 1965 from Fort Hays State University, 
where he developed his interest in mam- 


mals under the influence Gene Fleharty. He 
obtained his Ph.D. in 1970 from the Uni- 
versity of Kansas. His major professor at 
Kansas was Knox Jones, who played a strong 
role in his professional development. His 
doctoral research was supported by a Na- 
tional Science Foundation Traineeship and 
a Watkins Natural History grant. 

Elmer served as instructor of biology at 
Kearney State College in Nebraska in 1965- 
1966. After receiving the Ph.D., he joined 
the faculty of the University of Minnesota, 
where he is presently Curator of Mammals 
in the Bell Museum of Natural History and 
Professor in the Department of Ecology, 
Evolution, and Behavior. In addition, he 
has served as Director of Graduate Studies 
in both the Ecology Graduate Program and 
Zoology Graduate Program of the Depart- 
ment of Ecology, Evolution, and Behavior. 
He was Director of the Bell Museum from 
1990 to 1992. 

Elmer’s research has spanned a broad 
range of topics, including distribution, life 
history, ecology, systematics, physiology, 
and biochemistry. Among his important 
contributions are studies on the evolution 
of the enzyme systems involved in ascorbic 
acid biosynthesis in vertebrates conducted 
jointly with Robert Jenness and investiga- 
tions on the relationship of vegetative cover 
to microtine cycles and other aspects of 
grassland mammal communities with W. E. 
Grant, D. D. Baird, N. R. French, and oth- 
ers. The geographic focus of his work has 
been on mammals of central and western 
United States, and he was a participant in 
the U.S.I.B.P. Grassland Biome studies. He 
also has conducted substantial field work in 
Mexico, Argentina, Australia, and Antarc- 
tica. His publications include coauthorship 
of two books, The True Prairie Ecosystem 
and Handbook of Mammals of the North- 
central States, the chapter Community 
Ecology in Biology of New World Microtus, 
and the section on mammals in Minnesota’s 
Endangered Flora and Fauna. His work has 
been supported by grants from many 
sources, including the National Science 


66 LAYNE AND HOFFMANN 


Foundation, Society of Sigma Xi, Nongame 
and Minerals divisions of Minnesota Nat- 
ural Resources, Institute of Museum Ser- 
vices, and the Legislative Commission on 
Minnesota Resources. 

Elmer became a member of the ASM in 
1963 and, besides the presidency, has served 
as a Director, Vice-president, Managing Ed- 
itor, Journal Editor, and Editor for Special 
Publications. He has been a member of a 
number of standing committees and chaired 
the Membership, Editorial, and Develop- 
ment committees. Among his goals as pres- 
ident were to facilitate more open discus- 
sion of matters of concern to the 
membership, increase participation of 
women on committees and in other busi- 
ness of the society, and encourage election 
of younger members to the Board of Direc- 
tors. He is a Life Member of the South- 
western Society of Naturalists and member 
ofa number of other international, national, 
and regional scientific and environmental 
organizations. 


James Hemphill Brown: 1990-1992 


James H. Brown (Fig. 5) was born on 25 
September 1942 at Ithaca, New York, and 
grew up in rural upstate New York. His 
mother fostered his early interest in keeping 
pets and making natural history collections 
and when he was 8 or 9 years old introduced 
him to W. J. Hamilton, Jr. His association 
with Bill Hamilton during his elementary, 
high school, and university undergraduate 
years greatly influenced his development as 
a scientist. Between the ages of 10 and 12, 
he became acquainted with Kyle Barbeh- 
enn, then a graduate student of W. Robert 
Eadie, and regularly accompanied him in 
his field work on small rodent populations 
near the Browns’ home. This experience, 
particularly Kyle’s patience in answering 
questions and treatment of him as a fellow 
scientist, contributed greatly to shaping 
Jim’s interests in mammalogy and ecology. 
Jim and his wife, the former Astrid R. Ko- 


dric, were married in 1965 and have a son, 
Kevin, and daughter, Karen. Astrid is a sci- 
entific colleague as well as wife, and she and 
Jim have collaborated in a number of stud- 
ies. They have mutual interests in hiking; 
camping; reading; and collecting American 
Indian baskets, rugs, and pottery. 

Jim did his undergraduate work at Cor- 
nell, receiving the B.A. (with honors in zo- 
ology) in 1963. He received the Ph.D. in 
zoology from the University of Michigan in 
1967. Emmet Hooper was his major pro- 
fessor, but he also worked closely with Bill 
Burt and William Dawson. Supported by a 
Rackham Postdoctoral Fellowship from the 
University of Michigan, he did physiolog- 
ical ecology research with George A. Bar- 
tholomew at the University of California at 
Los Angeles in 1967-1968. 

He held faculty positions at the Univer- 
sity of California at Los Angeles (1968- 
1971), the University of Utah (1971-1975), 
and the University of Arizona (1975-1987). 
In 1987 he was appointed Professor of Bi- 
ology at the University of New Mexico. 
During his university career he has directed 
the work of 12 master’s students, 29 Ph.D. 
students, and 7 postdoctoral students. Jim’s 
primary research interests are the patterns 
and processes that influence the abundance, 
distribution, and diversity of species. Al- 
though mammals, particularly desert ro- 
dents, have been the principal subjects of 
his research, he has also worked on birds, 
fishes, insects, and plants. Among his stud- 
ies that have received broad recognition is 
the long-term experimental field investiga- 
tion of the interactions between granivorous 
mammals, birds, ants, and seed-producing 
plants in the Chihuahuan Desert of south- 
eastern Arizona. He also has worked in many 
other parts of the Southwest, as well as Mex- 
ico. In addition to numerous journal papers 
and chapters in books, he is coauthor of 
Biogeography and coeditor of Foundations 
of Ecologyand Biology of the Heteromyidae. 

He became a member of ASM in 1965 
and, in addition to the presidency, has served 
as a Director, Vice-president, and member 


PRESIDENTS 67 


James H. Brown James L. Patton 
(1990-1992) (1992-1994) 


Fic. 5.—Above: Presidents of the ASM from 1990 to 1994, James H. Brown and James L. Patton. 
Below (I-r): Early ASM presidents V. O. Bailey and C. H. Merriam with fellow members of the 
Bureau of Biological Survey, T. S. Palmer and A. K. Fischer, in the field at Lone Pine, Owens Valley, 
California, 13 June 1891. 


68 LAYNE AND HOFFMANN 


of a number of standing committees. As 
president he promoted a thorough reorga- 
nization of the editorial policies and format 
of the Journal of Mammalogy and initiated 
the policy of evaluating presentation of stu- 
dent papers at the annual meeting. He has 
been an active participant in other scientific 
societies, serving as Vice-president of the 
Ecological Society of America, President of 
the American Society of Naturalists, Coun- 
cil member of the Society for the Study of 
Evolution, and in editorial capacities for 
Ecology, Evolutionary Ecology, and the 
Journal of Biogeography. He served as a 
member of the Scientific Advisory Board of 
the American Museum of Natural History’s 
Southwestern Research Station and on the 
Population Biology and Physiological Ecol- 
ogy panels of the National Science Foun- 
dation. 

He is a recipient of the C. Hart Merriam 
Award from the ASM and a Fellow of the 
American Association for the Advance- 
ment of Science. In addition, he was award- 
ed a Certificate of Merit from the South- 
western and Rocky Mountain Division of 
the American Association for the Advance- 
ment of Science and a Guggenheim Fellow- 
ship in 1991-1992. 


James Lloyd Patton: 1992-1994 


James L. Patton (Fig. 5) was born in St. 
Louis, Missouri, on 21 June 1941. His fa- 
ther was a physician who served in the U.S. 
Army during World War II and the Korean 
War. Jim spent much of his early years as 
an “army brat” at various military bases in 
the United States and Germany. He was 
married to Carol Porter in 1966. They com- 
bined their honeymoon with the ASM 
meeting in Long Beach, which was both his 
first national meeting and the first time he 
presented a professional paper. 

Jim’s initial scientific interest was an- 
thropology, which was his undergraduate 
major at the University of Arizona. He 


planned to continue on in anthropology for 
his master’s, but after being exposed to a 
course in evolutionary genetics taught by 
William Heed he switched from anthro- 
pology to zoology and began work on chro- 
mosomal inversion polymorphisms in des- 
ert fruit flies (Drosophila). During this 
period, he met and began to interact with 
Al Gardner, one of Lendell Cockrum’s grad- 
uate students at the time, and as a result 
“discovered” small desert mammals and 
went on to complete both his master’s and 
doctorate on cytogenetics of pocket mice. 

He received all of his degrees from the 
University of Arizona: the B.A. (with dis- 
tinction) in 1963, M.S. in 1965, and Ph.D. 
in 1969. During his doctoral work, he held 
a National Defense Education Act Title IV 
Fellowship. 

In 1969, he was appointed Assistant Pro- 
fessor in the Department of Zoology and 
Assistant Curator in the Museum of Ver- 
tebrate Zoology of the University of Cali- 
fornia, Berkeley. He was promoted to As- 
sociate Professor and Associate Curator in 
1974 and Professor and Curator in 1979. 
He has been Associate Director of the Mu- 
seum of Vertebrate Zoology since 1982 and 
was Acting Director in 1988-1989 and 1992. 

Jim’s research has had two major themes: 
population genetics and geographic diver- 
gence in pocket gophers in western United 
States and systematics of neotropical mam- 
mals. However, he has also published on 
such diverse topics as chromosome evolu- 
tion in caecilians, biochemical relationships 
of Galapagos giant tortoises, genetic varia- 
tion in Galapagos finches, and distribution 
patterns of amphibians and reptiles in 
southern Peru. In addition to numerous 
journal papers, he is the author or coauthor 
of numerous papers and chapters in sym- 
posium volumes and books, including 
Mammal Studies of South America, Pat- 
terns of Evolution in Galapagos Organisms, 
Annual Review of Ecology and Systematics, 
Evolution in the Galapagos, and Mamma- 
lian Dispersal Patterns. 

Jim became a member of the ASM in 


PRESIDENTS 69 


1963. In addition to the presidency, he 
served on the Board of Directors, as First 
and Second Vice-president, and as Editor 
for Reviews of the Journal of Mammalogy. 
Posts held in other scientific societies in- 
clude Councilor and member of the Edi- 
torial Board, Society of Systematic Zoology; 
Second Vice-president of the Society for the 
Study of Evolution and Associate Editor of 
Evolution; and Associate Editor of Geneti- 
ca. He also has served on the Board of Di- 
rectors of the Charles Darwin Foundation; 
Board of Overseers of the Museum of Com- 
parative Zoology; several National Science 
Foundation panels and committees; as well 
as the editorial boards of the University of 
California Publications in Zoology, Current 
Mammalogy, and Israel Journal of Zoolo- 
gy. He is a Charter Member of the Society 
for Conservation Biology. 

Jim received the C. Hart Merriam Award 
from the ASM in 1983 in recognition of his 
outstanding contributions to mammalogy 
and is a Fellow of both the California Acad- 
emy of Sciences and American Association 
for the Advancement of Science. His honors 
also include a Distinguished Teaching 
Award from the University of California, 
Berkeley, and appointment as Distin- 
guished Visiting Scientist in the Museum of 
Zoology of the University of Michigan and 
Miller Professor in the Miller Institute for 
Basic Research of the University of Cali- 
fornia, Berkeley. He is a Research Associate 
in the Department of Mammalogy of the 
American Museum of Natural History and 
the Museum of Southwestern Biology of the 
University of New Mexico. 


Acknowledgments 


We especially thank the living past-presidents 
of ASM for providing us with background ma- 
terial on their careers in mammalogy and regret 
that space limitations prevented us from includ- 
ing many of the interesting details in the bio- 
graphical sketches. However, all material sup- 
plied will be placed in the society archives. We 


also thank Judith Eger for reviewing the account 
for R. L. Peterson; Elizabeth Peterson for pro- 
viding information on her father, W. P. Taylor; 
the Mammal Department of the American Mu- 
seum of Natural History for the source materials 
for the accounts of H. E. Anthony and R. G. Van 
Gelder; K. Grimes, Public Affairs Office, Amer- 
ican Museum of Natural History, and W. Deiss 
and E. Glenn, Smithsonian Institution Archives, 
for providing photographs; and the editors of this 
volume for their patience in awaiting completion 
of the manuscript. 


Literature Cited 


ALDRICH, J. W. 1977. In memoriam: Hartley Harrad 
Thompson Jackson (1881-1976). Journal of Mam- 
malogy, 58:691-694. 

ANoNYMous. 1923. American Society of Mammal- 
ogists articles of incorporation. Journal of Mam- 
malogy, 4:271-272. 

1942. Marcus Ward Lyon, Jr. 1875-1942. 

The American Midland Naturalist, 27(3):i-xv. 

. 1958a. Chairman of museum’s mammal de- 

partment retires after forty-seven years on staff. 

American Museum of Natural History (news re- 

lease), 13 April 1958. 

1958b. Harold Anthony retires. The Grape- 


vine (American Museum of Natural History), 15: 
1-2. 


. 1970. Noted mammalogist dies in California. 
American Museum of Natural History (news re- 
lease), 30 March 1970. 

. 1988. Emmet T. Hooper. Peromyscus News- 

letter, No. 6:9-11. 

. 1992. Emmet T. Hooper. [unidentified news- 
paper obituary, July 2, 1992]. 

BARBOUR, T., B. LAWRENCE, W. E. SCHEVILL, S. L. 
WASHBURN, AND M. B. Coss. 1943. Glover Morrill 
Allen 1879-1942. Journal of Mammalogy, 24:297- 
304. 

BEHLE, W. H. 1977. In memoriam: Stephen David 
Durrant (1902-1975). Journal of Mammalogy, 58: 
111-118. 

Cottam, C. 1971. TAYLOR, WALTER PENN (1888-). 
Pp. 317-318, in Leaders of American Conservation 
(H. Clepper, ed.). Ronald Press, New York, 353 pp. 

EGER, J., AND L. MITCHELL. 1990. Randolph Lee Pe- 
terson: 1920-1989. Journal of Mammalogy, 71:707- 
713. 

FINDLEY, J. S., AND J. K. Jones, JR. 1989. Eugene 
Raymond Hall: 1902-1986. Journal of Mammalogy, 
70:455-458. 

Go.tpman, E. A. 1935. Edward W. Nelson—natu- 
ralist, 1855-1934. Auk, 52:135-148. 

Grecory, W. K. 1930a. William Diller Matthew 
1871-1930. Natural History, 30:664-666. 

19306. William Diller Matthew, paleontol- 

ogist (1871-1930). Science, 1878:642-645. 

. 1931. A review of William Diller Matthew’s 


70 LAYNE AND HOFFMANN 


contributions to mammalian palaeontology. Amer- 
ican Museum Novitates, 473:1-22. 

GRINNELL, H. W. 1943. Bibliography of Clinton Hart 
Merriam. Journal of Mammalogy, 24:436-457. 

Hai, E. R. 1939. Joseph Grinnell (1877 to 1939). 
Journal of Mammalogy, 20:409-417. 

HOFFMEISTER, D. F. 1969. A history of the American 
Society of Mammalogists. Program of the 50th An- 
niversary Celebration of the American Society of 
Mammalogists, pp. 8-12. 

Huser, W. 1940. Witmer Stone (1866-1939). Jour- 
nal of Mammalogy, 21:1-4. 

Jackson, H.H.T. 1947. Association and correspon- 
dence with Goldman. Journal of Mammalogy, 28: 
109-113. 

. 1948. An earlier American mammal society. 
Journal of Mammalogy, 29:1-4. 

[Jones, C.]. 1990. Additions to the obituary of E. 
Raymond Hall. Journal of Mammalogy, 71:117. 
Lantis, M. 1954. Edward William Nelson. Anthro- 
pological Papers of the University of Alaska, 3(1): 

5-16. 

Layne, J. N., AND J.O. WHITAKER, JR. 1992. William 
John Hamilton, Jr.: 1902-1990. Journal of Mam- 
malogy, 73:693-706. 


LEHMANN, V. W. 1972. Walter P. Taylor, 1888-1972. 
Journal of Wildlife Management, 36:1379-1380. 
Litre, L. 1968. Alfred Brazier Howell, 1866-1961. 

Journal of Mammalogy, 49:732-742. 

Muut, I. 1990. William Henry Burt: 1903-1987. 
Journal of Mammalogy, 71:113-116. 

Oscoop, W.H. 1943. Clinton Hart Merriam — 1855- 
1942. Journal of Mammalogy, 24:421-436. 

SALT, G. W., AND R. L. Rupp. 1975. Tracy Irvin 
Storer (1889-1973)—an appreciation. Journal of 
Mammalogy, 56:538-552. 

SANBORN, C.C. 1948. Wilfred Hudson Osgood: 1875- 
1947. Journal of Mammalogy, 29:95-112. 

SETZER, H. W. 1977. A. Remington Kellogg—1892- 
1969. Journal of Mammalogy, 58:251-253. 

Snow, J. C. 1987. The road taken by a woman of 
science. Beacon [magazine], Sept. 20. Pp. 7-8,10. 
Taytor, W. P. 1947. Edward A. Goldman. Journal 

of Mammalogy, 28:114. 

TyLer, W. M. 1943. In memorium: Glover Morrill 
Allen. Auk, 60:163-168. 

Youna, S. P. 1947. Edward Alphonso Goldman: 
1873-1946. Journal of Mammalogy, 28:91-109. 
ZAHNISER, H. 1942. Vernon Orlando Bailey. 1894— 

1942. Science, 96:6-7. 


AWARDEES 


J. MARY TAYLOR AND DUANE A. SCHLITTER 


Introduction 


| P iiceceaaee by the ASM of outstanding 
persons in the field of mammalogy 
began in the first year of the society’s ex- 
istence with establishment of Honorary 
Membership. A second award, the C. Hart 
Merriam Award, was created 54 years later 
with the goal of recognizing exceptional 
contributions to the discipline of mammal- 
ogy within the past decade. This award was 
the first to bear the name of an earlier ASM 
member, himself an exemplar of the intent 
of this award. The Hartley H. T. Jackson 
Award, also named for a pre-eminent mam- 
malogist, was established only 3 years later 
to honor ASM members who have provided 
long and outstanding service to the society. 
Today, the H. H. T. Jackson Award is by 
its very nature the only one of the three 
awards whose recipients must be ASM 
members. The awardees of these three pres- 
tigious forms of recognition and their many 
contributions to mammalogy and the ASM 
are discussed in the following accounts. 


Honorary Members 
Honorary Membership, the most es- 


teemed recognition by the ASM, was first 
bestowed on Joel Asaph Allen in the same 


wl 


year that the society was founded. This 
award was established to acknowledge a 
“distinguished record of achievement’’ to 
the science of mammalogy. The recipient 
receives a certificate signed by the President 
of the society. In 1957 the procedure was 
formalized by the establishment of an Hon- 
orary Membership Committee consisting of 
the five most recent past presidents. The 
committee member in his or her 7th year 
after leaving office chairs the committee for 
two years. A successful nominee requires 
unanimous approval by the Committee, 
recommendation by the Board of Directors, 
and majority approval by the members 
present at the annual meeting. 

Although not awarded every year, 58 
mammalogists have had this honor con- 
ferred on them through 1992, and on oc- 
casion more than one person may be rec- 
ognized in a given year. Of these, 20 are 
past presidents, 7 are also recipients of the 
Jackson Award, and 1 of the Merriam 
Award. Recipients come from the following 
countries: United States (38), England (4), 
Germany (3), Russia (3), France (2), Nor- 
way (1), Spain (1), Japan (1), Denmark (1), 
China (1), Poland (1), Finland (1), and Mex- 
ico (1). 

The average age at which recipients have 


ie: TAYLOR AND SCHLITTER 


received Honorary Membership is 70, rang- 
ing from 48 (Sokolov) to 86 (Stejneger). 


Joel Asaph Allen, 1919 


Born 19 July 1838 in Springfield, Mas- 
sachusetts; A.B., Lawrence Scientific School, 
Cambridge; Ph.D. (honorary), University of 
Indiana; died 29 August 1921 (Journal of 
Mammalogy, 3:254-258, 1922) (Fig. 1). 

J. A. Allen descended from families who 
traced their New England origins to the ear- 
ly 17th Century. Raised on a farm, Allen 
showed an early interest and aptitude for 
studies of nature. At each higher level of 
academic opportunity, Allen was fortunate 
enough to study with someone who en- 
couraged his interest in nature until, finally, 
he won a position at Lawrence Scientific 
School in Cambridge, Massachusetts, where 
he studied under Professor Louis Agassiz. 
This relationship with Agassiz and the Mu- 
seum of Comparative Zoology (MCZ) was 
the beginning of 70 years of museum as- 
sociations. 

The first of numerous field trips was an 
expedition to Brazil with Agassiz in 1865. 
After nearly a year in northern and eastern 
Brazil, and in possession of large quantities 
of all orders of vertebrates, mollusks and 
other invertebrates, as well as samples of 
the flora, Allen returned to Cambridge. Af- 
ter a short break on the farm to recover his 
health, Allen left again for the field to collect 
on Lake Ontario and in Michigan, Indiana, 
and Illinois. He returned to the MCZ to 
become curator of birds and mammals. 
Within a year he left for a collecting trip to 
Florida, and in 1871 he took a nine-month 
swing through the Great Plains and Rocky 
Mountains as far as the Great Salt Lake. 
Again, Allen’s collections included large 
numbers of mollusks, insects, crustaceans, 
Recent and fossil fishes, as well as the usual 
birds, bird eggs, and mammals. In 1873 he 
was appointed scientific chief of the survey 
of the Northern Pacific Railroad in North 
Dakota and Montana on behalf of MCZ and 


the Smithsonian Institution. Because of the 
presence of hostile Indians, use of firearms 
and side trips were prohibited and collecting 
was difficult. On one occasion, he was es- 
corted by General George Custer and 1,400 
troops. With the exception of a short trip 
to Colorado in 1882, primarily for purposes 
of restoring his health, his return from the 
railway survey to Cambridge in late 1873 
marked the end of field work for Allen. 

In 1885 Allen accepted a position as Cu- 
rator of birds and mammals at the Ameri- 
can Museum of Natural History. The mu- 
seum had entered a period of scientific focus 
intended to match the already famous ex- 
hibitions. During the next 36 years, Allen 
was to become an outstanding scientist and 
editor. From a combined 14,300 specimens 
of birds and mammals in 1885, Allen saw 
the collection grow to nearly 250,000 spec- 
imens by 1920. At the same time, he edited 
37 volumes of the Bulletin and 22 volumes 
of the Memoirs, a total of 21,368 pages, 
between 1885 and 1917. During this inter- 
val, Allen published 23 papers on birds and 
168 on mammals, describing and naming a 
total of 724 new taxa. 

Allen was a leader in the founding of the 
American Ornithologists’ Union and edited 
The Auk for 30 years. He virtually wrote 
The Code of Nomenclature of the American 
Ornithologists’ Union and was a member of 
the Commission for Zoological Nomencla- 
ture from its formation in 1910 until his 
death. He founded the Audubon Society. 
Receiving numerous medals and honors, 
Allen was a member of nearly all of the 
leading scientific societies of the world. He 
was a member of the National Academy of 
Sciences and an honorary member of the 
Zoological Society of London, British Or- 
nithologists’ Union, and the New Zoologi- 
cal Society. 

Although plagued by periods of poor 
health, Allen continued his incredible pace 
until his death. His erudition and produc- 
tivity inspired universal affection and rev- 
erence. No more suitable person could have 
been honored as the first honorary member 


AWARDEES 73 


Max Weber? M. R. Oldfield Thomas 
(1919) (1928) (1928) 


William Berryman Scott‘ Alfred W. Anthony* 
(1929) (1936) (1936) 


da 32% 
Leonhard Stejneger® Gerrit S. Miller, Jr.' Angel Cabrere Latorre® 
(1937) (1941) (1947) 


Fic. 1.—Honorary members of the ASM, 1919-1947. Courtesy of: «Artis Library, University of 
Amsterdam; *Department of Library Sciences, American Museum of Natural History; ‘Department 
of Geological and Geophysical Sciences, Princeton University; ‘Natural History Museum, San Diego, 
California; ‘Photographic Collection, Smithsonian Institution; ‘Biographical Files, Smithsonian In- 
stitution; ‘National University of La Plata, Argentina. 


74 TAYLOR AND SCALITTER 


of the newly formed American Society of 
Mammalogists. He was also a Charter 
Member. Following his death, the Ameri- 
can Museum of Natural History named its 
new mammal hall the Allen Hall of North 
American Mammals. 


Edouard-Louis Trouessart, 1921 


Born 25 August 1842 in Angers, France; 
undergraduate degree, University of Poi- 
tiers, 1864; M.D., University of Paris, 1870; 
died 30 June 1927 (Journal of Mammalogy, 
$23924 1927), 

Trouessart was born into an academic 
family. His father, Joseph, was a professor 
of physics in the Faculty of Sciences at Poi- 
tiers. From the beginning, Edouard-Louis 
was interested in zoology and medicine. 
Upon graduation, he was sent with the 
French army as a physician to the 36th Reg- 
iment, Ist Battalion of the Vienne Mobile 
Guard, in the war with Germany (1870- 
1871). When peace was declared, he re- 
turned to the Maine-et-Loire district to hos- 
pital practice and work with the relief com- 
mittee. At the same time, he began his 
studies of natural history. Asa result of these 
activities, he was asked to collaborate in the 
production ofa Parisian journal of medicine 
and zoology, serving in this role from 1871 
to 1882. 

In 1882, he was appointed Director of the 
Museum of Natural History of Angers. He 
expanded activities of the museum in An- 
gers to include new public programs and 
natural history courses in the public schools. 
As such, he had a great impact on this town 
with his energy and advanced ideas. How- 
ever, budget cuts ultimately caused Troues- 
sart to leave Angers in 1885. He moved to 
Paris, where he began to produce a series of 
memoir volumes in natural history on a va- 
riety of subjects, including parasites, marine 
biology, mammals, birds, and medicine. 

In 1906, Trouessart was named Professor 
of mammalogy and ornithology at the Na- 
tional Museum of Natural History, Paris. 


In Paris he was able to continue production 
of monographs on a variety of subjects. The 
government also charged him with other 
duties, among them a study of the organi- 
zation and utilization of zoos in Belgium, 
Germany, and Holland, and a review of 
hunting in the colonies. 

Trouessart was a prolific author on a va- 
riety of subjects in medicine and zoology. 
Among his best know works are the mul- 
tivolume Les Mammiferes vivants et fossiles 
(1879-1907), series of papers entitled Les 
Acariens parasites et les acariens marins 
(1880-1907), La Faune des Mammiferes de 
France (1885), Les Microbes, les ferments et 
les moisissures (1886), La Géographie Zoo- 
logique (1890), Catalogue des Mammiferes 
vivants et fossiles (3 vols.) (1898, 1904), La 
Faune des Mammiféres de l’Algerie, du 
Maroc et de la Tunisie (1905), La Faune de 
Mammifeéres d’Europe (1909), and Cata- 
logue des Oiseaux d’Europe (1912). He col- 
laborated on the Grand Encyclopédie, Revue 
Scientifique, Nature, and Revue Générale des 
Sciences. 

Among the many honors and awards giv- 
en to Trouessart were the Knight of the Le- 
gion d’honneur and Knight of the Mérite 
Agricole. He was a corresponding member 
of the Zoological Society, London, and an 
officer of the Academy of Paris. He was 
awarded the Gold Medal from the Exposi- 
tion Universelle de Paris (1889) and the 
Dollfus Prize from the Société Entomolo- 
gique de France (1895). Although mam- 
malogists know Trouessart best for his many 
contributions on mammals, his breadth of 
study and level of productivity in both med- 
icine and zoology were even more impres- 
sive and unequalled in the modern day. 


Michael Rogers Oldfield Thomas, 1928 


Born 21 February 1858 in Millbrook, 
Bedfordshire, England; no university de- 
grees; died 18 June 1929 (Journal of Mam- 
malogy, 10:280, 1929) (Fig. 1). 

Oldfield Thomas began his career at age 


AWARDEES Tes. 


18 as aclerk at the British Museum, which 
then housed the natural history collections. 
After taking two years of lectures given by 
Thomas H. Huxley, Thomas’ hopes were 
realized in 1878 when he was appointed As- 
sistant in the Department of Zoology at the 
Museum, a position he held for 45 years. 

It was Dr. Albert Gunther, Keeper of Zo- 
ology, who steered Thomas to work on 
mammals. He spent most of his working 
life building up the collections of mammals 
in the Museum, taking them over from John 
Edward Gray, his predecessor from 1837 to 
1874. During Oldfield Thomas’ career, the 
biological collections were moved from the 
British Museum at Bloomsbury to the new 
Natural History Museum in South Ken- 
sington. It was then that Thomas started the 
enormous task of cataloging the mammal 
collections, beginning in 1888 with the Mar- 
supialia and Monotremata. 

Thomas became receptive to the new 
concept (developed in the 1890s by C. Hart 
Merriam and other American mammalo- 
gists) regarding the importance of series of 
specimens rather than the typological ap- 
proach. He responded accordingly by ac- 
quisition and study of widespread collec- 
tions. Thomas and his wife supported 
mammal collectors from all over the world 
and also financed collecting expeditions. 

Thomas is the author of almost eleven 
hundred published works, mostly descrip- 
tions of mammals, for which he proposed 
2,900 new taxa. Most of his publications 
appeared in the Annals and Magazine of 
Natural History or the Proceedings of the 
Zoological Society of London. He was elect- 
ed a Fellow of the Royal Society in 1901. 

Although officially retiring in 1923, Tho- 
mas continued his work. However, his wife 
died in mid-1928 and Oldfield Thomas ter- 
minated his life a year later. To this day, 
mammalogists owe him a debt, not only for 
his attempt to standardize skull measure- 
ments and tooth nomenclature, but for 
helping to frame the foundation that estab- 
lished the field of systematic mammalogy 
for the 20th Century. 


Max Weber, 1928 


Born 5 December 1852 in Bonn, Ger- 
many; M.D., University of Bonn, 1877; died 
7 December 1937 (Journal of Mammalogy, 
18:389-390, 1937) (Fig. 1). 

Max Weber studied biology and medicine 
at the University of Berlin and University 
of Bonn. In 1879, Weber accepted a posi- 
tion in Holland, first as Prosector of anat- 
omy at the University of Amsterdam. After 
a year, he moved to the University of 
Utrecht as Lecturer in anatomy. He re- 
turned to Amsterdam as Professor of anat- 
omy, a post he held until his retirement in 
1922. 

Weber became involved with various ex- 
peditions to the Dutch colonies. He was 
leader of the Netherlands Deep Sea Expe- 
dition of the ship Siboga to the East Indies. 
Weber studied the fishes from these expe- 
ditions and published on the zoogeography 
of the fauna. At the same time he became 
interested in cetaceans and, after several 
visits to whaling stations, completed a series 
of papers on the anatomy of whales. He 
established a program of salvage of stranded 
whales on the Dutch coast and utilized the 
carcasses for his anatomical studies. Be- 
cause the university was near the Amster- 
dam zoo, Weber was able to salvage many 
unique zoo animals for his dissections. The 
synoptic coverage of his efforts continued 
to expand. Weber studied the hairs of mam- 
mals, correlations between brain weight and 
body weight, and squamation. 

After making two trips to the East Indies, 
Weber went to South Africa in 1894. Nu- 
merous samples were brought back for his 
systematic and anatomical research. After 
a number of cooperative false starts on a 
compendium of the order Mammalia, We- 
ber completed a volume which he published 
in 1904. His Sdugetiere was a result of the 
accumulation of many years of data on 
anatomy, systematics, and paleontology, and 
it served as a single reference for many years. 
A more expanded second edition with two 
volumes was published in 1927. Although 


76 TAYLOR AND SCALITTER 


others collaborated with Weber on the sec- 
ond edition, it was still primarily his work. 
It was the first and most complete review 
of the field of mammalogy to date. This 
volume alone was sufficient to ensure Max 
Weber immortality in the field of mam- 
malogy. 


Henry Fairfield Osborn, 1929 


Born 8 August 1857 in Fairfield, Con- 
necticut; B.A., Princeton University, 1877; 
died 6 November 1935 (Journal of Mam- 
malogy, 17:84, 1936) (Fig. 1). 

Osborn was the son of William Henry 
Osborn, President of the Illinois Railroad. 
He attended private schools and Princeton 
University. Upon graduation, he was sent 
to England for additional studies at the Roy- 
al College of Science, London, and Cam- 
bridge University. 

After graduation from Princeton in 1877, 
Osborn began research in paleontology. In 
1881, he was appointed to the faculty of 
Princeton as an Assistant Professor of nat- 
ural science. Between 1883 and 1890, he 
was a Professor of paleontology at Princeton 
but, like most students of fossils, he divided 
his time equally with studies of anatomy. 
As a result of his anatomical research, he 
made significant contributions in the field 
of neurology. He received the De Costa chair 
of biology at Columbia University in 1890 
and held this chair until 1910. At about the 
same time (1891), he was appointed Curator 
of Vertebrate Paleontology at the American 
Museum of Natural History. He subse- 
quently served as President of the Museum 
from 1908 to 1932 and was directly re- 
sponsible for the museum’s emergence as a 
premier exhibit and research institution. He 
directed expeditions to various parts of 
North America, the Gobi Desert, Egypt, In- 
dia, and Samos Island. Osborn also served 
as vertebrate paleontologist on the geolog- 
ical surveys of both the United States (1900- 
1924) and Canada (1900-1904). 

Osborn received numerous honors and 


awards, including honorary degrees from 
Trinity College (1901), Princeton Univer- 
sity (1902), Cambridge University (1904), 
Columbia University (1907), the Univer- 
sity of Christiania (1911), Yale University 
(1923), Oxford University (1926), New York 
University (1927), Union College (1928), 
and the University of Paris (1931). 

During his career Osborn published near- 
ly 1,000 titles, many of which were lengthy 
memoirs in vertebrate paleontology. Among 
his best known works are: From the Greeks 
to Darwin (1894), The Age of Mammals 
(1910), The Origin and Evolution of Life or 
the Theory of Action, Reaction, and Inter- 
action (1917), Evolution and Religion (1923), 
The Titanotheres of Ancient Wyoming, Da- 
kota, and Nebraska (1929), and Cope: Mas- 
ter Naturalist (1931). At the time of his death 
at age 78, Osborn was actively completing 
a monograph on the Proboscidea. He ex- 
hibited the same enthusiasm and energy for 
all aspects of life and his work, but especially 
his studies on fossils, right up to his death. 


Alfred Webster Anthony, 1936 


Born 25 December 1865 in Cayuga Coun- 
ty, New York; Colorado School of Mines; 
died 14 May 1939 (Fig. 1). 

From New York, Anthony moved to Col- 
orado at age three. He followed in his fath- 
er’s profession as a mining engineer, moving 
to California and Baja California in search 
of gold. But, at the same time, he pursued 
his interest in birds and mammals. He made 
trips to islands off the Mexican coast where 
he studied seals, continued searching for gold 
in Alaska and Oregon, and even farmed for 
10 years in Oregon. 

In 1920, he became director of the San 
Diego Museum of Natural History. In 1924, 
he resigned to embark on a five-year col- 
lecting trip to Guatemala. Failing health 
caused him to return to San Diego. Except 
for a few short trips, he rarely strayed from 
San Diego for the remainder of his life. He 


AWARDEES 77 


did make a short visit to Ensenada, Mexico, 
to study the southern sea otter. 

Anthony was not a prolific writer, pre- 
ferring that others should study and report 
on his collections. His greatest contribution 
was the size and scope of the large collec- 
tions, primarily birds and mammals, that 
he made during his life. Being a mining en- 
gineer, he also made very important collec- 
tions of minerals while searching for gold. 

It should be noted that Alfred Anthony 
fathered a son, Harold E. Anthony, who was 
to became an eminent mammalogist in his 
own right at the American Museum of Nat- 
ural History, President of the American So- 
ciety of Mammalogists, and an Honorary 
Member like his father. In addition to his 
son, Alfred Anthony had a reputation of 
starting many other young budding natu- 
ralists in pursuit of careers in natural his- 
tory. He was a Charter Member of the ASM. 


William Berryman Scott, 1936 


Born 12 February 1858 in Cincinnati, 
Ohio; A.B., Princeton University, 1877, 
Royal School of Mines, London, Cambridge 
University; Ph.D., Heidelberg University, 
1880; died 29 March 1947 (Fig. 1). 

The great-great-great-grandson of Ben- 
jamin Franklin through his mother, Scott 
was born into a family of clergy and theo- 
logians, professors, and authors. Upon re- 
ceiving his doctorate in Germany, Scott re- 
turned to Princeton as an instructor in 
geology. He was awarded the Blair Chair in 
geology in 1884, and he held this position 
until his retirement in 1930. Although pri- 
marily interested in geology and paleontol- 
ogy, Scott also had conducted extensive re- 
search in embryology while at Cambridge 
and Heidelberg universities, publishing three 
monographs on newts and lampreys as a 
result of this work. His main research focus, 
however, was his studies of mammalian fos- 
sils of North and South America. Between 
1877 and 1897, he led 11 major collecting 


and exploratory trips to South Dakota, 
Montana, and Wyoming. In addition to his 
numerous short publications in many jour- 
nals, Scott published several books, includ- 
ing his well-known History of Land Mam- 
mals of the Western Hemisphere (1913), The 
Theory of Evolution (1917), and Physiog- 
raphy (1922). His textbook An Introduction 
to Geology (1897) had three editions, the 
last appearing in 1930. He was editor and 
author of fifteen quarto volumes on the re- 
sults of the “Princeton University Expedi- 
tions to Patagonia.” 

Scott received numerous honors during 
his lifetime. The University of Pennsylva- 
nia conferred an LL.D. degree on him in 
1906. He received Sc.D. degrees from Har- 
vard University (1909), Oxford University 
(1912), and Princeton University (1930). He 
was awarded the E. K. Kane Gold Medal 
of the Geographic Society of Philadelphia 
(1905), the Wollaston Gold Medal of the 
Geological Society of London (1910), the F. 
V. Hayden Medal of the Academy of Nat- 
ural Science, Philadelphia (1926), the Mary 
Clark Thompson Gold Medal of the Na- 
tional Academy of Sciences (1931), the 
Walker Grand Prize of the Boston Society 
of Natural History (1934), the Penrose 
Medal of the Geological Society of America 
(1936), and the Daniel Giraud Elliot Medal 
of the National Academy of Sciences (1940). 
He was a member of the National Academy 
of Sciences. 


Leonhard Stejneger, 1937 


Born 30 October 1851 in Bergen, Nor- 
way; Frederic’s University, Christiania, 
Norway; died 28 February 1943 (Journal of 
Mammalogy, 24:295, 1943) (Fig. 1). 

Steyneger received his undergraduate and 
postgraduate education at Frederic’s Uni- 
versity in his native Norway. Upon com- 
pletion of his education, he left Norway for 
the United States where, in 1881, he ac- 
cepted a position in the U.S. National Mu- 


78 TAYLOR AND SCHLITTER 


seum, a part of the Smithsonian Institution. 
In 1884 he became Assistant Curator of 
Birds, a position he held until becoming Cu- 
rator of Reptiles in 1889. In 1911 he became 
Head Curator of Biology, although continu- 
ing in his studies of reptiles and amphibians. 

Stejneger was associated with an out- 
standing cadre of naturalists who were 
working in Washington, D.C. at this time. 
Field work associated with the new Biolog- 
ical Survey was widespread and very active, 
resulting in large quantities of vertebrates 
coming to the National Museum. Leonard 
took advantage of the many opportunities 
to study these collections of birds, mam- 
mals, reptiles, and amphibians. His first love 
was herpetology, so as often as possible he 
accompanied some of the early expeditions 
of the Biological Survey as herpetologist and 
studied the herpetological collections made 
by many of the others. 

Stejneger’s contribution to mammalogy 
results primarily from his activities as a 
member of the Fur Seal Commission. He 
first became involved with fur seals as a 
member of the team sent in 1895 to the 
Commander Islands by the U.S. Fish Com- 
mission. The report from this team has 
served as the primary program for manag- 
ing fur seal resources of the North Pacific. 

Stejneger’s reputation as an all-round nat- 
uralist can be judged by his election to Hon- 
orary Membership in the ASM and to fellow 
of the American Ornithologists’ Union. 


Gerrit Smith Miller, Jr., 1941 


Born 6 December 1869 in Peterboro, New 
York; A.B., Harvard University, 1894; died 
24 February 1956 (Journal of Mammalogy, 
37:309, 1956) (Fig. 1). 

Miller, a shy and sensitive boy, grew up 
on a large estate in central New York and 
attended private schools. He developed a 
great interest in the animals living in the 
forests and fields of the estate. His great 
uncle, Greene Smith, who lived on the es- 


tate, was interested in birds and probably 
also greatly influenced Miller. 

After graduating from Harvard Univer- 
sity, Miller joined an aunt for a summer tour 
of Europe to attend the Wagnerian festival 
at Beireuth. Music was always to be a very 
important part of Miller’s life. Later, in 1894, 
he joined the Biological Survey in Wash- 
ington, D.C., where he remained working 
for C. Hart Merriam until 1898. At that 
time, he took a position as Assistant Curator 
of Mammals at the United States National 
Museum. He became Curator in 1909 and 
remained in that position until he retired at 
age 70 in 1940. During the intervening years, 
Miller became one of the outstanding mam- 
malogists of his time. Miller was a Fellow 
of the American Association for the Ad- 
vancement of Science, and a member of the 
American Academy of Arts and Sciences, 
the American Philosophical Society, and the 
Academy of Natural Sciences. 

Miller was married twice. Each of his 
wives was to influence his life greatly. His 
first wife, Elizabeth Page, was an older 
woman with three children when she mar- 
ried Miller in 1897. She was a quiet, reclu- 
sive woman. Rather than socialize, she and 
Miller found pleasure in scholarly activities. 
Miller spent his free time in his library. Af- 
ter a long illness, she died in 1920. During 
a trip to a meeting in Honolulu, Miller met 
Anne Chapin Gates. They were married in 
the summer of 1921. The Millers moved 
from Virginia to a hill overlooking the Na- 
tional Zoo. They socialized, and Miller came 
out of his private shell and interacted with 
people. He spent many hours observing pri- 
mate behavior at the zoo and even pub- 
lished on his observations. He was well 
known in mammalogy as an author of more 
than 400 papers, including monographs and 
the following books: The Families and Gen- 
era of Bats; Catalogue of the Land Mam- 
mals of Western Europe in the British Mu- 
seum; List of North American Land 
Mammals in the United States National 
Museum, 1911; List of North American Re- 
cent Mammals, 1923. 


AWARDEES 75!) 


Ernest Evan Thompson Seton, 1941 


Born 14 August 1860 in South Shields, 
England; Royal Academy School of Paint- 
ing and Sculpture, London; Julian Acade- 
my, Paris; died 23 October 1946. 

Born in England of Scottish parents, Se- 
ton moved to Canada at age five when his 
father settled on a farm in rural Ontario. 
When he was nine, Seton moved to Toron- 
to. However, four years of rural life had 
instilled in young Seton a love for nature 
that was to continue to grow throughout his 
life. Although his father was determined that 
Seton should be an artist, he was equally 
determined to become a naturalist. During 
a year of formal study at the Royal Academy 
in London in 1880, he was granted permis- 
sion to use the Natural History Library at 
the British Museum, a singular waiver of an 
inflexible rule. He returned to rural Mani- 
toba for further studies of wildlife. In late 
1883 he moved to New York City for a brief 
time to write short stories, but he soon re- 
turned to Manitoba. While in New York, 
he made numerous contacts relating to his 
sketches and drawings. Back in Manitoba, 
his arthritis began to impede his travels by 
foot so, in 1890, Seton left for Paris to study 
at the Julian Academy. His studies were in- 
terrupted by a trip to New Mexico in 1892 
to hunt wolves. 

During a voyage to France in 1894, Seton 
met Grace Gallatin. They were married in 
1896 and settled on an estate in New Jersey. 
At this time Seton was commissioned to 
illustrate Frank Chapman’s Bird Life. Se- 
ton’s career as an illustrator and writer flow- 
ered. He published his first collection of il- 
lustrated animal stories in 1898, Wild 
Animals I Have Known. The public clam- 
ored for his books. The Trail of the Sandhill 
Stag (1899), Lives of the Hunted (1901), 
Woodmyth and Fable (1905), Animal He- 
roes (1905), Life-histories of Northern Ani- 
mals (2 vols.) (1909), Wild Animals at Home 
(1913), Biography of a Grizzly (1918), Game 
Animals and the Lives They Live (1924), and 
Lives of Game Animals (4 vols.) (1925-1928) 


followed. Seton was also an accomplished 
writer and illustrator of popular children’s 
books. Among his best known are Two Lit- 
tle Savages (1903), Rolfin the Woods (1911), 
Woodcraft and Indian Lore (1912), Wild 
Animal Ways (1916), and Woodland Tales 
(1921). He was also a very successful lec- 
turer, giving animated lectures that cap- 
tured the attention of his audiences. 

From New York he moved to an estate 
in Connecticut, where he practiced some of 
his ideas on attracting waterfowl and raising 
furbearers. In 1930 he moved to a ranch 
near Santa Fe, New Mexico, where he lived 
the remainder of his life. He established the 
Seton Institute to promote an appreciation 
of traditional Indian customs and life. With 
his second wife, Julia M. Battree Moss, a 
noted Indian expert, he fathered a daughter 
at age 78. 

Seton was instrumental in establishing the 
Boy Scouts of America in 1910 and was 
honored as a Chief Scout. He received the 
John Burroughs Medal (1926) and the Dan- 
iel Girard Elliot Gold Medal (1928), the lat- 
ter from the National Academy of Sciences, 
for his book Game Animals and the Lives 
They Live. Seton was endowed with an 1n- 
fectious personality which allowed him to 
carry his messages of conservation and na- 
ture to large audiences. 


Rudolph Martin Anderson, 1947 


Born 30 June 1876 in rural Winneshiek 
County, Iowa; A.B. (1903) and Ph.D. (1906), 
University of Iowa; died 21 June 1961 
(Journal of Mammalogy, 42:444, 1961). 

Anderson had a passion for natural his- 
tory from his early youth. Birds were his 
first love, and his first publication at age 17 
was entitled The Marsh Hawk. His first 
book, The Birds of Iowa, was published in 
1907. He spent as much time as possible 
studying the natural history of birds, es- 
pecially nesting habits. 

The physically large and strong young 
Anderson was attracted to athletics, es- 


80 TAYLOR AND SCHLITTER 


pecially track and field where he won nu- 
merous medals and awards at the univer- 
sity. At this same time he was a member of 
the Cadet Corps, was sent to the Spanish- 
American War in 1898, and served in the 
Iowa and Missouri national guards between 
1900 and 1908. 

Anderson’s professional career began with 
a position as field agent and mammalogist 
on the American Museum of Natural His- 
tory’s expedition to Arctic Alaska and the 
Yukon and Northwest Territories from 
1908-1912. Anderson had found his call- 
ing. He was selected second in command of 
the Stefanson-Canadian Arctic Expedition 
from 1913-1916 and was appointed mam- 
malogist at the National Museum of Canada 
in 1913. He continued with the Museum 
and became Chief of the Biology Division 
in 1920, a position he held until his retire- 
ment in 1946. 

During his career, Anderson conducted 
or supervised field work and research at sites 
in all provinces and territories of Canada, 
many of them in Arctic and mountainous 
areas under extremely difficult conditions. 
Over the years, administrative duties cut 
severely into the time he had available for 
field work. However, he persisted by help- 
ing his assistants and students carry on the 
field work he loved so much. By 1929, An- 
derson had spent seven winters and ten 
summers north of the Arctic Circle. He ed- 
ited and partially wrote the 16 volumes and 
64 papers resulting from the Canadian Arc- 
tic Expedition. Anderson published 134 pa- 
pers and books. Noteworthy books are 
Methods of Collecting and Preserving Ver- 
tebrate Animals (1932), one of the first and 
most complete of its kind ever published, 
and Catalogue of Canadian Recent Mam- 
mals (1946). 

Anderson was a member of 11 profes- 
sional societies and associations and an 
honorary member or fellow of six more, in- 
cluding the American Association for the 
Advancement of Science and the Royal So- 
ciety of Canada. He was a Charter Member 
of the ASM. Anderson served as a consul- 


tant or committee member of many gov- 
ernmental agencies from Mines and Re- 
sources, Parks and Forests, and Wildlife 
Protection, to Library. 

Anderson was a gentle and quiet-spoken 
person with a keen sense of humor. As ev- 
idence, one need only read his account of 
Homo sapiens in the Catalogue of Canadian 
Recent Mammals. 


Angel Cabrera Latorre, 1947 


Born 19 February 1879 in Madrid, Spain; 
Ph.D., University of Madrid, 1902; died 7 
July 1960 (Journal of Mammalogy, 41:540, 
1960) (Fig. 1). 

Upon completion of his doctorate, Ca- 
brera accepted an honorary position as As- 
sistant Curator with the National Museum 
of Natural Sciences. He was to continue his 
association with this museum until 1925, 
when he moved to Argentina. Cabrera’s first 
task was a review of the mammals of the 
Iberian Peninsula. He published this work 
as Fauna Ibérica: Mamiferos (1914). He next 
began a review of genera of mammals, which 
he published from 1919 through 1925 as 
Genera Mammalium. In 1922, he produced 
the first Spanish manual of mammalogy 
(Manual de Mastozoologia). During this in- 
terval he began a study of the mammals of 
Morocco, part of which was a Spanish col- 
ony. This was published in 1932 as Los 
Mamiferos de Marruecos. 

Cabrera accepted a position at the Na- 
tional University of La Plata, Argentina, in 
1925. He was to focus the remainder of his 
career on neotropical mammalogy. From 
1927 to 1947, he was Professor and Head 
of the Department of Paleontology in the 
La Plata Museum and held the Chair of 
Professor of Zoology at the School of Ag- 
riculture and Veterinary Medicine of the 
University of Buenos Aires. His research 
and field work centered on fossil and Recent 
mammals of Argentina. In 1940 Cabrera 
published the first of his momentous re- 
views of South American mammals. His 


AWARDEES 81 


i. 


Theodore S. Palmer? Edward A. Preble” | William K. Gregory‘ 
(1951) (1952) (1954) 


4 


Albert R. Shadle* Magnus A. Degerbol* Stanley P. Young 
(1956) (1962) (1964) 


Z ss Nw / 4 ae 
Erna Mohr‘ Kazimierz Petrusewicz Charles S. Elton® 
(1966) (1972) (1973) 


Fic. 2.—Honorary members of the ASM, 1951-1973. Courtesy of: **The American Society of 
Mammalogists Records, Smithsonian Institution; ‘Department of Library Sciences, American Mu- 
seum of Natural History; ‘Roswell Park Memorial Institute; ‘Zoological Museum, University of 
Copenhagen; ‘Zoologisches Institut und Zoologisches Museum, Hamburg, Germany; *Ken Marsland. 


82 TAYLOR AND SGHLITTER 


Mamiferos Sudamericanos was a collabo- 
rative effort with José Yepes. This was fol- 
lowed by the first volume of Catalogo de los 
Mamiferos de la America del Sur (1958). 
Although Cabrera died in 1960 before the 
completion of the second volume, it was 
sufficiently complete so as to be published 
that same year. 

Cabrera was a prolific writer, authoring 
218 papers on mammals, 27 books on 
mammals, and more than 400 popular ar- 
ticles. In addition he often illustrated his 
work, especially his popular articles, with 
his own watercolors and other forms of art. 
His talent and hard work were rewarded 
with numerous honors and distinctions. 
Most significant were his election as Cor- 
responding Member (1907) and Honorary 
Foreign Member (1947) of the Zoological 
Society of London; Member of the Inter- 
national Commission on Zoological No- 
menclature (1930-1960); and elected Mem- 
ber of the Royal Academy of Physical and 
Natural Sciences, Madrid (1931-1960). He 
was a Charter Member of the ASM. 


Theodore Sherman Palmer, 1951 


Born 26 January 1868 in Oakland, Cal- 
ifornia; B.A., University of California, 
Berkeley, 1888; M.D., Georgetown Uni- 
versity, 1895; died 23 July 1955 (Fig. 2). 

Although employed initially as a banker, 
Palmer was attracted to natural history and 
accepted a position as field agent in the De- 
partment of Agriculture in 1889. A year lat- 
er he became the first assistant ornithologist 
and headed the Death Valley Expedition of 
1891. He moved over to the new Biological 
Survey in 1896 and continued with the Sur- 
vey in various positions until his retirement 
as a senior biologist in 1933. After retire- 
ment, Palmer worked for a time with the 
United States National Museum. 

During his nearly 40 years with the Bio- 
logical Survey, Palmer became an interna- 
tional expert on game protection and con- 
servation, publishing five books on the 


subject and helping draft numerous inter- 
national treaties and regulations covering 
migratory birds. He was one of the principal 
persons responsible for the first migratory 
bird treaty between the United States and 
Canada in 1916. In addition, he was deeply 
interested in nomenclature of birds and 
mammals. Using the vast library resources 
available to him in Washington, D.C., 
Palmer compiled his premier compendium 
of mammalian generic names, which was 
published as “Index Generum Mammal- 
ium” in North American Fauna number 23, 
1904. At the same time, he was a prolific 
contributor to scientific journals. 

Palmer was a Fellow of the American Or- 
nithologists’ Union, the American Associ- 
ation for the Advancement of Science, and 
the California Academy of Sciences. He was 
an active member of more than 27 other 
associations and societies, including four in 
Europe. 

Few, including many of his contempo- 
raries in ornithology and mammalogy, re- 
alize that Palmer had an equally distin- 
guished record in his hobby of philately. He 
was a prolific contributor to journals and 
magazines of philately and was recognized 
as an expert in the field by his philatelic 
peers. 

Like many of his colleagues who had 
trained as physicians, Palmer never prac- 
ticed medicine. Rather he merely used the 
degree as a means to study his first love, 
which was natural history. 


Edward Alexander Preble, 1952 


Born 11 June 1871 in Somerville, Mas- 
sachusetts; no university degrees; died 4 Oc- 
tober 1957 (Journal of Mammalogy, 38:546, 
1957) (Fig. 2). 

Preble could trace his ancestry in the 
United States to the early 17th Century of 
New England. Shortly after graduation from 
high school, in 1889, he took work as a 
plumber in Boston. After eight months of 
city life, he returned home to rural western 


AWARDEES 83 


Massachusetts. A fellow birder who had 
moved to Washington, D.C., to take a po- 
sition with C. Hart Merriam in the new D1- 
vision of Ornithology and Mammalogy 
made arrangements for Preble to join Mer- 
riam’s team. He began work on | April 1892 
with a trip to Texas with Vernon Bailey. 
Numerous trips for the new Biological Sur- 
vey followed to Maryland, Georgia, Oregon, 
Washington, and Utah. 

In the summer of 1900, Preble made the 
first of many survey trips to northern Can- 
ada and Alaska, a region which was to be 
the focus of most of his remaining years 
with the Survey. The results of these bio- 
logical surveys to northern regions were to 
be published as numerous numbers in North 
American Fauna during the next thirty years. 
In addition to Bailey, he worked with such 
Biological Survey notables as Merrit Cary, 
A. K. Fisher, E. T. Seton, Francis Harper, 
W. H. Osgood, W. L. McAtee, and J. A. 
Loring. Preble was a member, along with 
Osgood, of the famous Federal Commission 
to investigate the Pribilof Island fur seals in 
1914. 

Preble routinely was called upon by his 
colleagues to exercise his considerable edi- 
torial talents on their work. He had honed 
this talent by self study and considerable 
reading in a wide variety of fields in addition 
to nature. 

Upon retirement from government ser- 
vice in 1935, Preble became an Associate 
Editor of Nature Magazine. This second 
professional career offered him a forum to 
speak out freely and forcefully on issues of 
conservation, a topic that had begun to con- 
sume his interest and time. He became a 
prolific writer on the topic. Because few of 
his pieces were signed, the exact number of 
contributions is unknown, but nearly every 
issue had numerous articles or reports writ- 
ten by him. 

Preble was an unassuming individual who 
used the spoken word sparingly. Francis 
Harper reported that his “. . .outstanding 
traits were simplicity of character, forth- 
rightness, extraordinary patience and for- 


bearance, unswerving principles, and in- 
dependence... .” Inaddition to his honors, 
including charter membership in the ASM 
(which he valued very highly), he was a Fel- 
low of the American Ornithologists’ Union. 
He is best known for his numerous scientific 
contributions published in North American 
Fauna. 


William King Gregory, 1954 


Born 19 May 1876 in New York City, 
New York; A.B. (1900), A.M. (1905), and 
Ph.D. (1910), Columbia University; died 29 
December 1970 (Journal of Mammalogy, 
52:495, 1971) (Fig. 2). 

Joining the American Museum of Natural 
History as research assistant to Henry Fair- 
field Osborn and Editor of the American 
Museum Journal in 1899, Gregory was to 
spend much of his career at that institution. 
He continued as editor until 1901 and as 
Osborn’s assistant until 1913. His research 
interest was paleontology, and he was given 
a position as Assistant Curator in that de- 
partment in 1911. In addition he was a cu- 
rator in ichthyology and comparative anat- 
omy. In 1916 he accepted an official position 
as a paleontologist at Columbia University 
and became Professor in 1925. 

Gregory’s research focused on the evo- 
lution of the vertebrates, evolution of hu- 
mans and their dentition, and the relation- 
ship of humans and other anthropoids. With 
Osborn, he postulated that humans and 
higher anthropoids, especially the gorilla and 
chimpanzee, had a common, tailless ances- 
tor during the Tertiary. Gregory’s publica- 
tions were mostly on fossil mammals, es- 
pecially primates, but included such books 
on dentition as The Origin and Evolution of 
the Human Dentition (1922), The Dentition 
of Dryopithecus and the Origin of Man 
(1926), with Milo Hellman, and Our Face 


from Fish to Man (1929). 


Gregory was a member of 17 professional 
societies and associations, including the Na- 
tional Academy of Science, Zoological So- 


84 TAYLOR AND SCHLITTER 


ciety of London, and the Anthropologische 
Gesellschaft, Vienna. 


Lee R. Dice, 1956 


Born 15 July 1887, in Savannah, Georgia; 
B.A., Stanford University, 1911; M.A. 
(1914), and Ph.D. (1915), University of Cal- 
ifornia, Berkeley; died 31 January 1977 
(Journal of Mammalogy, 59:635-644, 
1978). 

Dice was raised on a farm in Prescott, 
Washington, and in his boyhood became 
deeply interested in natural history. He ini- 
tially went to Washington State University 
(then an Agricultural College) in Pullman, 
but decided to transfer to the University of 
Chicago where he came under the influence 
of Professor V. E. Shelford, who introduced 
him to ecology. It was then that Dice de- 
cided to become an ecologist. He was, how- 
ever, unable to continue at the University 
of Chicago for financial reasons, so went to 
Stanford University for his remaining un- 
dergraduate program. Asa doctoral student, 
Dice worked under Joseph Grinnell at the 
Museum of Vertebrate Zoology, University 
of California, Berkeley. After several brief 
jobs and Army service, Dice became Cu- 
rator of Mammals, Museum of Zoology, and 
Instructor of Zoology, University of Mich- 
igan, in 1919, and by 1942 had been made 
Professor. He remained at this institution 
throughout his career. Dice was the mentor 
and supervisor of many graduate students, 
several of whom later also became eminent 
mammalogists. 

In 1922 Dice began to work with Pero- 
myscus and kept several species in captivity. 
When geneticist Clarence C. Little became 
President of the University in 1925 and es- 
tablished the Laboratory of Mammalian 
Genetics (later renamed the Laboratory of 
Vertebrate Biology), he appointed Dice as 
an associate of the laboratory. From that 
time on Dice conducted his monumental 
studies of Peromyscus genetics, focusing es- 
pecially on the genetic nature of subspecific 


boundaries. He also became interested in 
human heredity and helped to establish the 
Heredity Clinic at the University and, as his 
research focus intensified there, his work on 
Peromyscus lessened. The Laboratory and 
Clinic were merged as the Institute of Hu- 
man Biology with Dice as its Director until 
retirement. Dice was the author of 138 pub- 
lications, nearly half of which were about 
Peromyscus. Throughout his career, his re- 
search was supported by many granting 
agencies. 

He was President of the Ecological So- 
ciety of America, the Society for Systematic 
Zoology, the Ecological Union (now the Na- 
ture Conservancy) and the American Soci- 
ety of Human Genetics. Dice was a Charter 
Member of the ASM, and from 1947-1951 
was Vice President. He died 20 years after 
retirement and is still remembered as the 
father of research on Peromyscus. 


Albert R. Shadle, 1956 


Born 18 July 1885 in Lockbourne, Ohio; 
B.A. (1913) and M.A. (1915), Ohio State 
University; Ph.D., Cornell University, 1933; 
died 23 May 1963 (Journal of Mammalogy, 
44:449, 1963) (Fig. 2). 

After earning the M.A. degree, Albert 
Shadle went to Cornell as an Assistant in 
Zoology for a year, then became an Instruc- 
tor for two years, and in 1918 was appointed 
Assistant Professor for one year. From there 
he moved to the Department of Experi- 
mental Biology, Roswell Park Memorial In- 
stitute in Buffalo, New York, where in 1920 
he was appointed Professor. He held that 
position until his retirement in 1956. He 
was head of the Department for all but four 
years of his tenure there. In 1956, he was 
made Emeritus Professor and Research As- 
sociate at the Institute. In 1959, he also be- 
came Associate Director of the Institute’s 
summer science program. 

Shadle was a person of broad research 
interests, working on the insect fauna of the 
Allegheny State Park, respiration in fresh- 


AWARDEES 85 


water clams, prostate cancer, pelvic changes 
during pregnancy and parturition, and ex- 
tensive growth and attrition of the incisors 
of rodents and lagomorphs. He was best 
known in the ASM for his deep commit- 
ment to the porcupine, and he studied its 
life history and reproductive biology exten- 
sively. He is remembered today as a cordial 
and outgoing man and one who could be 
counted on fora presentation on porcupines 
at almost every annual meeting of the ASM. 
He attracted a significant audience when he 
delivered lectures about their breeding ac- 
tivities, everyone hoping to find out just how 
porcupines did it! 

Shadle was a member of a number of so- 
cieties, including the New York Academy, 
Wildlife Society, Audubon Society, Society 
of Naturalists, and others. He was also sup- 
ported by the National Science Foundation. 

After his death, the ASM was informed 
that a fellowship in his name and that of his 
wife had been set up commencing in 1972 
to foster the research of Ph.D. students from 
the United States. Although administered 
and ultimately approved by the Buffalo 
Foundation, the ASM is given the oppor- 
tunity to select a finalist each year and to 
present the award. The Albert R. and Alma 
Shadle Fellowship provides several thou- 
sand dollars toward the support of the stu- 
dent’s research. At the end of the year, the 
student presents the results of the work at 
the plenary session of the annual meeting 
of the ASM. Dr. Shadle’s name and memory 
are carried on in this significant way. 


Francis Harper, 1959 


Born 17 November 1886 in Southbridge, 
Massachusetts; B.A. (1914) and Ph.D. 
(1925), Cornell University; died 17 Novem- 
ber 1972 (Journal of Mammalogy, 54:309, 
1973). 

Like many young biologists who were 
trained at the turn of the century, Francis 
Harper was interested in both birds and 
mammals. After receiving his first degree in 


1914, he took his initial trip north to Lake 
Athabaska. Other noteworthy trips to the 
north included his lengthy visit to southern 
Keewatin in 1947 and to the interior Un- 
gava Peninsula in 1953. During the inter- 
vening and later years, Francis managed to 
support himself with short periods of em- 
ployment and meager grants. He was noted 
for his outstanding editorial skills and pro- 
digious memory, asa writer of copious notes, 
and for his very strong opinions. During his 
lifetime, he had no long-term employment 
due to a self-professed inability to withstand 
close supervision. Rather, he chose to work 
mostly on projects funded by such groups 
as the Geological Society of Canada, the 
U. S. Biological Survey, the New York State 
Museum, the Boston Society of Natural 
History, the Penrose Fund, and the Amer- 
ican Philosophical Society. 

Francis Harper was a Guggenheim Fellow 
in 1950-1952 and was employed for a time 
by the Huyck Preserve in Rensselaerville, 
New York, after returning from Ungava 
Peninsula. In 1960 he moved to Chapel Hill, 
North Carolina, in order to resume his work 
on a favorite subject—the natural history 
and folklore of the Southeast, especially the 
Okeefenoke Swamp. 

Francis Harper was a Charter Member of 
the ASM and was corresponding secretary 
from 1931-1932. He was Honorary Life 
Elective Member of the American Orni- 
thologists’ Union and member of Phi Beta 
Kappa. 

During his lifetime Francis published 
about 135 titles on such subjects as mam- 
mals, birds, and other vertebrates, faunal 
zones, botany, conservation, Eskimos and 
Montagnais, folkore, and early naturalists. 
Particularly noteworthy are his lengthy pa- 
pers on Keewatin (The Barren Ground Car- 
ibou of Keewatin and The Mammals of Kee- 
watin) and Ungava Peninsula (Land and 
Fresh-water Mammals of the Ungava Pen- 
insula), the two volume treatise on the Bar- 
trams published in the Transaction of the 
American Philosophical Society in 1942 and 
1943, and The Travels of William Bartram 


86 TAYLOR AND SCHLIT PER 


(1958). His work on the Bartrams stands as 
a monument to his careful scholarship. 


Nagamichi Kuroda, 1959 


Born 24 November 1889 in Tokyo, Ja- 
pan; B.A. (1915) and Ph.D. (1924), Tokyo 
Imperial University; died 16 April 1978 
(Journal of Mammalogy, 59:908, 1978). 

After finishing his undergraduate studies 
in zoology in the College of Sciences, Ku- 
roda was commissioned in 1916 by the 
Government-General of Taiwan to conduct 
research on animals in Taiwan. From there 
he received a similar commission the next 
year from the Government-General of Ko- 
rea to do research on the birds of Korea. At 
the same time, he became involved in the 
preservation of natural areas as a member 
of the Society for Shimei. He became an 
examiner in 1919 in the society for the Min- 
istry of Internal Affairs and held the post 
until 1924. This position allowed him to 
travel to many sites of these memorial nat- 
ural areas. In 1921, he was appointed Chief 
of the Game Commission under the De- 
partment of the Imperial Household. He fo- 
cused on matters of game law, a newly de- 
veloping part of wildlife control in Japan, 
and gave numerous lectures on the subject 
throughout Japan. He held the post of Chief 
until 1940. From 1930 until 1937 he also 
served as Grand Master of Ceremonies and 
Commissioner of Game for the Imperial 
government. During the war years, Kuroda 
was commissioned to study various aspects 
of the natural resources of the Japanese Em- 
pire. Most of these studies were pursued 
through the auspices of the Ministry of Ag- 
riculture and Forestry. 

From an early age Kuroda was interested 
in birds, following in the footsteps of his 
maternal grandfather with an interest in 
ducks. At age 19, he published his first paper 
on ducks, and in 1912 he published Ducks 
of the World. Although never holding a uni- 
versity position, Kuroda had a major influ- 
ence on academia and scientific research in 


Japan. He was a founder of the Nippon Or- 
nithological Society in 1911 and ultimately 
served as President. Kuroda is considered 
to be the father of Japanese ornithology, but 
he was also a charter member of the ASM 
and helped found the Nippon Mammalog- 
ical Society in 1923. 

Because Kuroda was able to travel 
throughout the Empire, or to send others in 
his place, he acquired extensive collections 
of birds and mammals from parts of Man- 
churia, Korea, Taiwan, Philippines, Java, 
Celebes, Okinawa, and Japan. These col- 
lections formed the basis for numerous pub- 
lications on mammals, including such books 
as A Pictorial Book of Japanese Animals 
(1927), Outline of Vertebrate Animals— 
Mammals (1937), Catalogue of Japanese 
Mammals (1938), Colored Pictorial Book of 
Japanese Mammals (1940), and Classifi- 
cation System of Japanese Mammals with 
Diagrammatic Charts (1953). 

In addition to his extensive contributions 
to the taxonomy and biogeography of mam- 
mals of eastern Asia, Kuroda also published 
extensively on conservation, game laws and 
control of wildlife, and birds. He was equal- 
ly prolific as a writer of popular articles on 
natural history, especially on conservation 
of the fauna. He was a man of enormous 
and intense energy who left a legacy of the 
founding of two scientific professional so- 
cieties in Japan. 


Magnus Anton Degerbol, 1962 


Born 8 August 1895 in Sjorring, Thy, Jut- 
land, Denmark; A.B. (1912), Ph.D. (1921), 
and D.Sc. (1933), University of Copenha- 
gen; died 1977 (Journal of Mammalogy, 59: 
894-897, 1978) (Fig. 2). 

Magnus Degerbel was born in rural Den- 
mark of parents who directed a dairy co- 
operative. He showed an early interest in 
natural history. During his university stud- 
ies he came under the influence of Professor 
Herluf Winge, who also held a position at 
the university zoological museum. Winge 


AWARDEES 87 


had been studying an enormous collection 
of bones from excavations in Denmark and 
Greenland. After Winge’s death, Magnus 
became Curator of Mammals at the Mu- 
seum and continued these studies of the 
Pleistocene distribution of vertebrates, es- 
pecially mammals, in Scandinavia. He made 
detailed analyses of the morphology and 
distribution of various species. His bench- 
mark study of prehistoric versus Recent 
predators was used for his D.Sc. degree. 

In 1937, Magnus became Chief Curator 
of Vertebrates at the Zoological Museum. 
At the same time he started a program of 
exhibition growth, including new Arctic and 
African dioramas. Material for the African 
exhibits was obtained during Magnus’ ex- 
peditions to Central Africa in 1947. 

From 1927 to 1947, Degerbol added ever 
increasing teaching responsibilities to those 
of his museum duties. He enjoyed teaching, 
relishing the opportunity that it gave him 
to present his research results to a wide au- 
dience. This desire was also reflected in the 
increasing number of contributions he made 
to popular scientific journals. He was sought 
after for lectures to societies and appear- 
ances on radio programs. 

Magnus made four major international 
expeditions. The first was an expedition to 
his beloved Greenland in 1932. The next 
was the 1947 Danish Central African Ex- 
pedition. In 1952, he participated in the 
Galathea Expedition to the Campbell Is- 
lands, which was followed by a 1954 trip to 
the Andes of South America. 

Although small in physical stature, De- 
gerbol left a large mark in European Qua- 
ternary zoology and was a leading figure in 
Danish zoology. His impact on the Danish 
public through the zoological museum’s ex- 
hibits, and his lectures, radio programs, and 
popular articles was profound. 


Vladimir Georgievich Heptner, 1963 


Born 22 June 1901 in Moscow, Russia; 
D.Sc., Moscow State University, 1936; died 


5 July 1975 (Journal of Mammalogy, 56: 
728, 1975). 

Heptner began his active career In mam- 
malogy near the end of the illustrious career 
of S. I. Ognev at the Zoological Museum of 
Moscow State University. From the begin- 
ning of his career, Heptner participated in 
numerous expeditions to the far corners of 
the Union. He focused much of his field 
work on gerbils of Middle and Central Asia 
and Asia Minor. 

Heptner published more than 300 titles 
during his career. He also served as an editor 
or on the editorial board of four Russian 
journals and three foreign ones. He insti- 
gated the translation from English to Rus- 
sian of numerous books and edited the 
translations. At the same time, he was in- 
volved in academic activities and admin- 
istrative duties with the university and Mu- 
seum. Best known among his numerous 
early monographs and books are Mammals 
of the Kopet Dagh and adjacent plains 
(1929), General Zoogeography (1936), Ro- 
dents of Middle Asia (1936), Vertebrate an- 
imals of Badkhyz (1956), and Harmful and 
useful mammals of the protective forest zones 
(1950). 

With the initiation in 1962 of an English 
translation of Ognev’s multi-volume Mam- 
mals of the USSR and Adjacent Countries, 
an excellent work, although incomplete and 
now superseded, Heptner’s plan for a new, 
more complete Mammals of the Soviet 
Union took on an added degree of urgency. 
The first volumes covered large mammals 
and partially filled the gap left by Ognev. 
This monumental work is a fitting tribute 
to the life of one of Russia’s outstanding 
mammalogists, unequalled in the breadth 
and depth of his knowledge of the mam- 
malian fauna of Eurasia. 

Heptner was elected to honorary mem- 
bership in the Gesellschaft Naturforschen 
der Freunde zu Berlin, the Deutschen Ge- 
sellschaft fiir Sdugetierkunde, and the Zoo- 
logical Society of Czechoslovakia. He was 
also a member of numerous Russian soci- 
eties, including honorary membership in the 


88 TAYLOR AND SCHLITTER 


All-Russian Society of Wildlife Conserva- 
tion. 


Stanley Paul Young, 1964 


Born 30 October 1889 in Astoria, Ore- 
gon; B.A., University of Oregon, 1911, M.S. 
University of Michigan, 1915; died 15 May 
1969 (Journal of Mammalogy, 51:131-141) 
(Fig. 2). 

In his youth, he lived a free spirit life in 
Oregon, hunting, fishing and keeping small 
mammals captive to observe them. Al- 
though his undergraduate degree was in 
mining engineering, his interests changed in 
graduate school—first to geology and then 
to biology. Stanley Young’s first job was as 
a ranger with the U.S. Forest Service in Ar- 
izona. Soon thereafter, he went to the Bio- 
logical Survey as a U.S. Government Hun- 
ter in predator control. On one occasion, he 
inadvertently crossed into Mexico without 
credentials while tracking a wolf. He had to 
be rescued by the 25th U.S. Infantry, but 
not before being held captive for a week. 

Following a brief stint as Assistant In- 
spector of predator control for Arizona and 
New Mexico, he was appointed Assistant 
Leader and then Leader in predatory animal 
control for the Colorado-Kansas district. He 
was based in Denver and remained there 
until 1927, when he became Assistant Head 
of the Division of Predatory Animal and 
Rodent Control in Washington, D.C. There 
he held a number of increasingly responsi- 
ble positions, including Chief, Division of 
Game Management, and Chief, Division of 
Predator and Rodent Control. In 1939 he 
was appointed Senior Biologist, Branch of 
Wildlife Research, in the newly-merged Fish 
and Wildlife Service. Finally, in 1957 he 
became Director, Bird and Mammal Lab- 
oratories, U.S. National Museum, where he 
remained until his retirement in October, 
1959. In his latter positions he published 
regularly, mainly about predator control 
measures and techniques, and about life his- 
tories of large mammalian predators. Even 


after retirement, Stanley Young remained 
active in publication, facilitated by a col- 
laborative appointment at the U.S. Nation- 
al Museum. Volumes such as The Wolf in 
North America, The Bobcat of North Amer- 
ica, and The Clever Coyote (with H.H.T. 
Jackson) are a few of his books that are now 
considered classics. 

Young was a recipient of many honors, 
including Honorary Member of the Wildlife 
Society, Certificate of Appreciation from the 
Office of the Surgeon General of the U.S. 
Army, and the Distinguished Service Award 
from the Department of the Interior. Ten 
years after a retirement filled with writing, 
traveling, and tending to his rose garden, 
Young succumbed to a battle with cancer. 
His life came full circle when his ashes were 
returned to his birthplace in Oregon, where 
he had first savored the wilderness that be- 
came his lifelong interest. 


Erna Mohr, 1966 


Born 11 July 1894 in Hamburg, Ger- 
many; Dr.H.c., University of Munich, 1950; 
died 10 September 1968 (Journal of Mam- 
malogy, 50:232, 1969) (Fig. 2). 

In the beginning of her career, Erna Mohr 
studied fish and invertebrates and estab- 
lished a reputation in those disciplines. In 
her later years, she became world famous 
for her prodigious volume of scientific work 
on mammals, especially large mammals. Her 
interests were ubiquitous, covering all as- 
pects of mammalogy, including anatomy, 
taxonomy, behavior, ecology, general bi- 
ology, and natural history. Her ability to 
synthesize the voluminous amounts of in- 
formation available and her encyclopedic 
knowledge of international literature made 
the production of these works seem easy for 
her. But she was an indefatigable worker 
who continued at an incredible pace, even 
during months of illness near the end of her 
life. In all, she published more than 400 
titles. 


AWARDEES 89 


During most of her career, Mohr was Cu- 
rator of Mammals at the Zoologisches Mu- 
seum und Institut, Hamburg. The collec- 
tions and library gave her an opportunity 
to complete the series of review mono- 
graphs of such diverse groups of mammals 
as the rodents of Germany, European seals, 
European bison, wild boars, and porcu- 
pines. She initiated the studbooks for the 
European bison, Przewalski’s horse, and on- 
ager. 

During a life that spanned some of Ger- 
many’s worst times, Mohr overcame this 
adversity to lead Germany’s emergence as 
an international center of museum and zoo 
mammalogy. She is the only woman to be 
elected to Honorary Membership in the 
ASM. 


Klaus Zimmerman, 1966 


Born 7 July 1894 in Berlin, Germany; 
Ph.D., University of Rostock, 1929; died 5 
February 1967 (Journal of Mammalogy, 48: 
357, 1976; 50:232, 1979). 

After finishing his studies at the Bismarck 
Gymnasium in 1913, Zimmerman was faced 
with difficulties in finding employment be- 
cause of the state of the economy in Ger- 
many. He did not serve in World War I but, 
following family wishes, went to work for 
the family business of selling lumber in Ber- 
lin. After a year of military duty in 1926, 
he went back to his studies of zoology. In 
1929, he received his doctorate based on a 
dissertation on the systematics and geo- 
graphic variation in the Palearctic vespid 
wasp Polistes. He continued studying Hy- 
menoptera, Coleoptera, and Mollusca. 

During the period leading up to and dur- 
ing World War II, Zimmerman, like most 
German biologists, was involved with ex- 
ploration of the world and studies of the 
fauna and flora encountered. During this 
period, Zimmerman changed his research 
focus to small mammals. He began studies 
of Palearctic murids and the small mam- 
mals resulting from an expedition to Crete. 


He was also involved in a project attempt- 
ing to show whether young dogs and mice 
could synthesize vitamins in their appen- 
dix. 

In 1952 Zimmerman became Curator of 
Mammals at the Natural History Museum 
of the Humboldt Institut in Berlin and Pro- 
fessor at the University. He was to remain 
at this institution for the remainder of his 
career. From this position Zimmerman was 
able to continue his systematic studies of 
the Palearctic rodents, especially his anal- 
yses of geographic variation in Apodemus 
sylvaticus, Microtus oeconomus, and Mus 
musculus. In 1956, he participated in an 
expedition to northern China, and in 1963, 
at age 69, he went to the Tien-Shan region 
of Central Asia. His studies of the mammals 
from these expeditions contributed signifi- 
cantly to the knowledge of mammals of 
Central and Eastern Asia. 

In the later years of his life, Zimmerman, 
along with H. W. Stein, was involved in 
translating from Russian to German the nu- 
merous volumes of the Mammals of the So- 
viet Union series begun by B. G. Heptner. 
This was an immense project to which Zim- 
merman gave all of his energies so that these 
important volumes would be available to 
the widest possible international audience. 


George Gaylord Simpson, 1969 


Born 16 June 1902 in Chicago, Illinois; 
Ph.B. (1923) and Ph.D. (1926), Yale Uni- 
versity; died 6 October 1984 (Journal of 
Mammalogy, 66:207, 1985). 

George Gaylord Simpson was born in 
Chicago but moved to Colorado as a child 
and enrolled in the University of Colorado 
as an undergraduate before transferring to 
Yale. By 1926, with a newly obtained doc- 
toral degree, he was already considered an 
international authority on Mesozoic mam- 
mals. This interest took him to the British 
Museum (Natural History) for a year as a 
National Research Council Fellow in Bio- 
logical Sciences, and then to the American 


90 TAYLOR AND SCHEITTER 


Museum of Natural History for the next 32 
years. There he advanced from Assistant 
Curator of Paleontology to Curator of Fossil 
Mammals and Birds and Chairman of the 
Department of Paleontology and Geology. 
It was immediately after World War II that 
he organized this department, having just 
spent several years in service within Army 
intelligence. Concurrent with his leadership 
of the department, he was also a Professor 
at Columbia University in Zoology, holding 
that position through 1959. In 1959, he 
moved to Harvard University as Agassiz 
Professor of Vertebrate Paleontology, a po- 
sition he held until 1970. 

His forte in communication was through 
writing. He was a prolific writer, his earlier 
works, some as monographs, emphasizing 
his interest in Mesozoic and early Tertiary 
mammals. For all concerned with the major 
groups of mammals, Simpson’s monograph 
(1945) entitled The Principles of Classifi- 
cation and a Classification of Mammals be- 
came the primary reference worldwide. His 
expeditions to Patagonia, which he popu- 
larized in Attending Marvels and other 
books, and to various places in the United 
States resulted in extensive collections upon 
which this work was based. All this work 
reflected his rejection of plate tectonics as a 
mechanism of faunal dispersal. His Condon 
lectures, published in 1953, are a prime ex- 
ample of this. With his mathematician wife 
Anne Roe, he wrote the text Quantitative 
Zoology that identified his interest in ap- 
plying methods of biostatistics to research 
on fossil mammals. To anyone teaching in- 
troductory biology in the late 1950s, the 
book Life, of which he was the senior au- 
thor, was the most substantative and re- 
freshingly different introductory text avail- 
able. One of his works was classic in 
evolutionary biology: Tempo and Mode in 
Evolution (1944), which synthesized the en- 
tire field of evolutionary thought. Simpson 
was a leader in developing a synthetic the- 
ory of evolution in association with such 
other eminent scholars as J. Huxley, T. 
Dobzhansky, and E. Mayr. 


The enormous impact of his prolific works 
on evolutionary theory, systematics, and 
vertebrate paleontology (numbering rough- 
ly 1,000 titles) made him not only one of 
the most respected scientists of his time, it 
also garnered him many honors. He was 
elected to the National Academy of Sci- 
ences, recipient of numerous honorary de- 
grees and awards, and elected President of 
both the Society of Vertebrate Paleontology 
and the Society for the Study of Evolution. 

From 1967 until the time of his death, 
Simpson and his wife lived in Tucson where 
he held the position of Professor of Geo- 
sciences at the University of Arizona. It was 
there that he became involved in fostering 
graduate students. 


Kazimierz Petrusewicz, 1972 


Born 23 March 1906 in Minsk, Byelo- 
russia; Maritime Academy, Tczew, 1928; 
MLS. (1933), Sc.D. (1936), Stephen Bathory 
University, Vilnius; died 26 March 1982 
(Journal of Mammalogy, 63:543, 1982) (Fig. 
2). 

Originally trained in the merchant ma- 
rines, Petrusewicz continued his studies in 
natural sciences between voyages. Both 
graduate degrees were based on studies of 
spiders. His career was interrupted by World 
War II, when he fought with the under- 
ground Army. After the war he held a num- 
ber of significant posts in the new Polish 
government, helping to rebuild the ravaged 
countryside, economy, and educational sys- 
tem. In 1949, he was appointed Professor 
at the University of Warsaw. During the 20 
years he held that post, he trained more than 
50 doctoral students, as well as numerous 
master degree candidates. 

Petrusewicz helped found the Polish 
Academy of Sciences and established a De- 
partment of Ecology within the Academy in 
1952. This department was to become the 
Institute of Ecology in 1956. He headed that 
department and institute until 1973 and 
promoted it to world class status as a center 


AWARDEES a1 


of ecological research. He helped organize 
the International Biological Programme and 
chaired the Polish committee. The Polish 
Institute focused on studies of biological 
productivity. Petrusewicz was a leader in 
establishing the IBP Working Group on 
Small Mammals. He served as head of the 
group, editor of numerous reports and co- 
organizer of three international conferences. 

Petrusewicz published more than 140 pa- 
pers, including numerous books. He was 
elected to the Board of the International As- 
sociation for Ecology, honorary member of 
the British Ecological Society (1977), and 
Full Member of the Polish Academy of Sci- 
ences (1965). Clearly, Petrusewicz was an 
international figure in the field of ecology. 
Even more significantly, he single-handedly 
influenced the development of ecology in 
Poland from its infancy to international im- 
portance. 


Charles Sutherland Elton, 1973 


Born 29 March 1900 in Liverpool, En- 
gland; A.B., Oxford University, 1922; died 
1 May 1991 (Fig. 2). 

While an undergraduate at Oxford, Elton 
was selected as an Ecological Assistant to 
Sir Julian Huxley on the University expe- 
dition to Spitzbergen in 1921. Elton’s stud- 
ies of the ecology of the region’s animal life 
prompted him to return to the Arctic again 
in 1923, 1924, and 1930. Because he had 
extensive experience with Arctic animals, 
Elton was appointed as Biological Consul- 
tant to the Hudson’s Bay Company. His 
initial duties were to investigate variation 
in the number of furbearing mammals. 

In 1932, Elton helped establish the Bu- 
reau of Animal Population at Oxford Uni- 
versity. In 1936, he was appointed Reader 
in animal ecology and a Senior Research 
Fellow at Oxford, positions he held until his 
retirement in 1967. His experiences with 
environmental factors and their effects on 
mammal populations, especially those of 
rodents, were used during the war effort of 


World War II. Elton conducted extensive 
research on how to control populations of 
mice and rats and, thus, to conserve food 
in storage. 

Elton is best known for his numerous 
books, beginning with Animal Ecology 
(1927), followed by Animal Ecology and 
Evolution (1930), Voles, Mice and Lem- 
mings: Problems in Population Dynamics 
(1942), The Control of Rats and Mice (1954), 
The Ecology of Invasions by Animals and 
Plants (1958), and The Pattern of Animal 
Communities (1966). During his career, El- 
ton specialized in studies of food chains and 
cycles and the relationship of mammals to 
their environment and to other animals and 
plants. His early work on population cycles 
and numbers in the Arctic served as a basis 
for later research at the Bureau of Animal 
Population. It also won him the honor to 
be named the first editor of Journal of An- 
imal Ecology, begun by the British Ecolog- 
ical Society in 1932. 

Elton was awarded numerous medals and 
awards, the most noteworthy being the Gold 
Medal from the Linnean Society (1967), the 
Darwin Medal from the Royal Society 
(1970), the Tyler Ecology Award (1976), and 
the Edward W. Browning Award (Conser- 
vation) (1977). He was an honorary mem- 
ber of the American Academy of Arts and 
Sciences. 


Vladimir E. Sokolov, 1976 


Born | February 1928 in Moscow, Rus- 
sia; undergraduate degree (1950) and Doc- 
tor of Biological Sciences (1964), Moscow 
State University (Fig. 3). 

After he received his undergraduate de- 
gree, Vladimir spent three years at the Mos- 
cow Fur Institute and four years as Lecturer 
at the Moscow Institute of Fishery. He then 
joined the Biological Department at Mos- 
cow State University as Senior Lecturer for 
ten years and later became Professor of that 
department, a position he still holds. Cur- 
rently, he is also Head of the Department 


92 


Vladimir E. Sokolov 
(1976) 


Z. Kazimierz Pucek 
(1982) 


Bernardo Villa-Ramirez? 
(1986) 


TAYLOR AND SCALITTER 


et 7 oe RS - ig ‘ _ 
Oliver P. Pearson? Victor B. Scheffer 
(1979) (1981) 


a ae a 


Bjorn O. L. Kurten? John Edwards Hills 
(1983) (1985) 


Francis Petter Wuping Xia 
(1987) (1988) 


Fic. 3.—Honorary members of the ASM, 1976-1988. Courtesy of: **J. Mary Taylor; *Helsinki 
University, Photographic Service, Helsinki University Museum; ‘British Museum (Natural History). 


AWARDEES 93 


of General Biology, Russian Academy of 
Sciences, and Director of the A.N. Severt- 
zov Institute of Animal Evolutionary Mor- 
phology and Ecology. 

Vladimir Sokolov has also held numer- 
ous other honorary positions, including 
membership on the Presidium and Russian 
Academy of Sciences and President of Ther- 
iological Congresses I, II, and III (1974- 
1982). 

His research interests are mammalogy, 
ecological morphology, systematics, ecolo- 
gy, and nature conservation, and he teaches 
courses in vertebrate zoology, ecology, be- 
havior, and the environment. Vladimir has 
more than 500 publications to his name, 
including several books. His service on ed- 
itorial boards, such as Reports of the 
U.S.S.R. Academy of Sciences, Advances in 
Modern Biology, Acta Zoologica, and oth- 
ers, 1S extensive. 

The Order of Lenin (1982, 1988), USSR 
State Prize, and Order of ‘‘The North Star,” 
Mongolia, are but a few of the awards he 
has received. 


Oliver Payne Pearson, 1979 


Born 21 October 1915 in Philadelphia, 
Pennsylvania; B.A., Swarthmore College, 
1937; M.A. (1939) and Ph.D. (1947), Har- 
vard University (Fig. 3). 

Oliver Pearson, or Paynie as he is often 
called, credits his own training to such peo- 
ple as Robert K. Enders at Swarthmore and 
Francis Harper at the Academy of Natural 
Sciences. 

From a Research Assistant of the Acad- 
emy of Natural Sciences of Philadelphia for 
one year to Teaching Fellow at Harvard 
University for two, Paynie then became In- 
structor in Zoology at the University of Cal- 
ifornia, Berkeley, in 1947, and Assistant 
Curator of Mammals the next year. He rose 
through the professorial ranks, became Di- 
rector of the Museum of Vertebrate Zoology 
from 1967 to 1971, and briefly filled in as 
Acting Chairman of the Department of Zo- 


ology. Now he is Director Emeritus and 
Professor Emeritus. 

Hardly a year has gone by in his scientific 
career when he has not made a field trip to 
South America— Peru, Colombia, Bolivia, 
and most recently to Argentina or Chile. 
Paynie spent a year as Visiting Professor of 
Ecology at the University of Buenos Aires, 
where he inspired a number of his students 
to become professional biologists. He has 
done extensive field research, primarily on 
rodents of South America, and his 100 or 
so publications include many landmark pa- 
pers for South American mammalogy. Pay- 
nie and his co-worker wife, Anita, have done 
much to foster graduate student exchange 
between the Americas. 

His earlier work dealt mainly with repro- 
duction and physiology in birds as well as 
mammals. His emphasis has become in- 
creasingly ecological over the years. Paynie 
has been an inspiration and mentor for many 
students. 

As a long-term Trustee of the ASM, he 
helped to guide the growth of the Society’s 
endowment and also contributed the same 
expertise to the Cooper Ornithological So- 
ciety. Paynie received the Jackson Award 
in 1982 and is also Honorary Member of 
the Comité Argentino de Conservacion de 
la Naturaleza and Sociedad Argentina para 
el Estudio de los Mamiferos. 


Victor B. Scheffer, 1981 


Born 27 November 1906 in Manhattan, 
Kansas; B.S. (1930) and Ph.D. (1936), Uni- 
versity of Washington (Fig. 3). 

In 1937, Victor Scheffer joined the U.S. 
Biological Survey, which became the U.S. 
Fish and Wildlife Service three years later. 
His entire career was in this organization, 
and he retired in 1969. 

Initially, he was sent to the Aleutian Is- 
lands to conduct a wildlife survey. Next, he 
went to the Pribilof Islands where, from 
1940 to 1974, he made a long-term study 
of Alaskan fur seals, with intervals at the 


94 TAYLOR AND SCHMILTTER 


Rocky Mountain Forest and Range Exper- 
iment Station in Colorado. As a recipient 
of National Science Foundation support, he 
spent a year in Cambridge, England, to write 
the book, Seals, Sea Lions and Walruses. 
In 1964 he was a United States Observer 
on the first team to Antarctica under terms 
of the Antarctic Treaty of 1959. 

Victor Scheffer was given the Distin- 
guished Service Award by the U.S. Depart- 
ment of the Interior in 1965, and in 1977 
he was made Alumnus Summa Laude Dig- 
natus by the University of Washington. In 
1986 he was elected Honorary Member, So- 
ciety of Marine Mammalogy. For his book 
The Year of the Whale he received the Bur- 
roughs Medal of the John Burroughs Me- 
morial Association in 1970, and for 4 Voice 
for Wildlife he received the Joseph Wood 
Krutch Award of the Humane Society of 
the United States in 1975. He is the author 
of eleven books. 

Scheffer taught for a short time at the Uni- 
versity of Washington and at the Interna- 
tional College of the Cayman Islands. He 
was also a consultant for the National Oce- 
anic and Atmospheric Administration for a 
year. From 1973 to 1976, he was the first 
Chairman of the Marine Mammal Com- 
mission. He currently is living in Bellevue, 
Washington, where he has retired. 


Zdzislaw Kazimierz Pucek, 1982 


Born 2 April 1930 in Radzyn Podlaski, 
Lublin Palatinate, Poland; undergraduate 
studies, M. Curie-Sklodowska University, 
1952; Master’s degree, University of War- 
saw, 1954; Ph.D., M. Curie-Sklodowska 
University, 1961; Docent degree, Jagiellon- 
ian University, 1966 (Fig. 3). 

After his Master’s degree, Pucek became 
a Junior Scientific Worker at the Mammal 
Research Institute, Polish Academy of Sci- 
ences, and was made Director in 1962. After 
being awarded the Docent degree, he be- 
came Senior Research Worker at the Insti- 
tute, a position he still holds. Along with 


his research and administrative appoint- 
ments, he teaches mammalogy at several 
Polish universities and has supervised 15 
Masters’ and 19 Ph.D. theses. 

For 11 years Pucek was the Polish rep- 
resentative to the ITC, and he is currently 
the Chairman of the European Bison Spe- 
cialist Group of the SSC/IUCN. 

His research is primarily in biomorphol- 
ogy of shrews and rodents, small mammal 
ecology, and the fauna and protection of 
mammals in Poland. He is the author of 
five books and 140 papers and has been 
Editor-in-Chief of Acta Theriologica since 
1963. In 1990, he was elected Honorary 
Member of the All-Union Theriological So- 
ciety, Russia. 


Bjorn Olof Lennartson Kurtén, 1983 


Born 19 November 1924 in Vasa, Fin- 
land; undergraduate degree (1952) and Ph.D. 
(1954), University of Helsinki; died 28 De- 
cember 1988 (Fig. 3). 

Whether fossil or living, the biology of 
the organism was always paramount in Bj6rn 
Kurtén’s work, in which he emphasized 
functional morphology and paleontology of 
mammals—actually, a paleoecological fo- 
cus. This approach was evident as early as 
his doctoral dissertation, On the variation 
and population dynamics of fossil and Re- 
cent mammal populations. 

After 17 years as a Lecturer at the Uni- 
versity of Helsinki, Kurtén was Personal 
Professor of Paleontology from 1972 until 
his death. He was an inspiration to his stu- 
dents, some of whom came from England, 
the United States, Sweden, and Japan to 
study under his tutelage. Their claim is that 
he restricted them to minimal supervision 
to encourage their originality and indepen- 
dence. On occasion, he came to the United 
States for short periods as a Visiting Pro- 
fessor at the University of Florida and also 
at Harvard University. 

He bridged with enormous success the 
road between scientific and lay audiences 


AWARDEES 31s) 


by writing popular paleontological books 
and novels, such as The Cave Bear Story 
(1976). His inspiring lectures captured stu- 
dents and public alike. He participated in a 
Finnish television serial on the Ice Age just 
before his death. 

Throughout his life Kurtén received nu- 
merous awards, including UNESCO’s Kal- 
inga Award for the popularization of sci- 
ence. He also was elected Honorary Member 
of the Anthropological Association of 
Greece. Outstanding paleontologists, such 
as George G. Simpson and Stephen Jay 
Gould, consider Kurtén to be one of the 
finest paleontologists of all time. He was 
brilliant in his work and inspirational to all 
whose lives he touched. 


John Edwards Hill, 1985 


Born 11 June 1928 in Ashdown House, 
Forest Row, near East Grinstead, Sussex, 
England; no university degrees (Fig. 3). 

After receiving the Oxford Higher Schools 
Certificate in 1946, John joined the Royal 
Air Force as a Meteorological Observer. Two 
years later, he came to the British Museum 
(Natural History), Mammal Section, as As- 
sistant Experimental Officer, where he was 
promoted through the ranks until he be- 
came Principal Scientific Officer in 1977. 
He retired in mid-1988 but continues his 
professional work and association with the 
Museum. 

John’s distinguished career in mammal- 
ogy was strongly shaped by R. W. Hayman, 
his tutor. He was also strongly influenced 
by the taxonomic work of Sir John Ellerman 
and Sir Terrence Morrisson-Scott and is, 
himself, a descriptive taxonomist. He has 
described 57 new taxa of mammals, includ- 
ing a new family of bats (Craseonycteridae), 
and is the author of more than 120 scientific 
publications and five books. Early in his 
career he was a generalist in mammalogy, 
later specializing in the systematics and 
classification of the Chiroptera. He is re- 
sponsible for building the Museum’s col- 


lection of bats to become one of the most 
oustanding in the world. His interest also 
lies in the history of mammal collections 
housed in London and their literature. 

John Edwards Hill has several bats named 
after him, one of them named by Karl 
Koopman to honor both John Edwards Hill 
and another mammalogist of like name, 
John Eric Hill. John Edwards Hill collab- 
orated with the American mammalogist 
James D. Smith on the book Bats—A Nat- 
ural History, and with Gordon B. Corbett 
on A World List of Mammalian Species and 
Mammals of the Indomalayan Region: A 
Systematic Review, both invaluable refer- 
ences for systematic mammalogists. 


Bernardo Villa-Ramirez, 1986 


Born 4 May 1911 in Teloloapan, Guer- 
rero, Mexico; M.S., National University of 
Mexico, 1944; M.A., University of Kansas, 
1947; Doctor of Biology, National Univer- 
sity of Mexico, 1961 (Fig. 3). 

While still at the University of Kansas, 
Bernardo held the post of Assistant Profes- 
sor of Comparative Anatomy. He returned 
to Mexico after his Kansas degree and held 
the position of Assistant Professor of Zo- 
ology at the National University of Mexico 
for ten years. He was appointed Professor 
of Comparative Anatomy in 1960, was Head 
of the Section of Mammalogy from 1957- 
1967, and for three years was Head of the 
Department of Zoology. 

In the years following his graduate studies 
at Kansas under E. Raymond Hall, Bernar- 
do became a pioneer in guiding the devel- 
opment of mammalogy in Mexico and, 
through his teaching, has been the mentor 
of many students in this field. His own scope 
is broad, as befits his pioneering work. He 
is the author of more than 200 scientific 
papers (98 in mammalogy, with emphasis 
on bats), 94 technical papers, and 5 books. 
He established the first large scientific col- 
lection of mammals in Mexico and one of 
the country’s first national game reserves, 


96 TAYLOR AND SCAHLITTER 


and he helped develop the laws and licens- 
ing protocol for game hunting in Mexico. 

Bernardo endeared himself to the ASM 
decades ago for his faithful participation at 
annual meetings, often being the only per- 
son in attendance from outside the United 
States and Canada. He served on the ASM 
Board from 1956 to 1984 and was Vice 
President in 1965. His international per- 
spective and travels are reflected also in his 
many publications coauthored with inves- 
tigators outside of Mexico. He was the re- 
cipient of a John Simon Guggenheim fel- 
lowship in 1945-1947 and has received 
many other awards, including the Gerrit S. 
Miller Award in 1990 given by the 20th 
meeting of the North American Symposium 
on Bat Research. Bernardo was the first 
President of the Mexican Society for the 
Study of Marine Mammals, a founding 
member of the Marine Mammal Society, 
and honorary President of the Mexican As- 
sociation of Mammalogists. 


Francis Petter, 1987 


Born 28 July 1923 in Paris, France; 
D.V.M., University of Alfort, 1949; Sc.D., 
University of Paris, 1961 (Fig. 3). 

After receiving a veterinarian degree in 
1949, Francis Petter became Assistant in 
the Laboratory of Zoology (Mammalogy) at 
the Museum National d’Histoire Naturelle 
in Paris the same year. He specialized im- 
mediately on small rodents, their ecology, 
and epidemiology. He also began to inves- 
tigate the history of the relationship of man 
and domesticated animals. 

His earlier work took him to the Sahara, 
Iraq, and Madagascar, where he discovered 
and described several species of small mam- 
mals. Some of the parasites of these mam- 
mals were named for Petter. 

Immediately after receiving his doctoral 
degree, he was appointed Director of the 
Museum. In addition, he taught mammal- 
ogy and supervised theses at the Institute of 
Tropical Veterinary Medicine for many 


years. He was Secretary General of Mam- 
malia, founded in 1936. 

Petter is probably best known for his sys- 
tematic and ecological work on rodents of 
northern Africa. He is the author of close 
to 175 papers, also on Brazilian rodents and 
parasites, and on phylogeny based on elec- 
trophoretic analyses. 


Wuping Xia, 1988 


Born 19 May 1918 in Baixing County, 
Hebei Province, China; university degree, 
Yenching University, 1945 (Fig. 3). 

After his university training, Wuping Xia 
engaged in hydrobiological studies. How- 
ever, the Sino-Japanese war took its toll on 
scientific studies in China, especially mam- 
malogy. It was Xia, along with Professor T. 
H. Shaw and Professor H. S. Peng, who to- 
gether put the field of Recent mammalogy 
on firm footing in China during the follow- 
ing decade. Particular emphasis was placed 
on ecological studies of small mammals. The 
rapid growth of mammalogy in the follow- 
ing decades led to the establishment of the 
Mammalogical Society of China in 1980. 
Wuping Xia was the first President of the 
society, a position he continues to hold. The 
journal Acta Theriologica Sinica was found- 
ed the following year, and Xia is its Man- 
aging Editor. 

In 1980, Wuping became Director, 
Northwest Plateau Institute of Biology, Ac- 
ademia Sinica, then Honorary Director four 
years later, retiring in 1990. During his ten- 
ure, he established the Haibei Research Sta- 
tion of the Alpine Meadow Ecosystem, a 
field station located at 3,200 m in Quinhai 
Province. It is one of the highest research 
stations in the world for the study of grass- 
land and plateau biology. 

Wuping also has carried on an academic 
teaching career in the Academia Sinica, most 
recently as Professor. He is the author of 
seven books and more than 50 papers, 
mostly on rodent ecology and control. 


AWARDEES oF 


Karl F. Koopman? 
(1990) 


Karl F. Koopman, 1990 


Born | April 1920 in Honolulu, Hawaii; 
B.A. (1943), M.A. (1945), and Ph.D. (1950), 
Columbia University (Fig. 4). 

After graduate work, Karl became an In- 
structor in Biology at Queens College and, 
in 1958, moved to become Assistant Cu- 
rator in Mammalogy, Academy of Natural 
Sciences of Philadelphia, for one year. He 
then went to the Chicago Natural History 
Museum at the same level of appointment 
for two years. His real home professionally 
became the American Museum of Natural 
History, commencing in 1961, where he rose 
through the curatorial ranks in the Depart- 
ment of Mammals, retiring officially in 1986 
but still keenly active as Curator Emeritus. 

Karl’s chief interest is bats, especially Mi- 
crochiroptera, but he has a wealth of knowl- 
edge about all mammalian groups. As a for- 
mer student of Theodosius Dobzhansky, 
Karl carries forward a vast background in 
genetics and evolutionary biology. 

His collecting trips, chiefly for mammals 
and reptiles, have been largely in the equa- 
torial region and southern hemisphere, 
whereas his travels to professional meetings 
and museums have been worldwide. He is 
the author of many papers, particularly on 
bats, but also on primates and other ver- 
tebrates. 


Phillip Hershkovitz 
(1991) 


Fic. 4.—Honorary members of the ASM, 1990-1991. Courtesy of: «J. Mary Taylor. 


Karl serves the ASM in many capacities, 
including as a member of the Board of Di- 
rectors for many years. His services on the 
Nomenclature and Checklist committees 
have been unbounding, and his incredible 
scholarly attention to the presentations de- 
livered at the annual meetings of the ASM 
is unique. He has not only attended almost 
every meeting for more than 40 years, but 
is the guy in the front row of every session 
who asks such incisive questions! All this 
and more were recognized when he received 
the Jackson Award in 1988. 


Philip Hershkovitz, 1991 


Born 12 October 1909 in Pittsburgh, 
Pennsylvania; B.S. (1938) and M.S. (1940), 
University of Michigan (Fig. 4). 

Against all odds, Phil Hershkovitz sought 
training as a mammalogist in the depths of 
the Great Depression. As an undergraduate 
at the University of Michigan, he not only 
worked as a departmental assistant, he also 
took taxidermy jobs to support himself. In 
1932, he was hired to collect blind cave sal- 
amanders. Eager to trap mammals there as 
well, he asked Lee R. Dice, then Curator of 
Mammals, for traps. Dice was unable to 
supply them. When hitchhiking en route to 


98 TAYLOR AND SCALITIER 


Texas, he stopped in Chicago on a chance 
visit to the Field Museum of Natural His- 
tory. Colin Sanborn, then Curator of Mam- 
mals there, loaned him the traps. In ex- 
change, Phil sent the specimens to the Field 
Museum on this and subsequent field sea- 
sons. That stop at the Field Museum later 
shaped his future. 

As the depression worsened, Phil went to 
Ecuador, where he remained for five years, 
collecting an impressive array of mammals 
for the University of Michigan, living off 
the land as he did so. This collection was 
the basis for his Master’s degree work under 
William H. Burt, successor to Dice as Cu- 
rator of Mammals. He interrupted his doc- 
toral program to accept a prestigious trav- 
eling scholarship from the United States 
National Museum, and went to Colombia, 
where he collected mammals for that mu- 
seum for two years. 

After serving in World War II, he was 
offered a curatorial position at the Field Mu- 
seum and accepted it, fully cognizant that 
the return to his doctoral program was the 
sacrifice. He has remained there throughout 
his career, traveling to the Neotropics when- 
ever he could, sometimes for years at a time. 
It was there that he assembled invaluable 
collections of mammals upon which he has 
focused his research career. 

His highly productive career is singular 
in the magnitude of his scholarly contri- 
butions, especially in evolution and bioge- 
ography of South American mammals, and 
in the fact that he is the sole author of 99% 
of his 300 or more articles. He has been 
more influential in the arena of neotropical 
mammalogy than has anyone else in this 
century. He has published major mono- 
graphic revisions on every order of Recent 
mammals of South America. Although his 
work is often challenged, it serves as a great 
stimulus of ideas and of testing hypotheses. 

Hershkovitz became Research Curator at 
the Field Museum in 1961, formally retired 
in 1971, and continues today as Curator 
Emeritus. 


C. Hart Merriam Awardees 


The C. Hart Merriam Award was estab- 
lished in 1974 to recognize outstanding con- 
tributions to the discipline of mammalogy 
by a member of the society in more than 
one of the following areas: scientific re- 
search, education of mammalogists, and 
service to the ASM (Journal of Mammal- 
ogy, 55:694, 1974). The recipient is given a 
statuette of a bison cow that is cast in fi- 
berglass and painted in bronze. It is a copy 
of one made by John Paul Jonas of Jonas 
Brothers for an exhibit at the American Mu- 
seum of Natural History on the mammals 
of New York State. According to Sydney 
Anderson, who oversees the reproductions 
of this statuette, the reason it is a bison cow 
is that the larger bull would not fit on a 
bookshelf! 

The Merriam Award requires unanimous 
approval of the nominee by the committee 
and two-thirds approval by the Board of 
Directors. In 1977, the board decided that, 
in addition to the two-thirds affirmative vote 
by the Board, the nominee must receive the 
approval of three elected officers plus five 
senior Directors, thus requiring close to 
unanimity within this group. 

In 1981, the Board of Directors modified 
these criteria to reduce the emphasis on ser- 
vice, following the establishment of the 
Jackson Award, which is based on this in- 
tention. The Board revised the description 
of the Merriam Award to “The [Merriam] 
Award is to be made to a member of the 
Society, who, in his or her activities within 
the past ten years, has achieved a record of 
excellence in more than one of the following 
areas: scientific research, education of 
mammalogists, and service to the discipline 
of mammalogy”’ (unabridged minutes of the 
1981 Board of Directors’ Meeting). Fur- 
thermore, the policy established in 1979 of 
inviting the recipient of the Merriam Award 
to present a keynote address at the next an- 
nual meeting was reiterated. In 1989 the 
requirement of membership in the ASM was 


AWARDEES 99 


Terry A. Vaughan* . Robert J. Baker® 
(1979) (1980) 


a . yj < { ew 
John F. Eisenberg‘ Michael H. Smith 
(1981) (1985) 


-Y 


Timothy H. Clutton-Brock Guy G. Musser 
(1988) (1991) (1992) 


Fic. 5.—Merriam awardees of the ASM, 1979-1992. Courtesy of: *Department of Biological Sci- 
ences, Northern Arizona University; «J. Mary Taylor. 


100 TAYLOR AND SCHLITTER 


removed for nominees of this award. The 
criteria also were changed to “excellence in 
research and one or both of the other cat- 
egories” (Journal of Mammalogy, 70:880, 
1989). 

The first recipient of the Merriam Award 
was James N. Layne. Until 1991, when 
Timothy Clutton-Brock received the award, 
all recipients were from the United States 
and were members of the society. Fourteen 
individuals have received the award. Of 
these, seven have served as President of the 
society, two have received Honorary Mem- 
bership, and one received the H. H. T. Jack- 
son Award. Their average age at the time 
the award was bestowed is 48, ranging from 
38 to S56. 


Terry A. Vaughan, 1979 


Born 5 May 1928 in Los Angeles, Cali- 
fornia; B.A., Pomona College, 1950; M.A.., 
Claremont Graduate School, 1952; Ph.D., 
University of Kansas, 1958 (Fig. 5). 

Terry, who spent two years in the U.S. 
Army before going on for his Ph.D., was 
appointed Research Biologist in the De- 
partment of Range Science at Colorado State 
University immediately after he completed 
his doctorate. He held that position until 
1967, when he joined the Department of 
Biological Sciences, Northern Arizona Uni- 
versity, first as Associate Professor and then 
Professor of Zoology. He retired there in late 
1987. 

His deep interest and extensive research 
on the biology of bats began early in his 
career, his first publication being on hoary 
bats (Journal of Mammalogy, 34:256, 1953). 
He has published about 50 scientific papers, 
largely in the field of chiropteran morphol- 
ogy, but also on the biology of pocket go- 
phers and woodrats. Since its publication in 
1972, Terry Vaughn’s Mammalogy has gone 
through three editions and is still one of the 
most popular textbooks for the discipline. 

His research support includes the Na- 
tional Science Foundation, National Geo- 


graphical Society, U.S. Fish and Wildlife 
Service and International Biological Pro- 
gram. He is a member of the Ecological So- 
ciety, the Paleontological Society, the So- 
ciety for the Study of Evolution, and others. 

Terry’s formal service to the ASM has 
been as Editor for Feature Articles in the 
Journal of Mammalogy, 1966-1968, and as 
Second Vice President 1980-1982. He was 
a Visiting Professor of Zoology in Nairobi 
on two occasions, and most recently spent 
a year in Western Australia working on Me- 
gachiroptera. 

Terry continues to live in Rimrock, Ar- 
izona, in his retirement, having descended 
several thousand feet to escape snowy Flag- 
staff. 


Robert J. Baker, 1980 


Born 8 April 1942 in Warren, Arkansas; 
B.S., Arkansas A & M College, 1963; M.S., 
Oklahoma State University, 1965; Ph.D., 
University of Arizona, 1967 (Fig. 5). 

Since receiving his Ph.D., Robert J. Baker 
has been a faculty member in the Depart- 
ment of Biological Sciences at Texas Tech 
University. He is now the distinguished 
Horn Professor, Director of the Natural Sci- 
ence Research Laboratory, and Curator of 
Mammals and Vital Tissues at Tech, and is 
also Research Associate at the Carnegie Mu- 
seum of Natural History, and the Univer- 
sity of New Mexico, Albuquerque. 

Early in his career, he developed a deep 
interest in chromosomal evolution and is at 
the forefront of molecular genetics, in situ 
hybridization of chromosomal architecture, 
and the problems of contact zones between 
chromosomal races. His model is usually 
bats, although he uses murid rodents exten- 
sively as well. He teaches mammalogy, his- 
tology, cytology, general zoology, and var- 
ious research courses, and is the recipient 
of several awards for both teaching and re- 
search. 

Bob has received strong grant support 
from Texas Tech University and for 20 or 


AWARDEES 101 


more years from the National Science 
Foundation. The National Parks System and 
the Smithsonian Foreign Currency Program 
have also provided major support. His ex- 
tensive field work is primarily in the neo- 
tropics, but also in Tunisia and England. 

Since 1972, Bob has served long periods 
in editorial capacities for the Journal of 
Mammalogy. His society affiliations in- 
clude Society of Systematic Biologists, So- 
ciety for the Study of Evolution, and Texas 
Academy of Sciences. He has supervised 
more than 20 Master’s degree students and 
14 doctoral students. Bob is the author or 
coauthor of more than 175 papers, and he 
collaborates with a wide array of investi- 
gators. 


John Frederick Eisenberg, 1981 


Born 20 June 1935 in Everett, Washing- 
ton; B.S., Washington State University, 
1957; M.A. (1959) and Ph.D. (1962), Uni- 
versity of California, Berkeley (Fig. 5). 

John Eisenberg became Assistant Profes- 
sor of Zoology at the University of British 
Columbia in 1962, moved to the University 
of Maryland in 1964, and then in 1965 to 
the National Zoological Park (NZP) where 
he was Resident Scientist and then Head, 
Office of Zoological Research. Concurrently 
he was Associate, Department of Mental 
Hygiene at Johns Hopkins University, and 
Adjunct Professor of Zoology at the Uni- 
versity of Maryland. In 1979, he became 
Assistant Director, Animal Programs at the 
NZP and, in 1982, moved to the University 
of Florida to become Ordway Professor, 
Curator, and Eminent Scholar, Ecosystem 
Conservation. 

Early in his career, John began receiving 
significant recognition: Phi Beta Kappa, Phi 
Kappa Phi, President of the Animal Behav- 
ior Society in 1973, Fellow of the Animal 
Behavior Society and of the New York Zoo- 
logical Society, and fellowships from the 
National Science Foundation and the Na- 
tional Academy of Sciences. 


John is eminent worldwide in his field. 
His eclectic approach to mammalogy weaves 
together the fields of behavior, reproduc- 
tion, ecology, systematics, and evolutionary 
adaptations in a truly integrated approach. 
His service in the ASM includes member- 
ship on the Board of Directors and com- 
mittees. John is the author of more than 
125 papers and 1s a recipient of grants from 
the Smithsonian Institution, National Sci- 
ence Foundation, U.S. Department of In- 
terior, National Geographic Society, and 
others. His books, Mammalian Radiations 
and Mammals of the Neotropics, Volumes 
I and IT, are landmark publications in the 
field of mammalogy, and he has used vir- 
tually the full spectrum of mammals of the 
world as subjects of his papers. His field 
work has taken him throughout the world, 
and he has been mentor and supervisor of 
more than 21 graduate students and 10 
postdoctoral students. 


Michael H. Smith, 1985 


Born 30 August 1938 in San Pedro, Cal- 
ifornia; B.A. (1960) and M.A. (1962), San 
Diego State University; Ph.D., University 
of Florida, 1966 (Fig. 5). 

After his doctoral work, Mike was a Re- 
search Associate and then went through the 
ranks to a full professorship, all at the Uni- 
versity of Georgia. In 1973, he became D1- 
rector of its Savannah River Ecology Lab- 
oratory. He teaches population biology, 
vertebrate ecology, and population genetics, 
and supervises many undergraduate re- 
search projects. He has also supervised over 
15 graduate students. Mike has presented 
about 400 talks at universities, museums, 
and other institutions and professional 
meetings both here and in foreign countries. 

His research is focussed on both short- 
and long-term responses of biological 
systems to natural and man-made environ- 
mental changes. He is interested in the ge- 
netics of natural populations and its im- 
portance to regulation, conservation, and 


102 TAYLOR AND SCALITTER 


management of populations of both aquatic 
and terrestrial vertebrates. His research cuts 
across many fields, and its approach is com- 
prehensive. 

Mike is the author or coauthor of more 
than 175 papers, several chapters, books, 
and other publications. He has been prin- 
cipal or co-principal investigator of major 
grants from the AEC, EPA, National Re- 
search Council, and National Science Foun- 
dation, and he has received over 16 million 
dollars of support. 

Mike has served on the Board of Direc- 
tors of the ASM for about a dozen years. 
He belongs to several ecological societies, 
the American Fisheries Society, the Amer- 
ican Society of Ichthyologists and Herpe- 
tologists, the American Society of Natural- 
ists, and is a member of Sigma Xi and Phi 
Sigma. 


Jerry R. Choate, 1988 


Born 21 March 1943 in Bartlesville, 
Oklahoma; B.A., Kansas State College of 
Pittsburg, 1965; Ph.D., University of Kan- 
sas, 1969 (Fig. 5). 

After completing the doctorate at the 
University of Kansas under the tutelage of 
J. Knox Jones, Jr., Jerry was Assistant Pro- 
fessor at the University of Connecticut for 
two years. In 1971, he returned to Kansas 
as Assistant Professor at Fort Hays State 
University. He is now Professor at that uni- 
versity. Additionally, in 1973 he became 
Curator of Mammals and Director at the 
university’s Museum of the High Plains. In 
1980, he assumed administrative respon- 
sibilities for all museums on campus, which 
have been merged as the Sternberg Museum 
of Natural History. 

Jerry has received numerous honors, his 
most cherished (in addition to the Merriam 
Award) being the Southwestern Association 
of Naturalists’ Robert L. Packard Excel- 
lence in Education Award. He has been re- 
cipient of numerous grants from the Na- 
tional Science Foundation and other state 
and federal agencies. He has served on sev- 


eral ASM standing committees and was Re- 
cording Secretary of ASM from 1974 
through 1984. He presently serves as Chair 
of the ASM’s Trustees (a position he also 
holds with the Southwestern Association of 
Naturalists) and a member of the Board of 
Directors. 

His research interests include systemat- 
ics, biogeography, and natural history of 
mammals on the Great Plains. He is coau- 
thor of one book (with another currently in 
press) and 140 scientific papers. In addition, 
he is coeditor of the present volume. His 
greatest professional achievement, howev- 
er, has been 1n preparing undergraduate and 
masters-level students for Ph.D. studies in 
mammalogy. 


Timothy Hugh Clutton-Brock, 1991 


Born 13 August 1946 in London, En- 
gland; M.A. (1968), Ph.D. (1972), and Sc.D. 
(1986), Cambridge University (Fig. 5). 

Clutton-Brock’s graduate work began with 
specialties in archaeology and anthropolo- 
gy, and his doctoral work focused on com- 
parative social organization and ecology of 
colobus monkeys. This study took him to 
Tanzania and Uganda for field work. After 
one year as a postdoctoral fellow, he was 
appointed University Lecturer in Biology at 
the University of Sussex. He left in 1976 to 
become Senior Research Fellow in Behav- 
ioral Ecology at King’s College, Cambridge, 
and in 1981 he was appointed Advance Re- 
search Fellow in the Department of Zoology 
there. Two years later he was made Royal 
Society Research Fellow in Biology, and it 
was during that time that he was awarded 
a Sc.D. from Cambridge. In 1987 he was 
appointed University Lecturer in the De- 
partment of Zoology at Cambridge, and in 
1991 was promoted to Reader in Animal 
Ecology, an appointment he holds today. 

Tim’s research 1s focused primarily on the 
social organization, ecology, reproduction, 
and behavior of two mammalian groups: 
primates and red deer. Since 1986 he has 
been an author of 107 scientific papers and 


AWARDEES 103 


A 


Bryan P. Glass? 
(1980) 


Marie A. Lawrence? 
(1990) 


Murray L. Johnson 
(1986) 


John O. Whitaker, Jr. 
(1991) 


Fic. 6.—Hartley H. T. Jackson awardees of the ASM, 1980-1991. Courtesy of: «J. Mary Taylor; 


>’Chacma Inc., New York. 


three books, and he has served as editor of 
five other books. He has received major 
grant support, and has supervised research 
projects of 26 students, mainly at the doc- 
toral level. In 1980 he started the Large An- 
imal Research Group within the Depart- 
ment of Zoology at Cambridge. That same 
year he also became Chairman of the IUCN 
Deer Specialist Group. 

In recognition of his prodigious scholarly 
work, Tim has received several major hon- 
ors that include the Award for Best Book 
from the Wildlife Society of America in 
1983, the Scientific Medal from the Zoo- 
logical Society of London, and the Fellow 
of the Royal Society in 1993. 

Tim is married to Dafila Kathleen Scott, 
daughter of the well-known ornithologist, 
the late Sir Peter Scott. 


Guy G. Musser, 1992 


Born 10 August 1936 in Salt Lake City, 
Utah; B.S. (1959) and M.S. (1961), Uni- 
versity of Utah; Ph.D., University of Mich- 
igan, 1967 (Fig. 5). 

One year before receiving his Ph.D., Guy 
Musser was appointed Archbold Assistant 
Curator in the Department of Mammalogy 
at the American Museum of Natural His- 
tory. He has remained at that institution to 
the present day. Guy became Chairman of 
the Department in 1981, five years after be- 
ing promoted to Archbold Curator. Since 
1983 he also has held the appointment of 
Research Associate in the Department of 
Vertebrate Zoology at the National Muse- 
um of Natural History. 

Although born in the West and loving the 


104 TAYLOR AND SCHLITTER 


wilderness country of the region, Guy moved 
to the crowded core of New York City be- 
cause he was drawn to the resources of the 
mammal collections of the museum. He has 
made the large rodent collections his re- 
search tool in his lifelong commitment to 
the complexities of rodent systematics. As 
a graduate student, first of Stephen D. Dur- 
rant at Utah and later of Emmet T. Hooper 
at Michigan, Guy’s training and background 
helped him become outstanding in this field. 
He has worked in Costa Rica, the United 
States, and particularly in southeast Asia, 
gathering field data and specimens for anal- 
ysis. He has lived for years at camp sites in 
Sulawesi in an attempt to comprehend the 
subtleties of distributional limits of species 
in different altitudes and habitats. One only 
has to read his papers to realize the depth 
of his comprehension of environmental fac- 
tors that relate to distributions and habits 
of species of rodents. He has described a 
number of species and genera of rodents and 
has proposed several changes at the higher 
taxonomic levels. 

Guy’s publications are generally long and 
comprehensive papers, many of mono- 
graphic length. He is recognized interna- 
tionally for his outstanding contributions to 
the systematics of muroid rodents. 


Hartley H.T. Jackson Awardees 


The Hartley H. T. Jackson Award was 
established in 1977 and was first given to 
W. B. Davis in 1978. This award recognizes 
members of the society who have given long 
and outstanding service to the society (Jour- 
nal of Mammalogy, 58:709, 1977). The re- 
cipient is given a certificate that includes a 
sketch of Jackson, and a plaque that has the 
ASM pronghorn logo on it. The Jackson 
Award Committee was established with the 
guidelines that the committee should re- 
main small (5 members), it should be unan- 
imous in its recommendation, and the Board 
of Directors should support the committee’s 
nomination by a two-thirds majority if it is 


to be approved. The recipient is announced 
at the annual banquet. 

In 1981 the Board of Directors further 
decided that there should be no more than 
one recipient of the Jackson Award and of 
the Merriam Award in any given year, and 
that the awards need not be given each year 
if, in the opinion of the selection committee, 
suitable candidates are not available (un- 
abridged minutes of the 1981 Board of Di- 
rectors’ meeting). 

Since the inception of the Jackson Award, 
12 mammalogists have received it through 
1992. Of these, four are past presidents, sev- 
en have been elected Honorary Members, 
and one isa recipient of the Merriam Award. 
One woman (Marie Lawrence) has received 
the Jackson Award. Recipients have, by the 
nature of the award, all been members of 
the society for a long time and to date all 
have been from the United States. The av- 
erage age of the recipient at the time of re- 
ceiving the award has been 66, ranging from 
54 to 76. 


Bryan P. Glass, 1980 


Born 21 August 1919 in Mandeville, Lou- 
isiana; A.B., Baylor University, 1940; MLS., 
Texas A & M University, 1946; Ph.D., 
Oklahoma State University, 1952 (Fig. 6). 

Bryan spent his entire childhood in Chi- 
na, where he graduated from the China In- 
land Mission School, Chefoo, Shantung 
Province in 1953. He served in World War 
II, primarily as an intelligence officer in Chi- 
na with the 14th Air Force and OSS, and 
was awarded the Asiatic-Pacific Medal with 
two battle stars and a Presidential Unit Ci- 
tation. Bryan has spent his professional life 
at Oklahoma State University from 1946- 
1985, progressing through all the profes- 
sorial ranks; he became Director of the Uni- 
versity Museum in 1966. 

Bryan’s research focus is primarily on 
mammals, particularly on microchiropter- 
an bats. His publications reflect a special 
interest in distributional records, status, and 


AWARDEES 105 


in regional faunal surveys, primarily in 
Oklahoma, but he also made a survey of the 
mammals of Ethiopia and of a new national 
park in Brazil. 

Throughout his professional career, Bry- 
an Glass has given generously of his time 
and expertise to his university, his church, 
his city, and to the ASM. He is the recipient 
of Oklahoma State University’s Outstand- 
ing Service Award (1965) and Outstanding 
Teacher Award (1966), and recently was 
elected 2nd Vice-President at the 20th Bap- 
tist General Convention, and is Past Pres- 
ident of the Arts and Humanities Council 
in Stillwater. 

Bryan was elected Corresponding Secre- 
tary of the ASM in 1956, and from 1957 to 
1977 he served as Secretary-Treasurer. Dur- 
ing those 20 years, membership grew from 
1,500 to 3,900. He inaugurated the portrait 
file of Past Presidents and of group photo- 
graphs at annual meetings. Assisted by his 
wife, Carolyn, Bryan maintained the mail- 
ing list of members and subscribers and 
oversaw the printing of the program for the 
annual meeting each year, all in pre-com- 
puterization years. During his tenure, he was 
the major writer of the Society’s constitu- 
tion. 

Bryan’s tangible contributions to the ASM 
have led to both strength and growth of the 
society, but so have his undocumented con- 
tributions. Bryan is often one of the first to 
welcome student mammalogists at annual 
meetings and make them feel at ease by in- 
troducing them to fellow scientists. 


Murray L. Johnson, 1986 


Born 16 October 1914 in Tacoma, Wash- 
ington; B.A. (1935) and M.D. (1939), Uni- 
versity of Oregon School of Medicine (Fig. 
6). 

After postgraduate training in surgery at 
Union Memorial Hospital in Baltimore, 
Maryland, Murray joined the U.S. Naval 
Medical Corp and served for 3 years. He 
has been in the practice of medicine from 


1946 through 1983, becoming a certified 
member of the American Board of Surgery 
in 1948. 

Along with his medical practice, Murray 
has been a research biologist in mammal- 
ogy, spending almost 50% of his time in this 
field and, since his retirement, even more. 
From 1949 through 1983 he was Curator 
of Mammals at the Puget Sound Museum 
of Natural History, also chairing the Exec- 
utive Board there for many years. He was 
principal investigator in the Marine Mam- 
mal Program Project Chariot (AEC) through 
the Arctic Health Reseearch Center in An- 
chorage from 1959 through 1964. From 
1963 to 1983 he was Research Professor of 
Biology at the University of Puget Sound, 
held a number of National Science Foun- 
dation grants, and from 1984 to date has 
been an Affiliate in Mammalogy and Cu- 
rator of Mammals at Burke Memorial 
Washington State Museum, University of 
Washington. From 1989 to 1992, he has 
been a member of the Scientific Advisors, 
U.S. Marine Mammal Commission, and 
1984 to date the Secretary for the Foun- 
dation for Northwestern Natural History. 

Murray has been the invited participant 
in many scientific meetings, international as 
well as within North America. He is a mem- 
ber of numerous scientific organizations, 1n- 
cluding a Fellow of the American Associa- 
tion for the Advancement of Science. In 
1978, he was named Distinguished Citizen 
of the Year in Tacoma, Washington. He is 
the author of many papers on marine mam- 
mals and rodents, and some on reptiles and 
birds. He has special interest in blood pro- 
tein electrophoretic studies in mammalian 
taxonomy. His investigations are largely 
centered around the Pacific Northwest. His 
wife and strong supporter, Sherry, accom- 
panies Murray to every annual meeting of 
the ASM. 


Marie A. Lawrence, 1989 


Born 20 October 1924 in Poughkeepsie, 
New York; B.A., Vassar College, 1945; 


106 TAYLOR AND SCALITTER 


M.S.S., Smith College School for Social 
Work, 1952; M.A., New York University, 
1970; died 21 September 1992 (Fig. 6). 

Marie Lawrence began her career, not as 
a mammalogist or even as a biologist, but 
as a social worker in New York, a career 
she continued for almost 30 years. Her last 
position was Adjunct Associate Professor, 
New York University of Social Work, which 
she left in 1975. For the final two years, she 
was also Scientific Assistant, Department of 
Mammals, at the American Museum of 
Natural History. She held this position for 
nine years, during one of which she was also 
Assistant Professor of Zooarchaeology at 
Northwestern Archaeology Field School in 
Illinois. In 1982, she became Senior Sci- 
entific Assistant at the Museum, a position 
she held until her death. 

Although driven by a keen interest in 
zooarchaeology, Marie concentrated her re- 
search on Old World arvicoline rodents, 
megachiropteran nectar feeders, Myospa- 
lacine rodents, and the assessment of Me- 
dieval knowledge of mammalian natural 
history. 

Marie did yeoman’s service to produce 
Recent Literature in Mammalogy for 16 
years until the ASM discontinued it in 1985. 
She served on the Board of Directors and 
on several standing committees. She was the 
recipient of several prestigious awards, in- 
cluding a Ford Foundation Fellowship and 
the Margaret Mead/Kreiser Fellowship in 
Anthropology. She was not only the first 
woman to receive the Jackson Award, she 
was the first AfroAmerican to be honored 
by an award from the ASM. 


John O. Whitaker, Jr., 1991 


Born 22 April 1935 in Oneonta, New 
York; B.S. (1957) and Ph.D. (1962), Cornell 
University (Fig. 6). 

While still a graduate student, John 
worked as a field assistant during summers 
for the New York State Museum and the 
New York Conservation Department. Im- 
mediately following his doctoral work on 


Zapus hudsonius, under the direction of 
William J. Hamilton, Jr., John joined the 
Department of Life Sciences, Indiana State 
University, as Assistant Professor to teach 
vertebrate zoology, mammalogy, and other 
courses, including one on mammalian ec- 
toparasites. He now holds the rank of Pro- 
fessor. To date, John has been the mentor 
for more than 50 graduate students in both 
M.A. and Ph.D. programs. The diversity of 
thesis titles, as well as his more than 230 
publications, reflects his extraordinary di- 
versity of interests and expertise within the 
breadth of vertebrate biology and mam- 
malian parasites. He has written keys, an- 
alyzed diets, recorded new distributions, and 
studied herps and birds, as well as mam- 
mals, across a wide spectrum of research. 

John is the recipient of numerous grants 
and contracts that have sustained portions 
of the studies made by him and his students. 
He was elected a Fellow in the American 
Association for the Advancement of Sci- 
ence in 1968, a Fellow in the Indiana Acad- 
emy of Science in 1976, and was one of the 
first two people to be given an Indiana State 
University “Research and Creativity 
Award,” in 1981. 

Just as impressive as John’s contributions 
to the field of mammalogy and students in 
that field are his staggering contributions to 
the discovery and description of over 130 
new taxa of mammalian parasites, largely 
from North American mammals. His mem- 
bership in professional societies also mir- 
rors his breadth of interests and his extraor- 
dinary competence as an eclectic biologist. 


B. J. Verts, 1992 


Born 19 April 1927 in Nelson, Missouri; 
B.S., University of Missouri, Columbia, 
1954: M.S. (1956) and Ph.D. (1965); 
Southern Illinois University. 

B. J.’s doctoral thesis on the biology of 
the striped skunk was the basis of his first 
book of that name published in 1967. Ear- 
lier, however, he was author of several pa- 
pers in the Journal of Mammalogy and oth- 


AWARDEES 107 


er major journals, having published 15 
refereed scientific papers on a wide variety 
of mammals before receiving the Ph.D. 

His first position after earning the MLS. 
degree was as District Biologist, North Car- 
olina Wildlife Resources Commission, fol- 
lowed by that of Field Mammalogist and 
Project Leader, Illinois Natural History 
Survey, a position held during his tenure as 
a doctoral student. Upon receiving his doc- 
toral degree, Verts was appointed Assistant 
Professor, Department of Fisheries and 
Wildlife, Oregon State University, where he 
has remained throughout his career, ad- 
vancing to the rank of Professor. He spent 
one year as Visiting Professor at Pennsy]l- 
vania State University. At Oregon State 
University he also curated the collection of 
mammals, developing it into the best col- 
lection of mammals from Oregon at any 
institution in the state. His endeavors are 
especially valuable because Oregon has no 
significant museum of natural history. 

In 1979, B. J. married fellow mammal- 
ogist Leslie Carraway. They collaborate ex- 
tensively, not only in revision of B. J.’s in- 
valuable ““Keys to the Mammals of Oregon,” 
but on virtually half of B. J.’s publications 
since 1980. Currently, they are completing 
a book on the Mammals of Oregon, the first 
of its kind since Vernon Bailey’s book writ- 
ten in 1936. 

B. J..s work focuses heavily on small 
mammals of Oregon, especially life histo- 
ries and distributions. His interest in rabies 
and other diseases communicated by wild 
mammals is prevalent in his earlier publi- 
cations. He has a long-term interest in de- 
vising techniques, such as those of ageing, 
baiting, and sexing. He is a major contrib- 
utor to Mammalian Species. 

The deep commitment that B. J. has to 
the ASM is reflected in the extent to which 
he contributes to the society. He has served 
as Managing Editor, Journal Editor, and As- 
sociate Editor of the Journal of Mammal- 
ogy, as Editor and Associate Editor of Mam- 
malian Species, as Chairman of the Local 
Committee for the ASM’s 59th Annual 
Meeting, as Chairman of both the Merriam 


Award Committee and the Grants-in-Aid 
Committee, and as a member of 5 other 
committees. He served two terms on the 
Board of Directors. In addition, B. J. has 
served in leadership capacities in other sci- 
entific societies related to wildlife. 

B. J. has supervised 18 M.S. students and 
2 Ph.D. students particularly on projects fo- 
cusing on cottontail rabbits. Students under 
his guidance learn the art of scientific writing. 
B. J. is a rigorous master, having coauthored 
with D. E. Wilson and A. L. Gardner the 
ASM’s 1989 Guidelines for Manuscripts and 
taught courses on science writing and on 
manuscript preparation at Oregon State 
University. He has guided many authors in 
the Journal of Mammalogy in his editorial 
capacities. 


Conclusions 


Altogether, 76 mammalogists have been 
honored by the ASM between 1919 and 
1992 (Tables 1, 2 and 3). Of these, 24 are 
Charter Members (no Jackson or Merriam 
awardees are in this group). The recipients 
come from 13 countries and represent near- 
ly every discipline related to the biology and 
evolution of mammals. Edouard-Louis 
Trouessart, who was made an Honorary 
Member in 1921, was the first foreign re- 
cipient, and in 1966 Erna Mohr, also from 
Europe, became the first woman to be hon- 
ored by the ASM. The only person to re- 
ceive all three honors—the Merriam Award 
in 1977, the Jackson Award in 1983, and 
Honorary Membership in 1992—1is the late 
J. Knox Jones, Jr., who also had been Pres- 
ident of the society. 

Of the 58 persons who have been given 
Honorary Membership, 14 are still alive; of 
the 12 people to receive Jackson Awards, 9 
are living; of the 14 recipients of the Mer- 
riam Award, 13 are living. 

Two of these three honors keep alive the 
names of two eminent founders of the ASM. 
C. Hart Merriam, first President of the so- 
ciety and one who not only began the North 
American Fauna series but also had a pro- 


108 TAYLOR AND SCHEER 


TABLE |1.—Honorary Members of the American Society of Mammalogists. (P) Past President of 


ASM. 


Joel Asaph Allen (1919) 
Edouard-Louis Trouessart (1921) 
Max Weber (1928) 

M. R. Oldfield Thomas (1928) 
Henry Fairfield Osborn (1929) 
Edward W. Nelson (1930) (P) 

C. Hart Merriam (1930) (P) 
William Berryman Scott (1936) 
Alfred W. Anthony (1936) 
Leonhard Stejneger (1937) 

Gerrit S. Miller, Jr. (1941) 
Ernest E. Thompson Seton (1941) 
Marcus Ward Lyon, Jr. (1942) (P) 
Rudolph M. Anderson (1947) 
Angel Cabrere Latorre (1947) 

A. Brazier Howell (1951) (P) 
Theodore S. Palmer (1951) 
Edward A. Preble (1952) 

Hartley H. T. Jackson (1952) (P) 
William K. Gregory (1954) 

W. P. Taylor (1954) (P) 

Harold E. Anthony (1955) (P) 
Lee R. Dice (1956) 

Albert R. Shadle (1956) 

Francis Harper (1959) 
Nagmaichi Kuroda (1959) 
Magnus A. Degerbol (1962) 
Remington Kellogg (1963) (P) 
Tracy I. Storer (1963) (P) 


TABLE 2.— Recipients of the Merriam Award. 
(P) = Past President of ASM; (Hon.) = Honorary 
Member of ASM; (Jack.) = recipient of the Jack- 


son Award. 


James N. Layne (1976) (P) 

J. Knox Jones, Jr. (1977) (P) (Hon., Jack.) 
James S. Findley (1978) (P) 

Terry A. Vaughan (1979) 

Robert J. Baker (1980) 

John F. Eisenberg (1981) 

James L. Patton (1983) (P) 
Michael H. Smith (1985) 

William Z. Lidicker, Jr. (1986) (P) 
Hugh H. Genoways (1987) (P) 
Jerry R. Choate (1988) 

James N. Brown (1989) (P) 
Timothy H. Clutton-Brock (1991) 
Guy G. Musser (1992) 


V. G. Heptner (1963) 

E. Raymond Hall (1964) (P) 
Stanley P. Young (1964) 
William J. Hamilton, Jr. (1965) (P) 
Erna Mohr (1966) 

Klaus Zimmerman (1966) 
William H. Burt (1968) (P) 
William B. Davis (1968) (P) 
George Gaylord Simpson (1969) 
Robert T. Orr (1970) (P) 
Stephen D. Durrant (1971) (P) 
Kazimierz Petrusewicz (1972) 
Charles S. Elton (1973) 

Emmet T. Hooper (1976) (P) 
Vladimir E. Sokolov (1976) 
Oliver P. Pearson (1979) 

Victor B. Scheffer (1981) 
Donald F. Hoffmeister (1982) (P) 
Z. Kazimierz Pucek (1982) 
Bjorn O. L. Kurtén (1983) 

John Edwards Hill (1985) 
Bernardo Villa-Ramirez (1986) 
Randolph L. Peterson (1986) (P) 
Francis Petter (1987) 

Wuping Xia (1988) 

Karl F. Koopman (1990) 

Philip Hershkovitz (1991) 

J. Knox Jones, Jr. (1992) (P) 
Sydney Anderson (1992) (P) 


TABLE 3.—Recipients of the Hartley H. T. 
Jackson Award. (P) = Past President of ASM; 
(Hon.) = Honorary Member of ASM; (Mer.) = 
recipient of Merriam Award. 


William B. Davis (1978) (P) (Hon.) 
William H. Burt (1979) (P) (Hon.) 
Bryan P. Glass (1980) 

J. Knox Jones, Jr. (1983) (P) (Hon. Mer.) 
Oliver P. Pearson (1984) (Hon.) 

Sydney Anderson (1985) (P) (Hon.) 
Murray L. Johnson (1986) 

Donald F. Hoffmeister (1987) (P) (Hon.) 
Karl F. Koopman (1988) (Hon.) 

Marie A. Lawrence (1990) 

John O. Whitaker, Jr. (1991) 

B. J. Verts (1992) 


AWARDEES 109 


found effect on the development of the sci- 
ence of modern mammalogy; and Hartley 
H. T. Jackson, eleventh President of the so- 
ciety, who chaired the initial Organizing 
Committee of the society and served as its 
first Corresponding Secretary for six years. 


Acknowledgments 


Recognition of the invaluable assistance pro- 
vided by several people at The Cleveland Mu- 


seum of Natural History in the preparation of 
this chapter is due. First ofall, the extensive work 
of B. Hallaran, Executive Secretary, is deeply 
appreciated. So is the help of librarians W. Was- 
man and D. Condon. We also are grateful to 
many members of the ASM, who helped to sup- 
ply informational details. To all we owe a debt 
of gratitude in helping to bring this chapter to- 
gether. 


OTHER PROMINENT MEMBERS 


Davip M. ARMSTRONG, MurrRAy L. JOHNSON, AND 


RANDOLPH L. PETERSON 


Introduction 


his chapter is based on the observation 
that many of the mammalogists who 
have had enduring impacts on mammalogy 
in the past 75 years have not been honored 
formally by the ASM as Honorary Members 
or recipients of Merriam or Jackson awards; 
not all have served the society as senior of- 
ficers. Given the organization of this vol- 
ume, such individuals might have been 
overlooked. 

This chapter has had a sadly difficult his- 
tory because one of the original authors, 
Randolph L. Peterson, passed away as con- 
ceptualization of the chapter was in an early 
stage. It was Peterson who drafted the first 
list of noteworthy mammalogists who— 
having neither been honored previously by 
ASM nor served as a senior officer of the 
society — might go unmentioned in this vol- 
ume. Peterson listed 76 names, and then 
more were added. The list quickly became 
unmanageable; difficult decisions eventu- 
ally had to be made. 

We understood at the outset that this 
chapter was unlikely to please everyone— 
and indeed might please no one—because 
space alone limited numbers of individuals 
included. Limits imply choice, and choice 


110 


implies valuing, which no two mammalo- 
gists are likely to do in the same manner. 
There was early agreement that to be in- 
cluded an individual must be retired or de- 
ceased. Further, it was abundantly clear that 
treatment could not be comprehensive. 
Eventually, some organizational principles 
emerged: the chapter would be organized by 
decades, and biographies would be limited 
to no more than about five individuals who 
had left an indelible stamp on the mam- 
malogical ‘“‘character’’ of that decade. Fi- 
nally, based on the premise that one cannot 
really recognize importance or a “classic” 
until its enduring impact can be gauged, we 
have not presumed to extend our subjective 
analysis beyond the 1970s. With standards 
and procedures so obviously judgmental, 
who could fault us for having omitted a fa- 
vorite theriological character or a particu- 
larly inspirational academic “aunt” or “‘un- 
cle,’ an esteemed mentor or field tutor? 
We do not harbor any illusion that the 
history of the ASM is the history of Amer- 
ican mammalogy. Mammalogy was well es- 
tablished as a branch of natural history and 
biology well before 1919. Many would date 
the origin of American mammalogy from 


OTHER PROMINENT MEMBERS 111 


1858, with the publication of Spencer Ful- 
lerton Baird’s monumental Mammals, Vol- 
ume 8 of the Pacific Railroad Surveys. Oth- 
ers would dig deeper for roots, to Colonial 
naturalists like Mark Catesby and William 
Bartram, distinguished visitors like Sir John 
Richardson, or to the extraordinary zoolog- 
ical explorers and publicists ofa new nation: 
Lewis and Clark, George Ord, James DeKay, 
John Godman, Thomas Say, or Audubon 
and Bachman. The late 19th and early 20th 
centuries were times of extraordinary pro- 
ductivity (see, for example, Hoffmeister and 
Sterling, 1994; Wilson and Eisenberg, 1990), 
and eminent mammalogists left marks that 
still inspire and influence our work, among 
them Harrison Allen, J. A. Allen, W. H. 
Osgood, G. S. Miller, Jr., C. Hart Merriam 
(who continues to sign the register annually 
at meetings of ASM a full half-century after 
his death). 


The 1920s 


The roster of the organizational meeting 
of ASM in 1919 reads like a ““Who’s Who” 
of late 19th and early 20th century mam- 
malogy. Many of the luminaries present went 
on to give distinguished service to mam- 
malogy and ASM and are noted elsewhere 
in this volume. The decade in mammalogy 
was characterized by self-evaluation and def- 
inition, and dominated by the pioneers. 
Early numbers of the Journal of Mammal- 
ogy published earnest correspondence about 
taxonomic issues, the dubious value of 
common names for organisms neither com- 
monly seen nor much discussed by common 
folk, still-useful lists of desiderata for life 
history studies, and the relative exchange 
value of specimens of mice and mink. 
Browsing through early volumes of the 
Journal and minutes of early meetings, one 
readily agrees that we continue to stand on 
the shoulders of those giants and continue 
to earn interest on the intellectual capital 
they invested. Of nine authors in the in- 
augural number of the Journal (28 Novem- 


ber 1919), six served as President of the 
ASM, five eventually were named Honor- 
ary Members, and three received both of 
those recognitions. Here we note a few other 
individuals who left a mark during the first 
decade of ASM. 

Outram Bangs (1863-1932) was born in 
Watertown, Massachusetts, and graduated 
from the Lawrence Scientific School of Har- 
vard College in 1884. In the 1890s, Bangs 
published 50 papers on mammals. Ina sense, 
Bangs represents a sizable class of individ- 
uals, largely unsung—the local naturalists. 
Like dozens of other noteworthy local 
mammalogists of his era, he began to collect 
mammals asa child. Eventually he built one 
of the finest private collections in the U.S., 
which was purchased by Harvard’s Muse- 
um of Comparative Zoology in 1899, and 
Bangs was named Assistant in Mammalogy, 
although his research interests soon shifted 
to birds. 

Ned Hollister (1876-1924) was the orig- 
inal editor of the Journal of Mammalogy, 
setting the high standards for editorial qual- 
ity that are matched by few other scientific 
journals. Born in Delavan, Wisconsin, he 
collaborated with Ludwig Kumlien of Mil- 
ton College on Birds of Wisconsin (1903), 
accompanied Vernon Bailey on a Biological 
Survey expedition to Texas (1902), and 
worked with W. H. Osgood in Alaska (1903). 
He formally joined the staff of the Bureau 
of Biological Survey in 1904. Reputed to 
have a genius for museum work (Osgood, 
1925), he was appointed Assistant Curator 
of Mammals in the U.S. National Museum 
in 1909; in 1916 he became Superintendent 
of the National Zoological Park. In his brief 
career, Hollister collected 26 holotypes, 
named 162 taxa, and published 150 papers 
and monographs, including several works 
of enduring value, among them work on 
mammals of the Philippines (1913) and re- 
views of East African mammals in the U.S. 
National Museum (1918, 1919, 1924). 

A. H. Howell (1872-1940) was the only 
author in the inaugural number of the Jour- 
nal of Mammalogy who did not go on to 


12 ARMSTRONG ET AL. 


the presidency of ASM or election to hon- 
orary membership. However, his impact on 
systematic mammalogy continues to be 
great, largely because he provided (mostly 
in North American Fauna) the first mono- 
graphic treatments of a number of mam- 
malian genera: striped skunks (1901), spot- 
ted skunks (1906), harvest mice (1914), 
marmots (1915), flying squirrels (1918), pi- 
kas (1924), chipmunks (1929), and ground 
squirrels (1938). His biological survey of Al- 
abama (1921) was the only such product of 
the Bureau of Biological Survey outside the 
Mountain West. Mostly self-trained, How- 
ell farmed and worked as a stock-clerk be- 
fore being stimulated to a career in natural 
history through an association with the Lin- 
naean Society of New York. He received a 
temporary appointment in 1895 as assistant 
to Vernon Bailey for field work in the 
Northern Rockies and Pacific Northwest. 
He continued with the Bureau of Biological 
Survey (and the Fish and Wildlife Service) 
until his death 44 years later. 

H. H. Lane (1878-1965) was the original 
Recording Secretary of ASM, serving from 
1919 until 1932. Born in Bainbridge, In- 
diana, and educated at DePauw, Indiana, 
Cornell, and Chicago, he received a Ph.D. 
from Princeton in 1915. Lane taught at Hir- 
am College, the University of Oklahoma, 
and Phillips University before moving to 
the University of Kansas as Professor of 
Zoology and Paleontology in 1922. Mostly 
a paleontologist, he nonetheless influenced 
the classic generation of mammalogists at 
the University of Kansas, including Wil- 
liam Henry Burt, E. Raymond Hall, Claude 
W. Hibbard, and Jean M. Linsdale. 


The 1930s 


The 1930s saw progress in a number of 
areas of mammalogy, especially in mam- 
malian ecology, and some of the most no- 
table contributions remain classic autoeco- 
logical studies. 

Robert T. Hatt (1902-1989) served as 


Corresponding Secretary of ASM from 1932 
to 1934. Born in Lafayette, Indiana, and 
educated at Michigan and Columbia, Hatt 
spent several years at the American Muse- 
um of Natural History and then directed 
the Cranbrook Institute of Science from 
1935 to 1967, remaining as Senior Scientist 
until his retirement in 1971. Hatt’s enduring 
contributions included fine autecological 
studies, especially of squirrels (e.g., Hatt, 
1943), and work in anatomy (Hatt, 1932). 

Robert K. Enders (1899-1989) pursued 
an extraordinarily diverse career, centered 
on academic work at Swarthmore College. 
He conducted field work on Panamanian 
mammals for more than 40 years, from 1929 
to 1971. Although he served as Recording 
Secretary of ASM from 1933 to 1937, and 
in a variety of scientific organizations and 
agencies in leadership capacities, his most 
indelible mark on mammalogy may have 
been indirect, a consequence of his stew- 
ardship of the Rocky Mountain Biological 
Laboratory at Gothic, Colorado, as Director 
(1959-1968) and President (1969-1978). He 
also stimulated students, such as Oliver 
Pearson and Phil Myers, to pursue careers 
in mammalogy. 

Jean M. Linsdale (1902-1973) was part 
of that legendary “‘bumper-crop” of mam- 
malogists born in Kansas, and educated at 
the University of Kansas and the University 
of California, Berkeley, that included W. H. 
Burt and E. R. Hall. He may have described 
his most important legacy to vertebrate zo- 
ology best in the acknowledgments to his 
monumental work, The California Ground 
Squirrel (1946); among the list of students 
at the Hastings Natural History Reservation 
who contributed as observers were Lamont 
C. Cole, Carl Koford, Lloyd Tevis, P. Q. 
Tomich, G. A. Bartholemew, Jr., W. W. 
Dalquest, H. S. Fitch, W. V. Mayer, and C. 
G. Sibley. Linsdale spent his professional 
career with the Museum of Vertebrate Zo- 
ology, joining in 1922 the “fur book”’ pro- 
ject begun three years earlier by Grinnell 
and Dixon (Grinnell et al., 1937). His pains- 
taking work on the dusky-footed woodrat 


OTHER PROMINENT MEMBERS 113 


(Linsdale and Tevis, 1951) helped inspire 
the career of a younger great neotomologist, 
R. B. Finley, Jr. 

Olaus J. Murie (1889-1963) was born in 
Moorhead, Minnesota, and served from 
1920 to 1946 as a field biologist with the 
Bureau of Biological Survey, including work 
in the Canadian Arctic, Labrador, and the 
Aleutians. His work on the elk of Jackson 
Hole (begun in 1927) is an enduring classic, 
in part culminating in E/k of North America 
(O. J. Murie, 1951). A Field Guide to Animal 
Tracks (1954) remains an invaluable re- 
source for naturalists who would read sto- 
ries of mammals not in the library but in 
dust, mud, or snow. A confirmed conser- 
vationist, Murie retired from government 
service to help found The Wilderness So- 
ciety, of which he was President from 1950 
to 1957. 

Adolph Murie (1899-1974) pursued his 
distinguished research career at the Uni- 
versity of Michigan (where as recently as 
1968 a pair of his boots occupied a place of 
honor in a specimen case), the U.S. Fish 
and Wildlife Service, and the National Park 
Service. After classic studies of moose on 
Isle Royale (A. Murie, 1934), he began re- 
search on gray wolf—Dall sheep interactions 
in Mount McKinley National Park in 1939. 
The Wolves of Mount McKinley (A. Murie, 
1944) and The Grizzlies of Mount McKinley 
(reprinted, 1981) continue to inspire. Like 
his older brother Olaus, Adolph Murie was 
passionately committed to conservation and 
the ideal of national parks: ““The national 
park idea is one of the bright spots in our 
culture. The idealism in the park concept 
has made every American visiting the na- 
tional parks feel just a little more worthy” 
(A. Murie, 1981:241). 

Aldo Leopold (1887-1948) continues to 
enrich our science and our philosophy near- 
ly a half-century after his untimely death. 
It is difficult to know which decade deserves 
to be identified with his remarkable contri- 
butions. The publication of his seminal 
Game Management (1933) essentially re- 
defined the field as applied ecology, nudging 


it hard from folk-art toward science. A Sand 
County Almanac appeared posthumously 
(1949), with sensitive, sensible insights into 
ecological ethics that continue to inspire 
students and their elders alike. In another 
dimension of his enduring legacy, several of 
Leopold’s children went on to distinguished 
scientific careers, in wildlife biology (Stark- 
er), paleobotany (Estella), plant physiology 
(Carl), and earth sciences (Luna). 

Francis B. Sumner (1874-1945) had an 
extraordinary career, documented in a re- 
markable autobiography (1945), The Life 
History of an American Naturalist. Educat- 
ed at Minnesota and Columbia, he taught 
at the College of the City of New York, and 
worked on fish development as Director of 
the Biological Laboratory of the Bureau of 
Fisheries at Woods Hole. Remarks by Da- 
vid Starr Jordan about the importance of 
long-term studies of the effects of environ- 
ment on evolution inspired his mammalog- 
ical work, which was made possible by an 
appointment at the Scripps Oceanographic 
Institute. Thus began a remarkable career 
in mammalogy, centered on painstaking 
laboratory studies of the genetics of geo- 
graphic variation in species of Peromyscus 
(see Sumner, 1932). 


The 1940s 


In the 1940s, many of a generation of 
mammalogists saw military service in World 
War II. An earlier generation of scholars 
continued to work despite limited academic 
and agency budgets and rationing of such 
theriological essentials as paper, gasoline, 
and tires, producing works that must still 
be consulted daily, such as G. G. Simpson’s 
Principles of Classification and a Classifi- 
cation of the Mammals. 

Victor H. Cahalane (1901-1993) was a 
Director of ASM at various times from the 
1930s to the 1960s. Director of the Cran- 
brook Institute of Science from 1931-1934, 
his scientific career was spent mostly with 
the U.S. National Park Service, resulting in 


114 ARMSTRONG ET AL. 


such studies as his survey of Katmai Na- 
tional Monument (1959). Chief of the Bi- 
ology Branch from 1944-1955, he remained 
as a collaborator until 1970 while Assistant 
Director of the New York State Museum. 
Perhaps Cahalane’s most enduring contri- 
butions were in the genre of popular natural 
history. Mammals of North America (1947), 
with its charming illustrations by Francis L. 
Jacques (1887-1969), remains an important 
landmark in mammalogical publishing, and 
The Imperial Collection of Audubon Mam- 
mals (Cahalane, 1967) made Audubon and 
Bachman’s illustrations of mammals readi- 
ly available to the 20th century. 

Paul Errington (1902-1962) received his 
Ph.D. from the University of Wisconsin and 
spent his entire academic career at Iowa State 
University. He devoted much of his too— 
brief scientific career to a single species, the 
muskrat, a keystone in the glacial marshes 
of the Midwest, research that began “... 
with muddy feet on the family farm in east- 
central South Dakota” (Errington, 1967:x1). 
His central question was what determines 
numbers of free-living animal populations, 
a question pursued in remarkable depth, as 
“the study of predation is no field for snap 
judgments” (1967:xi). Muskrat Populations 
(1963) remains a standard reference, and 
Errington did not hesitate to apply lessons 
learned from muskrats to humankind, as he 
did in Of Men and Marshes (1957), and the 
posthumous works, Of Predation and Life 
(1967), and The Red Gods Call (1973). 

D. Dwight Davis (1908-1965) was born 
in Rockford, Illinois, and joined the Field 
Museum in 1930, rising from Assistant in 
Osteology to Curator of Anatomy. His 
memoir on the functional morphology of 
the giant panda is a landmark in mammal- 
ogy (Davis, 1964), setting a new standard 
for morphological studies of species. In- 
deed, Gould (1980) called Davis’s mono- 
graph “... probably the greatest work of 
modern evolutionary comparative anato- 
my.” 

Ian McTaggart Cowan was born in 1910 
in Scotland and educated at the universities 


of British Columbia and California. His dis- 
tinguished academic career at the Univer- 
sity of British Columbia was marked by 
honorary degrees from Simon Fraser Uni- 
versity and the universities of Alberta, Wa- 
terloo, British Columbia, and Victoria. 
Cowan’s study of geographic variation in 
native American sheep (1940) was a pains- 
taking example of the possibilities of deep 
insights from fragmentary material. With 
Charles Guiguet, the Curator of Birds and 
Mammals at the British Columbia provin- 
cial Museum, he authored The Mammals 
of British Columbia (Cowan and Guiguet, 
1956), which has gone through three edi- 
tions. 

Philip L. Wright was born in 1914 and 
reared in New Hampshire, earning his doc- 
torate from the University of Wisconsin in 
1940. His entire professional career was 
spent at the University of Montana, where 
he retired in 1985. Wright’s research was 
focused mostly on reproductive cycles of 
endotherms, and his enduring contributions 
to mammalogy include a number of pio- 
neering papers on reproductive cycles of 
mustelids (e.g., Wright, 1942), as well as 
more recent work to maintain Boone and 
Crockett Club records on big game mam- 
mals. 


The 1950s 


The 1950s were optimistic years typified 
not only by big projects—of which E. R. 
Hall and K. R. Kelson’s Mammals of North 
America surely stands as the grandest— but 
also by big questions, on the nature of pop- 
ulation regulation, for example. Through the 
decade governmental support of mammal- 
ogy increased in North America, resulting 
in patterns of funding and academic rewards 
that prevail today. 

A. W. F. Banfield (born in Toronto in 
1918) studied at the universities of Toronto 
and Michigan and served as a mammalogist 
in several Canadian governmental agencies, 
including the National Park Service, the 


OTHER PROMINENT MEMBERS 1 Us) 


Wildlife Service, and the National Museum. 
He was Director of the Museum of Natural 
Science from 1964 to 1969 and later taught 
at Brock University. His contributions to 
mammalogy included definitive studies of 
the caribou over three decades (Banfield, 
1951, 1961), a faunal survey of Banff Na- 
tional Park (1958), and his comprehensive 
The Mammals of Canada (1974). 

Donald R. Griffin (born in 1915 in South- 
ampton, New York) has had two distin- 
guished careers in mammalogy, either of 
which would have earned him a prominent 
place in this chapter, in any of several de- 
cades. His academic career began at Cornell. 
While at Harvard, he published his classic 
Listening in the Dark (1958), which—along 
with Echoes of Bats and Men (1959)—con- 
tinues to inspire chiropterologists. In 1965 
he moved to Rockefeller University. The 
Question of Animal Awareness (1976) de- 
fined the new field of cognitive ethology and 
posed anew questions that had been dis- 
missed as scientifically inaccessible a cen- 
tury earlier. A recent Festschrift for Griffin 
(Ristau, 1991) provided appropriate rec- 
ognition for a distinguished mammalogist. 

John J. Christian was born in Pennsy!l- 
vania in 1917 and educated at Princeton 
and Johns Hopkins. In a research career in 
various commercial, federal, and academic 
laboratories, he pursued intensive experi- 
mental studies of the relationships among 
population density, reproduction, and the 
endocrine system, especially the adreno-pi- 
tuitary axis (reviewed in Christian, 1963), 
stimulating renewed interest in field studies 
of fluctuations of numbers of small mam- 
mals. He received the Mercer Award from 
the Ecological Society of America in 1957 
and was a professor at SUNY Binghampton 
from 1969 until his retirement. 

John B. Calhoun was born in Elkton, Ten- 
nessee, in 1917, and educated at the Uni- 
versity of Virginia and Northwestern. He 
taught at Emory, Ohio State, and Johns 
Hopkins. His research focused on principles 
of population dynamics, and he realized that 
“derivation of these principles requires more 


data than can be obtained by the efforts of 
a single individual” (Calhoun, 1956). In 
1947 he organized and initiated the North 
American Census of Small Mammals 
(NACSM), sponsored first by the Rodent 
Ecology Project at Johns Hopkins, later by 
Jackson Laboratory at Bar Harbor, Maine, 
and finally by the National Institutes of 
Mental Health (where Calhoun moved in 
1954). NACSM inspired volunteer field- 
work across the continent for a dozen years. 
By using consistent protocols, it not only 
developed a very large data set but under- 
scored the importance and the difficulties 
of achieving a quantitative understanding 
of mammalian distributions in space and 
time. Calhoun’s (1963) monograph on the 
ecology and sociology of the Norway rat was 
a landmark in considering in evolutionary 
and ecological terms the sociopathology of 
mammalian populations, both rats and peo- 
ple. 

Carl B. Koford (1915-1980) was selected 
in 1939 by Joseph Grinnell and Alden H. 
Miller to study the California condor with 
the support of the National Audubon So- 
ciety. Associated throughout his career 
mostly with the Museum of Vertebrate Zo- 
ology, Koford’s work was characterized by 
extraordinary attention to detail and thor- 
ough pursuit of connections and relation- 
ships. Fortunately, he turned these skills to 
understanding the ecology of the black-tailed 
prairie dog, providing (Koford, 1958) a clas- 
sic study of the species in the context of the 
dynamic and overused, but poorly known, 
ecosystem in which it is a kind of keystone. 
Fortunately, too, he invested his mono- 
graph with passionate concern for conser- 
vation that—in concert with the voices of 
such other committed mammalogists as 
Victor Cahalane, the brothers Murie, and 
E. R. Hall—finally is beginning to bear fruit. 


The 1960s 


The 1960s saw the advent of new tools 
and concepts like digital computers, mul- 


116 ARMSTRONG ET AL. 


tivariate statistics, and the use of ““biosys- 
tematic” characters in mammalogy. How- 
ever, several of the landmarks of the decade 
were broad summaries in their fields, in- 
cluding J. A. King’s edited Biology of Pero- 
myscus, Anderson and Jones’ edited Recent 
Mammals of the World, and Walker’s 
Mammals of the World. 

Barbara Lawrence (born in Boston in 
1909) was educated at Vassar College and 
was associated with the Museum of Com- 
parative Zoology at Harvard from 1931 un- 
til her retirement in 1976. In addition to 
important work on mammals of New En- 
gland, the Caribbean, and Central America, 
Lawrence collaborated with William Bos- 
sert to produce a ground-breaking multi- 
variate morphometric study of North 
American Canis (Lawrence and Bossert, 
1967) that demonstrated the power of new 
kinds of statistics in gaining insights into 
complex evolutionary and ecological ques- 
tions. 

E. Lendell Cockrum (born in 1920 in Ses- 
ser, Illinois) published a comprehensive 
systematic work on the mammals of Kansas 
(1952) and went on to pursue a distin- 
guished academic career at the University 
of Arizona. One of his most influential con- 
tributions to mammalogy was his textbook, 
Introduction to Mammalogy (1962), which 
served a generation of students. Cockrum 
also co-authored textbooks in general zo- 
ology and general biology and produced ma- 
jor studies of mammals of Organ Pipe Na- 
tional Monument (e.g., Cockrum, 1981). 

B. Elizabeth Horner (born in 1917 in 
Merchantville, New Jersey) received her 
Ph.D. from the University of Michigan in 
1948 and taught zoology at Smith College 
from 1940 until her retirement in 1982. In 
1970, she was named Myra M. Sampson 
Professor of Biological Science. Her mam- 
malogical contributions included classic 
studies of the biology of rodents, especially 
ecomorphology of Peromyscus (e.g., Hor- 
ner, 1954) and marsupials. 

W. Frank Blair (1912-1985) was born in 
Dayton, Texas, and educated at the uni- 


versities of Tulsa and Florida, as well as the 
Laboratory of Vertebrate Biology at the 
University of Michigan. Perhaps best known 
for his work in herpetology at the University 
of Texas, he left an indelible stamp on the 
development of mammalogy in several 
ways, and over a period sufficiently long that 
it is difficult to ascribe his influence to a 
particular decade. His works on the biotic 
provinces of Oklahoma (Blair and Hubbell, 
1938) and Texas (Blair, 1950) are still valu- 
able, and Vertebrates of the United States 
(Blair et al., 1957) was consulted by gen- 
erations of mammalogists. He was among 
the first ecologists to develop mark-recap- 
ture methods in studies of population ecol- 
ogy. Moreover, his leadership of the United 
States International Biological Program in 
the late 1960s and into the 1970s (see Blair, 
1977; Mares and Cameron, 1994) allowed 
deep insights into the functional role of 
mammals in ecosystems, and facilitated in- 
ternational cooperation among mammalo- 
gists that continues to expand. 

Ernest P. Walker (1891-1969) first made 
a mark on zoology with a 1913 book on 
birds of Wyoming. His monumental mam- 
malogical project, Mammals of the World, 
began in 1930 while he was Assistant Di- 
rector of the National Zoological Park, and 
continued for 30 years, resulting in the stan- 
dard semi-technical reference on the extant 
genera of mammals, now in its fifth edition 
(Nowak, 1991). The work was painstakingly 
thorough and attempted to include a pho- 
tograph of a representative species 1n each 
genus. The first edition (Walker et al., 1961) 
included a remarkable third volume, a clas- 
sified bibliography of the literature of mam- 
malogy, based in large part on the “Recent 
Literature” section of the Journal of Mam- 
malogy, which remains an efficient entry to 
the literature of mammalogy to about 1960. 
Walker’s original dedication was “‘To the 
MAMMALS, GREAT AND SMALL, who 
contribute so much to the welfare and hap- 
piness of man, another mammal, but re- 
ceive so little in return, except blame, abuse, 
and extermination.” 


OTHER PROMINENT MEMBERS Tt7 


The 1970s 


The investigational and analytic tools of 
the 1960s bore rich fruit in the 1970s. It is 
too early to guess just which works will turn 
out to be classics, of course, but the decade 
had more than its share of classic workers, 
many of whom figure prominently in other 
chapters in this volume. 

Rollin H. Baker (born in Cordova, Illi- 
nois, in 1916) was educated at the Univer- 
sity of Texas, Texas A&M University, and 
the University of Kansas. He established a 
reputation as an ornithologist with his 
monograph on the avifauna of Micronesia 
(1951), but his professional efforts at the 
University of Kansas, and later at Michigan 
State University, soon focused on mammals 
of Mexico and Michigan. He and his stu- 
dents did pioneering work on the biosys- 
tematics of Sigmodon, and his monumental 
Michigan Mammals (1983) is a paragon of 
state mammal books. Baker retired in 1981. 

Karl Kenyon (born in 1918 in La Jolla, 
California) was educated at Pomona and 
Cornell. After service in the U.S. Navy, he 
taught at Mills College. In 1947, he joined 
the U.S. Fish and Wildlife Service, under 
Victor B. Scheffer at the Fur Seal Laboratory 
(later the Marine Mammal Laboratory), 
pursuing a distinguished research career that 
made him the preeminent authority on the 
biology of the sea otter. His monograph on 
the biology of the species (Kenyon, 1969) 
will remain a classic of its genre. 

Ralph M. Wetzel (1917-1984) received 
his Ph.D. from the University of Illinois in 
1949. His professional career was spent 
mostly at the University of Connecticut, en- 
riched by research appointments at the U.S. 
National Museum. He retired in 1982 and 
moved to a courtesy appointment at the 
University of Florida State Museum. Wet- 
Zel’s well-known work in the Gran Chaco 
of Paraguay began in 1972. It was there that 
he discovered that the Chacoan peccary 
(Catagonus wagneri), previously known only 
from pre-Hispanic, subfossil deposits, re- 
mained alive (Wetzel, 1977), perhaps en- 


couraging a younger generation of mam- 
malogists to turn toward South America with 
the heightened sense that really remarkable 
discoveries remain to be made. 

Charles H. Southwick was born in Woo- 
ster, Ohio, in 1928, graduated from the Col- 
lege of Wooster, and earned master’s and 
doctoral degrees from the University of 
Wisconsin. After faculty appointments at 
Hamilton College, Ohio University, and 
Johns Hopkins (and research appointments 
at Oxford and Stanford), he moved to the 
University of Colorado in 1979 and retired 
there in 1993. Southwick’s research career 
is focused on population and behavioral 
ecology. He continues to make fundamental 
contributions to our knowledge of mam- 
malian species as diverse as grasshopper 
mice, pikas, and mule deer, but his enduring 
legacy surely will be in understanding the 
biology of species of Macaca. His longitu- 
dinal research effort on Indian populations 
of rhesus macaques (reviewed in Fa and 
Southwick, 1988), now over three decades 
long and continuing, may be unequalled for 
any species in the history of mammalogy. 
Further, he has shared his deep insights into 
the problems and prospects for global en- 
vironmental conservation through texts such 
as Ecology and the Quality of Our Environ- 
ment (Southwick, 1976) and Global Ecology 
(Southwick, 1988). 

William A. Wimsatt (1917-1987) was ed- 
ucated at Cornell and spent most of his ac- 
ademic career there. His research career fo- 
cused on the ecology and physiology of 
reproduction in eastern bats, especially My- 
otis lucifugus, and he pioneered techniques 
and insights (see Wimsatt and Kallen, 1957) 
that have since been applied to numerous 
other species. His edited series, Biology of 
Bats (1970a, 19706, 1977), brought togeth- 
er a vast quantity of information and atten- 
dant literature and made it accessible to a 
new generation of chiropterologists. 

Robert L. Rausch was born in 1921 in 
Marion, Ohio. From Ohio State University 
he received a bachelor’s degree in 1942 and 
a D.V.M. in 1945. He then earned an M.S. 


118 ARMSTRONG ET AL. 


from Michigan State University in 1946 and 
a Ph.D. from the University of Wisconsin 
in 1949, in parasitology and wildlife man- 
agement. He joined the Arctic Health Re- 
search Center of the U.S. Public Health Ser- 
vice, serving as Chief of the Zoonotic Disease 
Section from 1950 until its closure in 1974. 
Rausch was Adjunct Professor at the Uni- 
versity of Alaska from 1967 to 1974 and 
Professor of Zoology from 1974 to 1975. 
He served as Professor of Parasitology at 
the University of Saskatchewan from 1975 
to 1978 and then moved to the University 
of Washington, where he was Professor of 
Pathobiology in the School of Medicine and 
Adjunct Professor of Zoology until his re- 
tirement in 1992. As a mammalogist, 
Rausch established an international repu- 
tation for his systematic insights on Arctic 
mammals (e.g., Rausch, 1953) and received 
honorary degrees from the universities of 
Saskatchewan, Alaska, and Ziirich. Rausch’s 
wife, Virginia (Reggie), is a scientist in her 
own right and a frequent collaborator on 
joint projects (e.g., Rausch and Rausch, 
1975). 

Claude W. Hibbard (1905-1973) was born 
in Toronto, Kansas, and educated at the 
universities of Kansas and Michigan. He 
worked and taught at Kansas from 1928 to 
1946 and then moved back to Ann Arbor, 
where he pursued a highly productive career 
as an energetic and insightful student of 
Pliocene and Pleistocene faunas of the Great 
Plains, with a strong emphasis on mam- 
mals. His most lasting scientific contribu- 
tions were the development and use of a 
technique for collecting microfossils (de- 
scribed by Zakrzewski and Lillgraven, 1994). 

Walter W. Dalquest (born 1917) is difh- 
cult to identify with any particular decade, 
for his career has been long and diversely 
productive. Educated at the University of 
Washington and Louisiana State, he pub- 
lished comprehensive faunal treatments of 
mammals of Washington (1948) and San 
Luis Potosi (1953) and went on to a distin- 
guished academic career at Midwestern State 
University, Texas, making important con- 


tributions to the study of vertebrates (es- 
pecially mammals and fishes) of south-cen- 
tral United States and Mexico. Over the 
years, his research focused increasingly on 
fossil vertebrates, especially those of Plio- 
cene and Pleistocene localities. A well-de- 
served Festschrift (Horner, 1984) celebrated 
his contributions to students and science. 


A Final Word 


Given the diversity and purview of mam- 
malogy and mammalogists and the richness 
of research during the past three-quarters of 
a century, the foregoing survey can hardly 
hope to be definitive; indeed, it can be little 
more than suggestive. There was not even 
full agreement among the authors on whom 
to include. Peterson would have included 
more Canadians and chiropterologists, 
Johnson more northwesterners and theriol- 
ogists from beyond North America, and— 
unrestrained by wiser colleagues—Arm- 
strong would have been biased toward his 
own local heroes and mentors. 

Whether one agrees with our commis- 
sions or omissions is hardly the point, how- 
ever. Surely one cannot do science without 
understanding the process, and the process 
is a distinctly human enterprise, burdened 
with the full weight (and blessed with the 
full possibility) that “human” implies. If we 
see farther than our predecessors, it surely 
is because we stand on their shoulders. 


Literature Cited 


Baker, R.H. 1951. The avifauna of Micronesia. Uni- 
versity of Kansas Publications, Museum of Natural 
History, 3:1-359. 

1983. Michigan mammals. Michigan State 
University Press, East Lansing, xx + 642 pp. 

BANFIELD, A. W. F. 1951. The barren-ground cari- 
bou. Department of Resources and Development, 
Ottawa, v + 52 pp. 

. 1958. The mammals of Banff National Park, 

Alberta, Canada. Bulletin, National Museum of Can- 

ada, 159:1-53. 

. 1961. A revision of the reindeer and caribou, 


OTHER PROMINENT MEMBERS 119 


genus Rangifer. Biological Series, National Museum 

of Canada, 66:vi + 1-137. 

1974. The mammals of Canada. University 
of Toronto Press, Toronto, xxv + 438 pp. 

Biair, W. F. 1950. The biotic provinces of Texas. 
Texas Journal of Science, 2:93-117. 

1977. Big biology, the US/IBP. Dowden, 
Hutchison, and Ross, Stroudsburg, Pennsylvania, x 
+ 261 pp. 

Barr, W. F., A. P. BLarr, P. BRopKorRB, F. R. CAGLE, 
AND G. A. Moore. 1957. Vertebrates of the United 
States. McGraw-Hill Book Company, New York, 
819 pp. 

Bair, W. F., AND T. H. HuBBELL. 1938. Biotic dis- 
tricts of Oklahoma. The American Midland Natu- 
ralist, 20:425-454. 

CAHALANE, V.H. 1947. Mammals of North America. 
MacMillan, New York, x + 682 pp. 

. 1959. A biological survey of Katmai National 

Monument. Smithsonian Miscellaneous Collections, 

138:1-246. 

(ED.). 1967. The imperial collection of Au- 
dubon mammals. Hammond, Inc., Maplewood, New 
Jersey, xvi + 307 pp. 

CALHOUN, J. B. (ED.). 1956. 1951 annual report— 
North American Census of Small Mammals. Ad- 
ministrative Publication, U.S. Department of Health, 
Education, and Welfare, I + 164 pp. 

CALHOUN, J. B. 1963. The ecology and sociology of 
the Norway rat. Publications, U.S. Public Health 
Service, 1008:viii + 288 pp. 

CuRISTIAN, J. J. 1963. Endocrine adaptive mecha- 
nisms and the physiologic regulation of population 
growth. Pp. 189-353 in Physiological mammalogy 
(W. V. Mayer and R. G. Van Gelder, eds.). Academic 
Press, New York, 1:1-381. 

CockruM, E. L. 1952. Mammals of Kansas. Uni- 
versity of Kansas Publications, Museum of Natural 
History, 7:1-303. 

1962. Introduction to mammalogy. Ronald 

Press Company, New York, 455 pp. 

1981. Bat populations and habitats at the 
Organ Pipe National Monument. University of Ar- 
izona Press, Tucson, 31 pp. 

Cowan, I. McT. 1940. Distribution and variation in 
the native sheep of North America. The American 
Midland Naturalist, 24:505-580. 

Cowan, I. McT., AND C. J. GuiGuetT. 1956. The 
mammals of British Columbia. Handbook, British 
Columbia Provincial Museum, 1 1:1-413. 

DaLQuesT, W. W. 1948. Mammals of Washington. 
University of Kansas Publications, Museum of Nat- 
ural History, 2:1-444. 

1953. Mammals of the Mexican state of San 

Luis Potosi. Louisiana State University Press, 229 


pp. 

Davis, D. 1964. The giant panda: a morphological 
study of evolutionary mechanisms. Fieldiana: Zo- 
ology Memoirs, 3:1-339. 

ERRINGTON, P. L. 1957. Of men and marshes. Iowa 
State University Press, Ames, 150 pp. 

1963. Muskrat populations. Iowa State Uni- 

versity Press, Ames, x + 665 pp. 

1967. Of predation and life. Iowa State Uni- 

versity Press, Ames, xii + 277 pp. 


1973. The red gods call (C. Errington, ed.) 
Iowa State University Press, Ames, ix + 171 pp. 
Fa, J. E., AND C. H. SOUTHWICK (EDs.). 1988. Ecology 
and behavior of food-enhanced primate groups. Alan 

R. Liss Publishers, New York, 355 pp. 

Gou Lp, S. J. 1980. The panda’s thumb, more reflec- 
tions on natural history. W. W. Norton & Company, 
New York, 343 pp. 

GRIFFIN, D. R. 1958. Listening in the dark. Yale 
University Press, New Haven, 413 pp. 

. 1959. Echoes of bats and men. Anchor Books, 

Garden City, New York, 156 pp. 

1976. The question of animal awareness: 
evolutionary continuity of mental experience. 
Rockefeller University Press, New York, viii + 135 
pp. 

GRINNELL, J., J. S. Dixon, AND J. M. LINSDALE. 1937. 
Fur-bearing mammals of California: their natural 
history, systematic status, and relations to man. Uni- 
versity of California Press, Berkeley, 2 vols, 777 pp. 

Hatt, R.T. 1932. The vertebral columns of ricochet- 
al rodents. Bulletin of the American Museum of Nat- 
ural History, 63:599-738. 

1943. The pine squirrel in Colorado. Journal 
of Mammalogy, 24:311-345. 

HOFFMEISTER, D. F., AND K. STERLING. 1994. Origin. 
Pp. 1-21, in Seventy-five years of mammalogy (1919- 
1994) (E. C. Birney and J. R. Choate, eds.). Special 
Publication, The American Society of Mammalo- 
gists, 11:1-433. 

Ho uister, N. 1913. A review of the land mammals 
of the Philippine Islands in the United States Na- 
tional Museum. Proceedings of the United States 
National Museum, 46:299-341. 

. 1918. East African mammals in the United 

States National Museum. Part I. Insectivora, Chi- 

roptera, and Carnivora. Bulletin of the United States 

National Museum, 99:1-94. 

1919. East African mammals in the United 

States National Museum. Part II. Rodentia, Lago- 

morpha, and Tubulidentata. Bulletin of the United 

States National Museum, 99:1-175. 

. 1924. East African mammals in the United 
States National Museum. Part III. Primates, Artio- 
dactyla, Perissodactyla, Proboscidea, and Hyracoi- 
dea. Bulletin of the United States National Museum, 
99:1-151. 

Horner, B. E. 1954. Arboreal adaptations of Pero- 
myscus with special reference to use of the tail. Con- 
tributions from the Laboratory of Vertebrate Biol- 
ogy, University of Michigan, 61:1-84. 

Horner, N. V. (ED.). 1984. Festschrift for Walter W. 
Dalquest in honor of his sixty—sixth birthday. De- 
partment of Biology, Midwestern State University, 
Wichita Falls, Texas, 163 pp. 

Howe i, A. H. 1901. Revision of the skunks of the 
genus Chincha. North American Fauna, 20:1-62. 
1906. Revision of the skunks of the genus 

Spilogale. North American Fauna, 26:1-55. 

. 1914. Revision of the American harvest mice 

(genus Reithrodontomys). North American Fauna, 

36:1-97. 

1915. Revision of the American marmots. 

North American Fauna, 37:1-80. 


120 


. 1918. Revision of the American flying squir- 

rels. North American Fauna, 44:1-64. 

. 1921. A biological survey of Alabama. North 

American Fauna, 45:1-88. 

. 1924. Revision of the American pikas (genus 

Ochotona). North American Fauna, 47:1-57. 

1929. Revision of the American chipmunks 

(genera Tamias and Eutamias). North American 

Fauna, 52:1-157. 

. 1938. Revision of the North American ground 
squirrels with a classification of the North American 
Sciuridae. North American Fauna, 56:1-256. 

Kenyon, K. W. 1969. The sea otter in the eastern 
Pacific Ocean. North American Fauna, 68:xii1 + I- 
352. 

Kororp, C. B. 1958. Prairie dogs, whitefaces, and 
blue grama. Wildlife Monographs, 3:1-78. 

LAWRENCE, B., AND W. H. BossertT. 1967. Multiple 
character analysis of Canis lupus, latrans, and famili- 
aris, with a discussion of the relationships of Canis 
niger. American Zoologist, 7:223-232. 

Leopo.tp, A. S. 1933. Game management. C. Scrib- 
ner’s Sons, New York, xxi + 481 pp. 

1949. A Sand County almanac, and sketches 
here and there. Oxford University Press, New York, 
xill + 226 pp. 

LInSDALE, J. M. 1946. The California ground squir- 
rel. University of California Press, Berkeley, xi + 
475 pp. 

LINSDALE, J. M., AND L. B. Tevis, Jk. 1951. The 
dusky-footed woodrat. University of California Press, 
Berkeley, vii + 664 pp. 

Mares, M. A., AND G. N. CAMERON. 1994. Com- 
munity and Ecosystem Ecology. Pp. 348-376, in 
Seventy-five years of mammalogy (1919-1994) (E. 
C. Birney and J. R. Choate, eds.). Special Publica- 
tion, The American Society of Mammalogists, 11: 
1-433. 

Muri, A. 1934. The moose of Isle Royale. Miscel- 
laneous Publications, University of Michigan Mu- 
seum of Zoology, 25:1-44. 

. 1944. The wolves of Mount McKinley. Fau- 

na of the National Parks of the United States, 5:x + 

1-238. 

. 1981. The grizzlies of Mount McKinley. Sci- 
entific Monograph Series, National Park Service, U.S. 
Department of the Interior, 14:xvi + 1-251. 

Murig, O. J. 1951. The elk of North America. Stack- 
pole Publishing Co., Harrisburg, Pennsylvania, 376 


p. 
. 1954. A field guide to animal tracks. Hough- 
ton Mifflin, Boston, xxii + 374 pp. 
Nowak, R. M. (ED.). 1991. Walker’s Mammals of the 


ARMSTRONG ET AL. 


World, Fifth ed. Johns Hopkins University Press, 
Baltimore, 1:1-642, 2:643-1629. 

Oscoop, W. H. 1925. Ned Hollister. Journal of 
Mammalogy, 6:1-12. 

Rauscu, R. L. 1953. On the status of some Arctic 
mammals. Arctic, 6:91-148. 

Rauscu, R. L., AND V. R. RAuscH. 1975. Taxonomy 
and zoogeography of Lemmus spp., with notes on 
laboratory-reared lemmings. Zeitschrift fiir Sauge- 
tierkunde, 40:8-34. 

RistAu, C. A. (ED.). 1991. Cognitive ethology: the 
minds of other animals, essays in honor of Donald 
R. Griffin. Erlbaum Associates, Hillsdale, New Jer- 
sey, xx + 332 pp. 

SouTtHwick, C. H. 1976. Ecology and the quality of 
our environment. Second ed. Van Nostrand Rein- 
hold, New York, 426 pp. 

(ED.). 1988. Global ecology. Sinauer Associ- 
ates, Sunderland, Massachusetts, 323 pp. 

SuMNER, F. B. 1932. Genetic, distributional, and evo- 
lutionary studies of the subspecies of deermice (Pero- 
myscus). Bibliographica Genetica, 9:1-106. 

1945. The life history of an American nat- 
uralist. Jaques Cattell Press, Lancaster, Pennsylva- 
nia, vil + 298 pp. 

WALKER, E. P., ET AL. 1961. Mammals of the world. 
Johns Hopkins University Press, Baltimore, 1:1-644, 
2:647-1500, 3:1-769. 

WETZEL, R. M. 1977. The Chacoan peccary Cata- 
gonus wagneri (Eusconi). Bulletin of Carnegie Mu- 
seum of Natural History, 3:1—-36. 

Witson, D. E., AND J. F. EISENBERG. 1990. Origin 
and applications of mammalogy in North America. 
Current Mammalogy, 2:1-35. 

Wimsatt, W. A. (ED.). 1970a. Biology of bats. Aca- 
demic Press, New York, 1:1—406. 

(ED.). 1970b. Biology of bats. Academic Press, 

New York, 2:1-—477. 

(ED.). 1977. Biology of bats. Academic Press, 
New York, 3:1-651. 

Wimsatt, W. A., AND F.C. KALLEN. 1957. The unique 
maturation response of the Graafian follicles of hi- 
bernating vespertilionid bats and the question of its 
significance. Anatomical Record, 129:115-131. 

WriGut, P. L. 1942. Delayed implantation in the 
long-tailed weasel (Mustela frenata), the short-tailed 
weasel (Mustela cigonanti), and the marten (Martes 
americana). Anatomical Record, 83:341-349. 

ZAKRZEWSKI, R. J., AND J. A. LILLEGRAVEN. 1994. 
Paleomammalogy. Pp. 200-214, in Seventy-five years 
of mammalogy (1919-1994) (E. C. Birney and J. R. 
Choate, eds.). Special Publication, The American 
Society of Mammalogists, 11:1-433. 


ACADEMIC PROPINQUITY 


JOHN O. WHITAKER, JR. 


Introduction 


here have been at least three major pa- 
pers on the history of North American 
mammalogy and the ASM (Hamilton, 1955; 
Hoffmeister, 1969; Storer, 1969). However, 
none of these papers presented information 
on the “roots” or “‘academic genealogy” of 
North American mammalogists. The 75th 
anniversary of the birth of the ASM is a 
good time to examine this topic. The idea 
arose from a paper given by J. Knox Jones, 
Jr., at the 1985 annual meeting of ASM at 
Orono, Maine. It was titled ““Genealogy of 
Twentieth Century Systematic Mammalo- 
gists in North America: The Descendants 
of Joseph Grinnell,” and was subsequently 
published (Jones, 1991). Jones indicated that 
the descendants of Joseph Grinnell at the 
University of California, Berkeley, along 
with a major subcenter founded by E. Ray- 
mond Hall at the University of Kansas, ac- 
counted for an academic dynasty that in- 
cluded perhaps 75% of North American 
systematic mammalogists. Elmer Birney, 
president of ASM in 1989, suggested this 
topic be examined in more detail and other 
“dynasties” be included as a chapter in a 


121 


history of the society to be presented in con- 
junction with its 75th anniversary. This 
study is an attempt to trace the roots of 
mammalogy in North America during the 
first 75 years of the society. 

The base data for this paper are given 
(Table 1) asa listing of many mammalogists 
who “have made their mark or are making 
their mark” on North American mammal- 
ogy. Most are or were associated with the 
ASM. I have drawn heavily from Jones 
(1991) for the material on the Grinnell dy- 
nasty but there is no other published set of 
data to which one can go for related infor- 
mation. It had to be obtained by word of 
mouth and through correspondence. At- 
tempts were made to include as many of the 
more active North American mammalo- 
gists in this table as possible. However, not 
all could be included in the text, and little 
information could be obtained for some. I 
hope that omissions and oversights will not 
detract too greatly from the overall picture. 
The data accumulated should serve to in- 
dicate the source of our collective roots. The 
earliest group listed, the field mammalogists 


122 WHITAKER 


assembled for the United States Biological 
Survey by C. Hart Merriam, is not an ac- 
ademic group, but nevertheless made a ma- 
jor impact on North American mammalo- 
gy. Three major academic groups are 
included: the Harvard Group (Agassiz/Al- 
len), the Berkeley/Kansas Group (Grinnell/ 
Hall), and the Cornell Group (Hamilton). 
Besides those obtaining their training from 
members of these groups, there are some 
smaller groups (Florida, Purdue, Tulane, 
Wisconsin), and a number of mammalogists 
have received their degrees in related fields, 
such as ecology, ornithology, wildlife, and 
genetics. 


I. The Merriam Group 


Before discussing the academically-ori- 
ented dynasties, it is important to mention 
the group formed in the latter part of the 
last century under Clinton Hart Merriam of 
the U.S. Biological Survey (Table 1, Section 
I). C. Hart Merriam was trained as an M.D. 
in New York and practiced medicine from 
1879 to 1885 (Storer, 1969). However, Mer- 
riam was a field naturalist at heart and had 
written early natural history books on the 
birds of Connecticut (1877) and the mam- 
mals of the Adirondacks (1884). In 1885, 
he became chief of the federal bureau that 
later became the U.S. Biological Survey. He 
gathered under him a staff of outstanding 
mammalogists that published numerous 
papers and books and greatly influenced the 
development of mammalogy in this cen- 
tury. Members of his team included Vernon 
Bailey, Albert K. Fisher, Edward A. Gold- 
man, Ned Hollister, Arthur H. Howell, Har- 
tley H. T. Jackson, W. L. McAtee, Edward 
W. Nelson, Wilfred Osgood, Theodore S. 
Palmer, and Edward A. Preble. Merriam 
sent collectors into the field, stimulated nu- 
merous studies of distribution of mammals, 
and initiated the North American Fauna se- 
ries, which included the first comprehensive 
taxonomic studies of North American 
mammals. Six individuals within this group 


became presidents of the ASM, including 
the first president, Merriam himself (Layne 
and Hoffmann, 1994). Some of C. H. Mer- 
riam’s underlings said C. H. stood for 
“Christ Himself.” 

Merriam produced nearly 500 publica- 
tions, and he and his colleagues in the U.S. 
Biological Survey published numerous pa- 
pers and books that were largely responsible 
for the growth and development of system- 
atic mammalogy in North America in the 
late 1800s and early 1900s. It must be re- 
membered, however, that these men gen- 
erally were not associated with academic 
institutions and therefore had no means to 
train students except by example and ap- 
prenticeship. 


IT, The Agassiz/Glover 
Allen Group (Harvard) 


The Harvard group also originated before 
the formation of the ASM (Table 1, Section 
II), and traces back to Louis Agassiz at the 
Museum of Comparative Zoology at Har- 
vard College. J. A. Allen (1838-1921), or- 
nithologist and mammalogist, studied un- 
der Agassiz before moving to the American 
Museum of Natural History in 1895, as did 
another of the early notable mammalogists, 
Gerrit Smith Miller, Jr., who graduated from 
Harvard in the class of 1894. Miller first 
worked for the U.S. Biological Survey, but 
in 1898 moved to the U.S. National Mu- 
seum where he remained until retirement 
in 1940. Agassiz was at the base of this ac- 
ademic line, but one of his students, Glover 
M. Allen, was Curator of Mammals at Har- 
vard’s Museum of Comparative Zoology 
and sponsored most of the early mammal- 
ogists from Harvard. Allen earned three de- 
grees from Harvard, including his Ph.D. in 
1904. Glover Allen produced some of the 
giants of our time—George A. Bartholo- 
mew, Jr., David E. Davis, Donald R. Grif- 
fin, Charles Lyman, and Oliver P. Pearson. 

George Bartholomew was one of the most 
eminent physiological ecologists in this 


PROPINQUITY 123 


TABLE |.— Academic genealogy of selected 20th TABLE |.— Continued. 


century North American mammalogists. 


Harold Reynolds 


I. C. Hart Merriam Group (U.S. Biological Barbara Lawrence Scheville 


Survey, Washington) 

C. Hart Merriam 
Vernon Bailey 
Albert K. Fisher 
Edward A. Goldman 
Ned Hollister 
Arthur H. Howell 
Hartley H. T. Jackson 
W. L. McAtee 
Edward W. Nelson 
Wilfred Osgood 
Theodore S. Palmer 
Edward A. Preble 
Stanley P. Young 


II. Harvard University (The Agassiz/Allen Group) 


Louis Agassiz 
Bryan Patterson 
Craig C. Black 
J. Sutton 
Lloyd E. Logan 
L. Kristalka 
I. Johnson 
Glover M. Allen 
George A. Bartholomew, Jr. 
Mark A. Chappell 
William R. Dawson 
Richard W. Hill 
Alan R. French 
Jack W. Hudson, Jr. 
James G. Kenagy 
Richard E. MacMillen 
Daniel K. Odell 
Thomas Poulson 
Barbara H. Blake 
Bruce Wunder 
David E. Davis 
John J. Christian 
Edward N. Francq 
Ronald E. Barry 
Frank B. Golley 
Rexford D. Lord 
Jan O. Murie 
Steven H. Vessey 
Donald R. Griffin 
Jack Bradbury 
Katherine Ralls 
Charles Lyman (Allen/Hisaw) 
Richard W. Thorington, Jr. (Ernst Mayr) 
Oliver Pearson (Allen/Hisaw) 
Daniel H. Brant 
Donald R. Breakey 
Gilbert S. Greenwald 
Stuart O. Landry 
Bert S. Pfeiffer 


J. A. Allen 
Herbert W. Rand 
Harold B. Hitchcock 


III. The Joseph Grinnell/E. Raymond Hall Group 


(Berkeley and the University of Kansas) 
Joseph Grinnell 
Seth Benson 
Robert L. Rudd 
Guy N. Cameron 
Peter Schramm 
Charles S. Thaeler 
Enrique P. Lessa 
Alan C. Ziegler (technically with W. B. 
Quay) 
W. H. Burt 
A. W. Frank Banfield 
Fred S. Barkalow 
Harold E. Broadbooks 
Robert K. Enders (Burt was mentor but 
not advisor) 
Lowell L. Getz 
Joyce Hoffman 
Donald H. Miller 
Harvey L. Gunderson 
Evan B. Hazard 
Timothy E. Lawlor 
Richard H. Manville (final examination 
chaired by Hooper) 
Illar Muul 
William O. Pruitt 
Dana P. Snyder 
Wendell E. Dodge 
Andrew Starrett 
Ian McTaggart Cowan 
Joseph F. Bendell 
Fred C. Zwickel 
Walter A. Sheppe 
William B. Davis 
Dilford C. Carter 
Patricia Dolan 
Richard K. Laval 
Donald A. McFarlane 
Ronald H. Pine 
Raul Valdez 
Paul W. Parmalee 
Randolph L. Peterson (Ph.D. with J. R. 
Dymond) 
Charles S. Churcher 
Judith L. Eger 
M. Brock Fenton 
Robert M. R. Barclay 
Gary P. Bell 
R. Mark Brigham 
Joe E. Cebek 


124 


TABLE 1.— Continued. 


WHITAKER 


James H. Fullard 
Robert M. Herd 
C. G. Van Zyll de Jong 


Lee R. Dice 


W. Frank Blair 
David L. Jameson 
Michael A. Mares 
Ruben M. Barquez 
Thomas E. Lacher, Jr. 
Ricardo Ojeda 
Michael R. Willig 
W. Howard McCarley 
Paul G. Pearson 
Richard D. Sage 
James R. Tamsitt 
Wallace D. Dawson 
Van T. Harris 
Don W. Hayne 
Paul C. Connor 
B. Elizabeth Horner 
Walter E. Howard 
Daniel B. Fagre 
John A. King 
Lee C. Drickamer 
C. Richard Terman 
Harley B. Sherman 
B. A. Barrington 
Joseph C. Moore 
Dale W. Rice 
Arthur Svihla 


E. Raymond Hall 


Ticul Alvarez-S. (Masters) 
Sydney Anderson 
Rollin H. Baker 
Donald P. Christian 
Peter L. Dalby 
James M. Dietz 
Gary A. Heidt 
Gordon L. Kirkland, Jr. 
John O. Matson 
Alan E. Muchlinski 
Howard J. Stains 
M. D. Bryant 
E. Lendell Cockrum 
Robert J. Baker 
John W. Bickham 
Luis Ruedas 
William J. Bleier 
J. Hoyt Bowers 
Robert D. Bradley 
Ira F. Greenbaum 
David Hale 
Philip Sudman 
Mike Haiduk 
Meredith Hamilton 
Rodney L. Honeycutt 


TABLE |.— Continued. 


Craig S. Hood 
David C. Kerridge 
Rick McDaniel 
Margaret A. O’Connell 
Calvin A. Porter 
Mazin B. Qumsiyeh 
Lynn W. Robbins (actual advisor was 
Francis Rose) 
Fred B. Stangl, Jr. 
Ron Van Den Bussche 
Terry L. Yates 
Joseph A. Cook 
Scott L. Gardner 
Sarah George 
Gregory D. Hartman 
Laura L. Janacek 
Dwight W. Moore 
David Reducker 
Brett R. Riddle 
Robert M. Sullivan 
Glen Bradley 
Russell P. Davis 
Bruce J. Hayward 
Keith Justice 
Peter L. Meserve 
James D. Layne 
C. Brian Robbins 
Robert G. Schwab 


Charles L. Douglas 
Stephan D. Durrant 


Richard M. Hansen 
Donald R. Johnson 
Keith R. Kelson 
M. Raymond Lee 
Fred Elder 
Mark L. McKnight 
William S. Modi 
Earl G. Zimmerman 
C. William Kilpatrick 
John V. Planz 


James S. Findley 


Kenneth W. Anderson 
Hal L. Black 
Michael A. Bogan 
William Caire 
Eugene D. Fleharty 
Patricia W. Freeman 
Kenneth N. Geluso 
Anthony L. Gennaro 
David J. Hafner 
Arthur H. Harris 
Clyde Jones 
John F. Pagels (co-chairs were Negus and 
Jones) 
Karen E. Petersen 
Daniel F. Williams 


PROPINQUITY 125 


TABLE |.— Continued. 


Don E. Wilson 
Robert B. Finley 
Donald F. Hoffmeister 

Wayne H. Davis 

Luis de la Torre 

Victor E. Diersing 

L. Scott Ellis 

John S. Hall 

W. Z. Lidicker, Jr. 
Blair A. Csuti 
K. T. DeLong 
Ayesha E. Gill 
Edward J. Heske 
David T. Krohne 
William F. Laurance 
Richard S. Ostfeld 
David O. Ribble 
Jeffy O. Wolff 

Charles A. McLaughlin 

Iyad A. Nader 

David J. Schmidly 
Paisley S. Cato (co-chaired with Clyde 
Jones) 
James N. Derr (co-chaired with John 
Bickham) 


Robert C. Dowler (co-chaired with John 


Bickham) 
Mark D. Engstrom 
James G. Owen 
Stephen A. Smith (co-chaired with Ira 
Greenbaum) 
William D. Severinghaus 
H. Duane Smith 
Richard G. Van Gelder 
David B. Wright 
Robert E. Wrigley 
J. Knox Jones, Jr. 
David M. Armstrong 
Kathleen A. Scott Fagerstone 
James C. Halfpenny 
Joseph F. Merritt (actual advisor was 
Olwen Williams) 
Elmer C. Birney 
Richard Lampe 
Lynn L. Rogers 
Robert M. Timm (actual advisor was 
Roger Price) 
John B. Bowles 
Alberto A. Cadena 
Jerry R. Choate 
Larry L. Choate 
G. Lawrence Forman 
Hugh H. Genoways 
Robert R. Hollander 
Thomas H. Kunz 
Edythe L. P. Anthony 


TABLE |.— Continued. 


Peter V. August 
Martha S. Fugita 
Allen Kurta 
Richard W. Manning 
Carleton J. Phillips 
Ronald W. Turner 
James Dale Smith 
Philip L. Krutzsch 
Charles A. Long 
George H. Lowery, Jr. 
Walter W. Dalquest 
Alfred L. Gardner 
Ronald M. Nowak 
Robert L. Packard 
Robert E. Martin 
Robert J. Russell 
Henry W. Setzer 
Duane A. Schlitter (actual advisor was 
Richard Highton, a herpetologist) 
Terry A. Vaughan 
Cindy Rebar 
O. J. Reichman 
Bernardo Villa-R. (Masters with Hall, Ph.D. 
from Univ. Mexico) 
Jose Ramirez Pulido 
John A. White 
John Eric Hill 
Emmet T. Hooper 
James H. Brown 
Michael A. Bowers 
Gerardo Ceballos 
James C. Munger 
Andrew T. Smith 
Michael D. Carleton 
Theodore H. Fleming 
Charles O. Handley, Jr. 
David G. Huckaby 
David Klingener 
James A. Lackey 
Guy G. Musser 
Albert Schwartz 
David H. Johnson 
A. Remington Kellogg (actual chair was 
William D. Mathew) 
Jean M. Linsdale 
Quentin P. Tomich 
Alden H. Miller (ornithologist) 
Richard F. Johnston (ornithologist) 
Gary Schnell (ornithologist) 
Troy L. Best 
Janet K. Braun 
Ronald K. Chesser 
E. Gus Gothran 
Michael L. Kennedy 
George D. Baumgardner 
Floyd W. Weckerly 


126 


TABLE |.— Continued. 


Robert D. Owen 
Carl B. Koford 
A. Starker Leopold 
Joseph G. Hall 
William J. Hamilton III 
Robert S. Hoffmann 
Fernando A. Cervantes-Reza 
Lawrence R. Heaney 
Donald L. Pattie 
Barbara R. Stein 
Merlin D. Tuttle 
John E. Warnock 
W. Christopher Wozencraft 
John H. Kaufmann 
Frank J. Bonaccorso 
Richard R. Lechleitner 
Frank A. Pitelka 
George O. Batzli 
Russell F. Cole 
Elizabeth A. Desy 
Richard Lindroth 
Stephen D. West 
Charles A. Reed 
Emily C. Oaks 
J. Mary Taylor 
Barry Thomas 
Marla L. Weston 
Robert T. Orr 


Tracy I. Storer (actual chair was Charles A. 


Kofoid) 
Walter P. Taylor 
Bryan P. Glass 
Stephen R. Humphrey 
Hector T. Arita 
Jacqueline Belwood 
Ralph Kirkpatrick 
Frederick H. Test 


IV. The Hamilton Group (Cornell University) 


William J. Hamilton, Jr. 
Roger W. Barbour 
Michael J. Harvey 
Marion Hassell 
Allen V. Benton 
Arthur H. Cook 
Robert A. Eadie 
Kyle R. Barbehenn 
Richard W. Dapson 
Harold G. Klein 
Jack W. Gottschang 
Everett W. Jameson 
Duncan Cameron, Jr. 
John D. Phillips, Jr. 
James N. Layne 
Harrison Ambrose 
William Platt 
Andrew A. Arata 


WHITAKER 


TABLE |.— Continued. 


Dan W. Walton 
Dale E. Birkenholz 
Llewellyn M. Ehrhart 
James V. Griffo 
John McManus 
Elizabeth S. Wing 
William O. Wirtz 
James L. Wolfe 

Robert J. Esher 

John G. New 
William G. Sheldon 
William Werner 
John O. Whitaker, Jr. 
Wynn W. Cudmore 
Thomas W. French 
Gwilym S. Jones 

Howard H. Thomas 
David Pistole 
Steven J. Ropski 


From Professors in Related Fields 
Ecology 


Marston Bates 
John W. Twente 


Arthur D. Hasler 
Kenneth B. Armitage 
Orlando A. Schwartz 


Charles Elton 
Dennis Chitty 
Rudy Boonstra 
Charles J. Krebs 
Michael S. Gaines 
Leroy R. McClenaghan 
Robert K. Rose 
Barry L. Keller 
Robert H. Tamarin 
Steven R. Pugh 
Francis C. Evans 
Lee H. Metzgar 
Stanley C. Wecker 
Richard R. Miller 


John T. Emlen 
Garrett C. Clough 
William A. Fuller 


Evelyn Hutchinson 
Donald Livingston 
Peter D. Weigl 
Robert H. MacArthur 
M. L. Rosenzweig 
Joel S. Brown 
Burt P. Kotler 
Cliff Lemon 
Gene D. Schroder 


John C. Neese 
Tim W. Clark 


TABLE |.— Continued. 


Eugene Odum 
W. Wilson Baker 
Gary W. Barrett 
Richard S. Mills 
Reed Fantin 
Clyde L. Pritchett 
William Prychodko 
Mary Etta Hight 


William Reeder 
Frank A. Iwen 


Victor Shelford 
S. Charles Kendeigh 

Robert M. Chew 

John A. Sealander, Jr. 
Donald W. Davis 
Philip S. Gipson 

Dana Snyder 

Ralph Wetzel 
Robert L. Martin 


Genetics 
Peter Brussard (ecological genetics) 
Gary F. McCracken 
Robert Lacey 


Theodosius Dobzhansky 
Karl F. Koopman 


W. B. Heed 
James L. Patton 
John C. Hafner 
Mark Hafner 
Philip Myers 
G. K. Creighton 
Robert Voss 
Duke S. Rogers 
Margaret F. Smith 
Donald O. Straney 
A. Christopher Carmichael 


Ethology 
M. W. Fox 
Marc Bekoff 
Joel Berger 


Peter Marler 
John F. Eisenberg 
Cheri Jones 
John G. Robinson 
R. Rudran 
Nicholas C. Smythe 
C. Wenimer 


Franz Sauer 
Michael H. Smith 
Mark C. Belk 
Donald W. Kaufman 
Paul L. Leberg 


PROPINQUITY 127 


TABLE 1|.— Continued. 


Susan McAlpine 
Paul R. Ramsey 
Kim T. Scribner 


Wildlife/Conservation 


Aldo Leopold 
James R. Beer 
Charles F. MacLeod 
Charles M. Kirkpatrick 
Thomas W. Hoekstra 
Russell E. Mumford 
Virgil Brack, Jr. 
David A. Easterla 
Harmon P. Weeks 


William H. Marshall 
John R. Tester 
Donald B. Siniff 
Douglas P. DeMaster 
J. Ward Testa 
Jeannette A. Thomas 


Robert A. McCabe 
Lloyd B. Keith 
Thomas A. Scott/Edward Kozicky 
Willard D. Klimstra 
B. J. Verts 
Leslie N. Carraway (actual advisor was 
Charles Warren) 
Joseph A. Chapman 
Kenneth L. Cramer 
George A. Feldhamer 


Entomology & Parasitology 

H. S. Fitch/Joseph Camin 
Richard B. Loomis 

Cluff Hopla 
Donald Gettinger (co-chaired with Michael A. 
Mares) 

Adrian Marshall 
Donald W. Thomas 


Anatomy/Physiology 
Howard Adelmann 
William A. Wimsatt 
Roy Horst 
Alvar W. Gustafson 
Gary G. Kwiecinski 


William J. McCauley 
Henry Mitchell 
G. Clay Mitchell 
Eugene H. Studier 
Roland K. Meyer (endocrinologist) 
William H. Elder 
Richard F. Myers 
Phillip L. Wright 
Clinton H. Conaway 


128 WHITAKER 


TABLE 1.— Continued. 


Larry N. Brown 
Milo E. Richmond 
Frederick J. Jannet 
John P. Hayes 
Rodney A. Mead 


Andrew V. Nalbandov (Univ. IIl., animal science) 
Glen C. Sanderson 


Alfred C. Redfield (Harvard, physiology) 
Peter R. Morrison 
Brian K. McNab 


Herpetology 


Robert Stebbins 
Paul K. Anderson 


Ornithology 


Arthur A. Allen 
Ralph S. Palmer 


Eugene Dustman 
Norman Negus 
Pat Berger 
Robert K. Chipman 
Jack A. Cranford 
Alicia T. Linzey 
Edwin Gould 
John F. Pagels (co-chaired with Clyde Jones) 
Aelita S. Pinter 
Carol N. Rowsemitt 
Thomas E. Tomasi 
Miles Pirnie 
Durwood L. Allen 
Frederick F. Knowlton 
Charles E. Harris 
L. David Mech 
Michael E. Nelson 
Rolf O. Peterson 
Fred A. Ryser, Jr. 
John R. Gustafson 
Herbert W. Rand 
Harold W. Hitchcock 


Miscellaneous 
William King Gregory (palaeontologist) 
Albert E. Wood 
Bjorn Kurten (palaeontologist) 
Phillip M. Youngman 
William F. Porter 
Paul F. Steblein 
S. David Webb (palaeontologist) 
Kenneth T. Wilkins 
Training in Other Professions 
Physicians 
H. Allen 
Elliot Coues 
Murray L. Johnson 


TABLE |.— Continued. 


Marcus Ward Lyon, Jr. 

Edgar A. Mearns 

C. Hart Merriam 

George Wislocki 
Veterinarians 

Denny J. Constantine 


Training in Museum or Field, No Ph.D. 
Rudolph M. Anderson 
Harold E. Anthony 
Benjamin P. Bole, Jr. 
Philip M. Blossom 
Victor Cahalane 
T. Donald Carter 
J. Kenneth Doutt 
Alfred J. Godin 
George F. Goodwin 
Arthur M. Greenhall 
Philip Hershkowitz 
A. Brazier Howell 
Laurence M. Huey 
Carl W. Kenyon 
Thomas J. McIntyre 
Gerrit S. Miller, Jr. 
John Paradiso 
Victor B. Scheffer 
Ermest Thompson Seton 
Albert R. Shadle 
Viola S. Shantz 
G. H. H. Tate 
Lloyd P. Tevis 
Hobart M. Van Deusen 
Ernest P. Walker 


country and trained a number of students 
at UCLA. Davis and Lyman have been ex- 
tremely influential in studies of hibernation: 
Davis at Penn State and North Carolina 
State; Lyman at Harvard. Griffin has had 
immense effect on studies of bat echoloca- 
tion and behavior from positions at Har- 
vard, Cornell, and Rockefeller University. 
Oliver Pearson of Berkeley is an ecological 
physiologist, well known for his work with 
poison glands of shrews, mammalian re- 
production, and ecology and systematics of 
South American mammals. Pearson, like 
William J. Hamilton, Jr., was greatly influ- 
enced by Francis Harper. Harper had earlier 
been a high school teacher, but was editing 
for the American Philosophical Society and 
frequently used the library at the Philadel- 


PROPINQUITY 129 


phia Academy of Science. Oliver Pearson 
used the library in conjunction with his work 
for Robert Enders and thereby came in con- 
tact with Harper, who had obtained his 
Ph.D. from Cornell in 1925 with the her- 
petologist, Albert Hazen Wright. All five of 
these Glover Allen-progeny have now pro- 
duced academic offspring of their own. The 
influence of Harvard on the development 
of North American mammalogy cannot be 
overestimated. 


ITI. The Joseph Grinnell/ 
E. R. Hall Group 
(Berkeley and Kansas) 


Early in this century, another intellectual 
dynasty was born on the West Coast, at 
Berkeley (Table 1, Section III). It was fos- 
tered by Annie Montague Alexander, who 
played an outstanding role in the develop- 
ment of mammalogy at Berkeley (H. Grin- 
nell, 1958). She was the founder and a life- 
long patron of the Museum of Vertebrate 
Zoology at Berkeley. She early developed a 
love for travel, hunting, and the natural sci- 
ences. Alexander also befriended C. Hart 
Merriam, and collected or purchased many 
of the bears that were studied by him; she 
supported and led three collecting expedi- 
tions to Alaska (1906, 1907, and 1908). 

Alexander had thought for some time 
about establishing a museum at the Uni- 
versity of California. When she returned 
from Alaska in the autumn of 1906 she be- 
gan serious discussions with Merriam about 
this. She had come to realize how fast the 
native game birds and mammals of the west 
were disappearing and felt specimens (in- 
cluding skeletons) should be preserved, as 
was happening in the east. At this time she 
happened to meet Joseph Grinnell, and was 
impressed with his “energy and enthusiasm 
and the neat and scholarly way in which his 
records were kept.” She mentally noted him 
as a possible coworker. 

Upon returning from her 1907 Alaska ex- 
pedition, Alexander presented her plan for 


the establishment of a museum of verte- 
brate zoology at The University of Califor- 
nia to President Benjamin Wheeler. The re- 
gents accepted her plan and a contract 
establishing the museum was signed on 23 
March 1908, with Joseph Grinnell appoint- 
ed as its director for | year. 

Many letters were exchanged between Al- 
exander and Grinnell in order to ensure the 
greatest possible usefulness for the museum. 
Alexander preferred that young biologists 
be enlisted, ‘““men with their accomplish- 
ments ahead of, rather than behind them,” 
and that the time of staff members should 
be divided between curatorial, field, and re- 
search work. There was effort to obtain bal- 
ance between specimens for research and for 
display in order to kindle popular interest 
in natural history. Alexander contributed 
monthly sums from 1908 to 1919, then she 
presented $200,000, plus another $225,000 
in 1936, as perpetual endowments. How- 
ever, she also gave many smaller amounts 
through the years until her death, and con- 
tributed hundreds of specimens collected by 
herself and her lifelong friend, Louise Kel- 
logg. 

The University of California wanted 
Grinnell to teach freshman Zoology, but Al- 
exander objected. She wanted his time spent 
on research and development of the mu- 
seum. However, Grinnell did become editor 
of the Condor in 1908 and continued in this 
position until his death in 1939. Head- 
quartering the Condor at Berkeley provided 
practice in editing to numerous students. 

Joseph Grinnell was born in 1877 in the 
Indian Territory, about 40 miles from Ft. 
Sill, in present-day Oklahoma. His family 
settled in California after his father’s retire- 
ment. Grinnell earned the bachelor’s degree 
from Throop Polytechnic Institute, which 
eventually became the California Institute 
of Technology, in 1897. He earned the M.A. 
and Ph.D. degrees from Leland Stanford, 
Jr., College in 1901 and 1913. This insti- 
tution was named for its benefactor, Leland 
Stanford, Jr., and later became Stanford 
University. His major professor or at least 


130 WHITAKER 


one of them was Charles Henry Gilbert 
(Hall, 1939). Grinnell taught at Throop 
Polytechnic for a time before becoming Di- 
rector of the Museum of Vertebrate Zoology 
in 1908. He held this post for 30 years, until 
shortly before his death at 62 in 1939. Grin- 
nell had styled himself after C. Hart Mer- 
riam; thus the roots of the Grinnell Dynasty 
go back partly to Merriam. However, the 
roots also reached back to another giant in 
vertebrate zoology of the time, David Starr 
Jordan. Jordan was primarily an ichthyol- 
ogist, but had broad interests in other ver- 
tebrates as well. Jordan did his undergrad- 
uate work at Cornell, where it is said that 
he camped out on campus. He earned an 
M.D. at Indiana Medical College in 1875, 
and a Ph.D. from Butler University (Indi- 
anapolis) in 1878. Jordan was President of 
Indiana University from 1885 to 1891, and 
in 1891 he became the first President of 
Leland Stanford, Jr., College. 

Grinnell was an excellent mammalogist 
and ornithologist, and an expert on birds 
and mammals of the West Coast, especially 
California. He was very shy, but an ener- 
getic worker in the field. His shyness man- 
ifested itself, for example, in instinctively 
placing his own hand behind his back when 
a newcomer offered to shake it. He was an 
excellent scientist, editor, and museum cu- 
rator. Emmet T. Hooper, one of Grinnell’s 
students, said that Grinnell would drive on 
trips into the field and would point out in- 
teresting geological, vegetative, or faunal 
features. On the return trip, however, he 
would let a student drive while he sat in the 
back, in order to work up his field notes and 
even start work on the papers to be pub- 
lished from the specimens and data ob- 
tained. 

During his tenure at Berkeley, Grinnell 
advised numerous graduate students in or- 
nithology and mammalogy, and also some 
in herpetology, but not all were his students 
in the strict sense that he was their major 
advisor. Charles A. Kofoid also played a 
major role in the education of many Berke- 
ley graduate students. Berkeley students 


fanned out over the land; they have played 
a major role in systematic mammalogy, and 
in vertebrate zoology as a whole throughout 
the world. Some of Grinnell’s better known 
students, not all of whom he directed to the 
doctoral degree, were the following (Table 
1, Section ITI). 


Seth Benson and Alden H. Miller (Berkeley) 

William H. Burt, Lee R. Dice, Emmet T. 
Hooper, and Fred R. Test (University of 
Michigan) 

Ian McTaggart Cowan (University of Brit- 
ish Columbia) 

William B. Davis and Walter P. Taylor 
(Texas A&M University) 

E. Raymond Hall (Berkeley and University 
of Kansas) 

John Eric Hill (American Museum of Nat- 
ural History) 

David H. Johnson and Remington Kellogg 
(U.S. National Museum) 

Jean M. Linsdale (Hastings Natural History 
Reservation) 

Robert T. Orr (California Academy of Sci- 
ence) 

Tracy I. Storer (University of California at 
Davis) 


Burt, Davis, Hall, Hooper, Kellogg, Orr, 
Storer, and Taylor each served as President 
of the ASM. Cowan served as Vice Presi- 
dent. Many members of this group estab- 
lished centers of learning of their own, from 
which additional students were trained, but 
others were in positions where having stu- 
dents was not an option. Some of the centers 
of learning and many of Grinnell’s progeny 
are discussed below. 

Berkeley.—Alden Miller was an orni- 
thologist on the staffat Berkeley and became 
director of the Museum following Grinnell’s 
death. He and Seth Benson, another Grin- 
nell student, were much involved in the 
training of students in mammalogy at 
Berkeley. Today the fine tradition of mam- 
malogy at Berkeley is continued by Oliver 
Pearson (a Harvard product), William Z. 
Lidicker, Jr. (a Grinnell “‘grandson’’), and 
James L. Patton (the incumbent ASM pres- 


PROPINQUITY 131 


ident). Patton studied under W. B. Heed, a 
geneticist, at the University of Arizona. 

University of Michigan. —Four of Grin- 
nell’s students, William H. Burt, Lee R. Dice, 
Emmet T. Hooper, and Fred H. Test, joined 
the staff at the University of Michigan, thus 
creating a major center for mammalogical 
training there. Burt sponsored a number of 
students, including A. W. Frank Banfield, 
Fred S. Barkalow, Lowell L. Getz, Timothy 
E. Lawlor, Richard H. Manville, and Illar 
Muhl. Students of Lee R. Dice included W. 
Frank Blair, Wallace Dawson, Don W. 
Hayne, B. Elizabeth Horner, and John A. 
King. Students of Emmet Hooper included 
James H. Brown, Michael D. Carleton, 
Theodore H. Fleming, Charles O. Handley, 
Jr., David Klingener, Guy G. Musser, and 
Albert Schwartz. Robert K. Enders deserves 
special note as he obtained his degree at 
Michigan, and then taught at Swarthmore 
where he was one of the great inspirational 
teachers. From Swarthmore he inspired Da- 
vid E. Davis, Philip Myers, and Oliver Pear- 
son to enter the field. 

University of British Columbia. —Ian 
McTaggart Cowan, born in Scotland, estab- 
lished his career at the University of British 
Columbia. Dennis Chitty, a student of 
Charles Elton (Oxford), and Cowan trained 
Charles Krebs, formerly of Indiana Uni- 
versity and now also of UBC. Krebs stu- 
dents include Michael Gaines, Barry Keller, 
and Robert Tamarin. J. Mary Taylor was 
also at UBC for many years. 

Texas A&M University.— At Texas A&M, 
a program developed under the leadership 
of William B. Davis and Walter P. Taylor, 
both Grinnell students. Some of Davis’s 
most notable students were Dilford Carter, 
Bryan P. Glass (Oklahoma State Universi- 
ty), and Randolph Peterson (Royal Ontario 
Museum at Toronto). Peterson’s students 
included C. G. Van Zyll de Jong, Judith 
Eger, and Brock Fenton. Fenton has estab- 
lished an excellent program in chiropteran 
biology at York University, York, Ontario. 
Dilford Carter returned to curate the mam- 
mal collection at Texas A&M, then moved 


to Texas Tech University. David Schmidly, 
a student of Donald F. Hoffmeister at IIli- 
nois, now serves as Curator of Mammals at 
Texas A&M. 

University of Kansas.—An outstanding 
program arising from the Grinnell dynasty 
was begun by E. Raymond Hall at the Uni- 
versity of Kansas. The Grinnell contingent 
of mammalogists would not be nearly as 
spectacular if it were not for Hall; thus it 
appears best to title this the Grinnell/Hall 
dynasty rather than simply the Grinnell dy- 
nasty. Hall earlier spent 15 years at Berke- 
ley, where he advised some students of 
Grinnell after Grinnell’s death. Hall’s first 
Ph.D. students were trained at Berkeley as 
well. Hall produced a large number of stu- 
dents, many of whom started programs at 
other institutions. To date, five of Hall’s 
academic “‘sons”’ (Anderson, Durrant, Fin- 
dley, Hoffmeister, and Jones) and six of his 
“grandsons” (Birney, Brown, Genoways, 
Lidicker, Van Gelder, and Wilson) have 
served as President of the ASM. Most of 
Hall’s students are indicated in Table 1, but 
those who established major Ph.D. pro- 
grams in their own right are: 


Rollin H. Baker, first at Kansas and later at 
Michigan State 

E. Lendell Cockrum at Arizona 

Stephen D. Durrant at Utah 

James S. Findley at New Mexico 

Donald F. Hoffmeister at Illinois 

J. Knox Jones, Jr., first at Kansas then at 
Texas Tech 

George H. Lowery at Louisiana State 

Terry A. Vaughan at Northern Arizona 


Rollin Baker, a student of Hall’s, and John 
King, a student of Dice’s, thus both “‘grand- 
sons” of Grinnell, trained a large number 
of students at Michigan State, including 
Donald P. Christian, Gary A. Heidt, and 
Gordon L. Kirkland, Jr. Mammalogy con- 
tinues at Michigan State today under the 
leadership of Donald O. Straney and Rich- 
ard W. Hill. 

From Cockrum’s program at Arizona 
came Robert J. Baker, who has established 


132 WHITAKER 


a major research program at Texas Tech, 
where he has trained a number of students, 
including John W. Bickham, Ira F. Green- 
baum, Rodney L. Honeycutt, and Terry L. 
Yates. 

At Utah, Stephen Durrant sponsored 
Richard M. Hansen, Keith R. Kelson, and 
M. Raymond Lee. Lee in turn sponsored 
Earl G. Zimmerman at the University of 
Illinois. An interesting sidelight related by 
Kelson is that Durrant, although a senior 
professor, had not yet finished his work on 
a doctorate at Kansas under Raymond Hall 
when he presided at Kelson’s Ph.D. final. A 
year later, Durrant came to Kansas for his 
final oral defense of the Ph.D. thesis and 
was examined by Kelson. 

Another major program arose under the 
tutelage of James S. Findley at the Univer- 
sity of New Mexico. Some of Findley’s out- 
standing students are Michael A. Bogan, 
William Caire, Eugene D. Fleharty, Patricia 
(Trish) Freeman, Arthur H. Harris, Clyde 
Jones, Daniel F. Williams, and Don E. Wil- 
son. Findley was subsequently joined at New 
Mexico by J. Scott Altenbach, Terry L. 
Yates, and James H. Brown, all Grinnell 
descendants. 

At least four faculty members associated 
directly or indirectly with Grinnell pro- 
duced outstanding students at the Univer- 
sity of Illinois. Faculty members were Don- 
ald H. Hoffmeister, M. Raymond Lee, 
George O. Batzli, and Lowell L. Getz along 
with ecologist S. Charles Kendeigh, a stu- 
dent of Victor Shelford. Some of the stu- 
dents of Hoffmeister are Wayne H. Davis 
(University of Kentucky), John S. Hall (Al- 
bright College), William Z. Lidicker, Jr. 
(Berkeley), David J. Schmidly (Texas A&M), 
H. Duane Smith (Brigham Young), and R. 
G. Van Gelder (American Museum). Mark 
L. McKnight (U.C. Davis) and Earl G. Zim- 
mermann (North Texas State University) 
were students of Lee. Richard Lindroth 
(University of Wisconsin) was a student of 
Batzli, Joyce Hoffman (Illinois Natural His- 
tory Survey) was a student of Getz, and Dana 


Snyder (University of Massachusetts) and 
Ralph Wetzel (University of Connecticut) 
were students of Kendeigh. 

One of Hall’s most productive students, 
J. Knox Jones, Jr., trained many fine stu- 
dents, first at Kansas, then at Texas Tech 
University, where he became Dean of the 
Graduate School and Vice President for Re- 
search. A team of six mammalogists on the 
faculty was assembled at Texas Tech, each 
with a Ph.D. from a different university — 
Arizona (Robert J. Baker), Texas A&M 
(Dilford C. Carter), Kansas (J. Knox Jones, 
Jr.), New Mexico (Clyde Jones), Oklahoma 
(first Ronald K. Chesser and currently Rob- 
ert D. Owen), and Pittsburgh (Michael R. 
Willig). All are academic descendants of Jo- 
seph Grinnell. 

Some of Jones’ most accomplished stu- 
dents are David M. Armstrong (University 
of Colorado), Elmer C. Birney (University 
of Minnesota), Jerry R. Choate (Fort Hays 
State University, Hays, Kansas), Hugh H. 
Genoways (Carnegie Museum and Univer- 
sity of Nebraska), Thomas H. Kunz (Boston 
University), Carleton J. Phillips (Hofstra 
University and Illinois State University), 
and James D. Smith (Fullerton State Uni- 
versity, California). Some of Jones’ notable 
academic grandsons are Joseph F. Merritt 
whose mentor was Armstrong (officially Ol- 
wen Williams), Robert M. Timm with Bir- 
ney (officially Roger Price, an entomolo- 
gist), and Edyth Anthony and Allen Kurta 
with Kunz. At Kansas, Hall was replaced 
by Robert S. Hoffmann, and subsequently 
Jones and Hoffmann were followed by Rob- 
ert M. Timm and Norman R. Slade. Ken- 
neth B. Armitage and Michael H. Gaines 
also have advised many students at Kansas 
as that center continues to train mammal- 
ogists. 

The major centers of mammalogical in- 
struction established by the first two gen- 
erations of Grinnell students are indicated 
in Fig. 1. Four major centers of learning 
were established by Grinnell’s first genera- 
tion students at British Columbia, Kansas, 


PROPINQUITY 199 


TORONTO N. ARIZONA LOUSIANA ARIZONA UTAH 
& 
Peterson Vaughan Lowery Cockrum Durrant Jones 


TEXAS A&M UNIV. OF MICHIGAN 
BRITISH 
Davis & COLUMBIA Burt, Dice 
Taylor Cowan & Hooper 
BERKELEY 
Grinnel 
& Miller 


KANSAS 
TX TECH 


Hoffmeister 


N. MEXICO MICHIGAN 
STATE 
Rollin 
Findley Baker 


BERKELEY 
& KANSAS 


Hall 


Fic. 1.—Outline of the main branches of the Grinnell Academic tree through the second generation 


students. 


Michigan, and Texas A&M, whereas nine 
were established by the second generation, 
most through E. Raymond Hall at Kansas. 

Other particularly successful students of 
Hall were Sydney Anderson at the Ameri- 
can Museum of Natural History, R. M. No- 
wak with the U.S. Fish and Wildlife Service, 
Henry W. Setzer who retired from the 
Smithsonian, and Terry A. Vaughan who 
for many years was at the University of 
Northern Arizona. Ticul Alvarez and Ber- 
nardo Villa-R. obtained the Masters degree 
with Hall, but have provided the backbone 
of mammalogy in Mexico. Villa-R. even- 
tually obtained the Ph.D. at the University 
of Mexico. 

There are many other academic relatives 
whose Grinnellian attachments are not as 
obvious, but are nonetheless very real. For 
example, Robert S. Hoffmann, now at the 
Smithsonian Institution, is a “great-grand- 
son.” His major professor at Berkeley was 
A. S. Leopold, who started working with 
Grinnell but finished with Alden H. Miller 
after Grinnell’s death. However, Miller’s 


advisor was Grinnell! An academic pro- 
gram has developed at Oklahoma with Gary 
D. Schnell and has produced Ronald K. 
Chesser, Troy L. Best, Janet K. Braun, Mi- 
chael L. Kennedy, and Robert D. Owen. 
Schnell’s Ph.D. is from Kansas, with Rich- 
ard F. Johnston, an ornithologist, serving 
as mentor. However, Johnston’s Ph.D. is 
from Berkeley and his major professor was 
Grinnell’s “‘son’”’ Miller. Michael A. Mares 
at Oklahoma studied under W. F. Blair at 
Texas, whose doctorate was completed un- 
der L. R. Dice at Michigan. Grinnell was 
Dice’s mentor, although not his major ad- 
visor, at Berkeley. 

Several Grinnellites are currently at the 
U.S. National Museum of Natural History 
or with the Fish and Wildlife Service in 
Washington, D.C. They include Michael D. 
Carleton, Alfred L. Gardner, Charles O. 
Handley, Jr., Robert S. Hoffmann, Ronald 
M. Nowak, and Don E. Wilson. 

In Mexico, the principals in the growth 
of systematic mammalogy were Ticul Al- 
varez and Bernardo Villa-R. Both earned 


134 WHITAKER 


their masters degrees at Kansas while study- 
ing with Hall. In Canada, Ian McTaggart 
Cowan and Donald L. Pattie in the west 
and, in the east, A. W. Frank Banfield, Ran- 
dolph L. Peterson, M. Brock Fenton, and 
Robert E. Wrigley are all Grinnell descen- 
dants. 

The Grinnell group has had tremendous 
impact on the ASM. Grinnell himself served 
as president in 1937-1938. Since 1940, when 
Walter P. Taylor was elected the 12th pres- 
ident, only three of the presidents in the 
succeeding 52 years—E. A. Goldman, W. J. 
Hamilton, Jr., and Hamilton’s academic 
“son” James N. Layne—are academically 
unrelated to Joseph Grinnell. 

Every recording secretary since 1938 has 
been a Grinnellite, as have all but three ed- 
itors of the Journal of Mammalogy since 
1941, including one unbroken string for the 
past 27 years. 


IV. The William J. Hamilton, Jr., 
Group (Cornell) 


The other large and important North 
American dynasty in mammalogy is that of 
William J. Hamilton, Jr., at Cornell Uni- 
versity (Table 1, Section IV). While the 
Grinnellian dynasty centered around sys- 
tematic mammalogy, the Hamiltonian dy- 
nasty centered around mammalian ecology 
and natural history. 

Hamilton received his B.S., M.S., and 
Ph.D. degrees in vertebrate zoology from 
Cornell University under A. H. Wright, ap- 
parently with much “unofficial’’ guidance 
from Francis Harper. Francis Harper was a 
teacher in a Long Island school class when 
Hamilton was reportedly “‘cutting up.”’ 
Harper asked Hamilton what bird he was 
holding and Hamilton correctly identified 
it as an immature female rose-breasted gros- 
beak. That brought Hamilton and Harper 
into lifelong friendship. Hamilton’s inter- 
ests were in life history and ecology of ver- 
tebrates, with specialties in food habits, re- 
production, and to some degree, parasites. 


He believed in obtaining as much infor- 
mation as possible from all animals sacri- 
ficed, and in working with the common- 
place rather than always with the exotics. 
In that way one could better obtain ade- 
quate data to make generalizations. He 
passed these interests and philosophies on 
to his students. 

James N. Layne, who taught at the Uni- 
versity of Florida, and at Cornell, and is 
now at Archbold Biological Station, Lake 
Placid, Florida, was an academic “‘son”’ of 
Hamilton’s. He has done much work on 
reproduction and development of mam- 
mals. This tradition has also been carried 
on by Harrison Ambrose and Andrew A. 
Arata, both academic “sons” of Layne. Also, 
Layne has had a longtime interest in para- 
sites, especially fleas. Some other students 
of Layne who worked with ecology and be- 
havior of mammals are James V. Griffo, 
Elizabeth Wing, Llewellyn Ehrhart, Dale 
Birkenholz, John McManus, and James 
Wolfe. Wolfe is now Dean of Graduate 
Studies, Emporia State University in Kan- 
sas, after several years as Executive Director 
of the Archbold Biological Station. Wolfe 
has produced “offspring” of his own, in- 
cluding Robert J. Esher, currently at Mis- 
sissippi State University. James V. Griffo is 
at Fairleigh Dickinson University, and Bir- 
kenholz is at Illinois State University. Wing 
is presently Curator of Zooarcheology, Flor- 
ida Museum of Natural History. William 
Platt started a Ph.D. with Layne at Cornell, 
but finished with Harrison Ambrose when 
Layne moved from Cornell to the Archbold 
Biological Station. Dan W. Walton, a stu- 
dent of Andrew Arata, is presently with the 
Australian Biological Resources Study, and 
is an editor of, and contributor to, the re- 
cently published mammal tome of the Fau- 
na of Australia series. John McManus died 
a few years after he received his Ph.D., but 
was extremely productive while at Fairleigh 
Dickinson University. 

Everett W. (Bill) Jameson, Jr., has carried 
on the tradition of parasite work far beyond 
his graduate student days with Hamilton 


PROPINQUITY 135 


where this interest began. Jameson is well 
known among parasitologists for his work 
on fleas and mites. Two of his ‘“‘sons” are 
John Phillips, a Research Biologist at the 
San Diego Zoological Society; and Duncan 
Cameron, at York University near Toronto. 
Allen H. Benton, now retired from the New 
York State University at Fredonia, is an- 
other of the Hamilton students who became 
interested in parasites, greatly furthering our 
knowledge of fleas. 

Roger Barbour carried on the tradition of 
studies in vertebrate natural history. For 
many years, Barbour was at the University 
of Kentucky, where he and Wayne Davis 
teamed up to teach, train students, and do 
research. Davis is a student of Donald Hoff- 
meister and therefore also a descendant of 
the Grinnellian dynasty. Michael J. Harvey, 
an academic “grandson” of Hamilton, is 
presently department head at Tennessee 
Tech University. Another is Marion D. 
Hassell, who taught at Murray State Uni- 
versity until his recent death. Harrison Am- 
brose and Jim Griffo were undergraduate 
students inspired by Roger Barbour. 

John O. Whitaker, Jr., was Hamilton’s 
last student in mammalogy. He took a po- 
sition at Indiana State University, which 
became a satellite for continuing studies of 
food habits of vertebrates and ectoparasites 
of mammals in the Hamiltonian tradition. 
He teamed up with a Grinnellian student 
trained by Burt, Russell E. Mumford, for 
long-term studies on the mammals of In- 
diana. Some of his students, the academic 
grandchildren of Hamilton, are now making 
their mark. Gwilym S. Jones (who took his 
master’s degree with Mumford) has estab- 
lished a center for vertebrate studies at 
Northeastern University in Boston. Tho- 
mas W. French is Assistant Director of the 
Massachusetts Department of Fish and 
Game. David Pistole is on the staff at In- 
diana University, Indiana, Pennsylvania. 

Robert W. Eadie, long associated with 
Hamilton at Cornell, had several students, 
including Kyle Barbehenn (EPA, Washing- 
ton), Richard W. Dapson (now in private 


industry), and Harold Klein (Plattsburg, 
NY). 

For many years, Jack W. Gottschang, a 
Ph.D. under Hamilton, has been at the Uni- 
versity of Cincinnati, where he chaired the 
Department of Biology and taught many 
students in the Hamiltonian tradition. 

And last, but not least, there is W. J. 
Hamilton III. “Young Bill’? took his Ph.D. 
at Berkeley with Grinnell’s ‘“‘son’’ Alden 
Miller, and is now at the University of Cal- 
ifornia, Davis. He has worked with behav- 
ior of primates, birds, and insects, and on 
growth and development of the red tree 
mouse. Early in his career, he worked on 
bird migration with Franz Sauer. 

Hamilton did not restrict his work to 
mammals, and likewise, many of his aca- 
demic descendants do not. Several have 
worked with parasites, notably Benton, 
Jameson, Layne, and Whitaker. Whitaker 
has also worked with herptiles and fish, and 
Layne with birds and herptiles. Ralph Yer- 
ger (Florida State University) and Margaret 
Stewart (State University of New York at 
Albany) are two of Hamilton’s students who 
work primarily with fish and herps, respec- 
tively. 

There are of course crossings of lines, and 
much inspiration at the undergraduate lev- 
el. Recording this type of contribution would 
be endless, but a few notable examples fol- 
low. Bill Jameson was a student of Hall’s at 
Kansas before going to Cornell. George Bar- 
tholomew got his M.A. with Alden Miller 
at Berkeley before going to Harvard. Robert 
K. Enders inspired David E. Davis, Oliver 
Pearson, and Philip Myers to pursue further 
studies. Jerry R. Choate at Fort Hays State 
University has inspired numerous students 
in mammalian systematics. Jerry has pro- 
duced 32 master’s students, at least 24 of 
whom have earned or are candidates for the 
Ph.D. These include Mark D. Engstrom, 
Sarah B. George, Cheri A. Jones, Nancy D. 
Moncrief, Philip D. Sudman, Michael P. 
Moulton, Lynn W. Robbins, Jerry W. Dra- 
goo, and Brett R. Riddle. James B. Cope 
(Earlham College, Richmond, Indiana) is 


136 WHITAKER 


another of the outstanding undergraduate 
teachers. He was originally inspired as an 
undergraduate student by Bill Hamilton and 
went on to teach at Earlham college at Rich- 
mond, Indiana. Earlham has no graduate 
program, but the influence on bat biology 
exerted through Cope and his students is 
considerable. Some of Cope’s undergradu- 
ate students at Earlham were Richard F. 
Myers (who influenced Thomas H. Kunz at 
the undergraduate level), Wilson Baker, 
Nixon Wilson, Anthony F. DeBlase, Steven 
R. Humphrey, Charles Thaeler, and Rich- 
ard Mills. 

Of course there has been continuous ex- 
change between the Hamilton and Grinnell/ 
Hall schools. Some outstanding workers that 
were influenced by Hamilton as undergrad- 
uates at Cornell, then went on to study un- 
der Grinnellian descendants are William Z. 
Lidicker, James H. Brown, Norman O. Ne- 
gus, and Edwin Gould. E. W. Jameson start- 
ed in the Grinnell school and did his Ph.D. 
with Hamilton. Earl G. Zimmerman, an 
eventual Grinnellite, began his productive 
career while working as an undergraduate 
student (and publishing his first paper) with 
Whitaker at Indiana State. 


V. Other Groups 


There are a few other centers of learning 
that have produced students in the field of 
mammalogy. These tend to be smaller, but 
have made many excellent contributions to 
the field. 

Florida. —A group of biologists has come 
together in recent years at the University of 
Florida, and Florida now can be thought of 
as a center for training mammalogists. Stev- 
en Humphrey (a student of Bryan Glass at 
Oklahoma State University), John H. Kauf- 
mann (Grinnellite via A. S. Leopold), and 
John F. Eisenberg (student of behaviorist 
Peter Marler) are there. James N. Layne 
(Archbold Biological Station) has been in- 


fluential in the development of this group. 
This group is supported by paleontologist 
S. David Webb, and ornithologists J. C. 
Dickinson and Franz Sauer. Jackie Belwood 
(student of Stephen Humphrey), Cheri Jones 
(student of John Eisenberg), Paul Pearson 
(student of Archie Carr), and Michael H. 
Smith obtained their training there. Mike 
Smith has headed the Savannah River Ecol- 
ogy Laboratory at Aiken, South Carolina, 
for many years. 

Purdue. —Purdue University has had its 
influence on mammalogy, earlier under 
Durwood L. Allen and Charles M. Kirk- 
patrick, both essentially conservationists, 
and later under two of Kirkpatrick’s stu- 
dents, Russell E. Mumford and Harmon P. 
Weeks. Some of the more notable students 
from this group are L. David Mech and Rolf 
O. Peterson, two wolf biologists, and Virgil 
Brack, Jr., a bat biologist. 

Tulane.—Norman C. Negus and James 
S. Findley grew up together in suburban 
Cleveland, Ohio. Their ‘Bible’? was Ham- 
ilton’s Mammals of the Eastern United 
States (1943). Negus studied under Eugene 
Dustman, an ornithologist, at Ohio State. 
Findley ended up heading the Kansas sub- 
group at New Mexico, and Negus then es- 
tablished a mammal center at Tulane, with 
Jack A. Cranford, Edwin Gould, John F. 
Pagels (co-advised with Clyde Jones), Aelita 
J. Pinter, and Thomas E. Tomasi among his 
students. Negus now heads a research group 
at the University of Utah. 

Wisconsin. —The University of Wiscon- 
sin has also served as a center, although 
neither of the two principals, John T. Emlen 
and Roland K. Meyer, is a mainstream 
mammalogist. Meyer is an endocrinologist 
and Emlen is a preeminent ecologist. Phillip 
L. Wright emerged from this program and 
established a program in mammalogy at the 
University of Montana. He was joined there 
for a time by Robert S. Hoffmann, who also 
had students at the University of Kansas 
and is now at the Smithsonian Institution. 
Garrett C. Clough and William A. Fuller 


PROPINQUITY 13y/ 


were students of Emlen, and John E. War- 
nock, Rodney A. Mead, and Tim W. Clark 
are notable mammalogists from the Wis- 
consin and Montana programs. 

Other Sources. — Many “‘mammalogists”’ 
have entered the field from other fields but 
are now working primarily with mammals. 

Several physicians have made names for 
themselves in mammalogy, one being C. 
Hart Merriam. Others include Marcus Ward 
Lyon, Jr., who wrote Mammals of Indiana 
in 1936 and who is also a past president of 
the ASM; Murray L. Johnson, who received 
his M.D. from Oregon Medical School; and 
George Wislocki, an anatomist at Harvard 
Medical School. Denny G. Constantine, who 
has made many valuable contributions con- 
cerning bat rabies, is a veterinarian. 

A number of individuals have received 
degrees in ecology, then have done concen- 
trated work in mammalogy. For example, 
Dennis Chitty and Francis C. Evans worked 
with Charles Elton at Oxford, Michael Ro- 
senzweig with Robert MacArthur at Penn- 
sylvania, E. V. Komarek with W. C. Allee 
at the University of Chicago, and Wilson 
Baker and Gary Barrett with Eugene Odum 
at the University of Georgia. Charles Krebs, 
in turn, worked with Dennis Chitty. 

Several individuals are associated with 
mammalogy from a wildlife biology back- 
ground, including both the principles of the 
Purdue group, Durwood L. Allen and 
Charles M. Kirkpatrick, and also Willard 
D. Klimstra, B. J. Verts, Joseph A. Chap- 
man, George A. Feldhamer, and Glen C. 
Sanderson. 

Several have entered mammalogy from a 
genetics background, such as Gary F. 
McCracken, who worked with Peter Brus- 
sard; and James L. Patton, who worked with 
W. B. Heed at the University of Arizona, 
where he also was closely associated with 
E. Lendell Cockrum. Karl F. Koopman took 
his Ph.D. with T. H. Dobzhansky at Co- 
lumbia. His doctoral dissertation, on nat- 
ural selection and reproductive isolation be- 
tween two closely related populations of 


Drosophila, was a classic of its day and fre- 
quently is cited in courses in evolution and 
genetics. Koopman has made numerous 
contributions on bats and is now an Hon- 
orary Member of ASM. 

Other examples given in Table I include 
William A. Wimsatt and Roy Horst, who 
worked with a morphologist; Duane A. 
Schlitter, Paul K. Anderson, Paul Pearson, 
and Kenneth Wilkins, who worked with 
herpetologists; and Albert E. Wood, who 
worked in palaeontology. 

There is another group of mammalogists 
who, similar to the Merriam group, did not 
have Ph.D.’s and thus did not have stu- 
dents, yet they have made major impacts 
on the field. These include individuals such 
as Rudolph M. Anderson, a long-time 
worker in Canada; G. H. H. Tate, who 
worked with mammals of eastern Asia and 
South America; Harold E. Anthony (mam- 
mals of North America); Hobart M. Van 
Deusen of the American Museum (mam- 
mals of New Guinea); and Phillip Hersh- 
kovitz of the Field Museum (South Amer- 
ican mammals). Karl Kenyon (marine 
mammals) and Olaus Murie (large carni- 
vores) are also high profile examples of this 
group. 

Present day mammalogists of North 
America come from a few major lineages 
and several other sources and backgrounds. 
The few earlier stems stimulated the field 
but the great diversity present today allows 
for diverse methods and ideas to be applied 
to problems in mammalogy and should help 
us to continue to make major intellectual 
advances. Systematics and life history 
studies led the way and are still exceedingly 
important, but today many other areas, no- 
tably genetics, behavior, ecology, physiol- 
ogy, conservation biology, and many other 
fields make their contributions. Although 
our roots to this point are relatively few, 
diversity continues to increase as specialists 
continue to add to the field of mammalogy, 
and the genealogy of mammalogists be- 
comes ever more complicated. 


138 WHITAKER 


Acknowledgments 


This paper would not have been possible with- 
out the cooperation of many individuals too nu- 
merous to mention. However, special thanks are 
due to Elmer C. Birney, James H. Brown, Donald 
F. Hoffmeister, J. Knox Jones, Jr., William Z. 
Lidicker, Jr., Oliver P. Pearson, and Don E. Wil- 
son, all of whom have read and greatly improved 
the manuscript. 


Literature Cited 


GRINNELL, H. W. 1958. Annie Montague Alexander. 
Grinnell Naturalists Society. Museum of Vertebrate 
Zoology, University of California, Berkeley, 27 pp. 

Hatt, E. R. 1939. Joseph Grinnell (1877 to 1939). 
Obituary. Journal of Mammalogy, 20:409-417. 

HamILTon, W. J., JR. 1943. Mammals of eastern 
United States. Comstock Publishing Co., Ithaca, New 
York, 432 pp. 

1955. Mammalogy in North America. Pp. 


661-668, in A century of progress in the natural 
sciences 1853-1953. California Academy of Sci- 
ences, San Francisco, 807 pp. 

HOFFMEISTER, D. F. 1969. The first fifty years of the 
American Society of Mammalogists. Journal of 
Mammalogy, 50:794-802. 

Jones, J. K., Jk. 1991. Genealogy of twentieth-cen- 
tury systematic mammalogists in North America: 
the descendants of Joseph Grinnell. Pp. 48-55, in 
Latin American mammalogy: history, biodiversity, 
and conservation (M. A. Mares and D. J. Schmidly, 
eds). University of Oklahoma Press, Norman, 468 
pp. 

LAYNE, J. N., AND R.S. HOFFMANN. 1994. Presidents. 
Pp. 22-70, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1-433. 

STORER, T. 1969. Mammalogy and the American So- 
ciety of Mammalogists, 1919-1969. Journal of 
Mammalogy 50:785-793. 

MEeErRRIAM, C. H. 1877. A review of the birds of Con- 
necticut, with remarks on their habits. Transactions 
Connecticut Academy of Arts and Sciences, 4:1—150. 

ME_ErRRIAM, C. H. 1884. The vertebrates of the Adi- 
rondack region: the Mammalia. Transactions of the 
Linnean Society of New York, 1:1-124. 


PUBLICATIONS 


B. J. VERTS AND ELMER C. BIRNEY 


Introduction 


Rn to the Bylaws and Rules 
adopted by the American Society of 
Mammalogists on 3 April 1919, “The ob- 
ject of the Society shall be the promotion 
of the interests of mammalogy by holding 
meetings, issuing a serial or other publica- 
tions, aiding research, and engaging in such 
other activities as may be deemed expedi- 
ent” (Article I., Sec. 2.). Of the budget ap- 
proved by the Board of Directors for 1992, 
$122,000 (74.1%) of the total $164,630 was 
allocated for expenses related directly to ed- 
itorial activities of the society. Throughout 
the 75 years of the existence of the society, 
no single activity has been of higher priority, 
received a larger share of the budget, 
or, arguably, had a greater impact on the 
development of the discipline than has pro- 
duction of the society’s publications, es- 
pecially the Journal of Mammalogy. It is 
the purpose of this chapter to provide a brief 
summary of the 75-year history of the pub- 
lications of the ASM, with special emphasis 
on trends observed in the content of the 
Journal of Mammalogy during this period. 


139 


Journal of 
Mammalogy 


American Society of Mammalogists 


The Journal of Mammalogy 


The Journal of Mammalogy has served 
the role of the serial publication authorized 
in the Bylaws and Rules since the ASM was 
founded. It also has functioned as an “‘of- 
ficial” publication of the society in that it 
includes announcements and minutes of 
meetings, lists of officers and committee 
members, and other announcements and 
communications. However, nowhere have 
we found that the Journal of Mammalogy 
ever was designated the official publication 
of the American Society of Mammalogists. 

The Journal of Mammalogy commenced 
publication on 28 November 1919, <8 
months after the society was founded. Vol- 
ume | (259 pages) consisted of five numbers 
(issues); the four published in 1920 almost 
certainly were intended to establish the Feb- 
ruary, May, August, and November pattern 
of publication, but each actually was pub- 
lished in the following month. Authors of 
the articles published in the first volume 
included some of the most renowned and 


140 VERTS AND BIRNEY 


| 72 
VOLUME 


Fic. 1.—Strata-surface graph of the number of 
pages devoted to feature articles, general notes, 
and other components in volumes 1-72 (1919- 
1991) of the Journal of Mammalogy. 


revered names in American mammalogy: 
Glover M. Allen, J. A. Allen, H. E. An- 
thony, Vernon Bailey, Lee R. Dice, James 
W. Gidley, Joseph Grinnell, G. Dallas Han- 
na, Francis Harper, Arthur H. Howell, A. 
Brazier Howell, Hartley H. T. Jackson, 
Stanley G. Jewett, C. Hart Merriam, Gerrit 
S. Miller, Jr., W. D. Matthew, Wilfred H. 
Osgood, John C. Phillips, Ernest Thompson 
Seton, Arthur de Carle Sowerby, H. L. Stod- 
dard, Walter P. Taylor, P. A. Taverner, and 
Edward R. Warren. Interestingly, only a sin- 
gle article was coauthored (by G. S. Miller, 
Jr., and James W. Gidley), only one was by 
a researcher from other than North America 
(by A. de C. Sowerby of England), and only 
eight (10.8%) of the 74 articles published 
were about mammals other than those in 
North America (one each on African car- 
nivores, cats, and monkeys; neotropical bats 
and cats; Asian bears; Japanese bats; and 
Brazilian tapirs). 

The initial issue of the Journal of Mam- 
malogy was a 51-page number consisting of 
7 feature articles (37.3 pages), 4 general notes 
(4.6 pages), 3 reviews and 49 references in 
a recent-literature section, an editorial com- 
ment (1.6 pages), and the Bylaws and Rules 
adopted on 3 April 1919 (2.6 pages) when 
the society was founded. Both feature arti- 
cles and general notes tended to be short; 
the former averaged 4.5 pages, the latter <1 
page. The comments by Editor Ned Hollis- 


ter consisted of a paragraph-long history of 
the organization of the society, a description 
of the scope of the Journal, solicitation of 
manuscripts for the Journal, a plea for 
members to recruit new members, an ac- 
knowledgment of Ernest Thompson Seton’s 
contribution of the sketch of the pronghorn 
for the front of the Journal, a report of the 
election of J. A. Allen as an Honorary Mem- 
ber, and acomment on the paper by C. Hart 
Merriam titled “Criteria for the recognition 
of species and genera.”” Volume 1, number 
4 contained a list of members, some of whom 
were listed subsequently as other than char- 
ter members (Journal of Mammalogy, 3: 
203-218, 1922). 

Although the basic composition of the 
Journal of Mammalogy was established at 
the onset, numerous changes have occurred 
in the proportion devoted to each of the 
sections. For example, during the first 3 de- 
cades of publication, general notes com- 
posed about 20—50 pages, irrespective of the 
total number of pages published in each vol- 
ume (Fig. 1). However, after about 1950, 
more and more space was devoted to gen- 
eral notes; in both 1988 and 1989, >290 
pages of general notes were published (Fig. 
1). Editorial policy was altered in 1990 to 
limit the number of general notes published 
as a means of enticing bibliographic services 
to include references to more of the shorter 
papers published in the Journal. The gen- 
eral-note format was abandoned commenc- 
ing with volume 73 (1992). 

Over the years, some minor evolution has 
occurred in components of the Journal of 
Mammalogy: “Editorial Comment” in vol- 
ume | (1919-1920) became “‘Correspon- 
dence” in volume 2 (1921) and remained 
so until volume 6 (1925) when it became 
‘Comment and News,” which in volume 
35 (1954) became ““Comments and News.” 
The “Recent Literature” section was an in- 
tegral part of the Journal from its inception 
through volume 50 (1969), published as a 
supplement to volumes 51-66 (1970-1985) 
of the Journal, then discontinued com- 
mencing with volume 67 (1986). Member- 


PUBLICATIONS 141 


ship lists were published in volumes 1 
(1919-1920), 3 (1922), 5 (1924), 11 (1930), 
15 (1934), 18 (1937), 21 (1940), 29 (1948), 
31 (1950), 35 (1954), 40 (1959), and 46 
(1965), and as supplements accompanying 
volumes 54 (1973), 59 (1978), 65 (1984), 
and 70 (1989). Other supplements were 
published irregularly and include three edi- 
tions of “Guidelines for manuscripts,” “Roles 
of standing committees,” “Survey of North 
American collections of Recent mammals,” 
and “Acceptable field methods in mammal- 
ogy.” The “Bylaws and Rules,” or, when 
amended, parts thereof, were included in 
several issues. 

Reviews of recent publications were in- 
cluded in the first issue and in most, but not 
all, subsequent issues of the Journal of 
Mammalogy. Until volume 17 (1936), re- 
views were included in the recent-literature 
section, but afterward were afforded a sec- 
tion of their own with the subheading “‘Re- 
views.” Reviews occupied 4-11 (¥ = 4.7) 
pages in volumes | 7-32 (1936-1951), 5-17 
(X = 11.7) pages in volumes 33-50 (1952- 
1969), 20-31 (X¥ = 23.6) pages in volumes 
51-66 (1970-1985), and 12-19 (¥ = 15.7) 
pages in volumes 67-72 (1986-1991). 
Greater emphasis was placed on the pub- 
lication of reviews commencing with vol- 
ume 73 (1992); 26 pages were devoted to 
reviews in that volume. 

An author-subject index is published in 
the last issue of each volume; however, the 
index to volume 52 (1971) was published 
as a supplement to the first number of vol- 
ume 53 (1972). Commencing with that in- 
dex and continuing to present, an alpha- 
betical (by last name of author or editor) 
listing of books reviewed in the volume fol- 
lowed by the page number on which the 
review may be found concludes each index. 
Also, commencing with volume 67 (1986) 
author and subject indices were separated. 

Announcements of the death of members 
of the American Society of Mammalogists 
were included in the Journal of Mammalogy 
for the first time in the fourth number of 
volume | (1919-1920). Like other com- 


ponents of the Journal of Mammalogy, death 
notices underwent considerable evolution. 
At the end of the first membership list, 
names of three deceased numbers were list- 
ed. The general-notes section of the same 
issue included a seven-line obituary for one 
of those listed (Thomas M. Owen) and a 
nearly page-long obituary for a Canadian 
naturalist and agency official (James M. Ma- 
coun) who apparently was never a member 
of the society. The second volume con- 
tained no death notices, but the third vol- 
ume (1922) contained a list of nine deceased 
members, including the three listed in vol- 
ume | (1919-1920); this appeared at the end 
of the new list of members. Seemingly, the 
intent initially was to include a list of all 
deceased members with each membership 
list, but the practice was abandoned after 
publication of the second such list. The first 
extensive obituary was the 7-page “‘appre- 
ciation’”’ for one of the founders of North 
American mammalogy, Joel Asaph Allen, 
published in volume 3 (1922); a second 
obituary for Allen was published in volume 
11 (1930) and, with a photograph, included 
>13 pages. However, no bibliography ac- 
companied the text of either. For about 25 
years, either lists of deceased members (usu- 
ally in bold-face type) published in the com- 
ments and news section or short (from 6- 
10 lines to a page or so) obituaries for de- 
ceased members were common. Sometimes 
a deceased member’s name appeared in one 
of the lists and an obituary for that member 
was published subsequently, but more often 
a deceased member was honored only once. 
Occasionally, obituaries for prominent 
members covered 3-5 pages or more and 
one that included a bibliography and cor- 
respondence (for President Edward A. 
Goldman) required 22 pages [volume 28 
(1947)]. Since about 1950, names of de- 
ceased members were listed in the com- 
ments and news section under the subhead- 
ing ““Deaths Reported.” Names were in 
boldface, but cities and states of residence 
and membership status (honorary, life, or 
emeritis), when included, were set in italic. 


142 VERTS AND BIRNEY 


Also, since about 1950, obituaries have been 
limited to past presidents and prominent 
mammalogists. In a few instances, a death 
notice or obituary has been included in the 
Journal of Mammalogy for a mammalogist 
or naturalist (usually foreign) for which there 
is no published record of their having been 
a member of the society. 

Miscellaneous items published in the 
Journal of Mammalogy from time to time 
include letters to the editor, letters from the 
president, publication policies and sugges- 
tions to authors, personal notices (mostly 
items for sale and items wanted), member- 
ship application forms, advertisements of 
society publications, and paid advertise- 
ments for equipment, supplies, and publi- 
cations of interest to mammalogists. One 
issue, the third of volume 11 (1930), con- 
tained 64 pages of papers resulting from a 
symposium on predatory animal control. 
The fourth issue of each volume commenc- 
es with a series of roman numbered pages 
(usually 8 pages, 2 of which are blank) that 
contain a list of editors, a reprinting of the 
verso of the front cover, and a reprinting of 
the contents of all four issues of the volume. 

The artwork of Seton graced the cover of 
the Journal of Mammalogy for a decade, 
but commencing with the first issue of vol- 
ume 11 (1931), a new cover designed by A. 
Brazier Howell and dominated by the head 
and cape of a pronghorn appeared. Howell’s 
artwork appeared on the cover through vol- 
ume 43 (1962). A new design depicting a 
standing pronghorn appeared on the cover 
of volumes 44-48 (1962-1967) and was fol- 
lowed by another head and cape view of the 
pronghorn in volumes 49-72 (1968-1991). 
Artwork for both cover designs was signed; 
“Hines” signed the former and the cryp- 
tographic signature on the latter is the ini- 
tials of Frances L. Jacques. No “‘Hines’”’ was 
listed as a member of the American Society 
of Mammalogists in membership lists pub- 
lished in 1959 or 1965, so likely the cover 
design used for volumes 44—48 (1962-1967) 
was drawn by a commercial artist. Jacques 
was an artist at the American Museum of 


Natural History and the James Ford Bell 
Museum of Natural History. A radical de- 
parture from the traditional green and black 
cover dominated by a pronghorn com- 
menced with volume 73 (1992). The prong- 
horn, although still present and still the art- 
work used in volumes 49-72 (1969-1991), 
no longer dominates the cover, but is rele- 
gated to a small circle. The central figure, 
consisting to date of artwork depicting some 
mammal, is unique to each issue. Green, 
although a different shade, remains featured 
on the somewhat thicker and smoother cov- 
er, but on the front, the lettering, a square 
enclosing the central figure, and the small 
circle enclosing the drawing of the prong- 
horn are white. On the back cover, large 
lettering and a rectangle containing a list of 
officers and directors also are white. Also, 
the first color plate for a research article was 
published in volume 73 (1992); however, 
the first and only other color plate published 
in the Journal was that of Rupicapra rupi- 
capra by F. Murr from Erna Mohr’s Sdau- 
getiere included in the review by R. H. Man- 
ville of that book published in volume 40 
(1959). 

Through volume 57 (1976), the entire 
Journal of Mammalogy was printed 1n sin- 
gle-column format. Commencing with vol- 
ume 58 (1977), literature-cited sections were 
printed in double-column format, but the 
text remained single column until volume 
73 (1992) when the space-saving and the 
easier-to-read double-column format was 
adopted. The Williams and Wilkins Com- 
pany, Baltimore, Maryland, printed the first 
37 volumes of the Journal of Mammalogy, 
but commencing with volume 38 (1957) of 
the Journal, Allen Press, Lawrence, Kansas, 
has served as the printer for all publications 
of the American Society of Mammalogists. 

Throughout the history of the Journal of 
Mammalogy, all editorial services have been 
provided by members who volunteered; for 
the first 37 volumes (1919-1956) all edi- 
torial services were provided by one person, 
designated the “editor.” Subsequently, sev- 
eral systems of dividing the ever-growing 


PUBLICATIONS 143 


editorial responsibilities were employed 
(Table 1). The present system, stable for the 
last 14 volumes (60-73) consists of a man- 
aging editor (the editor of record) respon- 
sible for production of the Journal, a journal 
editor responsible for matters of style and 
presentation, several associate editors re- 
sponsible for conducting the review process 
and judging the scientific merit of manu- 
scripts, an editor for reviews responsible for 
soliciting and editing reviews of books and 
assembling and publishing a “books re- 
ceived” list of books submitted but not re- 
viewed, and an editor for advertising re- 
sponsible for personal notices and 
commercial advertising. Since its inception, 
only 62 mammalogists have served the 
Journal of Mammalogy in one or more ed- 
itorial capacities (Table 1); the length of ser- 
vice ranged from | to 16 years and averaged 
4.6 years. The Journal has had only 17 ed- 
itors of record; length of service averaged 
4.5 years (range, 1-7 years). 

From the onset, the Journal of Mam- 
malogy was provided free to all members 
and was available to institutions by sub- 
scription. Just as there has been an increase 
in number of pages published (Fig. 1), there 
has been an increase in both membership 
dues and subscription rate (Fig. 2). Since 
1953 (when publication of a summary of 
the annual budget in the Journal com- 
menced), neither subscription rate nor 
membership dues has kept pace with funds 
budgeted for publication of the Journal of 
Mammalogy (Fig. 2). Income from the J. 
A. Allen Memorial Fund and other invest- 
ments managed by the society’s trustees 
(Kirkland and Smith, 1994) make it pos- 
sible to continue to provide members and 
subscribers with a quality publication at a 
modest cost. 

In his initial solicitation of papers, the 
first editor, Ned Hollister, emphasized the 
need to make the Journal of Mammalogy 
an essential tool for workers in all phases 
of mammalogy. To ascertain the effective- 
ness of this and similar pleas by subsequent 
editors, we analyzed trends related to length, 


120 
100 
80 
35 
25 x 1,000 
$ 40 
15 
20 
(Se pee emcee 
aes 
{e) 
| 72 


VOLUME 


Fic. 2.—Line graphs of membership dues 
(heavy line) and subscription rates (light line) for 
volumes 1-72 (1919-1991) on left ordinate and 
histogram of funds budgeted by the Board of 
Directors (as published in the Journal of Mam- 
malogy) for production and distribution of vol- 
umes 34-72 (1953-1991) of the Journal of Mam- 
malogy on right ordinate. 


subject matter, and authorship of papers 
published in the Journal of Mammalogy by 
sampling alternate volumes from volume 1 
(1919-1920) to volume 71 (1990). Length 
measured to the nearest 0.1 page, the con- 
tinent of origin for species reported on, 
number and residence of authors, number 
of references cited, and major topic covered 
were recorded for each article published in 
volumes sampled. 

Editors and authors have maintained a 
diversity of topics among articles published 
in the Journal of Mammalogy; after an ini- 
tial paucity of papers on morphology, re- 
production, and behavior they have main- 
tained a more even balance among topics. 
Articles published as feature articles (Fig. 
3a) cover more diverse topics than general 
notes (Fig. 3b) as > 20% were on topics other 
than the six classifications that we used to 
present results of our analysis. Overall, ap- 
proximately one-fourth of all articles pub- 
lished as feature articles were devoted to 
ecology and life history, and, among general 
notes, the same proportion was devoted to 
articles describing distributions and new lo- 
cality records (Fig. 3). Since about 1964, the 
number of general notes devoted to distri- 


VERTIS AND BIRNEY 


144 


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145 


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VERTS AND BIRNEY 


146 


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TABLE |.— Continued. 


Publications Journal Associate Editor for Advertising 


Managing 


editor 


R. K. Rose 


editors reviews 


editor 
T. L. Best 


editor 


R. M. Timm 


editor 
R. J. Baker 


Volume 


Year 


B. D. Patterson 


. W. Freeman 
McBee 


74 


1993 


PUBLICATIONS 147 


butions and new locality records has de- 
clined steadily (Fig. 3b); manuscripts com- 
posed largely of descriptions of extensions 
of geographic ranges of taxa based on single 
locality records were specifically excluded 
from the Journal by publication policy com- 
mencing in 1988. No other major trends in 
the diversity of topics of articles published 
in the Journal are discernable. 

Authorship has remained largely North 
American; of 1,691 feature articles and 2,613 
general notes published in alternate vol- 
umes from volume | (1919-1920) to vol- 
ume 71 (1990), 1,555 (91.9%) and 2,456 
(94.0%), respectively, were written exclu- 
sively by North American authors. Never- 
theless, a trend toward more articles au- 
thored by researchers outside of North 
America seems to be becoming established. 
In volume 71 (1990), 23.8% of the 63 fea- 
ture articles and 30.8% of the 39 general 
notes were authored by one or more re- 
searchers from other continents. Although 
84.8% of the 2,613 general notes and 81.8% 
of the 1,691 feature articles were about 
North American taxa, a trend established 
after World War II toward publication of 
more articles on mammals from other con- 
tinents continues. In volume 71 (1990), 
34.9% of 63 feature articles and 38.5% of 
39 general notes were about mammals from 
other than North America. Coauthorship 
became an increasing trend for feature ar- 
ticles and general notes; however, three or 
more authors were rare before volume 27 
(1946) for feature articles, and volume 39 
(1958) for general notes (Fig. 4). In volume 
71 (1990), the last for which we separated 
papers by type, 84.6% of general notes and 
55.6% of feature articles were written by 
more than one author (Fig. 4). Thus, not 
only is the number of authors increasing, 
but both the scope and the clientele of the 
Journal of Mammalogy are becoming more 
international. 

After the initial volume, average length 
of feature articles (Fig. Sa) was 6-9 pages 
during most years until volume 45 (1964) 
when the average length began a steady climb 


148 VERTS AND BIRNEY 


All Other Topics 


Morphology 


Ecology and Natural History 


VOLUME 


Ecology and Natural History 


Distribution and Locality Records 


| 7I 
VOLUME 
Fic. 3.—Surface graph of proportions of the 
total number of articles on each of several topics 
in alternate volumes for volumes 1-71 (1919- 
1990) of the Journal of Mammalogy: a, feature 
articles; b, general notes. 


that reached a peak of > 14 pages in volume 
56 (1975). The peak was followed by a 
somewhat precipitous decline to a plateau 
of <10 pages. During only | year before 
volume 45 (1964) did the average length of 
general notes exceed | page (Fig. 5b), but 
after volume 45 (1964), average page length 
increased gradually to >4 pages in volume 
69 (1988), but declined to 3.0 in volume 71 
(1990) in response to efforts by editors to 
emphasize feature articles. 

The number of references cited per paper 
averaged <10 for feature articles and less 
than two for general notes in most volumes 
before volume 45 (1964; Fig. 6). However, 
commencing about 1945, the average num- 


q 
100 < 
% 
c; TI 
VOLUME 
100 7 oe 
%e 
er Ta 
VOLUME 


Fic. 4.—Surface graph of proportions of the 
total number of articles authored by one, two, 
and three or more authors in alternate volumes 
for volumes 1-71 (1919-1990) of the Journal of 
Mammalogy: a, feature articles; b, general notes. 


ber of references proliferated greatly, at- 
taining an apex of >35 for feature articles 
and > 15 for general notes published in most 
recent volumes. No doubt, the almost log- 
arithmic increase in number of references 
cited per paper in both feature articles and 
general notes was a response to both the 
greater need to document previous findings 
and the greater availability of information 
on all aspects of mammalogy (Anderson and 
Van Gelder, 1970). 

In the first volume, new taxa were de- 
scribed in 12 (36.4%) and new names were 
applied to named taxa in three (9.1%) of the 
33 feature articles. Describing and naming 
new taxa remained a common topic of ar- 


PUBLICATIONS 149 


| 7 


PAGES 
a 
ion 


| 7 
VOLUME 
Fic. 5.—Bar graphs of the average number of 
pages per article published as: in alternate vol- 
umes for volumes 1-71 (1919-1990) of the Jour- 
nal of Mammalogy: a, feature articles; b, general 
notes. 


ticles published in the Journal of Mam- 
malogy in the first 20 volumes; subsequent- 
ly, alpha taxonomy was the topic of <10% 
of the articles published. Overall, only 3.6% 
of 4,304 articles published in alternate vol- 
umes of the Journal of Mammalogy con- 
tained descriptions of new taxa. 

Rodents, bats, and carnivores, in that or- 
der, were the most popular topics of articles 
published as general notes in the Journal of 
Mammalogy (Fig. 7a). Fewer general notes 
on insectivores or on more than one order 
were published in the last 15 years that pa- 
pers were segregated by type. Among feature 
articles, however, trends toward publication 
of more and more articles on rodents and 
fewer and fewer articles on taxa representing 
more than one order of Mammalia were 
evident almost from the beginning of pub- 
lication of the Journal (Fig. 7b). A similar 
trend was noted in oral presentations at an- 
nual meetings (Gill and Wozencraft, 1994). 
Obviously, manuscripts containing infor- 
mation on more than one order of mam- 
mals were not converted to general notes as 
the proportion of general notes on multior- 
dinal topics also has declined in recently 
published volumes (Fig. 6a). Likely, the or- 


40 


REFERENCES 
De) 
oO 


C TI 
VOLUME 


Fic. 6.—Line graph of the average number of 
references cited per feature article (heavy line) 
and general note (light line) published in alter- 
nate volumes for volumes 1-71 (1919-1990) of 
the Journal of Mammalogy. 


dinal topic chosen reflects the abundance, 
diversity, and ease of catching and handling 
rodents and bats. 

From this brief analysis, we conclude that 
the Journal of Mammalogy has filled and 


All Other Orders 


VOLUME 


Chiroptera 


Rodentia 


| 
VOLUME ia 


Fic. 7.—Surface graph of proportions of the 
total number of articles devoted primarily to each 
of several orders of mammals (and to more than 
one order of mammals) published in alternate 
volumes from volumes I-71 of the Journal of 
Mammalogy: a, feature articles; b, general notes. 


150 VERTS AND BIRNEY 


continues to fill the role and scope that the 
founders of the American Society of Mam- 
malogists envisioned for it. The diversity of 
subjects and orders of mammals treated, and 
the diversity in length and depth of treat- 
ments remains its greatest strength. Likely, 
this strength is one of the major binding 
forces of the American Society of Mam- 
malogists. 


Mammalian Species 


Mammalian Species is the most recently 
established serial publication of the Amer- 
ican Society of Mammalogists. The objec- 
tive of Mammalian Species “is to provide 
a critically compiled, accurate, and concise 
summary of the present state of our biolog- 
ical knowledge (and ignorance) of a species 
of mammal in a standard format...” (In- 
structions for contributors to Mammalian 
Species, 1987). Each account includes a 
complete synonymy and sections in which 
context and content, diagnosis, distribu- 
tion, general characters, fossil record, form 
and function, ontogeny and reproduction, 
ecology, behavior, and genetics are consid- 
ered. A remarks section, commonly con- 
taining an explanation of complex nomen- 
clature, and an extensive literature cited 
section completes each account. One ac- 
count in each genus must contain a generic 
synonymy and context and content sec- 
tions. Most accounts contain a photograph 
or artist’s depiction of a representative of 
the species, photographs or line drawings of 
dorsal, ventral, and lateral views of the skull, 
and a map depicting the geographic distri- 
bution of the species. Some accounts con- 
tain photographs or line drawings of certain 
diagnostic features such as the baculum, 
phallus, specific teeth or parts of toothrows, 
and karyotype. The intention was to limit 
the length of accounts to 8 pages (printed 
double-column), but several accounts, es- 
pecially those on well-researched species, 
exceed that length. 

The concept of Mammalian Species was 
presented to the Board of Directors at the 


1968 meeting (Journal of Mammalogy, 49: 
844, 1968) and was approved by the board 
at the 1969 meeting as a publication “‘to be 
sold by subscription” (Journal of Mam- 
malogy, 50:908, 1969). At the latter meet- 
ing, the board budgeted $2,000 for initial 
publication of the series. An announcement 
in the same issue of the Journal (p. 913) 
indicated that the first account (on Macrotus 
waterhousil) would be mailed to all mem- 
bers with a price list and subscription form. 
The following year an announcement (Jour- 
nal of Mammalogy, 51:842, 1970) indicated 
that the cost of a subscription to Mam- 
malian Species would be $9.60 to members 
and $12.00 to nonmembers; the first fascicle 
of six accounts was published 16 June 1971. 
Although timing of publication and number 
of accounts per fascicle were variable during 
the first 10-12 years, during recent years, 
two fascicles consisting of 8-20 accounts 
each were published annually. The present 
cost of subscriptions for members and non- 
members is $10 per year; individual ac- 
counts may be purchased (accounts in same 
order: $2 each for five or fewer, $1.50 each 
for six—10, and $1 each for = 11) and special 
packages of accounts (grouped by region, 
taxa, or other classification) are available at 
25% discount. 

Initially, authors for Mammalian Species 
accounts were solicited from among those 
especially knowledgeable of a taxon, but, 
more recently, prospective authors have re- 
quested assignment of exclusive privileges 
to produce accounts on specific species. 
Currently, assignments are made by the 
managing editor for a period of 3 years with 
authors retaining the option of requesting 
an extension of | year to complete accounts 
in progress. On the matter of timely com- 
pletion of assignments, editors have been 
flexible, to a point. 

As of 23 April 1993, 443 accounts in- 
cluding 452 species had been published (nine 
accounts each covered two closely related 
species). Through the first 443 accounts, 
numbers of accounts by order of mammal 
was strongly correlated (7? = 95.04, n = 20) 


PUBLICATIONS ies 


with numbers of species classified by order 
(Anderson and Jones, 1984:5-8). Orders that 
deviate most within this relationship are 
Primates with accounts published for only 
3 (1.7%) of 180 species and Carnivora and 
Artiodactyla for which accounts have been 
published for 54 (20.1%) of 269 species and 
27 (14.6%) of 185 species, respectively. As 
215 (48.5%) of the 443 published accounts 
are on North American mammals north of 
Mexico (comprising 50.6% of the species 
native to the region—Jones et al., 1992), the 
series is particularly valuable for North 
American researchers. 

Not only was Mammalian Species the 
brainchild of Sydney Anderson, but he 
sought and obtained approval for the new 
publication, demonstrated the concept by 
writing the first account, and nurtured the 
publication by serving in an editorial capac- 
ity for 312 of the accounts published. During 
the first year of publication he even sold the 
subscriptions to Mammalian Species. 

Others who served Mammalian Species 
in a regular editorial capacity for the first 
443 accounts were D. F. Williams, T. E. 
Lawlor, B. J. Verts, J. K. Jones, Jr., A. L. 
Gardner, C. J. Phillips, T. L. Best, K. F. 
Koopman, G. N. Cameron, C. S. Hood, J. 
A. Lackey, and D. E. Wilson. Several others 
served as guest editors of single accounts 
when authorship constituted a potential 
conflict of interest. 


Monographs and Special 
Publications 


Three Monographs of the American So- 
ciety of Mammalogists were published, one 
each in 1926, 1927, and 1928. These were: 
number |, Anatomy of the Wood Rat by A. 
Brazier Howell; number 2, The Beaver by 
Edward R. Warren; and number 3, Animal 
Life of the Carlsbad Cavern by Vernon Bai- 
ley. Hartley H. T. Jackson served as editor 
for all three, but was assisted by Edward A. 
Preble, Ethel M. Johnson, and Emma M. 
Charters on the last volume. All volumes 


were published by the Williams and Wilkins 
Company, Baltimore, Maryland. Number | 
was priced at $5.00, numbers 2 and 3 at 
$3.00 each; members of the American So- 
ciety of Mammalogists were afforded an 8% 
discount. 

Anatomy of the wood rat consists of nine 
chapters in 225 pages that include 4 tables, 
37 line drawings (seven overprinted with 
red and blue), 3 plates (photographs), a 
3-page bibliography, and a 5-page index. 
The beaver consists of an introduction, ac- 
knowledgments, and 13 chapters in 177 
pages that include 78 illustrations (70 pho- 
tographs), a 5-page bibliography, and a 
3-page index. Animal life of the Carlsbad 
Cavern consists of eight chapters in 195 
pages that include 67 figures (62 photo- 
graphs, 2 maps, and 3 drawings by L. A. 
Fuertes), and a 9-page index; no bibliogra- 
phy was included. In addition to chapters 
on mammals, the volume contained chap- 
ters on birds, reptiles, and invertebrates. 

Strangely, minutes of the meetings of the 
Board of Directors or of the members at 
large published in the Journal of Mam- 
malogy in the years before publication of 
the monographs contain no mention of of- 
ficial sanction or other involvement of the 
society. However, the minutes of the eighth 
annual meeting contain a statement an- 
nouncing the forthcoming publication of the 
first monograph (Journal of Mammalogy, 
7:241, 1926). Advertisements of the avail- 
ability of the monographs appeared on the 
inside of the back cover of the Journal of 
Mammalogy for several years. 

The minutes of the meeting of the Board 
of Directors at the 44th annual meeting held 
at Ciudad Universitaria, Mexico City, D.F., 
Mexico, include the statement, “‘The reviv- 
al of a monograph series was approved” 
(Journal of Mammalogy, 45:668, 1964). 
However, no mention was made in those 
minutes or those of subsequent meetings 
regarding the decision not to continue the 
monographs series per se, but to initiate an 
entirely new series. According to J. K. Jones, 
Jr. (pers. comm., 8 August 1990), a member 


{52 VERTS AND BIRNEY 


of the committee involved in reestablishing 
a monograph series, the 25-year period be- 
tween publication of the third monograph 
and consideration of reestablishment of the 
series, and the desire to change the focus of 
the monograph series, were paramount in 
the decision. At a special meeting of the 
Directors at the 45th annual meeting, “A 
maximum of $8,000 was authorized for the 
publication of an acceptable manuscript for 
the first Special Publication of the Society” 
(Journal of Mammalogy, 46:731, 1965). 

The first Special Publication, The natural 
history and behavior of the California sea 
lion, by Richard S. Peterson and George A. 
Bartholomew, was published 5 December 
1967. On page 11 of this number the series 
was described as follows: “This series, pub- 
lished by the American Society of Mam- 
malogists, has been established for papers 
of monographic scope concerned with some 
aspect of the biology of mammals.”’ William 
H. Burt was editor of the initial number, 
and J. Knox Jones, Jr., James N. Layne, and 
M. Raymond Lee were listed as additional 
members of the Committee on Special Pub- 
lications. The original price of the 91-page 
clothbound book was $3.50. 

Eleven numbers in this series have ap- 
peared, the most recent being the present 
volume in 1994. Published numbers, au- 
thors or editors, and dates of publication of 
Special Publications, in addition to the first, 
are as follows: number 2, Biology of Pero- 
myscus (Rodentia), edited by John A. King, 
20 December 1968; number 3, The life his- 
tory and ecology of the gray whale (Eschrich- 
tius robustus), by Dale W. Rice and Allen 
A. Wolman, 30 April 1971; number 4, Pop- 
ulation ecology of the little brown bat, My- 
otis lucifugus, in Indiana and north-central 
Kentucky, by Stephen R. Humphrey and 
James B. Cope, 30 January 1976; number 
5, Ecology and behavior of the manatee (Tri- 
chechus manatus) in Florida, by Daniel S. 
Hartman, 27 June 1979; number 6, Loco- 
motor morphology of the vampire bat, Des- 
modus rotundus, by J. Scott Altenbach, 22 


August 1979; number 7, Advances in the 
study of mammalian behavior, edited by 
John F. Eisenberg and Devra G. Kleiman, 
11 March 1983; number 8, Biology of New 
World Microtus, edited by Robert H. Tam- 
arin, 12 September 1985; number 9, Dis- 
persal in rodents: a resident fitness hypoth- 
esis, by Paul K. Anderson, 30 March 1989; 
number 10, Biology of the Heteromyidae, 
edited by Hugh H. Genoways and James H. 
Brown, 20 August 1993; and number 11, 
Seventy-five years of mammalogy (1919- 
1994) edited by Elmer C. Birney and Jerry 
R. Choate, 1994. 

Although all monographic in scope, these 
11 Special Publications can be categorized 
by scientific content and organization. 
Numbers 1, 3, 4, 5, and 6 each concentrate 
on one mammalian species, contain 79-153 
(Y = 118) numbered pages, and typically 
concern natural history and related topics. 
Of these, number 6 focuses exclusively on 
locomotor morphology, thus is the most 
specialized in terms of topics covered. 
Numbers 2, 7, 8, and 10 (which has a dou- 
ble-column format) contain 593-893 (X = 
740) numbered pages and consist of several 
(14-22) contributed manuscripts on a topic 
selected by an organizing editor. Numbers 
2 and 8 focus on a particular genus of mam- 
mals and number 10 pertains to a family of 
mammals, whereas number 7 covers a gen- 
eral topic (animal behavior). Number 9 fits 
neither of these categories, thus is unique 
within the series in that it presents and ad- 
vocates a new hypothesis (on dispersal in 
rodents) and compares and contrasts it with 
competing hypotheses. Number 11 also is 
unique, reviewing 75 years of mammalogy, 
as influenced by the ASM. 

Several members of the American Society 
of Mammalogists have served as editor or 
managing editor of the books in the Special 
Publications series. In addition to editing 
the first number, William H. Burt also ed- 
ited number 2. Beginning with number 3, 
each number had both an editor and a man- 
aging editor; the former was responsible for 


PUBLICATIONS 153 


selection, content, and quality control, the 
latter for matters related to production. 
James N. Layne served as editor and J. Knox 
Jones, Jr. as managing editor for numbers 
3-6; Hugh H. Genoways (editor) and Tim- 
othy E. Lawlor (managing editor) edited 
numbers 7 and 8; and Elmer C. Birney and 
Carleton J. Phillips served in these two ca- 
pacities, respectively, for number 9. Addi- 
tionally, Jerry R. Choate served as editor 
and Don E. Wilson as managing editor for 
a brief period, and Michael A. Mares (edi- 
tor) Craig S. Hood (managing editor) edited 
volume number 10. Mares (editor) and Jo- 
seph F. Merritt (managing editor) were re- 
sponsible for number 11. 


Cumulative Indices and 
Miscellaneous Publications 


Four cumulative indices to the Journal of 
Mammalogy have been published to date, 
and a fifth is scheduled for publication. The 
first was a 20-year index to volumes 1-20 
(1919-1939) edited by Viola S. Schantz and 
Emma M. Charters; it consists of 219 pages 
and sold for $2.50 in paperback, $3.50 
clothbound, when published on | August 
1945. Each of the next three published in- 
dices covered a 10-year span: volumes 21- 
30 (1940-1949), 31-40 (1950-1959), and 
41-50 (1960-1969). The second index also 
was edited by Schantz and Charters, the third 
by Schantz and a committee of four others, 
and the fourth was prepared by James S. 
Findley and six additional members of the 
Index Committee. The fifth index is to cov- 
er a 20-year period (volumes 51-70) and is 
being prepared by Michael Carleton and the 
four or five other members of the 1983- 
1990 Index Committees. The cumulative 
index for the decade of the 1940s consists 
of 146 numbered pages, appeared on 27 Oc- 
tober 1952, and sold originally for $3.25 in 
paperback, $3.75 in clothbound. That for 
the 1950s has 150 pages, a publication date 
of 18 May 1961, and sold for $5.00 in cloth- 


bound only. The fourth cumulative index 
consists of 109 numbered pages, is dated 
only as 1974, and sold for $5.00 in cloth- 
bound only. Each of these indices contains 
a few (4-10) pages of introduction and ex- 
planation in addition to the numbered pages. 

Six indices to Mammalian Species have 
been published; these are to species ac- 
counts numbered 1-100, 1-200, 1-300, 1- 
400, 101-200, and 201-306. Except for the 
indices to accounts numbered 1-300 and 1- 
400, which lack author indices, each index 
contains systematic, generic, and author 
lists. These accounts were distributed to 
subscribers with fascicles containing appro- 
priately numbered accounts. 

In April 1981, the American Society of 
Mammalogists published a limited edition 
of a pamphlet titled “Career trends and 
graduate education in mammalogy,” by 
Gary W. Barrett and Guy N. Cameron. An 
announcement of the availability of publi- 
cation and a notice of publication of a quar- 
terly newsletter for graduate students ap- 
peared in the comments and news section 
of the Journal of Mammalogy (62:875, 
1981). 

In June-July 1985, the American Society 
of Mammalogists cosponsored with the 
Australian Mammal Society publication of 
a special issue (volume 8, numbers 3 and 
4) of Australian Mammalogy containing pa- 
pers presented at symposia at the 1984 joint 
meeting of the two societies in Sydney, New 
South Wales, Australia. Number 3 con- 
tained six papers from a symposium titled 
‘““Niche spaces and small mammal com- 
munities’; number 4 contained papers from 
two symposia: “A” titled “Form-function 
analyses: the teeth and skulls of carnivores” 
with five papers, and “B”’ titled ‘Studies in 
the biology of bats’? with nine papers. Each 
of the three sections was edited by a different 
pair of editors: the first by Barry Fox and 
Roger A. Powell, the second by Roger A. 
Powell and Michael Archer, and the third 
by Leslie S. Hall and Suzanne J. Hand. Each 
section also included a preface in which one 


154 VERTS AND BIRNEY 


of the editors summarized and synthesized 
material presented by the participants. Last- 
ly, an envelope attached inside the back 
cover contains a microfiche with appendices 
to a paper in the niche-space symposium 
and contains abstracts of other papers pre- 
sented at the joint meeting. The issue of 
Australian Mammalogy (volume 8, num- 
bers 3 and 4, 1985) was available from the 
secretary-treasurer of the American Society 
of Mammalogists for $10 for those attend- 
ing the joint meeting and $15 for others. 

For several years, the secretary-treasurer 
has published brochures that contain lists 
and descriptions of Special Publications and 
Mammalian Species accounts with appro- 
priate order forms. Another brochure titled 
“The science of mammalogy” includes a de- 
scription and a brief history of mammalogy 
in North America and of the American So- 
ciety of Mammalogists. Lastly, a brochure 
titled “Careers in mammalogy” contains 
brief descriptions of the types of work that 
mammalogists do and of career opportu- 
nities in mammalogy. The latter two bro- 
chures were produced by the Committee on 
Education and Graduate Students. All of 
the brochures are revised or updated from 
time to time. 


Acknowledgments 


Thanks are due L. N. Carraway and L. F. Al- 
exander for assistance with the analyses. This is 
Technical Paper No. 10,041, Oregon Agricul- 
tural Experiment Station. 


Literature Cited 


ANDERSON, S., AND J. K. JoNEs, JR. 1984. Introduc- 
tion. Pp. 1-10, in Orders and families of Recent 
mammals of the world (S. Anderson and J. K. Jones, 
Jr., eds.). John Wiley & Sons, New York, 686 pp. 

ANDERSON, S., AND R. G. VAN GELDER. 1970. The 
history and status of the literature of mammalogy. 
BioScience, 20:949-957. 

GmLL, A. E., AND W. C. WozENCRAFT. 1994. Com- 
mittees and annual meetings. Pp. 155-170, in Sev- 
enty-five years of mammalogy (1919-1994) (E. C. 
Birney and J. R. Choate, eds.). Special Publication, 
The American Society of Mammalogists, 11:1—433. 

Jones, J. K., JR., R. S. HOFFMANN, D. W. Rice, C. 
JONES, R. J. BAKER, AND M. D. ENGstTrRom. 1992. 
Revised checklist of North American mammals north 
of Mexico, 1991. Occasional Papers, The Museum, 
Texas Tech University, 146:1-23. 

KIRKLAND, G. L., JR., AND H. D. SMitH. 1994. Mem- 
bership and finance. Pp. 170-178, in Seventy-five 
years of mammalogy (1919-1994) (E. C. Birney and 
J. R. Choate, eds.). Special Publication, The Amer- 
ican Society of Mammalogists, 1 1:1—433. 


COMMITTEES AND ANNUAL MEETINGS 


AYESHA E. GILL AND W. CHRIS WOZENCRAFT 


Introduction 


he first meeting of the ASM was held 
3-4 April 1919 in Washington, D.C., 
2 years after the end of World War I. This 
was the year that Prohibition, the 18th 
Amendment to the United States Consti- 
tution, was ratified and that the great Mex- 
ican leader, Emiliano Zapata, was killed. 
This organizational meeting was attended 
by 60 members of a charter membership of 
over 250. After discussion and approval of 
by-laws and a constitution, six officers and 
10 councilors (now called the Board of Di- 
rectors) were elected. An editor was selected 
for the society’s Journal of Mammalogy that 
was to start publication that year. Five 
standing committees were formed: Publi- 
cations, Life Histories of Mammals, Study 
of Game Mammals, Anatomy and Phylog- 
eny, and Bibliography. The policy of the 
society set forth at its organizational meet- 
ing was “to devote its attention to the study 
of mammals in a broad way, including life 
histories, habits, evolution, palaeontology, 
relations to plants and animals, anatomy, 
and other phases.’ The annual dues were 
$3 (Kirkland and Smith, 1994). 

ASM currently (1993) has nearly 4,000 
members residing in 60 countries. The so- 
ciety has held a general membership meet- 
ing every year since 1919 except for 2 years 


i ers) 


Publications 

Life Histories 

Study of Gane Pamrals 
Anatorsy and Phylogeny 
Bibliography 


during World War IT (1943 and 1944), when 
only the directors met. General member- 
ship meetings have been held in Washing- 
ton, D.C., Canada, Mexico, and 30 states 
of the U.S. ASM has had 38 presidents be- 
tween 1919 and 1993 (Layne and Hoff- 
mann, 1994). During this period, 74 vol- 
umes of the Journal of Mammalogy, 10 
Special Publications and close to 450 Mam- 
malian Species accounts have been pub- 
lished (Verts and Birney, 1994). New stand- 
ing and ad hoc committees were formed and 
old ones phased out; the current number of 
standing committees is 23. 

Hartley H. T. Jackson, a young staff 
member of the U.S. Biological Survey, 
played a prominent role in planning for the 
establishment of the ASM (ASM 50th An- 
niversary Program, 1969; Hoffmeister, 
1969; Hoffmeister and Sterling, 1994). 


History of the Committees of ASM 


Many members have served the ASM by 
actively participating in the work of the so- 
ciety’s committees and thus have contrib- 
uted to its development and vigor. Standing 
committees have functioned since the in- 
ception of ASM to promote the goals of the 


156 GILL AND WOZENCRAFT 


society through ongoing activities between 
annual meetings. These committees and 
their chairpersons are appointed by the 
president. The committees have a two-fold 
purpose—to conduct affairs of the society 
and to accord members the responsibilities 
and rewards of active participation in it. An 
ad hoc committee was appointed in 1982 
under the presidency of J. Mary Taylor to 
evaluate the standing committees and ex- 
plore all facets of their roles in the society. 
Past and present members of standing com- 
mittees and other members of the society 
were contacted with questions pertaining to 
the committees. Through their responses, 
the ad hoc committee compiled detailed re- 
ports on standing committees, including 
their history, function, effectiveness, and 
recommendations of committee members 
on the continued need and role of each com- 
mittee. These reports are sent to members 
of the committees so that they can have a 
better understanding of the committee’s 
purpose and how to serve on it effectively. 
Shorter descriptions of the functions of the 
current standing committees were first pub- 
lished as a supplement to Vol. 68, No. 1 
(1987) of the Journal of Mammalogy (‘Roles 
of Standing Committees of the American 
Society of Mammalogists’’). We have up- 
dated the list of Standing Committees of the 
ASM from its inception to extend it to the 
present (1993) (Table 1). In addition to the 
40 standing committees formed during the 
society’s history, ad hoc committees have 
been established frequently to perform spe- 
cific tasks. They cease to exist when their 
charge is completed. Often, however, a 
standing committee develops from an ad 
hoc committee, if a more lasting need for 
its function is perceived by the Board of 
Directors. Current standing committees and 
their members are listed on the inside of the 
back cover of each issue of the Journal. 
The standing committees created during 
the history of ASM can be divided into cat- 
egories concerned with the promotion of 
mammalogy, the development of the soci- 
ety itself, or the interactions of ASM with 


non-mammalogists. Committees have dealt 
with publications and particular topics in 
mammalogy (such as physiology and anat- 
omy, ecology, and conservation); some have 
dealt with taxonomy in general and others 
with specific taxa. Committees have been 
created to encourage young mammalogists 
and to build the society, to honor and re- 
ward its outstanding members and other 
mammalogists, and to record its history. 
Committees exist to promote the interac- 
tion of the society’s members with other 
mammalogists and to present the society’s 
views on critical national and international 
issues affecting mammalogy. Brief descrip- 
tions of the committees involved in each of 
these areas of activity follow. 

Promotion of mammalogy. —The original 
Publications Committee formed in 1919 
evolved in 1930 into the Editorial Com- 
mittee, which remains active. It oversees 
production of the Journal of Mammalogy, 
Mammalian Species, Special Publications, 
and miscellaneous publications such as 
membership lists. It sets editorial policy for 
the ASM, nominates new editors for ap- 
proval by the Board of Directors, and man- 
ages the publication budget. The committee 
is composed almost entirely of current ed- 
itors, who can be divided into two groups: 
those involved in the review process and 
judging the scientific merit of papers and 
those involved in the technical production 
of the publications. As the Journal of Mam- 
malogy grew over the years, the need de- 
veloped for a committee to prepare the in- 
dex for each volume. The Index of Journal 
of Mammalogy Committee was formed in 
1947, chaired by Viola S. Schantz, who also 
served the society for 23 years (1930 to 1952) 
as Treasurer. The name of this committee 
was abbreviated to Index Committee in 
1972. Besides preparing the index for each 
volume of the Journal, it prepares summary 
indices. The Bibliography Committee, 
which existed for 67 years, compiled the list 
of Recent Literature in Mammalogy for 
many years before it ceased to exist. This 
information is now available through other 


TABLE |.— Standing committees of the ASM from its inception in 1919 to 1993. 


Year formed 
ASM President 


1919 
C. H. Merriam 


1920 

C. H. Merriam 
1921 

E. W. Nelson 


1922 

E. W. Nelson 
1927 

G. M. Allen 


1928 
G. M. Allen 


1930 
W. Stone 


1945 

R. Hall 
1947 

R. Kellogg 


1950 
T. I. Storer 


1953 

W. H. Burt 
1956 

W. B. Davis 


COMMITTEES AND MEETINGS 


Committee 


Publications 


Life Histories of 
Mammals 

Study of Game Mam- 
mals 

Anatomy and Phylog- 
eny 


Bibliography 
Conservation 


Marine Mammals 


Economic Mammal- 
ogy 
J. A. Allen Memorial 


Life Histories and 
Ecology 

Conservation of Land 
Mammals 


Nomenclature 


Editorial 


Membership 


Special Committee on 
Trapping Methods 


Ecology (including life 
histories and popu- 
lations) 

Economic Mammalo- 
gy and Conserva- 
tion 

Index of Journal of 
Mammalogy 

Means for Encourag- 
ing Young Mam- 
malogists 

Dues Status of Re- 
tired Members 

Honoraria for Gradu- 
ate Students 


Resolutions 


Original chairperson/members 


G. S. Miller, Jr./E. A. Preble, W. P. Tay- 


lor, H. H. T. Jackson 


C. C. Adams/R. M. Anderson, V. Bailey, 


H. C. Bryant 


C. Sheldon/G. B. Grinnell 


W. K. Gregory/J. C. Merriam, H. H. Don- 
aldson, A. Wetmore, H. von W. Schulte, 


J. W. Gidley 


T. S. Palmer/W. H. Osgood, H. H. T. 


Jackson 


W. H. Osgood/E. W. Nelson, J. Dwight 


E. W. Nelson/G. S. Miller, Jr., T. S. Palm- 
er, B. W. Evermann, R. C. Murphy, G. 


M. Allen 


A. K. Fisher/W. B. Bell, H. C. Bryant 


M. Grant/H. F. Osborn, C. Frick, G. B. 
Grinnell, H. E. Anthony 


W. P. Taylor/C. S. Adams, V. Bailey 


E. A. Preble/J. C. Phillips, T. S. Palmer 


W. H. Osgood/G. M. Allen, A. H. Howell, 
G. S. Miller, Jr., T. S. Palmer 

E. A. Preble/G. M. Allen, A. H. Howell, 
R. Kellogg, G. S. Miller, Jr., G. B. Wis- 


locki 


W. P. Harris, Jr./T. Gregory, V. Bailey, 
R. M. Anderson, M. R. Thorpe, J. Dix- 


on, W. P. Taylor 


W.E. Sanderson/C. C. Adams, E. A. Preb- 


le, W. A. Young 


D. L. Allen/F. S. Barkalow, Jr., C. D. H. 
Clarke, W. J. Hamilton, Jr., J. M. Lins- 


dale 


E. R. Kalmbach/D. L. Allen, R. M. An- 
derson, A. E. Borell, H. J. Coolidge, T. 


I. Storer, C. T. Vorhies 


V. S. Schantz/H. H. T. Jackson, D. H. 


Johnson, R. Kellogg 


D. E. Davis/F. S. Barlow, Jr., P. D. Dalke 


A. R. Shadle/J. K. Doutt, R. I. Peterson 


W. R. Eadie/F. S. Barkalow, Jr., S. D. 


Durrant, R. T. Orr 


K. R. Kelson/E. T. Hooper, W. V. Mayer, 
S. D. Durrant, G. C. Rinker 


157 


Year 
ended 


1930 
LOD, 
1922 


1948 


1985 


1922 


active 


1953 


1929 


1947 


active 


active 


active 


active 


1947 


1948 


1948 


1972 


1951 


1951 


active 


active 


158 


GILL AND WOZENCRAFT 


TABLE 1.— Continued. 


Year formed Year 
ASM President Committee Original chairperson/members ended 
1957 Honorary W. H. Burt/E. R. Hall, W. J. Hamilton, active 
W. B. Davis Membership Jr. 
1960 International A. De Vos/W. O. Pruitt, Jr., H. M. Van active 
S. D. Durrant Relations Deusen 
1962 Anatomy and L. C. Dearden/K. L. Duke, M. Hilde- 1983 
E. T. Hooper Physiology brand, P. H. Kurtzsch, P. R. Morrison, 
W. B. Quay 
1966 Historian D. F. Hoffmeister 1986 
R. G. Van Gelder 
1971 Grants In Aid J.S. Findley/R. Horst, H.M. Van Duesen, active 
J. N. Layne J. L. Wolfe 
1971 Program B. E. Horner/L. N. Brown, O. P. Pearson, active 
J. N. Layne M. H. Smith, H. M. Van Duesen 
1972 Information S. Anderson/L. de la Torre, H. H. Gen- active 
J. K. Jones, Jr. Retrieval oways, R. S. Hoffmann, C. Jones, D. R. 
Patten, J. L. Patton, H. W. Setzer 
Index D. E. Wilson/R. D. Fisher, C. Jones, J. L. active 
Paradiso, R. H. Pine, H. W. Setzer, R. 
W. Thorington, Jr. 
Systematic J. R. Choate/J. H. Brown, E. T. Hooper, active 
Collections M. L. Johnson, C. Jones, J. L. Patton, 
T. A. Vaughan 
1974 Merriam Award J. K. Jones, Jr./C. C. Black, W. H. Burt, active 
S. Anderson J. F. Eisenberg, M. E. Hight, T. A. 
Vaughan, J. Whittaker 
1976 Legislation and Regu- H.H.Genoways/M.M. Alexander,S.An- active 
W. Z. Lidicker, Jr. lations derson, M. A. Bogan, J. R. Choate, R. 
C. Dowler, C. A. Hill, A. M. Johnson, 
C. Jones, T. J. McIntyre, J. L. Paradiso, 
R. L. Peterson 
1977 Jackson Award R. L. Peterson/W. H. Burt, J. S. Findley, active 
W. Z. Lidicker, Jr. D. F. Hoffmeister, C. Jones 
Mammal Slide J. A. Lackey/P. V. August, S. J. Bleiweiss, active 
Library P. L. Dalby, D. C. Gordon, H. L. Gun- 
derson, J. G. Hall, G. C. Hickman, L. 
L. Master, J. S. McCusker, G. L. Tweist 
1978 Education and Grad- G. W. Barrett/A. E. Baker, G. N. Cam- active 
R. S. Hoffmann uate Students eron, A. F. DeBlase, S. R. Humphrey, 
K. A. Shump, Jr. 
1982 Checklist K. Koopman/J. H. Calaby, F. Dieterlen, active 
J. M. Taylor R. S. Hoffmann, J. H. Honacki, J. G. 
Mead, G. G. Musser, P. Myers, R. W. 
Thorington, Jr. 
1986 Archives D. F. Hoffmeister, Historian; W.C. Woz- active 
D. E. Wilson encraft, Archivist 
1989 Development S. R. Humphrey/S. Anderson, J. R. active 
E. C. Birney Choate, H. H. Genoways, W. Z. Lidick- 


er, Jr., R. L. Peterson, D. J. Schmidly, 
J. M. Taylor, R. G. Van Gelder, M. R. 
Willig, D. E. Wilson 


COMMITTEES AND MEETINGS 159 


TABLE |.— Continued. 


Year formed 


ASM President Committee 
1990 Animal Care 
E. C. Birney and Use 


Year 
Original chairperson/members ended 
T. H. Kunz/R. J. Baker, T. Carter, J. R. active 


Choate, J. A. Cranford, G. Glass, I. F. 
Greenbaum, L. R. Heaney, G. R. Mich- 
ener, T. H. McIntyre, D. K. Odell, R. 
S. Ostfeld, A. Pinter, V. Scheffer, S. D. 
Thompson, R. A. Van Den Bussche 


Source: Journal of Mammalogy, Volumes 1-73, Supplement to Vol. 68, No. 1 (1987). 
Summary: In 1993, 23 active ASM committees, 17 extinct. 
See Layne and Hoffmann (1994) for additional information on presidents. 


means, such as computerized literature 
searches. 

The Mammal Slide Library Committee 
was established in 1977 to provide low-cost 
slides of mammals, often in natural habi- 
tats, principally for educational purposes. It 
now also stresses use of its slides for world- 
wide conservation efforts. The committee 
solicits, selects, and catalogs slides, and ad- 
vertises their availability to potential users 
world-wide. By 1993 over 1,000 different 
slides depicting 756 species in 19 orders were 
available. Over 100,000 duplicate slides 
were sold between 1978 and 1993. 

Many of the ASM committees that dealt 
with specific topics in mammalogy no lon- 
ger exist. These include the Life Histories 
of Mammals Committee (1919-1927), 
which evolved into the Life Histories and 
Ecology Committee (1927-1947) and final- 
ly into the Ecology Committee (including 
life histories and populations) (1947-1948). 
One committee focused on morphology, the 
Anatomy and Phylogeny Committee (1919- 
1948), and again in 1962-1983 (Anatomy 
and Physiology) but, although lasting for 
much of the society’s history, is no longer 
in existence. One special topics committee 
has proved remarkably resilient. The Ma- 
rine Mammals Committee, established in 
1921 when the society was just 2 years old, 
is still functioning. It provides the society 
membership with information about ma- 
rine mammalogy, including conservation 
and legislative issues, spearheads resolu- 


tions and legislation involving marine 
mammals, and serves as a liaison between 
ASM and the Society for Marine Mam- 
malogy (SMM). Committee members fre- 
quently are active in both ASM and SMM. 
The committee is particularly active on leg- 
islative issues regarding marine mammals. 

Several of the society’s committees have 
been concerned with economic mammalogy 
and conservation, of which one still exists. 
The earliest of these committees, the Study 
of Game Mammals, was initiated in 1919 
and lasted only 4 years; the Conservation 
Committee (1920-1922) also was short- 
lived. The successor to these committees, 
however, lasted much longer. The Com- 
mittee on Economic Mammalogy lasted 33 
years (1921-1953). The Committee on Eco- 
nomic Mammalogy and Conservation had 
a brief life (1947-1948), but the Committee 
on Conservation of Land Mammals, estab- 
lished in 1927, is one of the oldest active 
committees of the society. It monitors state, 
national, and international governmental 
activities and other activities that relate to 
conservation of land mammals, and it ad- 
vises officers and members of the society on 
issues of concern. The committee responds 
to these issues via formal resolutions to the 
membership, letters to responsible individ- 
uals or agencies, and other appropriate 
means. It serves as a clearinghouse for in- 
formation, leads or facilitates collective or 
individual responses to conservation issues, 
and, in a related function, establishes and 


160 GILL AND WOZENCRAFT 


maintains liaison with other conservation 
groups. Conservation always has been a 
concern of the society and, today, with the 
increasing loss of genetic variability in the 
world, remains a major concern. 

The society has four active committees 
concerned with taxonomy and systematic 
collections. The earliest, the Nomencla- 
ture Committee, was formed in 1928 to 
give advice to members of ASM on prob- 
lems pertaining to nomenclature and to 
answer any taxonomic questions that 
members might pose. About 1977 the 
committee also assumed an advisory re- 
lationship with the International Com- 
mission of Zoological Nomenclature. It 
screens applications involving North Amer- 
ican mammals to ascertain whether the facts 
as presented are both correct and complete 
and provides an opinion on what the general 
effect of the requested ruling will be on tax- 
onomic and nomenclatural practice. 

The Systematic Collections and the In- 
formation Retrieval Committees both were 
formed in 1972. The former was an out- 
growth of an ad hoc committee established 
at the request of the National Science Foun- 
dation to evaluate mammal collections for 
support by the NSF Biological Research Re- 
sources Program. The original charge in- 
cluded advising the society on matters per- 
taining to systematics and systematic 
collections and reviewing criteria for ap- 
praising collections. It also reviewed pro- 
posals submitted to granting agencies for 
monetary support of systematic collections. 
The present role of the committee focuses 
on the general objective of promoting prop- 
er maintenance of systematic collections. 
The committee has established minimal 
standards for proper maintenance of collec- 
tions, and it serves, on behalf of the society, 
as an informal inspecting and accrediting 
agency for the curatorial status of collec- 
tions. It also is responsible for surveys of 
collections of Recent mammals published 
periodically in the Journal of Mammalogy. 

The birth of the Information Retrieval 
Committee is indicative of the revolution 


that has occurred in information-retrieval 
systems and computers. Its original charge 
was to examine the feasibility of developing 
a national data-retrieval system for Recent 
mammal collections and, if possible, to de- 
velop funding for such a system. The com- 
mittee’s activities have involved develop- 
ing standardized documentation and 
retrieval methods, producing a publication 
on automatic data processing, and provid- 
ing information on computerization of 
mammal collection data. The evolving 
interests of the committee include, but 
expand beyond, collection-based informa- 
tion to bibliographic data and other natural- 
history data bases in mammalogy. 

The Checklist Committee was estab- 
lished in 1980 to provide advice on Mam- 
mal Species of the World, edited by J. H. 
Honacki et al. (1982) and published by Al- 
len Press and the Association for System- 
atics Collections. The first edition was pre- 
pared by 189 professional mammalogists 
from 23 countries and was coordinated by 
the Checklist Committee, with R. S. Hoff- 
mann as the Project Coordinator. During 
the last 10 years it has become the inter- 
national standard for mammalian taxono- 
my. The dynamics of mammalian taxono- 
my demand periodic updates of this vast 
amount of information. ASM and the 
Checklist Committee assumed responsibil- 
ity for the maintenance of this data base and 
its periodic revisions. The committee serves 
as both scientific consultant to editors and 
final arbitrator of nomenclatural, or other, 
decisions on content. The material has been 
transferred from a text-based manuscript to 
an information retrieval data base to facil- 
itate future updates and to enhance the abil- 
ity of the user to interactively access this 
information. The second edition of Mam- 
mal Species of the World (Wilson and Reed- 
er, 1993) was published in cooperation with 
the Smithsonian Institution Press. 

Development of the society. —The society 
has established five committees to encour- 
age young mammalogists, and four of these 
are still extant. The Committee on Hono- 


COMMITTEES AND MEETINGS 161 


raria (originally Honoraria for Graduate 
Students), formed in 1953, selects graduate 
students to be honored for their research in 
mammalogy. At present, three awards are 
given: the Anna M. Jackson, the A. Brazier 
Howell, and the American Society of Mam- 
malogists awards. Recipients are awarded 
an honorarium to attend the annual meet- 
ing, where they present results of their re- 
search at a plenary session. The Grants- 
in-Aid Committee was created in 1971 to 
solicit applications and select recipients for 
grants-in-aid of research and a nominee for 
the Albert R. and Alma Shadle Fellowship 
in Mammalogy. The grants are presently 
given to 11 students, with a monetary limit 
of $1,000 per student. The highest ranking 
student is honored with the B. Elizabeth 
Horner Award and gets a bonus of $100. 
The Shadle Fellowship, usually about 
$3,000, is awarded annually by the Buffalo 
Foundation and is intended to promote a 
professional career for a mammalogy stu- 
dent showing great promise. The ASM 
Grants-in-Aid Committee nominates the 
student and an alternate. Nominees for this 
award do not have to be members of the 
ASM but must be citizens of the United 
States. The recipient of the Shadle Fellow- 
ship is invited to speak on his or her re- 
search at an annual ASM meeting. 

The Committee on Education and Grad- 
uate Students was formed in 1978, with the 
purpose of assisting students of mammal- 
ogy to make more informed choices of ca- 
reer, to improve their scientific expertise, 
and to find employment in the discipline. 
These aims are achieved through prepara- 
tion of brochures and reports and through 
sponsoring of workshops related to educa- 
tion of mammalogists, career opportunities, 
research support, and other topics of inter- 
est to students. The Development Com- 
mittee began as an ad hoc committee in 
1989. It has raised monies for the Future 
Mammalogists Fund, established by R. L. 
Peterson in 1985, through a variety of 
means, including contributions to patron 
membership ($1,000), the Seventy-five Year 


Club ($75), and other individual contribu- 
tions. The initial goal of raising $100,000 
for the Fund was achieved in 1991. In 1993, 
at the recommendation of this committee, 
the Board of Directors initiated a major pro- 
gram of planned giving, including living 
trusts, pooled income funds, and wills and 
bequests. 

The Membership Committee and the 
Program Committee are instrumental in 
maintaining the society. The Membership 
Committee was established in 1930 to en- 
courage persons with an interest in mam- 
mals to become members of the society. In 
recent years an emphasis has been placed 
on making new members feel ‘‘at home” in 
the society and encouraging retention of 
members through writing welcoming letters 
to new members, pursuing any problems in 
the mailing of journals, and writing to 2-year 
delinquents to determine their reasons for 
dropping membership. In 1990-1991 the 
committee wrote to non-member authors 
who submitted manuscripts to the Journal 
(171 persons), inviting them to join the so- 
ciety; 14% of them did, accounting for 6% 
of the new members at that time. 

The Program Committee was established 
in 1971 in response to the need for a more 
effective method of selecting sites for and 
improving the organization of the annual 
meeting. It promotes the annual meeting 
and assists in its organization and conduct. 
This committee selects the basic format of 
the meeting, solicits, reviews, and selects 
symposia and workshops, and develops 
guidelines to increase the effectiveness of 
presentation of papers and posters. It also 
explores possibilities for special meetings, 
such as joint meetings with other mammal 
societies, and participates in planning the 
content and format of such meetings. It ad- 
vises local committees and provides liaison 
between successive local committees. The 
Program Committee constantly re-evalu- 
ates the organization of annual meetings, 
based on the accumulated experiences of 
local host committees. 

The society takes pride in honoring in- 


162 GILL AND WOZENCRAFT 


dividuals who have made outstanding con- 
tributions to mammalogy and has set up a 
number of committees to evaluate and se- 
lect these outstanding mammalogists. 

The Honorary Membership Committee 
was formed in 1957 to recommend candi- 
dates for honorary membership in ASM in 
recognition of distinguished service to the 
science of mammalogy, but honorary mem- 
bers have been chosen since the first meet- 
ing of the society, when Joel A. Allen was 
selected (Hoffmeister, 1969). The commit- 
tee is comprised of the five most recent past 
presidents with the chair held for a 2-year 
period by the second-most senior member. 
The C. Hart Merriam Award was estab- 
lished in 1974 to provide recognition for 
outstanding contributions to mammalogy 
by a member of the society. It was named 
in honor of one of the foremost early North 
American mammalogists, who also was first 
president of the society. The award is now 
given in recognition of excellent scientific 
research and either education of mammal- 
ogists or service to mammalogy (Anony- 
mous, 1992). Nominations for the Merriam 
Award are open to all mammalogists, re- 
gardless of country or membership in ASM. 
The Jackson Award Committee was estab- 
lished in 1977 to provide recognition of per- 
sons who have rendered long and outstand- 
ing service to the society. The committee 
evaluates nominations received and, based 
on supporting documentation, makes its 
recommendation of a recipient to the Board 
of Directors. These awards need not be made 
each year, as they are reserved for truly wor- 
thy candidates. 

The position of Historian was created in 
1966 when it was realized that important 
historical material of value to the society 
was not being preserved. Donald F. Hoff- 
meister has served as the society’s historian 
since the inception of this committee. The 
material originally was preserved in the Mu- 
seum of Natural History, University of II- 
linois, but in 1986 the decision was made 
to move all materials to the care of the Ar- 
chives of the Smithsonian Institution. At 


that time the Archives Committee was 
formed, consisting of Hoffmeister, who con- 
tinued as Historian, and W. Chris Wozen- 
craft as Archivist. The archives include in- 
formation on annual meetings; minutes of 
board meetings and business meetings; pho- 
tographs of past presidents, honorary mem- 
bers, and some other award recipients; an 
official set of the Journal of Mammalogy 
and other society publications; and a variety 
of miscellaneous materials, including cor- 
respondence of many past presidents. 

Interactions of ASM with the broader so- 
ciety. — Throughout its history, ASM has as- 
sumed a responsible role in expressing its 
views on issues relating to mammals and 
mammalogists, partly by means of resolu- 
tions passed by the membership of ASM. A 
Resolutions Committee was established in 
1956 primarily to avoid the problem of 
hastily submitted and poorly drafted last- 
minute resolutions on subject matter of di- 
rect concern to the society. The purpose of 
this committee is to provide a mechanism 
for the society to express its views and to 
try, collectively, to influence local, national, 
and world issues relating to mammals. These 
views are expressed in the form of resolu- 
tions proposed by ASM committees or 
members, or, sometimes, initiated by the 
Resolutions Committee itself, which re- 
views proposed resolutions with the pro- 
posers and other knowledgeable persons to 
ensure their accuracy and appropriateness. 
The committee decides through which 
agencies or channels to send resolutions that 
have been approved by a majority vote of 
the membership. Reports of this committee 
are available in the abridged minutes of ASM 
meetings in issue number 4 of each volume 
of the Journal of Mammalogy. 

A partial listing of resolutions passed by 
ASM is available from the archivist. Con- 
servation issues have been and remain an 
overriding concern of ASM and this is re- 
flected in its resolutions. Conservation and 
protection of large mammals especially, such 
as cetaceans, carnivores, and artiodactyls, 
have been the subject of many resolutions. 


COMMITTEES AND MEETINGS 163 


The society consistently has urged the pro- 
tection of national parks, threatened habi- 
tats, and endangered species and popula- 
tions, including recent resolutions on the 
conservation of biological diversity and 
support for the National Institute for the 
Environment. It has opposed the use of in- 
humane methods of predator control and 
those that poison the environment. The so- 
ciety has advocated forcefully the humane 
treatment of mammals in the wild and in 
captivity, with resolutions ranging from op- 
position to den hunting to destroy wolves 
in Minnesota and Wisconsin to support for 
humane and professional maintenance of 
mammals in captivity. An ad hoc commit- 
tee on Animal Care and Welfare updated 
acceptable field methods and proper labo- 
ratory care in the collection and use of wild 
mammals in research and teaching. This 
committee became the standing committee 
on Animal Care and Use in 1992. The so- 
ciety has taken a stand against scientific cre- 
ationism, supported biological surveys and 
studies of threatened populations and of the 
effects of animal introductions, and has 
hailed the establishment of new societies 
such as the Mexican Society of Mammal- 
ogists (1984) and the Society for Marine 
Mammalogy (1984). 

ASM also acts through its Legislation and 
Regulations Committee, established in 
1976, to bring its expertise to bear on this 
area. The committee was established in re- 
sponse to the need for monitoring and pro- 
viding input into the rapidly burgeoning 
state and federal legislation and regulations 
in such areas as endangered species, steel- 
trapping regulations, and use of animals for 
experimental purposes of direct concern to 
mammalogy. The committee also interacts 
with the legislative monitoring group of 
AIBS. Members of the committee are fa- 
miliar with federal and state agencies and 
operations and they attend agency hearings 
when necessary. 

The International Relations Committee 
was formed in 1960 to maintain and en- 
hance communication between ASM and 


mammalogists outside North America. It is 
truly an international committee with many 
of its members living outside North Amer- 
ica. It maintains liaisons with counterpart 
societies Overseas, participates in the Inter- 
national Theriological Congresses (ITC), 
which it helped initiate, and organizes joint 
meetings with other societies. ITC has held 
meetings in Russia, Czechoslovakia, Fin- 
land, Canada, Italy, and Australia. Joint 
meetings have been held with mammal so- 
cieties in Australia, Argentina, and China. 
The International Relations Committee 
maintains names and addresses of mammal 
societies and mammalogists throughout the 
world. It facilitates exchange of material on 
mammalogy, encourages foreign colleagues 
to attend annual ASM meetings, and fosters 
good relations between mammalogists in- 
ternationally. Members outside North 
America provide a link for international ac- 
tivities and serve as representatives of ASM 
in their own countries. 


The History of ASM Annual 
Meetings 


Population dynamics. —The membership 
of the ASM quickly grew from its initial 252 
members, breaking the 1,000 mark in 1930, 
but throughout the 1930s and most of the 
1940s the number of members returned to 
a range between 800 and 900 (Hoffmeister, 
1969). In 1948 the membership again ex- 
ceeded 1,000, passed 2,000 in 1963, and 
topped 3,000 in 1968; in recent years the 
membership figures have hovered around 
3,700 (Secretary-Treasurer’s annual report, 
1993). There presently are five categories of 
membership in ASM: annual (at $30 one of 
the best bargains in any professional soci- 
ety), life (currently $750), patron (currently 
$1,000), honorary, and emeritus. Emeritus 
membership was established in 1951 for 
persons who had regular membership in the 
society for 25 years or more, were in good 
standing, and requested such membership 
(Hoffmeister, 1969). In 1993, ASM had 636 


164 GILL AND WOZENCRAFT 


Number of papers, ASM meetings 
(1920-1990) 


240 


60 


0 


1920 1930 1940 


1950 


1960 1970 1980 


Fic. 1.—The number of papers and posters presented at ASM meetings, 1920-1990. 


life, 16 patron, 13 honorary, 147 emeritus, 
and 2,938 annual, for a total of 3,750 mem- 
bers. 

The number of papers and, more recently 
posters, presented by persons attending an- 
nual meetings has remained relatively con- 
stant over the years at 42-55% of the total 
number of registered participants. Although 
the general trend has been an increase in the 
number of papers presented, mirroring the 
increase in membership, this increase was 
gradual during the first four decades, in- 
creased more rapidly in the fifth decade, and 
increased dramatically in the mid-1970s 
(Fig. 1). There was a doubling of the number 
of papers in the first half of the 1970s, a high 
that was maintained from that time on, with 
active participation of graduate students be- 
coming a major factor in oral and, later, 
poster presentations and greatly affecting the 
atmosphere and emphasis of the meetings. 

The membership of ASM always has been 
skewed toward males. It is difficult to obtain 
a reliable estimate of the sex ratio of biol- 
ogists at annual meetings, so two approxi- 
mate methods were employed here. The 


number of females in annual meeting pho- 
tographs was counted, although this method 
has the bias of including some non-mem- 
bers. This was especially a factor for some 
of the earlier meetings, where non-members 
frequently appeared with their spouses for 
the photographs. This estimate was com- 
pared to the number of papers presented at 
annual meetings for which the first author 
was female (Fig. 2). Although women have 
increased significantly in a society that was 
essentially male (with active support from 
wives) at the beginning, in 1991 they still 
constituted only 20-30% of the member- 
ship and were first authors on about the 
same percentage of papers. This increase in 
the number of women has occurred prin- 
cipally since the early 1970s and largely as 
a result of an influx of female graduate stu- 
dents, corresponding to the dramatic in- 
crease in the number of graduate students, 
in general, participating in meetings. Only 
one of the 38 presidents of the ASM (through 
1993) has been a woman, J. Mary Taylor, 
who was elected in 1982 (Layne and Hoff- 
mann, 1994). Four of the chairpersons of 


COMMITTEES AND MEETINGS 165 


Estimation of Female biologists 
at ASM annual meetings 


| First Author 


40 


30 


20 


Percent 


10 


1920 1930 1940 


1950 


—— Group Photo 


1960 1970 1980 


Fic. 2.—Estimation of the number of female biologists at ASM annual meetings, based on the first 
author of papers presented (bar graph) and the group photo (line drawing). 


the 23 current standing committees of the 
society are women, lower than the percent- 
age of women in the society. 

An ad hoc committee on Women and Mi- 
nority Issues is currently functioning and it 
is hoped that its activities will facilitate 
greater involvement of women and minor- 
ities in leadership positions in the society 
in the future. Even more stark than the 
skewed sex ratio of ASM members is the 
obvious lack of visible minority members 
in ASM. We do not have figures on minority 
membership in ASM, but it is minimal as 
judged by those attending annual meetings, 
where the participants are predominantly 
white. Minority membership in ASM may 
reflect low numbers of minorities in mam- 
malogy in the U.S. This immediately sug- 
gests the urgency of outreach by ASM to 
attract minority students to the study of bi- 
ology. 

Geographic distribution. —Throughout its 
history, ASM has attracted mammalogists 


from all states of the U.S. as well as from 
Canada and Mexico. Nearly all states typ- 
ically are represented at each annual meet- 
ing. Representation of each state and Can- 
ada and Mexico was determined by noting 
the home region of the first author of each 
paper at annual meetings (Fig. 3, Table 2). 
Canadians have participated in notable 
numbers since the founding of the society. 
During its 70-year history, 10 states, listed 
in decreasing order, have accounted for half 
of the participants at annual meetings: Cal- 
ifornia, New York, Texas, Michigan, Dis- 
trict of Columbia, Kansas, Massachusetts, 
Illinois, Florida, and New Mexico. This re- 
flects the strong foundation of the society 
in mammalian systematics collections, as 
these 10 states also contain the 10 largest 
collections in North America. It was not 
until the 21st meeting in 1939 that an an- 
nual meeting was held at an institution not 
dominated by a large museum, and, during 
the first 5O meetings, more than two-thirds 


166 GILL AND WOZENCRAFT 


ASM Annual Meetings 


Home state of First Authors 


No. First Authors 


CI] 4 to 46 
FA 46 to 115 
8 115 to 640 


Fic. 3.— Map of the distribution in the U.S. of first authors of papers given at ASM annual meetings, 


1920-1990. 


of the meetings were held in association with 
large systematics collections. Since the mid- 
1960s this trend is less noticeable, with a 
gradual shift away from ““museum”’ insti- 
tutions to more general academic settings. 
When started, the society was principally an 
East Coast organization, with only one of 
the first 20 meetings held west of the Ap- 
palachians. By the third decade, states from 
the Far West and Midwest were having a 
much greater influence at society meetings. 
The states with the fewest representatives 
at annual meetings are Alabama, Arkansas, 
Delaware, Idaho, New Hampshire, Rhode 
Island, West Virginia, Nevada, Hawaii, and 
Maine. Meetings have been rotated 
throughout the United States for many years, 
with only two areas, the Northern Great 


Plains states (no meetings) and the southern 
U.S. (seven states have not hosted meetings) 
being under-represented. A map of the states 
that have hosted annual meetings closely 
resembles the map of the home state of first 
authors (Fig. 3), with the states heavily rep- 
resented there all having hosted meetings. 
Geographic rotation of the meetings con- 
tributes significantly to participation by 
graduate students, who may find it difficult 
to attend distant meetings. 

Topical and taxonomic emphasis of pa- 
pers at annual meetings. —The 51st annual 
meeting of the society at the University of 
British Columbia in 1971 established the 
format for annual meetings that has been 
used since: organized around topics, with a 
plenary session and concurrent sessions 


COMMITTEES AND MEETINGS 167 


Topical emphasis of papers at 
ASM annual meetings, by decade 


ee] Genetics : 8 Systematics 
Ree Paleontology Es Morphology 


100% 


80% 


60% 


40% 


20% 


0% 
1930 


1940 


1950 


QM Conservation Peay Techniques 
EEE] Ecology 


1960 


ZZ Miscellaneou 
canal] Behavior 


1970 1980 1990 


Fic. 4.—Topical emphasis of papers given at ASM annual meetings, by decade, 1920-1990 (key: 


1930 represents papers from 1920-1929, etc.). 


throughout the meeting. The topical em- 
phasis of papers at ASM annual meetings 
was examined, based on the titles of papers 
in the program of the annual meetings. Pa- 
pers were placed in one of nine categories: 
genetics (including all types of biochemical 
analysis); systematics (including evolution 
and geographic variation); conservation 
(only those papers specifically identified as 
dealing with conservation); techniques; pa- 
leontology (papers dealing with fossil taxa); 
morphology (including reproductive biol- 
ogy, physiology, and anatomy); ecology (in- 
cluding community and population); be- 
havior; and a catch-all field, miscellaneous, 
for all papers that could not be assigned 
clearly to one of the above categories or that 
cut across several topics. 

Several trends can be seen from changes 
in the relative representation of these cat- 
egories during the last 70 years (Fig. 4). Ear- 
ly in our society’s history, broadly based 
papers that covered a variety of topics were 


TABLE 2.— Ranked sequences by largest num- 
bers of presentations of the 10 political units listed 
as address of first authors on papers and posters 
listed in the program at ASM annual meetings 
1920-1990. Political units included were U.S. 
states and the District of Columbia (U.S. postal 
zip code abberviations), Canada (CD), Central 
America, and Mexico (the latter two not yet in 
top 10). 


Meeting numbers 
(number of political units 
represented by first authors) 


1-10 11-20 21-30 31-40 41-50 51-60 61-70 
Rank (11) (26) (35) (39) (49) (50) (54) 


DG NY NY CA CA CA” CA 
NY DC MI MI NY MI TX 
PA MI CA NY MI NY _ KS 
Cl ‘CA De. IL 6 CD NY 
MA MA TX CD CD_ KS_— FL 
MD PA CD UT TX FL MN 
CD MD PA €O CO TIL CD 
MI CD IL AZ LA SC MA 
KS IN KS DC SC MN MI 
VT WA MD WI NM WA _ PA 


COMmANANINAMNHRWN 


—_ 


168 GILL AND WOZENCRAFT 


Taxonomic emphasis of papers at 
ASM annual meetings, by decade 


S| Rodents LIZZ \nsectivores Carnivores a Primates EEE} Ungulates 
Marine Es Marsupials — Other Bats ees] Mixed 


100% 


80% 


60% 


40% 


20% F- 


0% 
1930 1940 1950 


epee, 


1960 1970 1980 1990 


Fic. 5.—Taxonomic emphasis of papers given at ASM annual meetings, decade, 1920-1990 (key: 


1930 represents papers from 1920-29, etc.). 


a significant component of the meetings 
(miscellaneous category). As the field has 
become more specialized, broadly based pa- 
pers have decreased and are now only a small 
part of the papers presented at annual meet- 
ings. This situation is typical of the in- 
creased specialization in society, in general, 
and of teaching and research in academic 
institutions, in particular. 

One of the dominant foci for the estab- 
lishment of ASM, which was reflected in 
titles of papers, was concern for the conser- 
vation of mammals. The trend, as illus- 
trated here, reflects a decrease in titles spe- 
cifically identifiable as conservation issues; 
however, many of the papers in other cat- 
egories have implications to the field of con- 
servation biology and could appropriately 
be presented at conservation meetings. 
Morphology and physiology have always 
been a major component of annual meet- 
ings; within this category there is a general 


trend of a decreasing number of anatomy 
and an increasing number of physiology pa- 
pers. It appears that most papers on anat- 
omy are now presented within the broader 
framework of evolutionary or systematic 
theory. Genetics, a topic present since the 
earliest days of the society, did not become 
a significant part of the meetings until the 
1970s, increasing with the proliferation of 
biochemical and molecular techniques in 
mammalian research. The actual number of 
papers in this category may be much greater, 
as many authors may not have used key- 
words in their titles that would lead to in- 
clusion of their papers in this category; 
therefore, they were not counted in this sur- 
vey. Perhaps the most noticeable trend 
among papers at the meetings is the dom- 
inance of ecology and behavior, which start- 
ed about 1970. Roughly half the papers pre- 
sented in the last 20 years fall into these two 
categories. There are other professional so- 


COMMITTEES AND MEETINGS 169 


cieties that overlap with the ASM in cov- 
ering these two topics, but they have had 
no noticeable effect on the topical makeup 
of papers at the mammal meetings. 

The taxonomic emphasis of papers at 
ASM annual meetings also was examined 
based on titles from annual meeting pro- 
grams. Papers were placed in one of 10 cat- 
egories: Rodentia, Insectivora, Marsupialia 
(sensu lato), Carnivora (excluding pinni- 
peds), Primates, ungulates (Artiodactyla, 
Perissodactyla, Proboscidea), Chiroptera, 
marine mammals (pinnipeds, Cetacea, Sire- 
nia), other (groups not mentioned), and 
mixed (papers not identifiable with a par- 
ticular taxonomic group or those that deal 
with multiple groups). Papers were tallied 
by decade (Fig. 5). 

Several trends can be seen from changes 
in the representation of these categories dur- 
ing the last 70 meetings. Studies of Rodentia 
have increased from less than 20% of the 
total papers to nearly halfin the last decade. 
This category probably would be inflated 
more if those papers that deal with small 
mammal ecology in their titles (included 
here in mixed) were counted. The represen- 
tation of most other taxonomic groups re- 
mained rather consistent over the years, with 
the notable exception of the Chiroptera and 
the Primates. Chiroptera were poorly rep- 
resented in the early meetings, but now are 
a much larger portion, the number of papers 
on bats peaking in the 1960s. The Primates 
were well represented in early meetings but 
are nearly absent from the later half of the 
70-year span. Most poorly represented in 
terms of taxonomic diversity are Insectiv- 
ora, generally only included here in the 
mixed category. Another marked difference 
between the first and last decade is in the 
number of papers that cut across taxonomic 
boundaries or are on topics that are not re- 
stricted to specific taxa (e.g., animal welfare, 
trapping, remote sensing, and the like), 
which were close to 50% in the 1920s and 
are less than 20% in the 1980s. We believe 
that this change is a direct reflection of the 


increasing specialization of mammalogists 
throughout this time period, an effect also 
apparent in the increasing specialization of 
topics at annual meetings. 

During the 70-year period that ASM has 
had annual meetings, several more special- 
ized societies have developed in which some 
ASM members have joint membership. Pri- 
matologists and physical anthropologists 
have a professional history as long, if not 
longer, than ASM and some of the early 
founders of ASM were drawn from this 
group. After the first decade, however, per- 
haps with the initial retirement or with- 
drawal of the founding primate biologists, 
the society has failed to attract this subject 
matter at annual meetings. Three new so- 
cieties have developed in the last two de- 
cades. When bat biologists began to meet 
on an annual basis, there was a marked de- 
crease in the number of papers on the Chi- 
roptera at the annual meetings. Effects of 
the newly formed Society for the Study of 
Mammalian Evolution may have a similar 
effect on papers in systematics and evolu- 
tion. When the Society for Marine Mam- 
malogy was formed, however, it did not 
produce a corresponding drop in the num- 
ber of papers on marine mammals at ASM 
meetings. 

Many profound changes that affect the 
lives and work of mammalogists have oc- 
curred in the world during the first 75 years 
of the ASM. In 1927, Charles Lindbergh’s 
historic flight ushered in the age of com- 
mercial aviation. Now mammalogists fly all 
over the globe to conduct research and to 
interact with colleagues at ITC and other 
international meetings. The U.S. urban 
population exceeded the rural in 1920 and 
another major shift occurred in the 1940s, 
during World War II. Economic changes that 
accompanied these and later population 
shifts, such as new agricultural methods with 
intensive use of fertilizers, irrigation, pes- 
ticides, and herbicides, have had a dramatic 
impact on the habitats and well being of 
mammalian populations. The society con- 


170 GILL AND WOZENCRAFT 


tinues its struggle for the conservation of 
mammals, meeting these new challenges 
through the research of its members, edu- 
cation, and activities to influence legisla- 
tion. 

World War II deeply affected the mem- 
bership of ASM and was the only period 
during which meetings were not held each 
year. Major growth in the membership of 
ASM occurred in the 1950s and 1960s, with 
an increase in graduate students in the 1970s. 
The interests, as well as numbers, of ASM 
members have changed over the decades. 
There has been increased specialization in 
teaching and research, and in papers at an- 
nual meetings, no doubt mirroring the sit- 
uation in society at large. The revolution in 
biotechnology and information systems, al- 
though facilitating research and exchange of 
information, has contributed to this in- 
creased specialization. Some events, how- 
ever, appear cyclic: the teaching of evolu- 
tion was banned in Tennessee and the 
“monkey trial’’ was held in 1925. Now there 
is a renewed onslaught against the teaching 
of evolution, to which the society has re- 
sponded. 

Over the past seven and a half decades 
the society has continued to grow and 
change. An enduring curiosity about and 
concern for mammals, a determination to 
conserve natural habitats, and the plants and 
animals that live there, and a continuing 
enjoyment of the work itself, of field biol- 


ogy, and of the fascinating mammals we 
work with, sustains the fundamental spirit 
and camaraderie of the ASM. 


Literature Cited 


ANONYMous. 1992. Nominations for the Merriam 
Award. Journal of Mammalogy, 73:951-952. 

HOFFMEISTER, D.N. 1969. The first fifty years of the 
American Society of Mammalogists. Journal of 
Mammalogy, 50:794-802. 

HOFFMEISTER, D. N., AND K. STERLING. 1994. Origin. 
Pp. 1-21, in Seventy-five years of mammalogy (1919- 
1994) (E. C. Birney and J. R. Choate, eds.). Special 
Publication, The American Society of Mammalo- 
gists, 11:1-433. 

Honackl, J. H., K. E. KINMAN, AND J. W. KOEpPPL 
(EDs.). 1982. Mammal species of the world: a tax- 
onomic and geographic reference. Allen Press, In- 
corporated and The Association of Systematics Col- 
lections, Lawrence, Kansas, 694 pp. 

KIRKLAND, G. L., JR., AND H. D. SmitH. 1994. Mem- 
bership and finance. Pp. 171-178, in Seventy-five 
years of mammalogy (1919-1994) (E. C. Birney and 
J. R. Choate, eds.). Special Publication, The Amer- 
ican Society of Mammalogists, 1 1:1—433. 

LAYNE, J. N., AND R.S. HOFFMANN. 1994. Presidents. 
Pp. 22-70, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1—433. 

Verts, B. J., AND E. C. Birney. 1994. Publications. 
Pp. 139-154, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1-433. 

WILson, D. E., AND D. M. REEDER. 1993. Mammal 
species of the world: a taxonomic and geographic 
reference. 2nd ed. Smithsonian Institution Press, Blue 
Ridge Summit, Pennsylvania, 1,206 pp. 


MEMBERSHIP AND FINANCE 


GORDON L. KIRKLAND, JR. AND H. DUANE SMITH 


Introduction 


he American Society of Mammalogists 
has a deserved reputation as one of 
the most fiscally conservative and finan- 
cially successful scientific societies in North 
America. The society’s current dues of $30 
are among the lowest of major professional 
societies in biology. This reflects in large 
measure the substantial contribution made 
each year to the general operating fund by 
the society’s Reserve Fund, which is man- 
aged by the society’s three trustees. Ap- 
proximately one-fifth to one-quarter of each 
year’s general operating budget comes from 
income earned by the Reserve Fund, which 
had a net value in excess of $1,000,000 on 
1 June 1992. Funds transferred to the gen- 
eral operating account represent income de- 
rived principally from the investment of life 
membership payments and special be- 
quests. As we review the membership and 
financial history of the American Society of 
Mammalogists during its first 75 years, we 
will document and salute the foresight of 
the founding members in terms of estab- 
lishing the firm financial base upon which 
the society continues to operate. 


Wd 


TIME 


Membership Classes 


The American Society of Mammalogists 
has five classes of membership: active, life, 
patron, emeritus, and honorary. Active 
members pay annual dues and receive the 
Journal of Mammalogy and other corre- 
spondence, such as the ‘‘Call for Papers” 
and program of the annual meeting, from 
the society. 

An individual may become a life member 
by making a payment equal to 25 times the 
current annual dues. This may be a single 
payment or may be made in four equal an- 
nual installments. The dues structure for life 
memberships has remained unchanged since 
the founding of the society in 1919, at which 
time annual dues were $3.00 and life mem- 
berships were $75.00. Life members receive 
the Journal of Mammalogy for life or until 
they no longer wish to do so. Life members 
currently comprise 17% of total ASM mem- 
bership. By contrast, in 1920 only 2.5% of 
ASM members were life members. During 
the succeeding four decades, the percentage 
of life members fluctuated slightly but had 


172 KIRKLAND AND SMITH 


TABLE 1.— Pattern of growth in life member- 
ships. 


Total Life % life 
Year membership members members 
1920 443 11 2-5 
1930 1,005 62 6.2 
1940 898 51 Se) 
1950 1,232 49 4.0 
1960 1,765 115 6.5 
1970 BE315 342 10.3 
1980 3,862 530 13.7 
1990 3,661 611 16.7 


risen only to 6.5% in 1960. Since then, the 
proportion of life members has increased 
by about 3% per decade (Table 1). This in- 
crease may reflect the desire of many mem- 
bers to save money by becoming life mem- 
bers just before dues increases, which have 
been more frequent during the past three 
decades (Table 2). 

Patron members are individuals who 
make a $1,000 payment to the society with- 
in a one-year period. Such individuals are 
entitled to receive the Journal and all other 
ASM publications for life. Although this 
membership category has existed through- 
out the history of the society, the first patron 
membership was not purchased until 1990. 
Today, patron memberships represent the 
society’s best financial bargain if viewed 
from the perspective that today’s patron 
memberships can be obtained for the same 
payment of $1,000 as in 1919. If the cost 
of patron memberships had kept pace with 
increases in dues over the past 75 years, 
patron memberships would cost $10,000. 

The emeritus membership category was 
established in 1951. Individuals who have 
been active members for at least 25 years 
may request emeritus membership status. 
These individuals pay no dues and do not 
receive the Journal of Mammalogy, but they 
do continue to receive other ASM corre- 
spondence. Emeritus members also do not 
have voting rights at annual meetings. 

The highest honor bestowed by the so- 
ciety is honorary membership, which is con- 
ferred in recognition of distinguished ser- 


TABLE 2.— Annual dues and subscription rate 
changes for American Society of Mammalogists. 


Year Dues Subscriptions 
1919 $ 3.00 $ 3.00 
1947 4.00 4.00 
1952 4.00 6.00 
1959 4.00 7.00 
1967 4.00 9.00 
1968 5.00 9.00 
1969 7.00 9.00 
1971 7.00 11.00 
1974 7.00 15.00 
1975 12.00 17.00 
1977 16.00 17.00 
1978 16.00 23.00 
1986 20.00 28.00 
1988 23.00 33.00 
1993 30.00 45.00 


vice to mammalogy. Fifty-eight individuals 
have been thus honored. These individuals 
are chronicled in this volume by Taylor and 
Schlitter (1994). 

The American Society of Mammalogists 
has always had one of the highest benefits 
to dues ratios among professional societies. 
Annual dues were $3 in 1919 and have in- 
creased to only $30 today. Historically, the 
society has been reluctant to raise dues, and 
it has been able to maintain its modest dues 
because many of the services that other so- 
cieties pay for are provided to the ASM on 
a volunteer basis by its members. Thus, 
ASM dues largely go to pay the costs of 
publishing the Journal of Mammalogy. As 
a consequence, increases in dues over the 
years (Table 2) have largely mirrored in- 
creases in the costs of publishing the Journal 
(Table 3). 

The philosophy that has supported reten- 
tion of lower dues also has been applied to 
subscription rates. During the society’s first 
33 years subscription rates were the same 
as member dues, but in 1952 the subscrip- 
tion rate was increased to $6 per year while 
dues remained at $4 (Table 2). There has 
been a differential between dues and sub- 
scription rates since that time. With pro- 
ceeds from the Reserve Fund subsidizing 
society services to members, the subscrip- 


MEMBERSHIP AND FINANCE ies) 


tion rate is still comparable in value to the 
subsidized membership dues. Subscription 
rates have increased since 1967, when they 
were $9 per year, to the current $45 per year 
(Table 2). 


Membership History 


The American Society of Mammalogists 
had 252 charter members—1.e., individuals 
who joined the society in 1919. The first 
member, based on payment of dues, was 
Dwight D. Stone (3 April 1919). Ernest 
Thompson Seton was the first life member 
and seventh member overall. The first 
woman member was Viola S. Schantz, who 
served as the society’s treasurer from 1930 
to 1953. Annie M. Alexander was the so- 
ciety’s first woman life member. The last 
surviving charter member was Vasco M. 
Tanner, who died in 1989, 70 years after 
joining the society. 

The society grew rapidly during its early 
years. Membership more than doubled 
within the first three years to 527 in 1921 
(Fig. 1). The society reached the 1,000- 
member level (1,005) in 1930. Membership 
exceeded 1,000 members (1,017) the fol- 
lowing year, but the Depression had a sig- 
nificant negative impact on the society’s 
membership, which dropped to 931 in 1932 
and reached a low of 770 in 1935 (a decrease 
of 24% in four years). Membership re- 
mained below 1,000 throughout the re- 
mainder of the Depression and during the 
war years (Fig. 1). Numerous ASM mem- 
bers who served on active duty in World 
War II were carried on the society’s books 
as inactive members during the war years. 
All such members were required to reacti- 
vate their memberships by 31 January 1948 
or be dropped from membership. It was not 
until 1948 that membership again exceeded 
1,000 (1,071). Membership grew steadily 
during the next 15 years, finally surpassing 
2,000 in 1963, but it took only five more 
years to reach 3,000 (3,194 in 1968). This 
rapid increase in membership corresponded 


4500 


4000 


3500 


3000 


2500 


2000 


1500 


NUMBER OF MEMBERS 


1000 


500 


1920 1930 1940 1950 1960 1970 1980 1990 
YEAR 


Fic. 1.—Growth in the membership of the 
American Society of Mammalogists from 1919 
to: 1992. 


to the dramatic expansion of graduate train- 
ing in the 1960s and the establishment of 
many new programs in mammalogy by ASM 
members who received their Ph.D.s during 
the 1950s and 1960s. Although member- 
ship exceeded 3,900 in 1975, 1976, and 
1979, it has yet to reach 4,000. During the 
past decade, membership has stabilized at 
3,600-3,700 (Fig. 1). 


International Membership 


Despite its name, the American Society 
of Mammalogists is an international sci- 
entific organization with a strong contingent 
of members who reside outside the United 
States. The international nature of the so- 
ciety’s membership dates from its earliest 
years. For example, the first List of Mem- 
bers published in the Journal of Mammal- 
ogy (1922, vol. 3, number 3) contained the 
names of 50 members who resided in 19 
countries outside the United States and its 
territories. These individuals represented 9% 
of the society’s 555 members in 1922. As 
of October 1992 the society’s 718 non-U.S. 
members comprised 19% of total member- 
ship. These non-U.S. members resided in 


174 KIRKLAND AND SMITH 


70 countries. The society’s strong interna- 
tional focus is also reflected in the individ- 
uals elected to honorary membership in the 
ASM during its first 75 years. Of 58 indi- 
viduals thus honored, 17 (29.3%) were non- 
U.S. mammalogists, including Prof. E. L. 
Trouessart, Museum d’Histoire Naturelle 
in Paris, who was the second individual 
elected to honorary membership in 1921. 

The society’s International Relations 
Committee, which was established in 1960, 
has endeavored during the past decade to 
coordinate activities with mammal socie- 
ties in other countries. These efforts have 
resulted in four joint meetings between the 
ASM and mammal societies in Australia 
(1985), Mexico (1987), China (1988), and 
Argentina (1990). These meetings have pro- 
vided opportunities for many ASM mem- 
bers in those countries to participate in an 
ASM activity for the first time. 


Corresponding Secretary, 
Treasurer, and 
Secretary- Treasurer 


From 1919 to 1957 the membership of 
the society was served by the separate offices 
of Corresponding Secretary and Treasurer. 
Eleven individuals held the office of Cor- 
responding Secretary: H. H. T. Jackson 
(1919-1925), A. Brazier Howell (1925- 
1931), Francis Harper (1931-1932), Robert 
T. Hatt (1932-1935), William H. Burt 
(1935-1938), William B. Davis (1938- 
1941), Emmet T. Hooper (1941-1947), 
Donald F. Hoffmeister (1947-1952), Keith 
R. Kelson (1952-1954), George C. Rinker 
(1954-1956), and Bryan P. Glass (1956- 
1957). Six of these individuals (Jackson, 
Howell, Burt, Davis, Hooper, and Hoff- 
meister) subsequently served as presidents 
of ASM. The tenures of treasurers were lon- 
ger and only five individuals held this po- 
sition: Walter P. Taylor (1919-1920), J.W. 
Gidley (1920-1921), Arthur J. Poole (1921- 
1930), Viola S. Schantz (1930-1953), and 
Caroline A. Heppenstall (1953-1957). Wal- 


ter P. Taylor subsequently served the so- 
ciety as its president. 

In 1957 the offices of Corresponding Sec- 
retary and Treasurer were combined into a 
single office of Secretary-Treasurer in order 
to conduct the business affairs of the ex- 
panding society more efficiently. To date, 
four individuals have held this office: Bryan 
P. Glass (1957-1977), Duane A. Schlitter 
(1977-1980), Gordon L. Kirkland, Jr. 
(1980-1986), and H. Duane Smith (1986- 
present). 

The Secretary-Treasurer is the chief ad- 
ministrative officer of the ASM and is re- 
sponsible for the society’s day-to-day op- 
erations. Duties include managing the 
society’s general operating account and the 
accounts for Mammalian Species and Spe- 
cial Publications, maintaining membership 
records, corresponding with ASM members 
and others seeking information or assis- 
tance, printing and mailing the “Call for 
Papers” and the program for the annual 
meeting, assisting with preparation of the 
annual budget, arranging for the annual au- 
dit for the society’s financial records, send- 
ing mailing labels for the Journal of Mam- 
malogy and Mammalian Species to the 
printer, processing orders for Special Pub- 
lications, and distributing copies of the res- 
olutions passed at the annual meetings. 


Reserve Fund 


The founders of the society showed ex- 
ceptional foresight in establishing a mech- 
anism to invest life and patron membership 
dues and gifts to the society in a permanent 
fund, some of the income from which was 
to be used to subsidize various functions of 
the society. This provision was incorporat- 
ed in the society’s first By-laws and Rules 
adopted in April 1919. The year 1922 
marked the first major initiative to develop 
the permanent fund, namely the J. A. Allen 
Memorial Fund. The initial goal of that fund 
was $10,000. The campaign to achieve that 
goal was supervised by the J. A. Allen Me- 


MEMBERSHIP AND FINANCE L75 


TABLE 3.—Growth of the Reserve Fund and 
contributions to the general operating budget (val- 
ues rounded to nearest whole dollar). 


Reserve Fund Reserve Fund 
contribution contribution as 


Value of to annual % of 
Year Reserve Fund budget Reserve Fund 
1930 $ 11,070 $ 500 4.5% 
1940 $ 23,235 $ 500 2.2% 
1950 $93,517 $: 1,500 2.8% 
1960 $124,747 S 3/51 3.0% 
1970 $208,603 $11,506 5.5% 
1980 $475,370 $20,234 4.3% 
1990 $809,376 $39,000 4.8% 


morial Committee. The first contributions 
to the fund, by W. D. Matthew, T. G. Pear- 
son, J. T. Nichols, B. S. Bowdish, and C. 
W. Richmond, totalled $105.00. The Allen 
Fund grew rapidly. The value of the fund 
was $6,335.42 in 1924, $7,606.12 in 1925, 
$8,525.24 in 1926, and $9,156.01 in 1927. 
With the fund at $9,975 in 1928, a special 
collection was taken among members at- 
tending the annual meeting to raise $25, 
with the following contributing: A. Brazier 
Howell, H. H. Lane, Carl Hartman, C. C. 
Adams, Lee R. Dice, M. W. Lyon, Jr., R. 
T. Hatt, H. C. Raven, and A. W. Leighton. 
The fund officially reached its goal on 9 April 
1929 when the fund totalled $10,465.27. 
Two hundred and seventy-three contribu- 
tors had given $8,428.78, with the differ- 
ence of $1,848.90 representing interest and 
bond coupons. Upon achieving its goal, the 
Allen Memorial Committee was dissolved 
and the funds subsequently were managed 
by the society’s trustees. 

In 1923, the by-laws were amended to 
provide for three trustees to administer the 
permanent fund. Trustees are elected by the 
Board of Directors and serve three-year, ro- 
tating terms. The first three trustees were 
Henry Bannon, Childs Frick, and Charles 
Sheldon. Thanks to the efforts of these and 
subsequent trustees, the Reserve Fund has 
experienced sustained growth during the past 
70 years. In general, the value of the Reserve 
Fund has doubled each decade (Table 3). 


TABLE 4.— Relationship between funds trans- 
ferred by the Reserve Fund to support the general 
operating account and funds transferred to the 
Reserve Fund for investment. 


Mean annual 


transfer to Mean annual 


general transfer to 

operating Reserve Fund Ratio of A 
Decade account (A) (B) to B 
1930s $ 470.00 $ 549.10 0.86 
1940s $ 670.00 $ 747.10 0.90 
1950s $ 2,300.60 $1,720.90 1.34 
1960s $ 6,489.10 $2,278.20 2.85 
1970s $16,865.90 $5,386.00 3.13 
1980s $27,553.50 $4,790.60 3.75 


As the value of the Reserve Fund has grown, 
the amount of money transferred to the gen- 
eral operating account has increased; how- 
ever, when figured as a percentage of the net 
value of the Reserve Fund, the amount 
transferred annually has remained relative- 
ly constant, fluctuating between 2.2 and 5.5% 
(Table 3). 

Each year, funds are transferred between 
the Reserve Fund and the society’s general 
operating account. Money transferred to the 
fund accrues principally from life member- 
ship payments. The average amount trans- 
ferred annually to the Reserve Fund in- 
creased from the 1930s through the 1970s 
but decreased slightly in the 1980s (Table 
4). During the 1930s and 1940s the amount 
transferred annually to the Reserve Fund 
exceeded the amount received annually from 
the fund to support operations of the soci- 
ety, specifically publication of the Journal 
of Mammalogy; however, since then the 
amount transferred to the general operating 
account has exceeded the amount annually 
transferred to the Reserve Fund (Table 4). 
Throughout the past 60 years, the ratio of 
funds received from the Reserve Fund com- 
pared to money transferred from the general 
operating account to the Reserve Fund has 
increased steadily, so that in the 1980s more 
than five times as much was received from 
the Reserve Fund as was transferred to it 
(Table 4). 


176 KIRKLAND AND SMITH 


TABLE 5.—Growth of the Future Mammalo- 
gists Fund, 1985-1992. 


Year Balance Year Balance 

1985 $ 4,938 1989 $ 69,090 
1986 34,720 1990 71,644 
1987 51,468 199] 92,204 
1988 60,637 1992 128,000 


Davis (1969) prepared a comprehensive 
history of the Reserve Fund on the occasion 
of the society’s 50th anniversary. He ex- 
amined the growth of the “Permanent Fund” 
on a decade by decade basis and provided 
a more detailed analysis of the finances of 
the fund, including the composition of the 
fund’s portfolio by decade and strategies for 
investing the society’s funds in light of the 
prevailing economic climate. 

The American Society of Mammalogists 
has always been concerned about the sci- 
ence of mammalogy and about providing 
opportunities for its members. In 1985, this 
concern led the society to establish the Fu- 
ture Mammalogists Fund with the goal to 
raise a minimum of $100,000 for invest- 
ment. Interest from this investment will 
support young mammalogists who are just 
getting started professionally. The fund- 
raising efforts of the members and wise in- 
vestments by the trustees have been very 
successful. Reference to Table 5 shows that 
the fund began slowly with a balance of 
$4,938 in 1985, but has grown rapidly in 
recent years, surpassing the original goal be- 
tween 1991 and 1992. The 1992 balance, 
$128,000, constituted 13% of the society’s 
Reserve Fund. The proceeds are now being 
used to support young mammalogists from 
around the world. 


ASM Budgets 


Traditionally, the bulk of the society’s an- 
nual operating budget has been devoted to 
publishing the Journal of Mammalogy. The 
budgeted cost of publishing the Journal (in- 
cluding production costs, editorial expenses 


TABLE 6.— Comparison of the annual budgets 
of the American Society of Mammalogists and 
the costs of publishing the Journal of Mammalogy 
by decade. 


Mean annual Cost of 
cost of Journal as 
Mean annual producing % of 

Decade budget Journal* budget 
1920s $ 2,741.50 $ 2,440.00 89.0% 
1930s 3,080.56 2,155.56 89.4% 
1940s 3,600.00 3,240.00 90.0% 
1950s 11,817.86 9,800.00 83.0% 
1960s 23,108.70 18,765.70 81.2% 
1970s 76,306.80 65,222.40 85.5% 
1980s 150,957.30 116,443.90 77.1% 
1990s 161,693.33 119,133.33 73.7% 


* Includes costs of printing, distribution, editorial 
expenses, editorial honoraria, preparation of the index 
and Recent Literature in Mammalogy. 


and honoraria, and costs incurred by the 
bibliography and index committees) has ris- 
en from $1,600 in 1920 to $122,000 in 1992 
(a 7,525% increase). During that period, dues 
increased from $3.00 to $23.00 (a 667% in- 
crease). The cost of publishing the Journal 
averaged about 90% of the society’s annual 
budgets during its first three decades (Table 
6). In the 1950s annual budgets increased 
substantially (228%) compared to the pre- 
ceding decade, whereas the cost of publish- 
ing the Journal increased 202% (Table 6). 
This difference reflected increased costs of 
running the society’s executive office and a 
broader scope of society expenditures, in- 
cluding funds for graduate student hono- 
raria and dues to afhliate societies (e.g., 
membership in AIBS). As a consequence, 
in the 1950s expenditures for publishing the 
Journal averaged 83% of the annual budget. 
This percentage remained about the same 
in the 1960s (Table 6); however, the socie- 
ty’s budgets in the 1960s averaged about 
twice those of the preceding decade, as did 
costs of publishing the Journal of Mam- 
malogy (Table 6). In the 1970s, both av- 
erage annual budgets (230% increase) and 
costs of publishing the Journal (248% in- 
crease) more than doubled. As a conse- 
quence, the percentage of the annual budget 


MEMBERSHIP AND FINANCE 177 


devoted to publishing the Journal during 
the 1970s increased to 85.5%. 

The first budget in excess of $100,000 was 
approved for 1977. There was less than a 
two-fold increase in budgets and costs of 
publishing the Journal in the 1980s with the 
percentage contribution of publishing the 
Journal declining to 77% (Table 6). During 
the first three years of the 1990s, budgets 
have increased little over those for the 1980s 
(Table 6). 


Summary 


Members of the American Society of 
Mammalogists can take singular pride in the 
financial history of their society. Today’s 
members benefit from the financial acumen 
and foresight of the society’s founding 
members. In terms of its finances, the ASM 
is a model for other scientific and profes- 
sional societies, who in the past have con- 
tacted the society’s executive office for ad- 
vice on financial matters. Members of the 
ASM not only belong to the oldest and larg- 


est scientific society devoted to the study of 
mammals, they are members of a society 
whose astute and prudent financial man- 
agement over the years has made it one of 
the “best buys” among professional soci- 
eties. 


Acknowledgments 


We thank staff members of the Smithsonian 
Archives, especially W. Cox, for facilitating ac- 
cess to the society’s historical files. We also thank 
W. C. Wozencraft for his assistance in locating 
ASM archival materials. 


Literature Cited 


Davis, W. D. 1969. The American Society of Mam- 
malogists permanent fund: a special report of the 
trustees. Pp. 33-40, in The American Society of 
Mammalogists 50th anniversary celebration. Pro- 
cessed by the American Museum of Natural History, 
New York, New York. 44 pp. 

TAYLOR, J. M., AND D. A. SCHLITTER. 1994. Award- 
ees. Pp. 71-109 in Seventy-five years of Mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 11: 1-433. 


PART II 


INTELLECTUAL DEVELOPMENT OF 
THE SCIENCE OF MAMMALOGY 


TAXONOMY 


MArkK D. ENGSTROM, JERRY R. CHOATE, AND HUGH H. GENOWAyYS 


Introduction 


i) Rares aden has been termed the “theory 

and practice of classifying organisms” 
(Mayr and Ashlock, 1991:2), whereas sys- 
tematics is the broader study of the history 
and diversity of life. In practice, in distinc- 
tion between these disciplines is often 
blurred. In this review, we focus on the role 
of taxonomy and taxonomists in the de- 
velopment of the discipline of mammalogy 
in North America over the past 75 years, 
although we occasionally will slip into 
broader discussions of systematics where it 
has influenced the philosophical underpin- 
nings of taxonomy. For purposes of discus- 
sion and in practice, taxonomy also can be 
divided conveniently into two levels: mi- 
crotaxonomy—the methods and principles 
by which species are recognized and delim- 
ited; and macrotaxonomy—the methods and 
principles by which recognized kinds of or- 
ganisms are Classified (Mayr, 1982). 


Historical Perspective 


Development of mammalian taxonomy 
in North American was a natural conse- 
quence of exploration of the continent. Many 
of the early descriptions of new mammals 
from the East were made by Linnaeus and 


179 


his contemporaries based on specimens re- 
turned to Europe from the American col- 
onies. Some of the most important taxo- 
nomic contributions by American authors 
were taxonomic catalogues (reviewed by 
Hoffmeister and Sterling, 1994). However, 
American naturalists were largely respon- 
sible for taxonomic investigations resulting 
from exploration of the West, beginning with 
the Lewis and Clark expedition and cul- 
minating with the expeditions of Major Ste- 
phen Long, Zebulon Pike, Thomas Say, 
Maximilian Prince of Wied-Neuweid, John 
C. Fremont, and numerous others. The tax- 
onomic products of those expeditions in- 
cluded such monumental catalogues as 
Baird’s (1857) report on the mammals of 
North America, Coues and Allen’s (1877) 
review of North American rodents, Elliot’s 
(1904) checklist of mammals of North 
America and the West Indies, and Mearns’ 
(1907) Mammals of the Mexican boundary 
of the United States. 

Most of the taxonomic collections made 
by naturalists of the 19th Century were re- 
turned to museums in the East, notably the 
Charleston Museum, Peale’s Museum in 
Philadelphia, Museum of Comparative Zo- 
ology at Harvard College, American Mu- 
seum of Natural History in New York, 


180 


United States National Museum, Chicago 
Academy of Sciences, and Field Museum of 
Natural History in Chicago. Taxonomists 
associated with those museums were among 
the leaders in development of the science 
of mammalogy, as were naturalists associ- 
ated with the most influential universities 
of the day: Harvard, Yale, Michigan, Cor- 
nell, California, and others. California was 
especially important because it was there 
that Joseph Grinnell had begun a dynasty 
of mammalogists that persists even today 
(Jones, 1991; Whitaker, 1994). 

However, the most productive group of 
North American mammalogists of the day 
by far were those associated with the Bureau 
of the Biological Survey, the progenitor of 
the U.S. Fish and Wildlife Service (de- 
scribed by Hoffmeister and Sterling, 1994). 
Authors of monographic revisions pub- 
lished by bureau employees in its North 
American Fauna series read like a who’s 
who of North American mammalogy in the 
years preceding the origin of the ASM: 
Vernon Bailey; Edward A. Goldman; Ned 
Hollister; A. Brazier Howell; Arthur H. 
Howell; Hartley H. T. Jackson; C. Hart 
Merriam; E. W. Nelson; Wilfred H. Osgood; 
Edward A. Prebel. Of the monographs pub- 
lished before 1919, Osgood’s (1909) revi- 
sion of the genus Peromyscus arguably has 
stood the test of time better, and has stim- 
ulated more taxonomic studies, than any 
other. 

The ASM came into being at a time when 
much of the work of North American mam- 
malogists was directed at understanding the 
diversity of mammals on the continent. 
Most of the founding fathers of ASM were 
thus taxonomists, and taxonomists subse- 
quently have had a greater influence on the 
society than have mammalogists of any oth- 
er subdiscipline. 

For North American taxonomists from 
the mid-1800s until about the turn of the 
century, the predominant species concept 
was typological—species were held to be 
nearly fixed entities that varied about a fi- 
nite number of types. By this concept, root- 
ed in the classical philosophy of European 


ENGSTROM ET AL. 


systematics, species were delimited subjec- 
tively based on relative degree of morpho- 
logical difference and consisted of aggrega- 
tions of individuals that agreed with the 
author’s diagnosis. There was little appre- 
ciation of the distinction between variation 
due to gender, age, or individual and geo- 
graphic differentiation. During this period, 
most new forms were described as species 
despite the fact that the category of subspe- 
cies was already in common use in orni- 
thology. Designation of morphologically 
distinct forms as species was understand- 
able in that most early collections consisted 
of specimens from widely separated local- 
ities and the concept of geographic variation 
was poorly understood. The extensive col- 
lections amassed under the auspices of the 
Bureau of the Biological Survey (among 
others), however, eventually demonstrated 
the pervasiveness of geographic variation 
and intergradation among many nominal 
“‘species.’’ Gradually, the practice of sorting 
apparently distinctive specimens into spe- 
cies was replaced by a broader view of spe- 
cies aS interrelated groups of populations 
united by reproductive ties. Taxoncmists 
shifted from describing and classifying ob- 
jects (specimens) to attempting to describe 
the living diversity of populations that those 
specimens represented. Perhaps the most 
notable early example of this shift was Os- 
good’s (1909) revision of Peromyscus. Os- 
good reduced the number of recognized spe- 
cies of deer mice from 130 to 43 (see review 
by Carleton, 1989) and, in one instance, 
combined 28 nominal species into the taxon 
he recognized as Peromyscus maniculatus. 
Many of the former species names were re- 
tained as formal subspecies, and the practice 
of recognizing polytypic species (consisting 
of two or more subspecies), in use since the 
late 1800s, thus became entrenched. 
Change from a typological or strict mor- 
phological concept of species to recognition 
of polytypic species composed of morpho- 
logically distinct, intergrading subspecies 
was gradual and was not universally ac- 
cepted by the time of formation of the ASM 
in 1919. For example, the first issue of the 


TAXONOMY 181 


Journal of Mammalogy contains a staunch 
defense of a morphological species concept 
by Merriam (1919). He stated (p. 7) that 
“the criterion of intergradation is one of the 
most pernicious that has ever been intro- 
duced into the systematic study of animals 
and plants .. .” and, quoting a previous ar- 
ticle in Science (p. 9), “forms which differ 
only slightly should rank as subspecies even 
if known not to intergrade, while forms 
which differ in definite, constant and easily 
recognized characters should rank as species 
even if known to intergrade.”’ This philos- 
ophy, coupled with samples inadequate to 
demonstrate the full range of intra- and in- 
terpopulational variation, led him (Merri- 
am, 1918) to recognize two genera and 78 
species of brown bears, all now considered 
to represent a single species (Hall, 1984). 
Interestingly, a rejoinder by Taverner (1920: 
126) in the first volume of the Journal of 
Mammalogy advocated the essentials of 
what later would become known as the bi- 
ological species concept: “‘the species is a 
definite entity and its essential quality is its 
genetic isolation.” 

Many of the taxonomy publications of the 
1920s were by employees of the Bureau of 
the Biological Survey; however, the most 
important taxonomic catalogue of the pe- 
riod was by Gerrit S. Miller, Jr. (1924), Cu- 
rator of Mammals at the U.S. National Mu- 
seum, who updated his earlier (Miller, 1912) 
list of North American mammals. Several 
taxonomic revisions were published during 
this decade, most notably those by A. B. 
Howell (1926, 1927), Jackson (1928), Miller 
and Allen (1928), and A. H. Howell (1929). 
Most of the taxonomic publications of the 
period were monographic in extent. 

Taxonomic work in the 1930s was dom- 
inated less than that of the previous decade 
by employees of the Bureau of the Biological 
Survey. An increasing number of mam- 
malogists at academic institutions and at 
museums other than the United States Na- 
tional Museum began to have an impact. 
One of the most important taxonomic re- 
visions of the period was the monograph on 
squirrels by A. H. Howell (1938). Other re- 


visionary studies emanated from the Field 
and American museums of Natural History 
and dealt largely with Latin American 
mammals (e.g., Sanborn, 1937; Tate, 1933). 
During this decade, there was an increasing 
tendency for taxonomic work to be less than 
monographic in extent and to focus on in- 
dividual species rather than genera or higher 
categories (e.g., Nelson and Goldman, 1933). 

Most North American mammalogists 
would agree that the taxonomic highlight of 
the 1940s was Simpson’s (1945) The Prin- 
ciples of Classification and a Classification 
of Mammals. Few generic revisions were 
published during the decade, as an increas- 
ing number of taxonomic studies focused 
on geographic variation within species (e.g., 
Hooper, 1943). 

The 1950s was a watershed decade for 
mammalian taxonomy in North America. 
An important taxonomic catalogue (North 
American Recent Mammals, by Miller and 
Kellogg, 1955) was published early in the 
decade only to be overshadowed by another 
(The Mammals of North America, by Hall 
and Kelson, 1959). Hall and Kelson’s mon- 
umental two-volume work quickly became 
a veritable landmark in mammalogy in that 
it summarized everything then known about 
the distribution and taxonomy of native 
mammals in North America. Much of the 
explosion of taxonomic research (especially 
on relationships within genera and geo- 
graphic variation within species—see dis- 
cussion of subspecies, beyond) was a direct 
result of studies leading to or stimulated by 
publication of this epic monograph. In place 
of faunal studies, numerous taxonomic re- 
visions were published during the 1950s. 
Some of the best known of those revisions 
were by Goldman (1950), Hall (1951), Hoff- 
meister (1951), Hooper (1952), Handley 
(1959), Moore (1959), and Van Gelder 
(1959). The number of studies of variation 
within species continued to climb, that by 
Findley (1955) serving as an example. 

The explosion of taxonomic literature on 
North American mammals that began in 
the 1950s continued in the 1960s. Taxo- 
nomic catalogues published during the de- 


182 ENGSTROM ET AL. 


cade included those of Hershkovitz (1966) 
on living whales, and Anderson and Jones 
(1967) on mammals of the world. Taxo- 
nomic revisions continued to be numerous, 
a few examples being those of Lidicker 
(1960), Packard (1960), Russell (1968a, 
19685), Davis (1968, 1969, 1970), Musser 
(1968), and Lawlor (1969). Increasingly, 
these revisions were of small genera and 
were less than monographic in length—a 
phenomenon possibly resulting in part from 
the increasing difficulty in finding outlets for 
lengthy, monographic manuscripts. 

The 1970s witnessed publication of few 
taxonomic catalogues (one example being 
Varona’s 1974 catalogue of Antillean mam- 
mals), but a large number of both “Mam- 
mals of ...’ books and taxonomic revi- 
sions. A sample of the many taxonomic 
revisions of the period includes those by 
Choate (1970), Findley and Traut (1970), 
Zimmerman (1970), Genoways and Jones 
(1971), Hooper (1972), Pine (1972), Smith 
(1972), Thaeler (1972), Birney (1973), 
Gardner (1973), Genoways (1973), Carle- 
ton (1977), Eger (1977), Hennings and Hoff- 
mann (1977), Yates and Schmidly (1977), 
Hoffmeister and Diersing (1978), Williams 
(1978), Carleton and Eshelman (1979), Haf- 
ner et al. (1979), Silva-Taboada (1979), and 
Williams and Genoways (1979). By the end 
of the decade, new techniques for taxonom- 
ic analysis (Baker and Hafner, 1994; Ho- 
neycutt and Yates, 1994) and changing pri- 
orities at academic institutions and funding 
agencies were beginning to take a toll on 
faunal studies and taxonomic revisions, of- 
ten relegating both to the category of long- 
term, low priority projects. 

The 1980s began with publication of 
Hall’s (1981) long-awaited update of The 
Mammals of North America. As noted by 
Jones (1982:718) in his review of this mon- 
umental taxonomic catalogue, “It is unlike- 
ly that any other American mammalogist 
would have undertaken, or will undertake 
again, such a gigantic task.” Another useful 
catalogue published during the decade was 
Anderson and Jones’ (1984) revised syn- 


opsis of mammals of the world. During the 
previous decade, an enthusiastic cadre of 
young mammalian taxonomists had begun 
developing in Mexico, and the 1980s were 
marked by the beginnings of taxonomic 
products from this group (e.g., Arita and 
Humphrey, 1988; Ceballos and Galindo, 
1984; Ramirez-Pulido et al., 1986). It seems 
likely that a substantially greater percentage 
of the taxonomic papers on Latin American 
mammals will be authored by Latin Amer- 
ican mammalogists in decades to come. Fi- 
nally, the decade was marked by Koop- 
man’s (1984) classification of bats and a 
multitude of taxonomic reviews, many of 
the latter employing genetic techniques or 
focusing on species or species groups. A few 
examples were the studies by Carleton 
(1980), Huckaby (1980), Engstrom and Wil- 
son (1981), George et al. (1981), Patton and 
Smith (1981), Patton et al. (1981), Honey- 
cutt and Williams (1982), George et al. 
(1982), Grifhths (1982), Hafner (1982), 
Rogers and Schmidly (1982), Heaney and 
Timm (1983), Baker (1984), van Zyll de 
Jong (1984), Sullivan (1985), Sullivan et al. 
(1986), Webster and Handley (1986), Baker 
et al. (1988), George (1988), Robbins and 
Sarich (1988), Voss (1988), Baker et al. 
(1989), Carleton and Musser (1989), van 
Zyll de Jong and Kirkland (1989), and Woz- 
encraft (1989a, 1989b). 

The 1990s show promise of a continua- 
tion of the existing emphasis on microtax- 
onomic studies employing modern genetic 
methods plus development of a much great- 
er emphasis than in the past on macrotax- 
onomy. Early examples of the research that 
will characterize the decade include the 
studies by Patton and Smith (1990), Hafner 
(1991), Johnson and George (1991), Rogers 
and Engstrom (1992), and Wall et al. (1992). 


Biological Species Concept 
The empirical demonstration of species 


as natural aggregates of populations delin- 
eated from related species by reproductive 


TAXONOMY 183 


MW Families 1! Genera Species  () Subspecies 


3000 + 
2500 
2000 


1500 + 


500 + 


Hall and Kelson, 1959 Hall, 1981 


Miller and Rehn, 1902 


Miller, 1924 

Fic. 1.—Total number of families, genera, spe- 
cies, and subspecies of North American mam- 
mals recognized as valid in major taxonomic 
summaries of the 20th Century. 


gaps led to formulation of a biological spe- 
cies concept: ““Species are groups of inter- 
breeding natural populations that are re- 
productively isolated from other such 
groups” (Mayr, 1969:26). Viewing species 
as natural, objective entities rather than 
classes of objects had its genesis in the late 
1700s and was accepted by ornithologists 
and ichthyologists by the turn of the cen- 
tury. Mammalogists were more conserva- 
tive, but the concept (with its common rec- 
ognition of polytypic species) had taken hold 
by the 1920s. Coupled with this philosoph- 
ical shift was the ascendancy of the neo- 
Darwinian school of “‘new systematics” 
(Huxley, 1940), led by R. A. Fisher, J. B. S. 
Haldane, S. Wright, T. Dobzhansky, E. 
Mayr, G. G. Simpson, V. Grant, and others 
from the 1930s through the 1960s. This phi- 
losophy reasserted the fundamental impor- 
tance of taxonomy and systematics. With 
its concern for microevolutionary processes 
underlying intraspecific genetic variation 
and the generation of diversity, this school 
focused on issues of population genetics, 
geographic variation, and speciation. Sev- 
eral classic generic revisions of mammals 
were written during this period, with an em- 
phasis on discerning patterns of geographic 
variation and taxonomic limits of species 
rather than on primary descriptions. 


Abandonment of a typological or strict 
morphological concept and recognition of 
geographically variable, polytypic species, 
led to a clarification and simplification of 
the classification of North American mam- 
mals at the species level. In the 35-year pe- 
riod between 1924 and 1959, the 1,441 spe- 
cies of North American mammals admitted 
by Miller (1924) were reduced to 1,003 (Hall 
and Kelson, 1959), despite the description 
of numerous new, valid species (Figs. 1 and 
2). This led Hall and Kelson (1959:vi) to 
remark that “The decrease in number of 
species results from many of the named 
kinds having been reduced from specific to 
subspecific status in the past thirty years. 
Certainly the number of species listed in the 
present work is still too large, many geo- 
graphically adjacent pairs of nominal spe- 
cies will prove to be only subspecies of one 
and the same species when adequate spec- 
imens are studied from geographic areas be- 
tween the known areas of occurrence of the 
two kinds.’ Unfortunately, the descriptive 
efforts of some mammalian taxonomists 
soon were directed to the formal recognition 
of taxa below the level of species, and an 
explosion of new subspecies ensued (see fol- 
lowing section on subspecies). Conversely, 
from 1902 to 1924 the number of recog- 
nized genera and families increased by a 
factor of about one-half, due mostly to a 
less inclusive view of higher taxa; this num- 
ber has remained relatively stable since that 
time (Fig. 1). 

In the enthusiasm for polytypic species as 
a taxonomic device to address the problem 
wrought by the proclivity of some early tax- 
onomists to name every local variant as a 
species, application of the biological species 
concept sometimes was overly conserva- 
tive. In the never-ending search for real or 
inferred intergrades, several subtle but dis- 
tinct species were subsumed under the 
headings of single species. Thus, Merriam’s 
(1919:7) admonition rings true: “‘it [the cri- 
terion of intergradation to delimit species] 
has often resulted in bringing together forms 
between which intergradation has not only 


184 ENGSTROM ET AL. 


not been demonstrated, but which in many 
cases never existed ...”’ Moreover, some 
authors came to view any evidence of hy- 
bridization as proof of intergradation and 
conspecificity (see discussion of Hall, 1981, 
in Patton and Smith, 1990). That the num- 
ber of distinct species of North American 
mammals currently is underestimated has 
become increasingly evident with the ap- 
plication of modern genetic and morpho- 
logical techniques to studies of geographic 
variation and speciation. Recent systematic 
studies often have revealed that many pur- 
portedly intergrading taxa actually repre- 
sent protected, reproductively isolated gene 
pools (Baker, 1984; Baker et al., 1985; Bir- 
ney, 1976; Carleton, 1989; Genoways and 
Choate, 1972; Patton and Smith, 1990; 
Schmidly et al., 1988; Zimmerman, 1970). 
The number of recognized species of North 
American mammals declined from 1,003 to 
887 between 1959 and 1981 (Fig. 1), as pre- 
viously predicted by Hall and Kelson (1959). 
Between 1981 and 1993 the number de- 
creased again to 866 (Wilson and Reeder, 
1993). Included in that total, however, is 
the long-awaited systematic review of brown 
bears (Hall, 1984), wherein the number of 
species was reduced from 78 to 1. Discount- 
ing the 77 species names belatedly placed 
in synonymy by Hall, the number of ad- 
mitted species actually rose by 56 during 
this period despite the fact that discovery 
of hitherto unknown species of mammals 
slowed to a trickle (Fig. 2). We anticipate 
that the number of recognized species will 
continue to rise as our view of species is 
refined, as more specimens become avail- 
able, as geographic coverage improves, and 
especially as multidisciplinary techniques 
are applied to studies of geographic varia- 
tion in a wider variety of taxa (see also 
Carleton, 1989). To the casual observer, 
these changes probably will appear to result 
from a frictionless pendulum perpetually 
swinging between “lumpers” and “‘split- 
ters.”” Instead, we would argue that these 
oscillations represent significant progress in 
our understanding of the composition and 


—— Subspecies --~-~- Species 


Fic. 2.—Number of species and subspecies of 
North American mammals described between 
1900 and 1990. Data for 1900 to 1977 were com- 
piled from Hall (1981), and those for 1977 to 
1990 were taken from The Zoological Record. 


distribution of North American mammals 
during the past 75 years. 

Although the biological species concept 
was, and continues to be, the dominant con- 
cept applied by North American mammal- 
ogists, it is by no means universally ac- 
cepted. Space precludes a full review of this 
ongoing debate, but a few comments may 
be pertinent. For operational reasons, phe- 
neticists dispute the idea that species are 
objective units bound by reproductive con- 
tinuity. Instead, they reiterate the nomi- 
nalist claim that the only objective unit in 
nature is the individual, and that all collec- 
tive higher categories (including species) are 
human constructs (Sokal and Crovello, 
1970). This claim appears intuitively false 
when applied to sympatric species of sex- 
ually reproducing taxa, such as mammals 
or birds (Mayr, 1969). It does, however, 
highlight the difficulty of applying the bio- 
logical species concept to allopatric and al- 
lochronic populations where the potential 
for interbreeding and intergradation must 
be inferred, or in geographically contiguous 
populations among which gene flow is min- 
imal (Ehrlich and Raven, 1969). In these 
cases, biological species indeed are subjec- 
tive constructs, and the erection of polytyp- 
ic species as a taxonomic device runs the 


TAXONOMY 185 


risk of underestimating or misrepresenting 
the number of independent evolutionary 
units. More recently, Wiley (1978, 1981) 
restated Simpson’s (1961) concept of evo- 
lutionary species. ““An evolutionary species 
as a single lineage of ancestor-descendant 
populations which maintains its identity 
from other such lineages and which has its 
own evolutionary tendencies and historical 
fate” (Wiley, 1981:25). This concept stress- 
es that species are bound by unique com- 
mon ancestry, whether or not reproductive 
continuity is evident, and adds the missing 
element of common evolutionary history to 
the biological species concept (Brooks and 
McLennan, 1991). This broader definition 
provides a conceptual means of delineating 
natural species, although operationally it 
sometimes is no less subjective than the bi- 
ological species concept. For example, faced 
with a monophyletic set of allopatric pop- 
ulations, the taxonomist must now decide 
if these populations represent a single evo- 
lutionary lineage, instead of deciding 
whether or not they potentially could inter- 
breed. Nonetheless, in our view, this theo- 
retical concept more closely approximates 
real species-level units (i.e., actual evolu- 
tionary units as manifested by the organ- 
isms themselves). Its application in mam- 
malogy portends a more realistic view of 
species-level taxonomy and the process of 
speciation (but see alternative view in Mayr 
and Ashlock, 1991). Operational variations 
on this theme, such as the phylogenetic spe- 
cies concept (Cracraft, 1983; Donoghue, 
1985; McKitrick and Zink, 1988) also may 
prove useful but, strictly applied, run the 
risk of recognizing all apparently distinctive 
populations as species and a return to a ty- 
pological concept (for a recent application, 
see Engstrom et al., 1992). 


Subspecies Concept 
The history of the category of subspecies 


is closely tied to that of species. Subspecies 
came into regular use in North American 


mammalogy near the turn of the century, 
although the formal trinomen had been used 
by ornithologists since the mid- 1800s. Orig- 
inally conceived as a substitute for the am- 
biguous term variety (which had been used 
as a catch-all for a plethora of intra- and 
interpopulational miscreants), subspecies 
had the general connotation of geographic 
race (e.g., Osgood, 1909). As with species, 
they initially were viewed typologically and 
were defined on a morphological basis: a 
subspecies was a set of specimens that dif- 
fered from another set but not to the same 
degree as species. The general acceptance of 
polytypic species, and the potential for in- 
tergradation as a means of discerning spe- 
cies-limits, spurred use of the category as a 
means of characterizing morphologically 
distinct but intergrading sets of populations. 

Application of the trinomen initially was 
conservative and, until about 1920, about 
as many subspecies were named as new spe- 
cies (Fig. 2). With the onset of the “new 
systematics” in the 1930s and its focus on 
microevolutionary processes, considerable 
effort was expended by mammalogists in 
studying patterns of geographic variation, 
which were formally recognized using the 
trinomen. Unfortunately, rather than ex- 
amining the role of geographic differentia- 
tion in the generation of diversity, discovery 
of statistically distinct subspecies soon be- 
came a primary goal of some mammalian 
taxonomists and the “‘wild-goose chase”’ 
(Mayr, 1963:347) to find new subspecies was 
on. As a consequence, the rate of description 
of subspecies relative to species rose dra- 
matically during the period from 1930 to 
1960 (Fig. 2). The number of recognized 
subspecies of North American mammals 
nearly doubled during this time, whereas 
the number of species decreased by about a 
third (Fig. 1). These changes reflected both 
increased acceptance of the utility of sub- 
species as a taxonomic device and conser- 
vative application of a biological species 
concept. 

During this period, different authors had 
different concepts of subspecies, ranging 


186 ENGSTROM ET AL. 


from subjective geographic divisions of tax- 
onomic convenience to incipient species. For 
example, Mayr (1969) regarded subspecies 
as an arbitrary device to facilitate intraspe- 
cific classification and not as evolutionary 
entities, whereas Lidicker (1960) believed 
that the category should be reserved for phy- 
logenetically delimited subunits of species. 
Given that subspecies are subjective and at 
a point in a gradient between local popu- 
lations and species, the lower-limit of di- 
vergence at which they were recognized also 
varied greatly among authors. As noted by 
Lidicker (1960:161) “‘it is axiomatic that 
populations which consist of different in- 
dividuals are different. The ability to prove 
this difference statistically depends only on 
the size of the samples used and the per- 
ceptual ability of the investigator.”’ None- 
theless, some authors (Mayr et al., 1953; 
Simpson, 1961) advocated a 75% rule—if 
75% of the individuals in one population 
could be distinguished from all individuals 
of an adjacent population, ensuring a sta- 
tistically significant difference, the two could 
be formally recognized as subspecies (pre- 
sumably based on even a single character). 
In some species, where localized patterns of 
geographic differentiation were pro- 
nounced, numerous microgeographic races 
were described. Hence, Setzer (1949) rec- 
ognized 35 subspecies of Ord’s kangaroo rat, 
Dipodomys ordii, many occupying small 
geographic areas. In perhaps the most in- 
famous example, 213 subspecies of the 
pocket gopher, Thomomys umbrinus, were 
admitted in Hall and Kelson (1959). This 
latter case prompted Simpson (1961:173) to 
remark critically “those who enjoy this game 
may go on until every little colony of these 
gophers sports its own Linnaean name.” As 
a mere device for cataloguing geographic 
variants based on a few or single characters, 
recognition of subspecies often has little bi- 
ological meaning and results in formal rec- 
ognition of rankless groups, with no predic- 
tive value relative to additional characters 
(Barrowclough, 1982). In our view, the larg- 
est abuses of the category were made by 


authors who described subspecies based on 
small samples from limited geographic ar- 
eas without a thorough analysis of variation 
within the entire species. 

By 1950, subspecies of North American 
mammals had become an amalgam of old 
names not relegated to full synonymy, lo- 
calized variants, arbitrarily partitioned sec- 
tions of geographic clines, polytopic and mi- 
crogeographic races, discrete evolutionary 
units and, in some instances, subtle but dis- 
tinct species. Not surprisingly, infraspecific 
taxonomy of vertebrates came under heavy 
criticism during a debate on the utility of 
the category that raged largely in the pages 
of Systematic Zoology for 10 years, sparked 
by Wilson and Brown (1953). They noted 
(p. 100), “‘the subspecies concept is the most 
critical and disorderly area of modern sys- 
tematic theory” and advocated that the cat- 
egory be abandoned. For North American 
mammalogists, among the most influential 
contributions to this debate were those of 
Lidicker (1960, 1962), who defined subspe- 
cies as (1962:169) “a relatively homoge- 
neous and genetically distinct portion of a 
species which represents a separately evolv- 
ing, or recently evolved, lineage with its own 
evolutionary tendencies, inhabits a definite 
geographic area, is usually at least partially 
isolated, and may intergrade gradually, al- 
though over a fairly narrow zone, with ad- 
jacent subspecies.” This restrictive defini- 
tion has been widely cited although it 
probably is no coincidence that its most suc- 
cessful applications have been with geo- 
myoid rodents (Genoways, 1973; Lidicker, 
1960; Smith and Patton, 1988) in which 
gene flow among local populations often is 
restricted, pronounced local microgeo- 
graphic differentiation is commonplace, and 
geographic variation is partitioned hierar- 
chically. In other groups for which rates of 
gene flow are higher and geographic differ- 
entiation is less abrupt, taxa fitting the above 
definition most often would be regarded as 
distinct evolutionary species. 

After this debate, application of the sub- 
species category in mammalian taxonomy 


TAXONOMY 187 


became much more conservative and the 
rate of description of new subspecies ap- 
proximated that of species, as it had prior 
to 1920 (Fig. 1). Thus, between 1959 and 
1981, the number of recognized subspecies 
of North American mammals remained rel- 
atively stable (Fig. 2) owing to nearly equal 
rates of additions (new descriptions) and de- 
letions (relegation to synonymy of existing 
subspecies). Since 1981, the rate of descrip- 
tion of new subspecies of North American 
mammals has decreased to less than five per 
year, and there has been a tendency to attach 
less significance to the category (e.g., Wilson 
and Reeder, 1993). As an aside, critics of 
the subspecies category often have branded 
museum curators as the culprits who use 
subspecies aS a convenient device to aid 
them in arranging and subdividing groups 
of specimens in drawers. As curators who 
have spent many unproductive hours at- 
tempting to assign specimens to poorly de- 
fined, undiagnosable subspecies, which seem 
inevitably to be from geographically inter- 
mediate areas, we can assure the reader that 
arbitrarily defined infraspecific taxa are no 
boon to curatorial efficiency or order. 

The current state of the subspecies cate- 
gory in vertebrate taxonomy (and concom- 
itantly of recognized taxa at this level) is 
muddled. Some authors would abandon the 
category entirely (Cracraft, 1983; McKitrick 
and Zink, 1988), whereas several mam- 
malian systematists find a restricted concept 
useful in formally depicting discrete pat- 
terns of geographic variation (e.g., Patton 
and Smith, 1990). In our view, the real pur- 
pose of the trinomen is to describe formally 
patterns of geographic variation by calling 
attention to geographic discontinuities 
among distinctive, evolutionarily discrete 
subsets of populations. We anticipate that, 
as detailed multidisciplinary studies of geo- 
graphic variation are completed for more 
species, and as a conservative concept of 
subspecies is consistently employed, the 
number of recognized subspecies of North 
American mammals will decline substan- 
tially over the coming decades. 


Higher Level Taxonomy 


Schools of systematics and classifica- 
tion. —After the exploratory phase of tax- 
onomy of North American mammals in the 
late 19th and early 20th centuries, the focus 
of studies shifted more towards discerning 
systematic relationships among species. The 
new systematics emphasized studies at low- 
er taxonomic levels and down-played phy- 
logenetic research. Thus, during the period 
from 1930 to 1960 many comprehensive 
taxonomic studies of North American 
mammals focused on species and generic 
level revisions rather than on higher clas- 
sification. The guiding philosophical basis 
of this research was the somewhat intuitive 
school of evolutionary taxonomy champi- 
oned by E. Mayr, G. G. Simpson, and oth- 
ers. The goal was to discern genealogical 
relationships among taxa and then to rep- 
resent both genealogy and extent of phyletic 
divergence in the final classification. How 
these factors were to be weighed was up to 
the discretion of the investigator, and the 
process was said to be part art and part sci- 
ence (Simpson, 1961). Examples of this ap- 
proach include Simpson (1945) and Koop- 
man (1984). These classifications were 
meant to be inherently stable, utilitarian de- 
vices, consistent with what was known about 
evolutionary relationships and magnitude 
of evolutionary change. 

The seeming lack of objectivity of the 
evolutionary school triggered a change in 
systematic philosophy through develop- 
ment of the opposed phenetic and phylo- 
genetic schools of systematics in the 1950s 
and 1960s. These schools were largely re- 
sponsible for the revival of interest in mac- 
rotaxonomy that continues today. Early 
proponents of phenetics (or numerical tax- 
onomy) suggested that, because genealogies 
were difficult to reconstruct and phylogenies 
largely unknown, “natural” higher taxa were 
most objectively discerned by overall sim- 
ilarity (Sneath and Sokal, 1973; Sokal and 
Sneath, 1963). This operationalist (theory- 
free) school is concerned primarily with 


188 ENGSTROM ET AL. 


multivariate, numerical methodologies for 
representing empirical phenetic relation- 
ships, typically weighting all characters 
equally. The method has its genetic exten- 
sion in DNA hybridization, where overall 
similarity between species is calculated from 
average melting temperatures of hybrid 
DNA molecules. Exemplary studies in North 
American mammalogy that employed these 
techniques (but did not necessarily adhere 
to a strict view of the philosophy) include 
those by Findley (1972), Schnell et al. (1978), 
Freeman (1981), Brownell (1983), Owen 
(1988), and Kirsch et al. (1993). Criticisms 
of the use of phenetics in classification in- 
clude: that overall similarity often gives a 
distorted view of phylogenetic relation- 
ships, especially when shared primitive, 
convergent, or uniquely derived character 
states predominate; and that the method, 
although repeatable using the same char- 
acters, produces inherently unstable classi- 
fications likely to be altered when new at- 
tributes are examined. Although phenetic 
philosophy for construction of classifica- 
tions has not been widely accepted in mam- 
malogy, numerical methodology for analyz- 
ing patterns of variation, particularly at the 
microtaxonomic level, has become an in- 
tegral part of the repertoire of techniques 
used by mammalian taxonomists. 

At about the same time as the develop- 
ment of phenetics, the school of phyloge- 
netic systematics (or cladistics) arose and 
has produced a revolution in macrotaxon- 
omy. Stimulated by the writings of Hennig 
(1950, 1966), phylogenetics aims to fulfill 
the goal set by Darwin to base classifications 
directly on genealogy. Phylogenetic rela- 
tionships are based on propinquity of de- 
scent determined from special similarity of 
homologous characters (shared derived 
character states) rather than unweighted, 
overall similarity. Reconstructed phyloge- 
nies subsequently are translated directly into 
classifications. Space precludes a review of 
the development of this school, but much 
of the debate concerning its methodology 
and philosophy (which is far from uniform) 


appears in the pages of Systematic Zoology 
from the 1970s to the present and is sum- 
marized in the texts by Wiley (1981) and 
Eldredge and Cracraft (1980) (see also the 
primer by Wiley et al., 1991). North Amer- 
ican mammalogists have been bit players in 
the development of phylogenetics, although 
arguably the most important recent ad- 
vances in higher classifications of mammals 
have employed this method (at least to re- 
construct cladistic branching sequences). In 
particular, molecular systematists working 
on North American mammals who initially 
used phenetic methods almost exclusively 
now routinely apply cladistic parsimony to 
discern relationships. One only need peruse 
the pages of the Journal of Mammalogy or 
Systematic Biology (formerly Systematic 
Zoology) for the past 10 years to see the 
predominant influence of this school on 
vertebrate taxonomy and systematics. Pub- 
lications on North American mammals em- 
ploying this methodology are too numerous 
to cite, but a few exemplary studies include: 
Greenbaum and Baker (1978); Carleton 
(1980); Smith and Madkour (1980); Grif- 
fiths (1982); Hood and Smith (1982); Rog- 
ers et al. (1984); Owen (1987); Voss (1988); 
Baker et al. (1989); Miyamoto et al. (1989); 
Wozencraft (1989a); Wyss (1989); Hafner 
(1991); Pacheco and Patterson (1991); Lim 
(1993). 

Continued dialogue (often acrimonious) 
among these three schools of systematics 
has resulted in considerable refinement of 
taxonomic methodology. By partitioning 
historical evolution (descent with modifi- 
cation) into the separate components of 
phenetic divergence and genealogy, classi- 
fications no longer need rest on intuition 
and authority; instead, they are based on 
empirical evidence of change in character 
states. Thus, as Hooper (1968:33) noted: “A 
classification is a tentative thing; it is not 
sacred.” This change has resulted in a re- 
kindled interest in macrotaxonomy in gen- 
eral, and in the higher classification of mam- 
mals, in particular. It also has sparked a new 
interest in using classifications to test hy- 


TAXONOMY 189 


potheses about historical processes in bio- 
logical disciplines outside the field of sys- 
tematics (Brooks and McLennan, 1991). 

Higher classification. —The history of 
mammalian classification was reviewed by 
Gregory (1910), Simpson (1945), Szalay 
(1977), and Novacek (1982, 1990), and only 
a few highlights will be mentioned here. 
Since the turn of the century, much of the 
outstanding work by North Americans on 
classification of mammals has emanated 
from the Department of Vertebrate Pale- 
ontology of the American Museum of Nat- 
ural History. Before the formation of the 
ASM in 1919, the most comprehensive and 
influential mammalian classification was 
that of Gregory (1910: Table 1). Phyloge- 
netic in approach, Gregory was concerned 
with distinguishing between primitive and 
derived traits and with eliminating conver- 
gence (although these tenets were not always 
consistently followed in defining groups). 
Gregory’s classification was relatively high- 
ly resolved; an optimistic solution not shared 
by several later workers (including Simp- 
son, 1945), who more often regarded rela- 
tionships among most eutherian orders as 
an unresolved phylogenetic ““bush.”” Among 
several other groups, Gregory (1910) de- 
fined and defended the Archonta (including 
elephant shrews, tree shrews, bats, gliding 
lemurs, and primates), over which there has 
been much recent debate. Included in this 
synthesis (Gregory, 1910) is a fascinating 
historical review of mammalian classifica- 
tion that merits careful reading by anyone 
interested in the development of system- 
atics. 

Simpson (1945) later published what has 
been widely regarded as the standard clas- 
sification of mammals (Table 1). This work 
was more detailed than that of Gregory, in 
that all mammals were classified to genus. 
Until the last decade, the pervasive influ- 
ence of this monograph could be seen by 
touring the large museum and university 
collections of mammals in the United States, 
most of which were ‘“‘arranged according to 
Simpson (1945). Part of that influence 


stemmed from Simpson’s position as a lead- 
ing evolutionary theorist and his strong ad- 
vocation of intuitive, evolutionary taxon- 
omy. Many of his groups were based on his 
perception of phylogeny (e.g., recognition of 
the Ferungulata, including carnivores, un- 
gulates, and related orders to the exclusion 
of other mammals), although these groups 
were not justified by shared derived features 
and have not been well accepted. In fact, 
despite its comprehensiveness, there was 
little explicit discussion of characters on 
which the classification was based. For ex- 
ample, Simpson (1945:173) dismissed 
Gregory’s Archonta, without reference to 
characters or literature citations: “it is in- 
credible to me. . . that the primates are more 
closely related to bats than to the insecti- 
vores, and all recent research ... opposes 
that opinion.” 

Thirty years later, changes in systematic 
philosophy and discovery of new Mesozoic 
fossils led to a radical departure from Simp- 
son (McKenna, 1975; Table 1). This was 
the first major classification of mammals 
that used cladistic methodology to recon- 
struct phylogeny and it included explicit 
discussion of character state transforma- 
tions (especially dental homologies). Ini- 
tially, McKenna (1975) was criticized be- 
cause his classification was complex and 
because he erected a large number of new 
superordinal categories to reflect relative re- 
cency of common ancestry directly (Szalay, 
1977). However, as noted by Novacek 
(1982), his departure from “traditional” 
systematics by providing explicit consid- 
erations of alternative phylogenetic hypoth- 
eses has not received due credit. Some of 
McKenna’s (1975) more important depar- 
tures from Simpson include (Table 1): early 
branching of the Edentata from the rest of 
the eutherian mammals; resurrection of 
Gregory’s Archonta (sans the elephant 
shrews— Macroscelidea); phylogenetic as- 
sociation of Macroscelidea and lagomorphs; 
arrangement of whales (Cetacea) within a 
superordinal group including ungulates and 
their relatives but excluding carnivores. AI- 


190 


TABLE |.— Selected 20th century, higher-level 


ENGSTROM ET AL. 


classifications of extant mammals. 


Gregory, 1910 


Class Mammalia 
Subclass Prototheria 


Order Monotremata 


Subclass Theria 
Infraclass Metatheria 


Order Marsupialia 
Suborder Diprotodontia 
Suborder Paucituberculata 
Suborder Polyprotodontia 


Infraclass Eutheria 


Superorder Therictoidea 
Order Insectivora 
Suborder Lipotyphla 
Order Ferae 
Suborder Fissipedia 
Suborder Pinnipedia 
Superorder Archonta 
Order Menotyphla [includes Tupaidae, 
Macroscelidae] 
Order Dermoptera 
Order Chiroptera 
Order Primates 
Superorder Rodentia 
Order Glires 
Suborder Duplicidentata [Lagomorpha] 
Suborder Simplicidentata [Rodentia] 
Superorder Edentata 
Order Tubulidentata 
Order Pholidota 
Order Xenarthra 
Superorder Paraxonia 
Order Artiodactyla 
Superorder Ungulata 
Order Sirenia 
Order Hyraces 
Order Mesaxonia [includes Perissodactyla] 
Superorder Cetacea 
Order Odontoceti 
Order Mystacoceti 


Simpson, 1945 


Class Mammalia 
Subclass Prototheria 


Order Monotremata 


Subclass Theria 
Infraclass Metatheria 


Order Marsupialia 


Infraclass Eutheria 


Cohort Unguiculata 
Order Insectivora [includes Lipotyphla, 
Macroscelidae] 
Order Dermoptera 
Order Chiroptera 
Order Primates 


TABLE |.— Continued. 


Order Edentata 
Order Pholidota 
Cohort Glires 
Order Lagomorpha 
Order Rodentia 
Cohort Mutica 
Order Cetacea 
Cohort Ferungulata 
Superorder Ferae 
Order Carnivora 
Suborder Fissipedia 
Suborder Pinnipedia 
Superorder Protungulata 
Order Tubulidentata 
Superorder Paenungulata 
Order Proboscidea 
Order Hyracoidea 
Order Sirenia 
Superorder Mesaxonia 
Order Perissodactyla 
Superorder Paraxonia 
Order Artiodactyla 


McKenna, 1975 


Class Mammalia 
Subclass Prototheria 
Infraclass Ormithodelphia 
Order Monotremata 
Subclass Theria 
Infraclass Tribosphenida 
Supercohort Marsupialia 
Supercohort Eutheria 
Cohort Edentata 
Order Cingulata 
Order Pilosa 
Cohort Epitheria 
Magnorder Ernothena 
Order Macroscelidea 
Order Lagomorpha 
Magnorder Preptotheria 
Grandorder Ferae 
Order Carnivora 
Grandorder Insectivora 
Order Erinaceomorpha 
Order Soricomorpha 
Grandorder Archonta 
Order Scandentia 
Order Dermoptera 
Order Chiroptera 
Order Primates 
Grandorder Ungulata 
Mirorder Eparctocyona 
Order Tubulidentata 
Order Artiodactyla 
Mirorder Cete 
Order Cetacea 


TAXONOMY 191 


TABLE |.— Continued. 


Suborder Odontoceti 
Suborder Mysticeti 
Mirorder Phenacodonta 
Order Perissodactyla 
Order Hyracoidea 
Mirorder Tethytheria 
Order Proboscidea 
Order Sirenia 
Magnorder Preptotheria, incertae sedis 
Order Pholidota 
Cohort Epitheria, incertae sedis 
Order Rodentia 


Eutherian Mammals (Novacek, 1986) 


Subclass Theria 
Infraclass Eutheria 
Cohort Edentata 
Order Xenarthra 
Order Pholidota 
Cohort Epitheria 
Superorder Insectivora 
Order Lipotyphla 
Superorder Volitantia 
Order Dermoptera 
Order Chiroptera 
Superorder Anagalida 
Order Macroscelidea 
Grandorder Glires 
Order Rodentia 
Order Lagomorpha 
Superorder Ungulata 
Order Artiodactyla 
Order Cetacea 
Order Perissodactyla 
Grandorder Paenungulata 
Order Hyracoidea 
Mirorder Tethytheria 
Order Proboscidea 
Order Sirenia 
Cohort Epitheria incertae sedis 
Order Tubulidentata 
Order Carnivora 
Order Primates 
Order Scandentia 


Metatherian Mammals (Marshall et al., 1990) 


Subclass Theria 
Infraclass Metatheria 
Supercohort Marsupialia 

Cohort Ameridelphia 
Order Didelphimorphia 
Order Paucituberculata 

Cohort Australidelphia 
Order Microbiotheria 
Order Dasyuromorphia 
Order Peramelina 
Order Notoryctemorphia 
Order Diprotodontia 


though subsequent authors (e.g., Szalay, 
1977) have disagreed with some of Mc- 
Kenna’s (1975) interpretations of characters 
and methodology, this paper set the stage 
for a dynamic reinvestigation of higher-lev- 
el relationships in mammals. 

Szalay (1977) examined phylogeny of eu- 
therian mammals based on largely on tarsal 
morphology. His resulting classification was 
derived both from the proposed genealogy 
and his view of “‘adaptational history.”” He 
supported some of the same groups as Mc- 
Kenna (1975), such as the Archonta, the 
association of the Macroscelidea and Lago- 
morpha, and the existence of an ungulate 
supergroup, but was not convinced of the 
early derivation of edentates. 

A more recent phylogeny and classifica- 
tion of eutherian mammals was proposed 
by Novacek (1986; Table 1), reconstructed 
using a large suite of skeletal and soft ana- 
tomical characters (although his cladograms 
were based on characters of the skull). Al- 
though far from fully resolved, this is the 
most explicit statement and defense of eu- 
therian superordinal relationships to date. 
Therein (Table 1), he supported McKenna’s 
(1975) early derivation of edentates (in- 
cluding pangolins—Pholidota) from other 
eutherians, an ungulate superorder, and the 
association of Macroscelidea with lago- 
morphs and rodents. He was, however, un- 
able to find support for Gregory’s (1910) 
Archonta (tentative justification for this 
group based on penial morphology and 
structure of the tarsus is given in Novacek 
and Wyss, 1986; Novacek et al., 1988, but 
see comments in Novacek, 1993). 

Perhaps the most radical recent change in 
the higher classification of mammals is the 
subdivision of marsupials into several or- 
ders (Aplin and Archer 1987; Marshall et 
al., 1990; Ride, 1964; Szalay, 1982; Table 
1). In particular, the recognition of the South 
American Microbiotheridae as a member of 
the Australidelphia clade (Aplin and Ar- 
cher, 1987; Kirsch et al., 1991; Marshall et 
al., 1990; Szalay, 1982) is novel. 

The past 20 years have witnessed a re- 


192 ENGSTROM ET AL. 


markable improvement in the state of our 
knowledge concerning higher classification of 
mammals, aided immeasurably by the for- 
mulation of explicit, falsifiable hypotheses of 
monophyly and evolution of character states. 
Thus, the statement by Ammerman and 
Hillis (1992:230) that, ““Mammalogists to- 
day have less confidence in the branching 
order of the 18 orders of mammals than 
they did 100 years ago” is overly pessimis- 
tic. Analyses of molecular data hold con- 
siderable promise in the resolution of sev- 
eral of the seemingly intractable problems 
of mammalian phylogeny and interordinal 
relationships (Czelusniak et al., 1990; Ho- 
neycutt and Yates, 1994; Miyamoto and 
Goodman, 1986). Examination of congru- 
ence (or the lack thereof) among molecular 
and morphological data sets, however, sug- 
gests that this promise has yet to be fully 
realized (Novacek, 1989, 1990; Novacek et 
al., 1988; Wyss et al., 1987). We cautiously 
agree with McKenna (1987:82), referring to 
the congruence of amino acid sequences and 
morphology: “As with all information, there 
is a mixture of signal and noise, ..., but 
the situation seems to be getting quieter 
[italics ours].” 


Faunal Surveys 


One of the natural outgrowths of taxo- 
nomic work on mammals has been pro- 
duction of catalogues of mammals occur- 
ring in circumscribed geographic areas. The 
best of these catalogues have been written 
by practicing taxonomists. For the most part, 
these catalogues were not compiled within 
a “biodiversity” framework; however, they 
form the basis of our knowledge of mam- 
malian diversity and geographic distribu- 
tion. Mammalian faunal surveys have deep 
roots reaching back to the 19th Century to 
such classics as Harlan’s (1825) Fauna 
Americana, Richardson’s (1829) Fauna Bo- 
reali-Americana, DeKay’s (1842) Zoology 
of New- York, and Audubon and Bachman’s 
(1846 to 1854) The Viviparous Quadrupeds 


of North America. The monumental classic 
of the era was Baird’s (1857) review of 
mammals of North America. This publi- 
cation preceded by 100 years the classic of 
the next century, The Mammals of North 
America, by Hall and Kelson (1959). Both 
monographs stimulated considerable addi- 
tional taxonomic studies and faunal sur- 
veys. 

Faunal studies in the 10 years following 
the establishment of the ASM included those 
of Goldman (1920) for Panama, Howell 
(1921) for Alabama, and Bailey (1926) for 
North Dakota. The number of “Mammals 
of...’ monographs showed a marked in- 
crease during the 1930s, the most notable 
by Bailey (1932, 1936) for New Mexico and 
Oregon, Grinnell (1933) for California, and 
Goodwin (1935) for Connecticut. However, 
a sign of things to come was the publication 
of the first faunal studies by two of Grin- 
nell’s professional progeny (Burt, 1938, So- 
nora; Davis, 1939, Idaho). The 1940s, like 
the 1930s, were characterized by publica- 
tion of an increasing number of faunal stud- 
ies, a few of the best known being those of 
Bole and Moulthrop (1942) for Ohio, Ham- 
ilton (1943) for the eastern U.S., Anderson 
(1947) for Canada, Burt (1948) for Michi- 
gan, and Dalquest (1948) for Washington. 
Also published during this period was Hall’s 
(1946) Mammals of Nevada, which set the 
standard for subsequent mammalian sur- 
veys. 

Relatively few faunal studies were pub- 
lished in the 1950s, the most important be- 
ing the classic Biological Investigations in 
Mexico, by Goldman (1951). Additional ex- 
amples were the state faunas and regional 
surveys by Cockrum (1952) for Kansas, 
Durrant (1952) for Utah, Dalquest (1953) 
for San Luis Potosi, Baker (1956) for Coa- 
huila, and Bee and Hall (1956) for northern 
Alaska. Noteworthy faunal studies during 
the 1960s were those of Jackson (1961) for 
Wisconsin, Baker and Greer (1962) for Du- 
rango, Alvarez (1963) for Tamaulipas, Hall 
and Dalquest (1963) for Veracruz, Jones 
(1964) for Nebraska, Long (1965) for Wy- 
oming, Peterson (1966) for eastern Canada, 


TAXONOMY EOS 


Villa-R. (1967) for Mexico, and Goodwin 
(1969) for Oaxaca. Some of the more im- 
portant of the large number of ““Mammals 
of ...” books produced during the 1970s 
were those of Armstrong (1972) for Colo- 
rado, Anderson (1972) for Chihuahua, Ban- 
field (1974) for Canada, Lowery (1974) for 
Louisiana, Findley et al. (1975) for New 
Mexico, Youngman (1975) for the Yukon 
Territory, and Schmidly (1977) for Trans- 
Pecos Texas. Faunal studies published dur- 
ing the 1980s included those of Mumford 
and Whitaker (1982) for Indiana, Baker 
(1983) for Michigan, Jones et al. (1983) for 
the Great Plains, Schmidly (1983) for east- 
ern Texas, Hoffmeister (1986, 1989) for Ar- 
izona and Illinois, Caire et al. (1989) for 
Oklahoma, and Merritt (1987) for Penn- 
sylvania. 

Coincident with the formation of the 
Mexican Society of Mammalogy (AM- 
MAC), there has been an increasing trend 
for locally produced faunal surveys and 
identification guides in Mexico over the last 
decade, a few examples of which include 
Ceballos and Galindo (1984) for the valley 
of México, Ceballos and Miranda (1986) for 
Chamela, Jalisco, Ramirez-Pulido et al. 
(1986) for Mexico, Aranda and March 
(1987) for Chiapas, Coates-Estrada and Es- 
trada (1986) for Los Tuxtlas, Veracruz, and 
Alvarez-Castaneda and Alvarez (1991) for 
Chiapas. These studies herald the burgeon- 
ing local interest and expertise in the region 
of highest diversity of mammals in North 
America, and we anticipate an increasing 
number of faunal surveys in Mexico over 
the coming decades. 


Acknowledgments 


We thank B. K. Lim for his assistance in com- 
piling historical data and producing the figures. 


Literature Cited 


ALVAREZ, T. 1963. The Recent mammals of Tamau- 
lipas. University of Kansas Publications, Museum 
of Natural History, 14:363-473. 


ALVAREZ-CASTANEDA, S. T., AND T. ALVAREZ. 1991. 
Los murciélagos de Chiapas. Instituto Politecnico 
Nacional, México, D. F., 211 pp. 

AMMERMAN, L. K., AND D. M. Hiuis. 1992. A mo- 
lecular test of bat relationships: monophyly or di- 
phyly? Systematic Biology, 41:222-232. 

ANDERSON, R. M. 1947. Catalogue of Canadian Re- 
cent mammals. National Museum of Canada Bul- 
letin, Biological Series, 102:1-238. 

ANDERSON, S. 1972. Mammals of Chihuahua: tax- 
onomy and distribution. Bulletin of the American 
Museum of Natural History, 148:149-410. 

ANDERSON, S., AND J. K. Jones, JR. 1967. Recent 
mammals of the world: a synopsis of families. Ron- 
ald Press Company, New York, 453 pp. 

1984. Orders and families of Recent mam- 
mals of the world. John Wiley and Sons, New York, 
686 pp. 

APLIN, K. P., AND M. ARCHER. 1987. Recent ad- 
vances in marsupial systematics with a new syncretic 
classification. Pp. xv—xxii, in Possums and opos- 
sums: studies in evolution (M. Archer, ed.). Surrey 
Beatty and Sons Pty. Ltd., and Royal Zoological So- 
ciety of New South Wales, Sydney, 1:1-400. 

ARANDA, M., AND I. MarcH. 1987. Guia de los ma- 
miferos silvestres de Chiapas. Instituto Nacional de 
Investigaciones sobre recursos bioticos, Xalapa, Ve- 
racruz, 149 pp. 

ArITA, H. T., AND S. R. HUMPHREY. 1988. Revision 
taxonomica de los murciélagos del género Lepto- 
nycteris (Chiroptera: Phyllostomidae). Acta Zoolo- 
gica Mexicana, n. ser., 29:1-60. 

ARMSTRONG, D. M. 1972. Distribution of mammals 
in Colorado. Monograph of the Museum of Natural 
History, The University of Kansas, 3:1-415. 

AUDUBON, J. J., AND J. BACHMAN. 1846-1854. The 
viviparous quadrupeds of North America. V. G. Au- 
dubon, New York, 3 volumes (1:1-389; 2:1-334; 
3:1-348). 

BaiLEy, V. 1926. A biological survey of North Da- 
kota. North American Fauna, 49:1-226. 

1932. Mammals of New Mexico. North 

American Fauna, 53:1-402. 

1936. The mammals and life zones of Ore- 
gon. North American Fauna, 55:1-416. 

BairD, S. F. 1857. General report upon the zoology 
of the several Pacific railroad routes: mammals. Vol. 
8, pt. 1, in Reports of explorations and surveys to 
ascertain the most practicable and economical route 
for a railroad from the Mississippi River to the Pa- 
cific Ocean. Senate executive document no. 78, 
Washington, D.C., 757 pp. 

BAKER, R. H. 1956. Mammals of Coahuila, México. 
University of Kansas Publications, Museum of Nat- 
ural History, 9:125-335. 

1983. Michigan mammals. Michigan State 
University Press, East Lansing, 642 pp. 

BAKER, R. H., AND J. K. GREER. 1962. Mammals of 
the Mexican state of Durango. Publications of the 
Museum, Michigan State University, 2:1-154. 

BAKER, R. J. 1984. A sympatric cryptic species of 
mammal: a new species of Rhogeessa (Chiroptera 
Vespertilionidae). Systematic Zoology, 33:178-183. 

BAKER, R. J., AND M.S. HAFNER. 1994. Cytogenetics. 
Pp. 310-321, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 


194 ENGSTROM ET AL. 


Special Publication, The American Society of Mam- 
malogists, 1 1:1-433. 

BAKER, R. J., J. W. BIcKHAM, AND M. L. ARNOLD. 
1985. Chromosomal evolution in Rhogeessa (Chi- 
roptera: Vespertilionidae): possible speciation by 
centric fusions. Evolution, 39:233-243. 

Baker, R. J., C. S. Hoop, AND R. L. HONEyYCUTT. 
1989. Phylogenetic relationships and classification 
of the higher categories of the New World bat family 
Phyllostomidae. Systematic Zoology, 38:228-238. 

Baker, R. J., J. C. Patron, H. H. GENoways, AND J. 
W. BICKHAM. 1988. Genic studies of Lasiurus (Chi- 
roptera: Vespertilionidae). Occasional Papers, The 
Museum, Texas Tech University, 117:1-15. 

BANFIELD, A. W. F. 1974. The mammals of Canada. 
The University of Toronto Press, Toronto, 438 pp. 

BARROWCLOUGH, G. F. 1982. Geographic variation, 
predictiveness, and subspecies. Auk, 99:601-603. 

Beg, J. W., ANDE.R. HALL. 1956. Mammals of north- 
ern Alaska on the Arctic Slope. Miscellaneous Pub- 
lications, Museum of Natural History, University of 
Kansas, 8:1-309. 

BirNEY, E. C. 1973. Systematics of three species of 
woodrats (genus Neotoma) in central North Amer- 
ica. Miscellaneous Publications, University of Kan- 
sas, Museum of Natural History, 58:1-173. 

1976. An assessment of relationships and 
effects of interbreeding among woodrats of the Ne- 
otoma floridana species-group. Journal of Mam- 
malogy, 57:103-132. 

Bote, B. J., Jk., AND P. N. MoULTHROP. 1942. The 
Ohio Recent mammal collection in the Cleveland 
Museum of Natural History. Scientific Publication 
of the Cleveland Museum of Natural History, 5:83- 
181. 

Brooks, D. R., AND D. A. MCLENNAN. 1991. Phy- 
logeny, ecology, and behavior: a research program 
in comparative biology. The University of Chicago 
Press, Chicago, 434 pp. 

BROWNELL, E. 1983. DNA/DNA hybridization stud- 
ies of muroid rodents: symmetry and rates of mo- 
lecular evolution. Evolution, 37:1034-1051. 

Burt, W. H. 1938. Faunal relationships and geo- 
graphic distribution of mammals in Sonora, Mexico. 
Miscellaneous Publications of the Museum of Zo- 
ology, University of Michigan, 39:1-77. 

1948. The mammals of Michigan. The Uni- 
versity of Michigan Press, Ann Arbor, 288 pp. 

CarrRE, W., J.D. TyLer, B. P. GLAss, AND M. A. MARES. 
1989. Mammals of Oklahoma. University of Okla- 
homa Press, Norman, 567 pp. 

CARLETON, M. D. 1977. Interrelationships of popu- 
lations of the Peromyscus boylii species group (Ro- 
dentia, Muridae) in western Mexico. Occasional Pa- 
pers of the Museum of Zoology, University of 
Michigan, 675:1-47. 

1980. Phylogenetic relationships in neoto- 

mine-peromyscine rodents (Muroidea) and a reap- 

praisal of the dichotomy within New World Criceti- 
nae. Miscellaneous Publications, Museum of Zoology, 

University of Michigan, 157:1-146. 

. 1989. Systematics and evolution. Pp. 7-141, 

in Advances in the study of Peromyscus (Rodentia) 

(G. L. Kirkland, Jr., and J. N. Layne, eds.). Texas 

Tech University Press, Lubbock, 366 pp. 


CARLETON, M. D., AND R. E. ESHELMAN. 1979. A 
synopsis of fossil grasshopper mice, genus Onycho- 
mys, and their relationships to Recent species. Uni- 
versity of Michigan, Museum of Paleontology, Pa- 
pers on Paleontology, 21:1-63. 

CARLETON, M. D., AND G. G. Musser. 1989. System- 
atic studies of oryzomyine rodents (Muridae, Sig- 
modontinae): a synopsis of Microryzomys. Bulletin 
of the American Museum of Natural History, 191: 
1-83. 

CEBALLOS, G., AND C. GALINDO. 1984. Mamiferos 
silvestres de la Cuenca de México. Instituto de Ecolo- 
gia y Museo de Historia Natural de la Ciudad de 
México, México, D. F., 299 pp. 

CEBALLOS, G., AND A. MIRANDA. 1986. Los mami- 
feros de Chamela, Jalisco. Instituto de Biologia, 
Universidad Nacional Autonoma de México, Méx- 
ico, D. F., 436 pp. 

CHOATE, J. R. 1970. Systematics and zoogeography 
of Middle American shrews of the genus Cryptotis. 
University of Kansas Publications, Museum of Nat- 
ural History, 19:195-317. 

COATES-EsSTRADA, R., AND A. ESTRADA. 1986. Manu- 
al de identificacidn de campo de los mamiferos de 
la estacion de biologia “Los Tuxtlas.” Instituto de 
Biologia, Universidad Nacional Autonoma de Méxi- 
co, México, D. F., 151 pp. 

Cockrum, E. L. 1952. Mammals of Kansas. Uni- 
versity of Kansas Publications, Museum of Natural 
History, 7:1-306. 

Cougs, E., AND J. A. ALLEN. 1877. Monographs of 
North American Rodentia. Report of the U.S. Geo- 
logical Survey of the Territories (F. V. Hayden, U.S. 
Geologist-in-Charge), Washington, D.C., Volume 11, 
1,091 pp. 

CRACRAFT, J. 1983. Species concepts and speciation 
analysis. Current Ornithology, 1:159-187. 

CZELUSNIAK, J., ET AL. 1990. Perspectives from ami- 
no acid and nucleotide sequences on cladistic rela- 
tionships among higher taxa of Eutheria. Pp. 545- 
577, in Current mammalogy (H. H. Genoways, ed.). 
Plenum Press, New York, 2:1—577. 

DALQUEST, W. W. 1948. Mammals of Washington. 
University of Kansas Publications, Museum of Nat- 
ural History, 2:1-444. 

1953. Mammals of the Mexican state of San 
Luis Potosi. Louisiana State Universities Studies, 
Biological Science Series, 1:1-—229. 

Davis, W. B. 1939. The Recent mammals of Idaho. 
Caxton Printers, Ltd., Caldwell, Idaho, 400 pp. 

. 1968. A review of the genus Uroderma (Chi- 

roptera). Journal of Mammalogy, 49:676-698. 

. 1969. A review of the small fruit bats (genus 

Artibeus) of Middle America. The Southwestern 

Naturalist, 14:15-29. 

. 1970. A review of the small fruit bats (genus 
Artibeus) of Middle America. Part II. The South- 
western Naturalist, 14:389—402. 

DeKay, J. E. 1842. Zoology of New-York, or the 
New-York fauna; comprising detailed descriptions 
of all the animals hitherto observed within the state 
of New-York, with brief notices of those occasionally 
found near its borders and accompanied by appro- 
priate illustrations. Part |. Mammalia. Printed by 
W. A. White and J. Visscher, Albany, 146 pp. 


TAXONOMY 195 


DONOGHUE, M. J. 1985. A critique of the biological 
species concept, and recommendation for a phylo- 
genetic alternative. Bryologist, 88:172-181. 

DurRRANT, S. D. 1952. Mammals of Utah: taxonomy 
and distribution. University of Kansas Publications, 
Museum of Natural History, 6:1-549. 

Ecer, J. L. 1977. Systematics of the genus Eumops 
(Chiroptera: Molossidae). Life Sciences Contribu- 
tions, Royal Ontario Museum, 110:1-69. 

EuRLICcH, P. R., AND P. H. RAvEN. 1969. Differen- 
tiation of populations. Science, 165:1228-1232. 
ELDRIDGE, N., AND J. CRACRAFT. 1980. Phylogenetic 
patterns and the evolutionary process. Columbia 

University Press, New York, 349 pp. 

Extuiot, D. G. 1904. A checklist of mammals of the 
North American continent, the West Indies, and the 
neighboring seas. Field Columbian Museum, Pub- 
lication 105, Zoological Series, 6:1-761. 

ENGsTROM, M. D., AND D. E. Witson. 1981. System- 
atics of Antrozous dubiaquercus (Chiroptera: Ves- 
pertilionidae), with comments on the status of Bau- 
erus Van Gelder. Annals of Carnegie Museum, 50: 
371-383. 

ENGstTrom, M. D., O. SANCHEZ-HERRERA, AND G. 
URBANO-VIDALES. 1992. Distribution, geographic 
variation, and systematic relationships within Ne/- 
sonia (Rodentia: Sigmodontinae). Proceedings of the 
Biological Society of Washington, 105:867-881. 

FINDLEY, J. S. 1955. Speciation of the wandering 
shrew. University of Kansas Publications, Museum 
of Natural History, 9:1-68. 

FINDLEY J. S. 1972. Phenetic relationships among 
bats of the genus Myotis. Systematic Zoology, 21: 
31-52. 

FINDLEY, J. S., AND G. L. TRAuT. 1970. Geographic 
variation in Pipistrellus hesperus. Journal of Mam- 
malogy, 51:741-765. 

FINDLEY, J. S., A. H. Harris, D. E. WILSON, AND C. 
Jones. 1975. Mammals of New Mexico. Univer- 
sity of New Mexico Press, Albuquerque, 360 pp. 

FREEMAN, P. W. 1981. A multivariate study of the 
family Molossidae (Mammalia: Chiroptera): mor- 
phology, ecology, evolution. Fieldiana, Zoology, New 
Series, 7:1-173. 

GARDNER, A. L. 1973. The systematics of the genus 
Didelphis (Marsupialia: Didelphidae) in North and 
Middle America. Special Publications, The Muse- 
um, Texas Tech University, 4:1-81. 

Genoways, H. H. 1973. Systematics and evolution- 
ary relationships of spiny pocket mice, genus Lio- 
mys. Special Publications, The Museum, Texas Tech 
University, 5:1-368. 

Genoways, H. H., AND J. R. CHOATE. 1972. A mul- 
tivariate analysis of systematic relationships among 
populations of the short-tailed shrew (genus Blarina) 
in Nebraska. Systematic Zoology, 21:106-116. 

Genoways, H. H., AND J. K. Jones, JR. 1971. Sys- 
tematics of southern banner-tailed kangaroo rats of 
the Dipodomys phillipsii group. Journal of Mam- 
malogy, 52:265-287. 

GeorGE, S. B. 1988. Systematics, historical bioge- 
ography, and evolution of the genus Sorex. Journal 
of Mammalogy, 69:443-461. 

GEORGE, S. B., J. R. CHOATE, AND H. H. GENOWAYS. 
1981. Distribution and taxonomic status of Blarina 


hylophaga Elliot (Insectivora: Soricidae). Annals of 
Carnegie Museum, 50:493-513. 

GeorGE, S. B., H. H. GENoways, J. R. CHOATE, AND 
R. J. BAKER. 1982. Karyotypic relationships within 
the short-tailed shrews, genus Blarina. Journal of 
Mammalogy, 63:639-645. 

GOLDMAN, E. A. 1920. Mammals of Panama. Smith- 
sonian Miscellaneous Collections, 69:1—309. 

1950. Raccoons of North and Middle Amer- 

ica. North American Fauna, 60:1-153. 

1951. Biological investigations in Mexico. 
Smithsonian Miscellaneous Collections, 115:1-476. 

Goopwin, G. G. 1935. The mammals of Connecti- 
cut. State of Connecticut, State Geological and Nat- 
ural History Survey, Bulletin, 53:1-221. 

1969. Mammals from the state of Oaxaca, 
Mexico, in the American Museum of Natural His- 
tory. Bulletin of the American Museum of Natural 
History, 141:1-269. 

GREENBAUM, I. F., AND R. J. BAKER. 1978. Deter- 
mination of the primitive karyotype of Peromyscus. 
Journal of Mammalogy, 59:820-834. 

Grecory, W.K. 1910. The orders of mammals. Bul- 
letin of the American Museum of Natural History, 
27:1-524. 

GRIFFITHS, T. A. 1982. Systematics of the New World 
nectar-feeding bats (Mammalia, Phyllostomidae) 
based on features of the morphology of the hyoid 
and lingual regions. American Museum Novitates, 
2742:1-45. 

GRINNELL, J. 1933. Review of the Recent mammal 
fauna of California. University of California Publi- 
cations in Zoology, 40:71-234. 

HaFNneR, D. J., J.C. HAFNER, AND M.S. HAFNER. 1979. 
Systematic status of kangaroo rats, genus Microdi- 
podops: morphometric, chromosomal, and protein 
analyses. Journal of Mammalogy, 60:1-10. 

Harner, M S. 1982. A biochemical investigation of 
geomyoid systematics (Mammalia: Rodentia). Zeit- 
schrift fur Zoologische Systematik und Evolutions- 
forschung, 20:118-—130. 

. 1991. Evolutionary genetics and zoogeogra- 
phy of Middle American pocket gophers, genus Or- 
thogeomys. Journal of Mammalogy, 72:1-10. 

HALL, E. R. 1946. Mammals of Nevada. University 
of California Press, Berkeley, 710 pp. 

1951. American weasels. University of Kan- 

sas Publications, Museum of Natural History, 4:1- 

466. 


. 1981. The mammals of North America. Sec- 
ond ed. John Wiley and Sons, New York, 1:1-—600 
+ 90, 2:601-1181 + 90. 

1984. Geographic variation among brown 
and grizzly bears (Ursus arctos) in North America. 
Special Publication, Museum of Natural History, 
University of Kansas, 13:1-16. 

HALL, E. R., AND W. W. DALQugEst. 1963. The mam- 
mals of Veracruz. University of Kansas Publications, 
Museum of Natural History, 14:165-362. 

HALL, E. R., AND K. R. Ketson. 1959. The mammals 
of North America. Ronald Press Co., New York, 
1:1-546 + 79, 2:547-1083 + 79. 

HaMILTON, W. J., JR. 1943. The mammals of eastern 
United States: an account of Recent land mammals 


196 ENGSTROM Ef AL. 


occurring east of the Mississippi. Comstock Pub- 
lishing Co., Inc., Ithaca, New York, 432 pp. 

HANDLEY, C. O., Jr. 1959. Revision of American 
bats of the genera Euderma and Plecotus. Proceed- 
ings of the United States National Museum, 1 10:95- 
246. 

HARLAN, R. 1825. Fauna Americana, being a de- 
scription of the mammiferous animals inhabiting 
North America. Anthony Finley (publisher), Phila- 
delphia, 320 pp. 

HEANEY, L. R., AND R. M. Timm. 1983. Relationships 
of pocket gophers of the genus Geomys from the 
central and northern Great Plains. Miscellaneous 
Publications, Museum of Natural History, Univer- 
sity of Kansas, 73:1-59. 

HENNIG, W. 1950. Grundzuge einer Theorie der phy- 
logenetischen Systematik. Deutscher Zentralverlag, 
Berlin, 370 pp. 

1966. Phylogenetic systematics. University 
of Illinois Press, Urbana, 263 pp. 

HENNINGS, D., AND R. S. HOFFMANN. 1977. A review 
of the taxonomy of the Sorex vagrans species com- 
plex from western North America. Occasional Pa- 
pers of the Museum of Natural History, University 
of Kansas, 68:1-35. 

HERSHKOVITZ, P. 1966. Catalog of living whales. Bul- 
letin of the United States National Museum, 246:1- 
259. 

HOFFMEISTER, D. F. 1951. A taxonomic and evolu- 
tionary study of the pinion mouse, Peromyscus true. 
Illinois Biological Monographs, 21:1-104. 

1986. Mammals of Arizona. The University 

of Arizona Press, Tucson, 602 pp. 

. 1989. Mammals of Illinois. University of 
Illinois Press, Urbana, 348 pp. 

HOFFMEISTER, D. F., AND V. E. DiERSING. 1978. Re- 
view of the tassel-eared squirrels of the subgenus 
Otosciurus. Journal of Mammalogy, 59:402-413. 

HOFFMEISTER, D. F., AND K. B. STERLING. 1994. Or- 
igin. Pp. 1-21, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1433. 

HOoneYCUTT, R. L., AND S. L. WILLIAMS. 1982. Genic 
differentiation in pocket gophers of the genus Pap- 
pogeomys, with comments on intergeneric relation- 
ships in the subfamily Geomyinae. Journal of Mam- 
malogy, 63:208-217. 

Honeycutt, R.L.,ANDT.L. YATgEs. 1994. Molecular 
systematics. Pp. 288-309, in Seventy-five years of 
mammalogy (1919-1994) (E. C. Birney and J. R. 
Choate, eds.). Special Publication, The American 
Society of Mammalogists, | 1:1-433. 

Hoop, C.S., AND J. D. SMitH. 1982. Cladistical anal- 
ysis of female reproductive histomorphology in 
phyllostomatoid bats. Systematic Zoology, 31:241- 
251. 

Hooper, E. T. 1943. Geographic variation in harvest 
mice of the species Reithrodontomys humulis. Oc- 
casional Papers of the Museum of Zoology, Univer- 
sity of Michigan, 477:1-19. 

1952. A systematic review of harvest mice 

(genus Reithrodontomys) of Latin America. Miscel- 

laneous Publications of the Museum of Zoology, 

University of Michigan, 77:1-255. 


1968. Classification. Pp. 27-74, in Biology 
of Peromyscus (Rodentia) (J. A. King, ed.). Special 
Publication, The American Society of Mammalo- 
gists, 2:1-593. 

1972. A synopsis of the rodent genus Scoti- 
nomys. Occasional Papers of the Museum of Zool- 
ogy, University of Michigan, 665:1—32. 

Howe, A. B. 1926. Voles of the genus Phenacomys. 
North American Fauna, 48:1-66. 

1927. Revision of the American lemming 
mice (genus Synaptomys). North American Fauna, 
50:1-37. 

Howe i, A. H. 1921. A biological survey of Ala- 
bama. North American Fauna, 45:1-88. 

. 1929. Revision of the American chipmunks. 

North American Fauna, 52:1-157. 

. 1938. Revision of the North American ground 
squirrels, with a classification of the North American 
Sciuridae. North American Fauna, 56:1-—256. 

Huckasy, D. G. 1980. Species limits in the Pero- 
myscus mexicanus group (Mammalia: Rodentia: 
Muroidea). Contributions in Science, Natural His- 
tory Museum of Los Angeles County, 326:1-—24. 

Hux ey, J.S.(Ep.). 1940. The new systematics. Clar- 
endon Press, Oxford, 583 pp. 

JAcKSON, H. H. T. 1928. A taxonomic review of the 
American long-tailed shrews. North American Fau- 
na, 51:1-238. 

. 1961. Mammals of Wisconsin. University of 
Wisconsin Press, Madison, 504 pp. 

JOHNSON, M. L., AND S. B. GeorGE. 1991. Species 
limits within the Aborimus longicaudus species— 
complex (Mammalia: Rodentia) with a description 
of a new species from California. Contributions in 
Science, Natural History Museum of Los Angeles 
County, 429:1-16. 

Jones, J. K., JR. 1964. Distribution and taxonomy 
of mammals of Nebraska. University of Kansas Pub- 
lications, Museum of Natural History, 16:1-356. 

1982. [Review of] The mammals of North 

America, Second ed., by E. R. Hall. Journal of Mam- 

malogy, 63:717-718. 

1991. Genealogy of Twentieth-century sys- 
tematic mammalogists in North America: the de- 
scendants of Joseph Grinnell. Pp. 48-56, in Latin 
American mammalogy: history, biodiversity, and 
conservation (M. A. Mares and D. J. Schmidly, eds.). 
University of Oklahoma Press, Norman, 468 pp. 

Jones, J. K., Jk., D. M. ARMSTRONG, R. S. HOFFMANN, 
AND C. Jones. 1983. Mammals of the northern 
Great Plains. University of Nebraska Press, Lincoln, 
379 pp. 

Kurscu, J. A. W., R. E. BLetwetss, A. W. DICKERMAN, 
AND O. A. ReIG. 1993. DNA/DNA hybridization 
studies of carnivorous marsupials. III. Relationships 
among species of Didelphis (Didelphidae). Journal 
of Mammalian Evolution, 1:75-97. 

Kirscu, J. A. W., A. W. DICKERMAN, O. A. REIG, AND 
M. S. SprinGerR. 1991. DNA hybridization evi- 
dence for the affinity of the American marsupial 
Dromiciops australis. Proceedings of the National 
Academy of Sciences, USA, 88:10465-10469. 

Koopman, K. F. 1984. A synopsis of the families of 
bats—part VII. Bat Research News, 25:25-27. 

Lawior, T.E. 1969. A systematic study of the rodent 


TAXONOMY 197 


genus Ofotylomys. Journal of Mammalogy, 50:28— 
42. 

Lipicker, W. Z., Jr. 1960. Ananalysis of intraspecific 
variation in the kangaroo rat Dipodomys merriami. 
University of California Publications in Zoology, 67: 
125-218. 

1962. The nature of subspecific boundaries 
in a desert rodent and its implications for subspecific 
taxonomy. Systematic Zoology, 11:160-171. 

Lim, B. K. 1993. Cladistic reappraisal of Neotropical 
stenodermatine bat phylogeny. Cladistics, 9:147-165. 

Lona, C. A. 1965. The mammals of Wyoming. Uni- 
versity of Kansas Publications, Museum of Natural 
History, 14:493-758. 

Lowery, G.H.,Jr. 1974. The mammals of Louisiana 
and its adjacent waters. Louisiana State University 
Press, Baton Rouge, 565 pp. 

MARSHALL, L. G., J. A. CASE, AND M. O. WoODBURNE. 
1990. Phylogenetic relationships of the families of 
marsupials. Pp. 433-505, in Current Mammalogy 
(H. H. Genoways, ed.). Plenum Press, New York, 
2:1-577. 

Mayr, E. 1963. Animal species and evolution. The 
Belknap Press of Harvard University Press, Cam- 
bridge, Massachusetts, 797 pp. 

. 1969. Principles of systematic zoology. Mc- 

Graw-Hill Book Co., New York, 428 pp. 

1982. The growth of biological thought: di- 
versity, evolution, and inheritance. The Belknap Press 
of Harvard University Press, Cambridge, 974 pp. 

Mayr, E., AND P. D. AsHLock. 1991. Principles of 
systematic zoology. Second ed. McGraw-Hill Inc., 
New York, 475 pp. 

Mayr, E., E. G. LINsSLEy, AND R. L. Ustncer. 1953. 
Methods and principles of systematic zoology. Mc- 
Graw-Hill Book Co., New York, 336 pp. 

McKenna, M.C. 1975. Toward a phylogenetic clas- 
sification of the Mammalia. Pp. 21-46, in Phylogeny 
of the primates: a multidisciplinary approach (W. P. 
Luckett and F. S. Szalay, eds.). Plenum Publishing 
Co., New York, 483 pp. 

1987. Molecular and morphological analysis 
of high-level mammalian interrelationships. Pp. 55- 
93, in Molecules and morphology in evolution: con- 
flict or compromise? (C. Patterson, ed.). Cambridge 
University Press, Cambridge, Great Britain, 229 pp. 

MckKirtrick, M. C., AND R. M. Zink. 1988. Species 
concepts in ornithology. Condor, 90:1-14. 

Mearns, E. A. 1907. Mammals of the Mexican 
boundary of the United States ... Pt. 1, Families 
Didelphidae to Muridae. Bulletin of the United States 
National Museum, 56:1—530. 

MerrRIAM, C.H. 1918. Review of the grizzly and big 
brown bears of North America (genus Ursus) with 
description of a new genus, Vetularctos. North 
American Fauna, 41:1-36. 

1919. Criteria for the recognition of species 
and genera. Journal of Mammalogy, 1:6-9. 

Me_rRriItTT, J. F. 1987. Guide to the mammals of Penn- 
sylvania, University of Pittsburgh Press, Pittsburgh, 
Pennsylvania, 408 pp. 

Miiter, G.S., Jr. 1912. List of North American land 
mammals in the United States National Museum, 
1911. Bulletin of the United States National Mu- 
seum, 79:1-455. 


. 1924. List of North American Recent mam- 
mals. Bulletin of the United States National Muse- 
um, 128:1-673. 

MILter, G. S., JR., AND G. M. ALLEN. 1928. The 
American bats of the genera Myotis and Pizonyx. 
Bulletin of the United States National Museum, 144: 
1-218. 

MILLER, G. S., JR., AND R. KELttoGc. 1955. List of 
North American Recent mammals. Bulletin of the 
United States National Museum, 205:1-954. 

Miter, G. S., JR., AND J. A. G. REHN. 1902. Sys- 
tematic results of the study of North American land 
mammals to the close of the year 1900. Proceedings 
of the Boston Society of Natural History, 30:1-352. 

MryaAmorto, M. M., AND M.GoopMAN. 1986. Biomo- 
lecular systematics of eutherian mammals: phylo- 
genetic patterns and classification. Systematic Zo- 
ology, 35:230-240. 

MryaAmorto, M. M.,S. M. TANHAUSER, AND P. J. Larpis. 
1989. Systematic relationships in the artiodactyl tribe 
Bovini (Family Bovidae), as determined from mi- 
tochondrial DNA sequences. Systematic Zoology, 38: 
342-349, 

Moorg, J. C. 1959. Relationships among the living 
squirrels of the Sciurinae. Bulletin of the American 
Museum of Natural History, 118:157-206. 

Mumrorp, R. E., AND J. O. WHITAKER, JR. 1982. 
Mammals of Indiana. Indiana University Press, 
Bloomington, 537 pp. 

Musser, G. G. 1968. A systematic study of the Mex- 
ican and Guatemalan gray squirrel, Sciurus aureo- 
gaster F. Cuvier (Rodentia: Sciuridae). Miscella- 
neous Publications, Museum of Zoology, University 
of Michigan, 137:1-112. 

NELSON, E. W., AND E. A. GOLDMAN. 1933. Revision 
of the jaguars. Journal of Mammalogy, 14:221-240. 

Novacek, M. J. 1982. Information for molecular 
studies from anatomical and fossil evidence on high- 
er eutherian phylogeny. Pp. 3-41, in Macromolec- 
ular sequences in systematic and evolutionary bi- 
ology (M. Goodman, ed.). Plenum Press, New York, 
418 pp. 

. 1986. The skull of leptictid insectivorans and 

the higher-level classification of eutherian mammals. 

Bulletin of the American Museum of Natural His- 

tory, 183:1-112. 

1989. Higher mammal phylogeny: the mor- 

phological-molecular synthesis. Pp. 421-435, in The 

hierarchy of life (B. Fernholm, K. Bremer, and H. 

Jornvall, eds.). Elsevier Science Publishers B.V. 

(Biomedical Division), 499 pp. 

. 1990. Morphology, paleontology, and the 

higher clades of mammals. Pp. 507-543, in Current 

Mammalogy (H. H. Genoways, ed.). Plenum Press, 

New York, 2:1-577. 

. 1993. Reflections on higher mammalian phy- 
logenetics. Journal of Mammalian Evolution, 1:3- 
30. 

Novacek, M.J., ANDA. R. Wyss. 1986. Higher-level 
relationships of the Recent eutherian orders: mor- 
phological evidence. Cladistics, 2:257-287. 

Novacek, M. J., A. R. Wyss, AND M. C. MCKENNA. 
1988. The major groups of eutherian mammals. Pp. 
31-71, in The phylogeny and classification of the 


198 ENGSTROM ET AL. 


tetrapods, volume 2: mammals (M. J. Benton, ed.). 
Clarendon Press, Oxford, 329 pp. 

Oscoop, W. H. 1909. Revision of the mice of the 
American genus Peromyscus. North American Fau- 
na, 28:1—285. 

Owen, R. D. 1987. Phylogenetic analyses of the bat 
subfamily Stenodermatinae (Mammalia: Chirop- 
tera). Special Publication, The Museum, Texas Tech 
University, 26:1-65. 

. 1988. Phenetic analyses of the bat subfamily 
Stenodermatinae (Chiroptera: Phyllostomidae). 
Journal of Mammalogy, 69:795-8 10. 

PACHECO, V., AND B. D. PATTERSON. 1991. Phylo- 
genetic relationships of the New World genus Stur- 
nira (Chiroptera: Phyllostomidae). Bulletin of the 
American Museum of Natural History, 206:101-121. 

PACKARD, R. L. 1960. Speciation and evolution of 
the pygmy mice, genus Baiomys. University of Kan- 
sas Publications, Museum of Natural History, 9:579- 
670. 

Patton, J. L., AND M. F. SmitrH. 1981. Molecular 
evolution in Thomomys: phyletic systematics, para- 
phyly, and rates of evolution. Journal of Mammal- 
ogy, 62:493-500. 

1990. The evolutionary dynamics of the 
pocket gopher, Thomomys bottae, with emphasis on 
California populations. University of California 
Publications in Zoology, 123:1-161. 

Patton, J. L., S. W. SHERWOOD, AND S. Y. YANG. 
1981. Biochemical systematics of chaetodipine 
pocket mice, genus Perognathus. Journal of Mam- 
malogy, 62:477-492. 

PETERSON, R.L. 1966. The mammals of eastern Can- 
ada. Oxford University Press, Toronto, 465 pp. 

Pine, R. H. 1972. The bats of the genus Carollia. 
Technical Monograph, Texas Agricultural Experi- 
ment Station, Texas A&M University, 8:1-125. 

RAMIiREZ-PULIDO, J., AND M. C. Britron. 1981. An 
historical synthesis of the Mexican mammalian tax- 
onomy. Proceedings of the Biological Society of 
Washington, 91:1-17. 

RAMiREZ-PULIDO, J., M. C. Brrrron, A. PERDOMO, AND 
A. Castro. 1986. Guia de los mamiferos de Méxi- 
co, referencias hasta 1983. Universidad Aut6noma 
Metropolitana, Unidad Iztapalapa, México, D. F., 
720 pp. 

RICHARDSON, J. 1829. Fauna boreali-Americana; or 
the zoology of the northern parts of British America: 
containing descriptions of the objects of natural his- 
tory collected on the northern land expedition under 
the command of Captain Sir John Franklin, R. N. 
Part first, containing the quadrupeds. John Murray 
(publishers), Ltd., London, 300 pp. 

Ripe, W.D.L. 1964. A review ofthe Australian fossil 
marsupials. Journal Proceedings of the Royal Society 
of Western Australia, 47:97-131. 

Rossins, L. W., AND V. M. SaricH. 1988. Evolu- 
tionary relationships in the family Emballonuridae 
(Chiroptera). Journal of Mammalogy, 69:1-13. 

Rocers, D. S., AND M. D. ENGstrom. 1992. Evo- 
lutionary implications of allozymic variation in trop- 
ical Peromyscus of the mexicanus species group. 
Journal of Mammalogy, 73:55-69. 

Rocers, D. S., AND D. J. ScomipLy. 1982. System- 
atics of spiny pocket mice (genus Heteromys) of the 
desmarestianus species group from Mexico and 


northern Central America. Journal of Mammalogy, 
63:375-386. 

Rocers, D. S., I. F. GREENBAUM, S. J. GUNN, AND M. 
D.ENGsTrRoM. 1984. Cytosystematic value of chro- 
mosomal inversion data in the genus Peromyscus. 
Journal of Mammalogy, 65:457-465. 

RussELL, R. J. 1968a. Evolution and classification of 
the pocket gophers of the subfamily Geomyinae. 
University of Kansas Publications, Museum of Nat- 
ural History, 16:473-579. 

19685. Revision of pocket gophers of the 
genus Pappogeomys. University of Kansas Publi- 
cations, Museum of Natural History, 16:581-776. 

SANBORN, C.C. 1937. American bats of the subfamily 
Emballonurinae. Field Museum of Natural History, 
Zoological Series, 20:321-354. 

SCHMIDLY, D. J. 1977. The mammals of Trans-Pecos 
Texas. Texas A&M University Press, College Sta- 
tion, 225 pp. 

. 1983. Texas mammals east of the Balcones 
Fault Zone. Texas A&M University Press, College 
Station, 400 pp. 

SCHMIDLY, D. J., R. D. BRADLEY, AND P. S. CaTo. 
1988. Morphometric differentiation and taxonomy 
of three chromosomally characterized groups of 
Peromyscus boylii from east-central Mexico. Journal 
of Mammalogy, 69:462-480. 

SCHNELL, G. D., T. L. Best, AND M. L. KENNEDY. 
1978. Interspecific morphologic variation in kan- 
garoo rats (Dipodomys): degree of concordance with 
genic variation. Systematic Zoology, 27:34-48. 

SETzER, H. W. 1949. Subspeciation in the kangaroo 
rat, Dipodomys ordii. University of Kansas Publi- 
cations, Museum of Natural History, 1:473-573. 

SILVA-TABOADA, G. 1979. Los murciélagos de Cuba. 
Editorial Academia, 423 pp. 

Smpson, G. G. 1945. The principles of classification 
and a classification of mammals. Bulletin of the 
American Museum of Natural History, 85:1-350. 

1961. Principles of animal taxonomy. Co- 
lumbia University Press, New York, 247 pp. 

SmitH, J. D. 1972. Systematics of the chiropteran 
family Mormoopidae. Miscellaneous Publications, 
Museum of Natural History, University of Kansas, 
56:1-132. 

SmiTH, J. D., AND G. MApxour. 1980. Penial mor- 
phology and the question of chiropteran phylogeny. 
Pp. 347-365, in Proceedings of the fifth international 
bat research conference (D. E. Wilson and A. L. 
Gardner, eds.). Texas Tech University Press, Lub- 
bock, 434 pp. 

Smitu, M. F., AND J. L. PATron. 1988. Subspecies 
of pocket gophers: causal bases for geographic dif- 
ferentiation in Thomomys bottae. Systematic Zool- 
ogy, 37:163-178. 

SNEATH, P. H. A., ANDR. R. SOKAL. 1973. Numerical 
taxonomy: the principles and practice of numerical 
classification. W. H. Freeman and Co., San Fran- 
cisco, 573 pp. 

SOKAL, R. R., AND T. J. CROVELLO. 1970. The bio- 
logical species concept: a critical explanation. The 
American Naturalist, 104:127-153. 

SOKAL, R. R., AND P. H. A. SNEATH. 1963. Principles 
of numerical taxonomy. W. H. Freeman and Co., 
San Francisco, 359 pp. 

SULLIVAN, R. M. 1985. Phyletic, biogeographic and 


TAXONOMY oe) 


ecological relationships among montane populations 
of least chipmunks (Eutamias minimus) in the 
Southwest. Systematic Zoology, 34:419-448. 

SULLIVAN, R. M., D. J. HAFNER, AND T. L. YATES. 
1986. Genetics of a contact zone between three 
chromosomal forms of the grasshopper mouse (ge- 
nus Onychomys): a reassessment. Journal of Mam- 
malogy, 67:640-659. 

SZALAY, F.S. 1977. Phylogenetic relationships and a 
classification of the eutherian Mammalia. Pp. 315- 
374, in Major patterns in vertebrate evolution (M. 
K. Hecht, P. C. Goody, and B. M. Hecht, eds.). 
Plenum Press, New York, 908 pp. 

1982. A new appraisal of marsupial phylog- 
eny and classification. Pp. 621-640, in Carnivorous 
marsupials (M. Archer, ed.). Royal Zoological So- 
ciety of New South Wales, Mosman, 2:397-804. 

Tate, G. H. H. 1933. A systematic revision of the 
marsupial genus Marmosa. Bulletin of the American 
Museum of Natural History, 66:1—250. 

TAVERNER, P. A. 1920. The test of the subspecies. 
Journal of Mammalogy, 1:124—-127. 

THAELER, C. S., JR. 1972. Taxonomic status of the 
pocket gophers, Thomomys idahoensis and Tho- 
momys pygmaeus (Rodentia: Geomyidae). Journal 
of Mammalogy, 53:417-428. 

VAN GELDER, R. G. 1959. A taxonomic revision of 
the spotted skunks (genus Spi/ogale). Bulletin of the 
American Museum of Natural History, 1 17:229-392. 

VAN ZYLL DE JONG, C. G. 1984. Taxonomic rela- 
tionships of Nearctic small-footed bats of the Myotis 
leibii group (Chiroptera: Vespertilionidae). Canadian 
Journal of Zoology, 62:2519-2526. 

VAN ZYLL DE JONG, C. G., AND G. L. KIRKLAND, JR. 
1989. A morphometric analysis of the Sorex cine- 
reus group in central and eastern North America. 
Journal of Mammalogy, 70:1 10-122. 

Varona, L. S. 1974. Catalogo de los mamiferos vi- 
vientes y extinguidos de las Antillas. Instituto de 
Zoologia, Academia de Ciencias de Cuba (Havana), 
139 pp. 

ViLLA-R., B. 1967. Los murciélagos de México. Ana- 
les de Instituto de Biologia, Universidad Nacional 
Autonoma de México, México, D.F., 491 pp. 

Voss, R.S. 1988. Systematics and ecology of ichthyo- 
myine rodents (Muroidea): patterns of morpholog- 
ical evolution in a small adaptive radiation. Bulletin 
of the American Museum of Natural History, 188: 
259-493. 

WALL, D. A., S. K. Davis, AND B. M. Reap. 1992. 
Phylogenetic relationships in the subfamily Bovinae 
(Mammalia: Artiodactyla) based on ribosomal DNA. 
Journal of Mammalogy, 73:262-275. 

WEBSTER, W. D., AND C. O. HANDLEY, JR. 1986. Sys- 
tematics of Miller’s long-tongued bat Glossophaga 
longirostris, with description of two new subspecies. 
Occasional Papers, The Museum, Texas Tech Uni- 
versity, 100:1-22. 


WHITAKER, J. O., JR. 1994. Academic Propinquity. 
Pp. 121-138, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1-433. 

Wiey, E.O. 1978. The evolutionary species concept 
reconsidered. Systematic Zoology, 28:88-92. 

1981. Phylogenetics: the theory and practice 
of phylogenetic systematics. John Wiley and Sons, 
New York, 439 pp. 

WILEY, E. O., D. SIEGEL-CAusEy, D. R. BROOKS, AND 
V. A. Funk. 1991. The compleat cladist: a primer 
of phylogenetic procedures. Special Publication, The 
University of Kansas, Museum of Natural History, 
19:1-158. 

WILLIAMS, D. F. 1978. Systematics and ecogeograph- 
ic variation of the Apache pocket mouse (Rodentia: 
Heteromyidae). Bulletin of Carnegie Museum of 
Natural History, 10:1-57. 

WILLIAMS, D. F., AND H. H. GeNoways. 1979. A 
systematic review of the olive-backed pocket mouse, 
Perognathus fasciatus (Rodentia: Heteromyidae). 
Annals of Carnegie Museum, 48:1-36. 

WILSON, D. E., AND D. M. REEDER. 1993. Mammal 
species of the world: a taxonomic and geographic 
reference. Second ed. Smithsonian Institution Press, 
Washington, D.C., 1,206 pp. 

WILSON, E. O., AND W. L. Brown, Jr. 1953. The 
subspecies concept and its taxonomic application. 
Systematic Zoology, 2:97-111. 

WOZENCRAFT, W.C. 1989a. The phylogeny of Recent 
Carnivora. Pp. 495-535, in Carnivore behavior, 
ecology, and evolution (J. L. Gittleman, ed.). Cornell 
University Press, Ithaca, New York, 620 pp. 

WOZENCRAFT, W.C. 19895. Classification of the Re- 
cent Carnivora. Pp. 569-593, in Carnivore behavior, 
ecology, and evolution (J. L. Gittleman, ed.). Cornell 
University Press, Ithaca, New York, 620 pp. 

Wyss, A. R. 1989. Flippers and pinniped phylogeny: 
has the problem of convergence been overrated? Ma- 
rine Mammal Science, 5:343-360. 

Wyss, A. R., M. J. NovAcek, AND M. C. MCKENNA. 
1987. Amino acid sequence versus morphological 
data and the interordinal relationships of mammals. 
Molecular Biology and Evolution, 4:99-116. 

YATES, T. L., AND D. J. ScHMIDLY. 1977. Systematics 
of Scalopus aquaticus in Texas and adjacent states. 
Occasional Papers, The Museum, Texas Tech Uni- 
versity, 45:1-36. 

YOUNGMAN, P. M. 1975. Mammals of the Yukon 
Territory. National Museum of Natural Sciences 
(Ottawa), Publications in Zoology, 10:1-192. 

ZIMMERMAN, E.G. 1970. Karyology, systematics and 
chromosomal evolution in the rodent genus Sig- 
modon. Publications of The Museum, Michigan State 
University Biological Series, 4:385-454. 


PALEOMAMMALOGY 


RICHARD J. ZAKRZEWSKI AND JASON A. LILLEGRAVEN 


Introduction 


Ne differences between neo- and pa- 
leomammalogy already existed as 
early as 1919, primarily because of the na- 
ture of the materials researched and the 
technologies that could be utilized. Even so, 
paleomammalogists have made major ad- 
vances toward a better understanding of 
mammalogy since the founding of the so- 
ciety. These contributions can be consid- 
ered under four areas: general, geological, 
biological, and a blending of the latter two. 
General advances include a significant in- 
crease in the number of individuals and in- 
stitutions working in paleomammalogy; a 
tremendous increase in the size of collec- 
tions, especially of smaller taxa, due to the 
development and modification of screen 
washing techniques; a better understanding 
of the fossilization process through tapho- 
nomic studies; and development of a com- 
prehensive bibliography. Geologically ori- 
ented advances include the use of improved 
biostratigraphic techniques, together with 
radiometric dating and magnetostratigra- 
phy, to increase our understanding of the 
sequential occurrence of mammalian fau- 
nas and decipher the complex geology of the 
highly deformed ranges and intermontane 
basins in the American West; and use of the 


plate tectonics model to explain biogeo- 
graphic distributions and patterns. Biolog- 
ically oriented advances include major im- 
provements in our understanding of the 
reptile-mammal transition and the defini- 
tion of ‘““mammal’’; important systematic 
studies of many mammalian taxa, using 
various taxonomic philosophies; and mul- 
titudinous studies of form, function, and 
phylogenetic relationships of particular 
groups of Cenozoic mammals. These have 
all been blended into important studies con- 
sidering the issues of tempo and mode in 
evolution and the cause of extinctions. 


Compartmentalization of 
Mammalogy 


By 1919, the discipline of mammalogy 
already had become compartmentalized into 
neo- and paleomammalogy. Osborn (1921), 
in the first article of Volume 2 of the Journal 
of Mammalogy, pointed out that paleo- 
mammalogists were constrained to dealing 
only with hard parts and, therefore, the types 
of studies that were undertaken usually were 
different from those of neomammalogists. 


200 


PALEOMAMMALOGY 201 


However, he further suggested that there 
should be more standardization of terms 
and approaches to research problems in 
mammalogy, as well as cooperative studies 
between neo- and paleomammalogists for 
their mutual benefit. He cited, as an ex- 
ample, the work on rodents by Miller and 
Gidley (1918). Unfortunately, such collab- 
orations have been infrequent (e.g., Carle- 
ton and Eshelman, 1979; White and Keller, 
1984). A classic exception is the work of the 
late John E. Guilday who, perhaps as well 
as anyone in this century, fused mammal- 
ogy and paleomammalogy (e.g., Guilday, 
1971). The less-than-expected level of in- 
terchange between paleo- and neomam- 
malogists probably relates to a perception 
that the former still are constrained to 
studying only hard parts in a geological con- 
text, whereas the latter have even more av- 
enues and methods of study available to 
them now than existed 75 years ago. Also, 
geophysical advances of the 20th Century 
that are critically important to geologically 
oriented paleomammalogists often have ap- 
peared scientifically irrelevant to neomam- 
malogists. Perhaps advances in specialized 
technology themselves have led to wider 
gulfs between subdisciplines of mammalo- 
gy. Be that as it may, paleomammalogists 
have made major contributions to the gen- 
eral field of mammalogy, and a small sam- 
pling of these is considered below. 
Analogous to the dichotomy between neo- 
and paleomammalogists, there exists sig- 
nificant compartmentalization among pa- 
leomammalogists. The splits result, in part, 
from interest and training, but also stem 
from the use of fossils in approaching geo- 
logical versus biological problems. Some 
paleontologists (who might prefer to be 
called mammalian biostratigraphers) study 
fossil mammals principally to determine the 
age of enclosing sediments to solve strati- 
graphic or structural problems; paleomam- 
malogists sensu stricto, like neomammal- 
ogists, typically are more interested in 
anatomical and functional problems and 
evolutionary implications associated with 


fossils. Many paleontologists, however, have 
attempted to work in both areas. 

Before considering contributions made in 
these two areas, we briefly examine impor- 
tant developments of a general nature that 
have led to enormous benefit both within 
geologically and biologically oriented paleo- 
mammalogy. These include an increase 
in the number of paleomammalogists, tech- 
niques in collecting fossils, understanding 
how particular associations of fossils come 
to be, and the development of a compre- 
hensive bibliography. 


General Advancements 


A Slow Start for American 
Paleomammalogy 


When the ASM was founded in New York 
on 3 April 1919, there were only two major 
centers of mammalian paleontology in the 
United States. One at the American Mu- 
seum of Natural History, led by Henry Fair- 
field Osborn, William Diller Matthew, and 
Childs Frick, the other at the University of 
California, Berkeley, where John C. Mer- 
riam had built a program. About the time 
the ASM was founded, Merriam became 
president of the Carnegie Institution of 
Washington. He was among the charter 
members of the society, together with col- 
leagues from New York and the Smithson- 
ian Institution. Merriam and Matthew were 
two of the first council members of the so- 
ciety. Merriam also served on the first Anat- 
omy and Physiology Committee, together 
with William King Gregory of Columbia 
University and Alexander Wetmore and 
James W. Gidley of the Smithsonian Insti- 
tution. Matthew served as President of the 
society in 1926, the only paleomammalogist 
to have done so. 

Why there were so few centers of mam- 
malian paleontology as late as 1919 is not 
clear. Perhaps it was, in part, a legacy from 
the days of Cope and Marsh, when the 1m- 


202 ZAKRZEWSKI AND LILLEGRAVEN 


petus was for the collection of large reptiles 
from Mesozoic deposits of the American 
West. This tendency extended into early 
parts of the 20th Century with collections 
made by Earl Douglass for the Carnegie Mu- 
seum in Pittsburgh and the Sternbergs for 
various Canadian institutions. Many of the 
paleontologists in the first quarter of the 20th 
Century were interested principally in lower 
vertebrates rather than mammals. In marked 
contrast, there exist today in North America 
about 120 institutions in which individuals 
perform research on fossil mammals; esti- 
mating conservatively, at least 20 of these 
research centers must be considered major. 
Such increase in interest since 1919 must, 
in itself, be considered a huge advance in 
American paleomammalogy. 


Finding the Tiny 


Philip D. Gingerich (1986), in lamenting 
the demise of the paleontology program at 
his alma mater, aptly showed that theoret- 
ically oriented paleontology depends upon 
extensive personal experience based, in turn, 
upon a solid data base. For all paleontolog- 
ical endeavors, the fundamental objective 
elements of data are the fossils themselves, 
set within geological contexts. Early collec- 
tors of fossil mammals, perhaps influenced 
by their predecessors’ searches for dino- 
saurs, selectively looked for sites with ac- 
cumulations of large mammals. Although 
small mammals certainly were not inten- 
tionally ignored, quarrying techniques em- 
ployed by many early collectors were not 
conducive to discovery of minuscule fossils. 
A “fossil” to many of these individuals had 
to be at least six inches long, preferably 
bearing teeth. 

A change in attitude began about a decade 
after the founding of the ASM. In 1928, 
Claude W. Hibbard (a future director of the 
ASM) was hired as cook and camp caretaker 
for a field party from the University of Kan- 
sas led by Handel Tong Martin. The crew 


was returning to Edson Quarry (late Mio- 
cene, Sherman County, Kansas) for another 
summer of collecting. During the previous 
summer, Martin had found some fossil sal- 
amander bones and was asked to collect ad- 
ditional remains by an anatomist who was 
interested in studying the group. When Hib- 
bard went to the quarry, after finishing camp 
chores, Martin greeted him with a pair of 
tweezers and told him to collect all the small 
bone he could find on the spoil pile. Hibbard 
soon decided to expedite matters. Obtaining 
some window screen from the local rancher, 
he attached the screen to a wooden frame 
to produce a little box. Loose sediment from 
the spoil pile passed easily through the screen 
and the fossil bone was trapped by it and 
picked out. Hibbard thought he might hurry 
the process even more by the use of water. 
Thus, he took the sediment and his box to 
a nearby buffalo wallow and proceeded to 
agitate the box in the water. Within a few 
days, he had enough small material to fill a 
large matchbox. When Hibbard showed the 
material to Martin, the latter stated that there 
were enough small specimens in the box to 
keep paleontologists occupied for years. De- 
spite the innovation, Hibbard spent the re- 
mainder of the summer in the quarry, help- 
ing Martin collect “real” (1.e., large) fossils. 

Subsequently, Hibbard (1949) expanded 
on the washing technique and used it to 
accumulate tens of thousands of specimens 
from southwestern Kansas and northwest- 
ern Oklahoma. Thereby, he was able to doc- 
ument a sequence of faunas that reflected 
both phylogenetic and climatic change 
(Bayne, 1976; Zakrzewski, 1975). Subse- 
quent workers (e.g., McKenna, 1962) have 
modified the technique for massive collec- 
tion of fossils from other areas and ages. An 
example of the importance of widespread 
use of screen-washing techniques is the in- 
crease in our knowledge of Mesozoic mam- 
mals. When George Gaylord Simpson (1928, 
1929) published his comprehensive sum- 
mary of known Mesozoic mammals (based, 
in part, on his Ph.D. thesis), he worked with 


PALEOMAMMALOGY 203 


fewer than a thousand specimens, collected 
by standard quarrying from around the 
world. When William A. Clemens, Jr. (1963, 
1966, 1973) and Jason A. Lillegraven (1969) 
published their Ph.D. theses on latest Cre- 
taceous mammals, their specimens from 
only two local faunas numbered well into 
the thousands. Mammalian paleontology in 
North America, especially dealing with the 
Mesozoic, was never quite the same again. 
As Simpson (1971) stated, it “would not be 
possible now, as it was in 1871, 1888, and 
1928-1929 for one person to treat all avail- 
able material on Mesozoic mammals... .” 


Grasping the “How” of Fossil 
Accumulations 


Most workers are painfully aware of im- 
portant biases in the fossil record. Before 
useful scientific inferences can be drawn 
from paleontological data, one needs to 
know how the fossils themselves accumu- 
lated. Although inadequacies and biases in 
the fossil record have been appreciated for 
many years (e.g., Darwin, 1859), it has been 
only relatively recently that formal study of 
the process of fossilization (i.e., taphonomy) 
has been undertaken on a large scale. The 
majority of early taphonomic work was by 
the Russians, applied to faunas of lower ver- 
tebrates (Olson, 1980). Perhaps the seminal 
work in North America for explaining the 
occurrences of accumulations of large mam- 
mals is that of Michael R. Voorhies (1969) 
on the Verdigre Quarry in northeastern Ne- 
braska. Subsequent work by Anna K. Beh- 
rensmeyer and her colleagues (e.g., Beh- 
rensmeyer and Hill, 1980) have added much 
to the understanding of how deposits of fos- 
sil mammals might accumulate. James S. 
Mellett (1974) demonstrated that many mi- 
cromammal accumulations result from owl 
predation, a mechanism suggested earlier by 
Hibbard (1941). Subsequently, problems of 
origin of microvertebrate fossils have been 
addressed by various workers, such as Dod- 


son and Wexlar (1979) and Korth (1979). 
The subdiscipline of taphonomy is only in 
its infancy relative to understanding asso- 
ciations of fossil mammals. 


Unique Research Tool 


The development of a unique biblio- 
graphic research tool cannot be omitted from 
discussion of 20th Century progress in pa- 
leomammalogy; we refer to the Bibliogra- 
phy of Fossil Vertebrates (BFV) (Gregory et 
al., 1989, plus predecessor volumes involv- 
ing various editors, including Charles L. 
Camp). The BFV is published by the Society 
of Vertebrate Paleontology (which shares a 
large membership with the ASM), and pro- 
vides unparalleled, annual access to the 
breadth of world literature on fossil mam- 
mals. 


Geologically Directed 
Paleomammalogy 


Toward a More Useful Time Scale 


Original versions of the geologic time scale 
were developed using the law of superpo- 
sition in combination with the stage of evo- 
lution of marine invertebrates, mostly in- 
volving European rock sequences. Some of 
the sequences could be correlated with those 
in North America using marine inverte- 
brates. Where American continental and 
marine deposits interfingered, there was lit- 
tle problem in placing the terrestrial units 
into a scheme of relative chronology. How- 
ever, as workers moved on to the High Plains 
and into the structurally isolated intermon- 
tane basins of the American West, many 
mammal-bearing nonmarine stratigraphic 
units could not be placed easily into context 
within the standard time scale. As mam- 
mals often were the most abundant fossils 
in these strata, early workers sometimes 
named deposits after the most common 


204 ZAKRZEWSKI AND LILLEGRAVEN 


kinds. Names such as the Equus beds of 
Kansas and the Titanotherium and Oreodon 
beds of South Dakota were established. 
These ill-defined units were assigned to Eu- 
ropean-based Tertiary epochs through com- 
parative estimation of the stage of evolution 
of contained mammals. This procedure of- 
ten involved litthe more than guesswork, 
however, and it ultimately led to wide- 
spread misconceptions in correlation. All 
but one of the standard Tertiary epochs were 
based on marine fossils, and few North 
American continental deposits could be su- 
perpositionally related to marine strata. 
Clearly, a new method for dating and 
correlating the North American mammal- 
bearing continental units had to be devel- 
oped, independent of the standard Euro- 
pean marine sequence. 

Eventually, a committee was established 
to devise such a time scale independent of 
the marine standard. Work of the ‘““Wood 
Committee” led to the development of the 
North American Land-Mammal Ages 
(NALMAs; Wood et al., 1941), as reviewed 
by Hesse (1941). NALMAs were defined 
principally on the first occurrence of certain 
genera and the unique occurrences or con- 
sistent associations of others. Although last 
occurrences also were considered, these 
usually were given less weight because of 
potential complications to correlation re- 
sulting from relictual taxa. The original 
NALMAs applied only to Tertiary time. 
Subsequent to work by the Wood Com- 
mittee, Savage (1951) established the Ir- 
vingtonian and Rancholabrean NALMAs 
for the Pleistocene. Although all NALMAs 
originally were intended to be independent 
of the Lyellian, European-based Tertiary 
epochs, NALMAs inevitably became al- 
most synonymized with Lyellian epochs in 
the minds of geologists and paleontologists 
alike. Such mental linkages (e.g., Bridgerian 
= middle Eocene; Chadronian = early Oli- 
gocene; etc.) have proven highly unfortu- 
nate in the history of North American geo- 
logical research, being a source of much 
confusion in temporal correlation between 
vertebrate paleontologists and traditional 


geologists. Gradually, however, expanded 
use and reliability of radioisotopic dating 
techniques (starting most importantly with 
the pioneering work of Evernden et al., 
1964), in conjunction with data from fossil 
mammals has increased markedly the reli- 
ability of temporal correlation between 
North American nonmarine sequences and 
other parts of the world (see Savage and 
Russell, 1983). 

In 1973, asymposium on Vertebrate Pa- 
leontology and Geochronology was held in 
Dallas at the annual meeting of the Geo- 
logical Society of America. One outcome of 
the symposium was re-establishment of 
committees to refine the various NALMAs. 
After much trial, tribulation, and delay, their 
work resulted in publication of Cenozoic 
Mammals of North America, Geochronol- 
ogy and Biostratigraphy (edited by Wood- 
burne, 1987). 

The use of mammals for biostratigraphic 
purposes reached its acme in the decipher- 
ing of the complex Cenozoic history of 
mountain ranges and intermontane basins 
in western North America. Beginning late 
in the Cretaceous and continuing to the 
present time, most of this area has been 
subjected to major tectonism. Large seg- 
ments of the continental crust experienced 
important displacement, both horizontally 
and vertically. Erosion of zones of defor- 
mation provided sediments that accumu- 
lated to prodigious thicknesses 1n the inter- 
montane basins. Mammalian assemblages, 
involving all Cenozoic NALMAs, have 
proven to be of outstanding utility in the 
relative dating of structural and deposition- 
al histories of western North America. Per- 
haps there exist no better examples of the 
marriage between paleomammalogy and 
historical geology than the various works of 
Galusha and Blick (1971), Dorr et al. (1977), 
Skinner et al. (1977), and Wilson (1978). 


Mobile Continents and Oceanic Basins 


The advent of plate tectonics in the late 
1960s had a profound effect upon American 


PALEOMAMMALOGY 205 


paleomammalogy of the 20th Century. As 
imaginatively summarized by McKenna 
(1973, 1983), general recognition that major 
plates across the surface of the earth were 
mobile (and, by way of seafloor spreading, 
subduction, and collision, could change in 
shape and size through geologic time) rev- 
olutionized the discipline of historical bio- 
geography. The geological impact of plate 
tectonics upon historical biogeography can, 
without exaggeration, be compared to the 
importance of Darwinism within the bio- 
logical sciences. 

It is certainly true that the two editions 
(1915, 1939) of Matthew’s Climate and 
Evolution established the foundations of 
modern historical biogeography. Significant 
additional refinements in principles were 
provided by Simpson (e.g., 1952, 1953a). 
Further, influences on evolutionary thought 
of continental stabilist biogeographic view- 
points issuing from these two eminent 
American paleomammalogists were pro- 
found. Both workers had developed con- 
vincing biogeographical interpretations 
(principally involving fossil mammals) that 
seemingly did not require mobilized con- 
tinents, especially for geologic intervals as 
young as the Cenozoic. 

In essence, it took independent devel- 
opment and observational application of 
new techniques in geophysics (especially pa- 
leomagnetism) to shake the American com- 
munity of geoscientists into accepting the 
reality of highly mobile continents (and ac- 
tively evolving oceanic basins). Interesting- 
ly, much of the European community of 
paleontologists had accepted various forms 
of continental drift far in advance of most 
Americans, even though all proposed phys- 
ical mechanisms seemed inadequate for 
purposes of explanation. Once geophysical- 
ly established, however, American paleo- 
mammalogists jumped solidly onto the 
plate-tectonic bandwagon, and continental 
mobilism has been a fundamental compo- 
nent of their training and research ever since. 
Further, it has been accepted that plate tec- 
tonics is highly relevant in explaining dis- 
tributional patterns of particular groups of 


Cenozoic mammals, such as marsupials 
(e.g., Tedford, 1974; Woodburne and Zins- 
meister, 1984), and even of wholesale con- 
tinental exchanges (e.g., Dawson, 1980; 
Webb, 1985). 

Along with acceptance of a continental 
mobilistic perspective came appreciation of 
a whole series of new possible mechanisms 
(in supplement of Simpsonian corridors, fil- 
ter bridges, and sweepstakes routes) for ex- 
planation of geographic distributions of 
suites of fossils. Some processes involved 
passive transport of already-fossilized as- 
semblages (e.g., the “grounded Viking fu- 
neral ships” of McKenna, 1983), but most 
were pertinent to ancient groups of organ- 
isms at times during which they were still 
alive (e.g., continental ““Noah’s arks”’ of Mc- 
Kenna, 1973: ‘‘escalator counterflow,”’ 
‘hopscotch on the escalator,” and ““voyages 
to nowhere and return” of McKenna, 1983). 
As occurs all too often in the case of real 
progress in scientific understanding, recog- 
nition of the possibility of several of these 
cited mechanisms also has served to com- 
plicate interpretations of historical bioge- 
ography, especially in situations involving 
archipelagos. 


Biologically Directed 
Paleomammalogy 


So What Is a Mammal? 


If one studies only modern-day elements 
of earth’s biota, mammals can be differen- 
tiated easily from all other vertebrate groups. 
As one traces the paleontological history of 
Mammalia back into the middle Mesozoic, 
however, one-by-one the usual features used 
to define what a mammal /s appear in more 
and more primitive stages, becoming blurred 
to generally non-mammalian in therapsid 
ancestors. As a result, paleomammalogists, 
much more than neomammalogists, have 
given attention to definition of the Mam- 
malia, and to questions of phylogenetic re- 
lationships within the class. One result of 
such effort is an interesting paradox. On the 


206 ZAKRZEWSKI AND LILLEGRAVEN 


one hand, the reptile-mammal (or perhaps 
better, the therapsid-mammal or cyno- 
dont—mammal) transformation is better un- 
derstood anatomically than any other in- 
terclass transition within the Vertebrata. But, 
in contrast, and in large part because of the 
deadly combination of great Triassic diver- 
sity, extensive parallel evolution in many 
features, and a generally spotty Mesozoic 
fossil record, the phylogenetic path(s) from 
therapsids toward mammals is (are) ex- 
ceedingly uncertain (compare results, for 
example, among Crompton and Sun, 1985; 
Hopson and Barghusen, 1986; Miao, 1991; 
Rowe, 1988). 


Paleomammalogy and Systematics 


Most of the early paleomammalogists 
were typologists. Each morphological vari- 
ant seemed to demand at least a new specific 
(if not generic) name. Likewise, it seemed 
that scientific reputation and prestige for 
some workers was directly proportional to 
the number of taxa described and named. 
A classic example of this situation was pro- 
vided by E. D. Cope when he named the 
arvicoline genera Anaptogonia and Sycium. 
Anaptogonia, originally considered a sub- 
genus of Arvicola (Cope, 1871), was based 
primarily upon mls of the taxon, whereas 
Sycium was based on upper teeth (Cope, 
1899). Subsequently, Hibbard (1947) dem- 
onstrated that these two taxa were junior 
synonyms for the modern muskrat, Ondat- 
ra. Fortunately, a major advance within pa- 
leomammalogy during the 20th Century has 
been to step away from typological ap- 
proaches to science. 

As the flood of newly discovered fossils 
accumulated in museums, and as masses of 
data became available from new and di- 
verse fields of biological science (e.g., pop- 
ulation genetics), workers in the 1930s and 
1940s tried to integrate all aspects of the 
study of life, as dubbed the ‘“‘new synthesis.” 
Particularly important parts of this integra- 
tion were publication by Simpson of Tempo 
and Mode in Evolution (1944) and The Ma- 
jor Features of Evolution (1953b). The sem- 


inal paper on mammalian interrelation- 
ships 1s The Principles of Classification and 
a Classification of Mammals by Simpson 
(1945). Compiled before WW II, Simpson’s 
classification dealt with every mammalian 
genus known to him, taxonomically utiliz- 
ing the philosophy of the new synthesis. A 
rationale of his approach to the classifica- 
tion of mammals was presented at the 24th 
annual meeting of ASM at the American 
Museum of National History, and the work 
was reviewed in the Journal of Mammalogy 
by E. Raymond Hall (1946). Although in 
many places outdated, Simpson’s work re- 
mains an invaluable taxonomic reference; 
a more detailed compendium has yet to be 
published. 

Toward that end, however, McKenna 
(1975) has updated information for a com- 
prehensive revision of mammalian taxon- 
omy, with development of elaborately an- 
notated computer files. McKenna (1975) 
provided a first approximation of this mon- 
umental work, using cladistic philosophy as 
developed by Willi Hennig (1966). Mc- 
Kenna’s tentative classification remained 
above the level of family, and involved many 
new taxonomic terms that have not been 
readily accepted by the professional com- 
munity. 

Cladistics as a taxonomic philosophy is 
being used increasingly by paleomammal- 
ogists as seen in a recent special volume by 
the Systematics Association edited by Ben- 
ton (1988). A more detailed discussion of 
the cladistic method can be found in Eng- 
strom et al. (1994). Additional synthesis by 
attempting to combine morphological and 
molecular studies in the phylogeny of mam- 
mals can be found in the volumes edited by 
Szalay et al. (1993). 


Knowledge of the First Two-thirds of 
Mammalian History 


Tremendous strides have been made dur- 
ing the 20th Century in documentation of 
Mesozoic mammals. Because few Mesozoic 
mammals have yet proven their potential 


PALEOMAMMALOGY 207 


worth as biostratigraphic tools, most re- 
search on them has been taxonomic or of 
generally biologic nature. Published re- 
search on systematic paleontology of Me- 
sozoic taxa 1S expanding at an astounding 
rate (e.g., Cifelli, 1990; Clemens, 1973; Fox, 
1989), to the point that necessity for taxo- 
nomic and stratigraphic specialization in 
study of Mesozoic mammals has become a 
reality, as has long been the case for Ce- 
nozoic forms. Additionally, major features 
in the origin of tribosphenic molars have 
been worked out (e.g., Crompton, 1971): 
serious attempts have been made at deter- 
mining origins of mammalian metabolic 
pathways (e.g., McNab, 1978); and even 
study of major steps in Mesozoic mam- 
malian reproduction (e.g., Blackburn et al., 
1988) have been approached. Cladistic 
methodology has figured importantly with- 
in comparative studies of detailed anatomy 
of Mesozoic mammals (e.g., Wible and 
Hopson, 1993), largely in pursuit of phy- 
logenetic analysis. Diverse forms of re- 
search (biological and geological) on Me- 
sozoic Mammalia hold promise for an 
unusually bright future. 


Unparalleled Expansion of New 
Biological Information on 
Cenozoic Mammals 


The extent of increased knowledge made 
available since 1919 on comparative anat- 
omy, biological function, paleogeographical 
distribution, and evolutionary relationships 
among Cenozoic mammals is no less than 
astounding. Whole new disciplines of pa- 
leobiological research, such as paleoneurol- 
ogy (e.g., Edinger, 1948; Jerison, 1973; Ra- 
dinsky, 1981), have come into existence. 
Major paleogeographic surprises, such as the 
discovery of North American pangolins 
(Emry, 1970), have occurred. Documenta- 
tion of highly specialized adaptive realms 
for mammalian life, such as origin of pow- 
ered flight (e.g., Jepsen, 1970; Novacek, 
1987) or entry into the sea (e.g., Barnes et 
al., 1985; Domning et al., 1986; Kellogg, 


1936; Repenning et al., 1979) has become 
available. 

Functional studies, varying from mech- 
anisms of mastication (e.g., Krause, 1982) 
to origins of arborealism (e.g., Jenkins, 1974) 
to recognition of the importance of body 
size in ancient mammals (e.g., Damuth and 
MacFadden, 1990), have burgeoned. Fi- 
nally, at least rough phylogenetic frame- 
works have been established for most mam- 
malian orders (e.g., Gazin, 1953; Novacek, 
1990; Prothero and Schoch, 1989; Schoch, 
1986; Simons and Kay, 1983; Wilson, 1986; 
Wood, 1955). Unquestionably, the greatest 
diversity and absolute volume of research 
in 20th Century paleomammalogy has been 
in the documentation of form, function, and 
phylogenetic relationships of particular Ce- 
nozoic taxa. 


The Blending of Geologically and 
Biologically Directed 
Paleomammalogy 


Tempo and Mode in Evolution 


Paleomammalogy can provide unique in- 
formation that is of key importance to the 
research of neomammalogists. Obvious ex- 
amples include paleobiogeographic histo- 
ries and minimum dates of evolutionary di- 
vergence of particular taxa. Potential for 
such useful applications has been recog- 
nized since the origin of paleontology as a 
science. More recently, however, new kinds 
of evolutionary inquiry have resulted from 
the blending of procedural advances de- 
rived jointly from the geological and bio- 
logical sciences. A few examples follow. 

Rates and mechanisms of evolutionary 
change involve questions that have in- 
trigued scientists since the appearance of 
Charles Darwin’s (1859) The Origin of Spe- 
cies. For nearly a century after its publica- 
tion, however, most questions remained 
vaguely posed, with little real progress being 
made toward understanding the detailed na- 
ture of evolutionary modification. Principal 


208 ZAKRZEWSKI AND LILLEGRAVEN 


underlying reasons involved an inadequate- 
ly documented fossil record combined with 
infancy of the science of genetics. Both areas 
were strengthened during the first 40 years 
of the 20th Century, setting the stage for the 
‘new synthesis.” It was in large part the 
greatly improved fossil record of mammals, 
developed through literally centuries of man- 
years of field and laboratory effort, and ex- 
ploited by Simpson (1944, 19535), that 
allowed integration of paleontological 
knowledge with paradigms derived from 
advances in population genetics. Better doc- 
umentation of morphological change 
through geologic time, as based on detailed 
studies of fossil mammals, allowed greater 
scientific focus on tempos of evolution. 
Simpson demonstrated, for example, that 
rates of mammalian evolution varied with- 
in and among taxa. He also noticed that 
paleontologically recognizable change oc- 
curred in spurts and starts, separated by what 
appeared to be extensive intervals of mor- 
phological stability. Simpson was a firm be- 
liever, however, in the essential gradualness 
of evolutionary change, and attributed much 
of the apparent irregularity in rates to strati- 
graphic and geographic imperfections and 
biases within the fossil record. 

More recently, questions of tempo and 
mode in evolution have been reconsidered 
by Eldredge and Gould (1972), using a more 
literal interpretation of the fossil record. 
They suggested that the apparent stasis 
within species, and the paucity of transi- 
tional forms between species, are real, and 
represent ways in which the allopatric mod- 
el of speciation would be expected to be 
reflected in the fossil record. Because of the 
apparent sudden appearance of new species 
in local stratigraphic columns above long 
sections of morphological stasis, they coined 
the term “‘punctuated equilibrium”’ for their 
concept. Although their suggestion origi- 
nally attempted to reconcile the fossil record 
with the concept of allopatric speciation, 
they expanded it subsequently to include 
other features as well, such as the restriction 


of virtually all evolutionary change to the 
process of speciation (Gould, 1985; Gould 
and Eldredge, 1977). The punctuated equil- 
ibrists have been opposed by many neo- 
Darwinists (e.g., Bown and Rose, 1987; 
Gingerich, 1985), who demonstrated strat- 
igraphically controlled gradual change be- 
tween mammalian species in the fossil rec- 
ord; such workers have come to be known 
as phyletic gradualists. Yet a third group 
reached a compromise position, suggesting 
that both patterns have operated, as already 
had been suggested in some cases by earlier 
workers (see Newman et al., 1985). 

Issues involved in the debate cited above 
were summarized by Barnosky (1987), who 
examined results of various studies on Qua- 
ternary mammals. He pointed out that the 
Quaternary should be an ideal geologic in- 
terval for the testing of competing models 
because both time- and species-resolution 
are highly determinable, at least compared 
to the Mesozoic or Tertiary. Case-histories 
cited by Barnosky (1987) demonstrate that 
some species transitions appear to follow 
patterns of punctuated equilibrium, where- 
as others seem to fit more closely models of 
phyletic gradualism. No matter where the 
truth eventually may be shown to lie, all of 
these highly focused studies have depended 
upon elevated standards of detailed, strat- 
igraphically documented collections made 
in the field at levels of thoroughness only 
imagined even when the new synthesis was 
being developed. 


The Spectre of Heterochrony in 
Homotaxy 


Huxley (1870) recognized the spectre of 
heterochrony [i.e., “temporal overlap of as- 
semblages assigned to successive, presumed 
non-overlapping ages, or assemblages as- 
signed to the same age being time trans- 
gressive or not precisely time-equivalent” 
(Flynn et al., 1984)]. Huxley (1870) also ap- 
preciated that the possibility of heterochro- 


PALEOMAMMALOGY 209 


ny cannot be eliminated through applica- 
tion of standard paleontological techniques 
alone. Wisely, he suspected that fully ho- 
motaxic faunas (i.e., taxonomically identi- 
cal assemblages), even when using the most 
closely spaced, stratigraphically controlled 
fossil collections, in reality, might be asyn- 
chronous. Therefore, it is possible that when 
comparing identically changing taxonomic 
assemblages between geographically sepa- 
rated areas, the usual assumption of syn- 
chrony of the assemblages may be incorrect. 
Instead, the geographically separated but 
homotaxic faunas may, for example, have 
been tracking, through time, shifting eco- 
logical regimes. Needless to say, anyone en- 
deavoring to study the tempo and mode of 
evolutionary change must be able to rec- 
ognize absolutely that no significant asyn- 
chrony exists between geographically sep- 
arated, homotaxic faunal assemblages. As 
discussed by Flynn et al. (1984), two recent 
advances from the geological sciences pro- 
vide capabilities, not available in the days 
of Huxley, to better evaluate the possibili- 
ties of heterochrony. 

One advance involves detailed study of 
the record of polarity reversals of earth’s 
magnetic field through orientation of fer- 
romagnetic minerals in individual fossil lo- 
calities (e.g., Lindsay et al., 1981). When 
polarity data are used in combination with 
other, independent dating techniques, it 1s 
often possible to identify particular brief in- 
tervals of earth’s magnetic polarity history. 
The other advancement has been with ra- 
dioisotopic dating, of which a multiplicity 
of suitable isotopes and variations in tech- 
niques is now known to exist. One particular 
variant that is especially promising for ap- 
plication to pre-Pleistocene, mammal-bear- 
ing units is the single-crystal, laser-fusion 
method, involving isotopes of argon 
(Swisher and Prothero, 1990). Through 
combination of detailed paleontology, mag- 
netostratigraphy, and high-resolution ra- 
dioisotopic dating, it is possible (Flynn et 
al., 1984) to recognize the existence of geo- 


graphic migration of ‘“‘age-defining” taxa 
through geologically significant intervals of 
time; however, no entire land-mammal fau- 
na has yet been shown to be heterochronic. 


The Nature of Extinction 


The phenomenon of extinction has in- 
trigued scientists since its possibility was 
first proposed by Hooke in the 1670s (Dott 
and Batten, 1971). Although extinctions 
have occurred throughout the history of life, 
the times of major (or mass) extinctions, 
reputedly concentrated at major geologic 
boundaries, have received the most atten- 
tion. Most of the recent study on extinctions 
by North American paleomammalogists has 
been on those in the proximity of the Cre- 
taceous/Tertiary (K/T) and Pleistocene/ 
Holocene (P/H) boundaries. In both cases, 
the majority of paleomammalogists has fa- 
vored a conservative (i.e., rather gradual- 
istic) point of view in explaining the ex- 
tinctions; others, however, have suggested 
more dramatic scenarios. 

A catastrophic perspective for the K/T 
boundary was presented initially by Alvarez 
et al. (1980), involving a presumed impact 
with earth of a major extraterrestrial body, 
probably an asteroid. Over the following de- 
cade, a variety of independent geological 
and paleontological evidence has been mar- 
shalled in support of the impact theory (see 
Izett, 1990). In simplest terms, the putative 
impact led to a kind of “nuclear winter” 
caused by fine-grained debris hurled into 
the atmosphere, initiating a complex series 
of events that essentially ended, through ex- 
tensive marine and terrestrial extinctions, 
the unique biota that was characteristic of 
late Cretaceous time. 

Most vertebrate paleontologists, in con- 
trast, have been unconvinced. Archibald and 
Bryant (1990), for example, have examined 
the entirety of the extensive vertebrate fau- 
nas (including aquatic, semiaquatic, and 
terrestrial species) as stratigraphically rep- 


210 


resented below, at, and above the presumed 
K/T boundary of northeastern Montana. 
This is the world’s only nonmarine section 
at which a detailed analysis of faunal change 
across the K/T boundary has been com- 
pleted. Observed faunal changes across the 
boundary not only run contrary to ecolog- 
ical predictions for the effects of a nuclear 
winter but, according to Archibald and Bry- 
ant, are not even necessarily consistent with 
environmental catastrophy. They suggest the 
possibility of a more protracted interval (in- 
volving various extinctions and replace- 
ments). Such change could have been allied, 
for example, to alteration of habitat across 
the broad, latest Cretaceous coastal plain, 
resulting from retreat from North America 
of the Western Interior Seaway. 

One suggested explanation of extinctions 
near the P/H boundary is that of overkill 
by invading humans (e.g., Martin, 1984). 
Chief evidence, at least in the Americas, 
involves the correlation, supported by ra- 
diometric dates, of extinctions of large un- 
gulates (and their contemporaneous pred- 
ators) with the first appearances of Man. 
Many archaeological sites across Eurasia and 
North America unequivocally document the 
prowess of late Pleistocene Man as a hunter, 
even of the largest contemporary mammals. 

The idea of Man as the principal culprit 
in P/H extinctions has not, however, en- 
joyed unanimous acceptance, and all inter- 
gradations of viewpoints exist. Some work- 
ers have been willing to accept certain 
limited extinctions as having resulted from 
human overkill (principally through habitat 
destruction), particularly on oceanic is- 
lands. Widespread avian extinctions, for ex- 
ample, are well documented in the Hawai- 
ian Islands (James et al., 1987) in association 
with arrival of the original Polynesians and 
their various commensals. Other workers 
(see Martin, 1967 for citations), in contrast, 
simply have found it difficult to accept the 
demise of vast herds of North American 
Pleistocene mammals at the hands of Man. 
This is especially true in light of the long 
coexistence of Man and mammalian mega- 


ZAKRZEWSKI AND LILLEGRAVEN 


faunas across Eurasia and Africa during all 
of Quaternary time. 

As presumed for the K/T boundary, many 
workers have suggested that habitat changes 
were responsible for extinction of the latest 
Pleistocene megafaunas. Although evidence 
associated with local habitat (or global cli- 
matic) change may not be obvious for the 
latest Pleistocene, it is clear that climates 
became cooler overall and more seasonal in 
the interior through the latter half of the 
Cenozoic. Such changes caused dramatic 
shifts in distributions and types of plant 
communities. For example, Webb (1983) 
demonstrated changes in dominance of 
North American ungulates from browsing 
to grazing forms during late Miocene time. 
Workers such as Guthrie (1984) suggested 
that climatic changes accounting for the 
Miocene shift continued into Plio—Pleisto- 
cene time, thereby ultimately decreasing the 
net annual quality and quantity of food re- 
sources available to the megafauna. Gra- 
ham and Lundelius (1984) suggested biotic 
disequilibrium as a possible reason for late 
Pleistocene extinctions. 

In any case, no matter how the physical 
evidence itself may be interpreted, mar- 
riages among detailed biostratigraphy, mag- 
netostratigraphy, radioisotopic dating, and 
even archaeology have led to greatly im- 
proved levels of focused inquiry associated 
with questions of causation in extinction. 
At least for the late Pleistocene and early 
Holocene, the levels of precision in dating 
made possible by '*C-technology have 
reached levels that make such age deter- 
minations of true relevance of biological 
considerations of extinction. 


Epilogue 


We have summarized what we consider 
to be a broad sampling of major contribu- 
tions by paleomammalogists to the field of 
mammalogy since the founding of the ASM 
75 years ago. Many of these advancements 


PALEOMAMMALOGY 2a 


have occurred in the last 25 years as new 
technologies, philosophies, and more work- 
ers have entered the field. As technologies 
continue to improve, philosophies mature, 
and information expands, we look forward 
to the spectacular additional progress that 
surely will be documented in the Centennial 
Volume of the society. 


Acknowledgments 


We thank B. H. Breithaupt, J.-P. Cavigelli, R. 
W. Graham, L. E. Lillegraven, and R. W. Wilson 
for their help in the preparation of this manu- 
script. We thank W. A. Clemens, Jr. and M. R. 
Voorhies for their critical reviews. 


Literature Cited 


ALVAREZ, L. W., W. ALVAREZ, F. ASARO, AND H. V. 
MicHEL. 1980. Extraterrestrial cause for the Cre- 
taceous-Tertiary extinction. Science, 208:1095-1108. 

ARCHIBALD, J. D., AND L. J. BRYANT. 1990. Differ- 
ential Cretaceous-Tertiary extinctions of non-ma- 
rine vertebrates from northeastern Montana. Geo- 
logical Society of America, Special Paper, 247:549- 
562. 

Barnes, L. G., D. P. DOMNING, AND C. E. RAy. 1985. 
Status of studies on fossil marine mammals. Marine 
Mammal Science, 1:15-53. 

Barnosky, A. D. 1987. Punctuated equilibrium and 
phyletic gradualism: some facts from the Quaternary 
mammalian record. Current Mammalogy, 1:109- 
147. 

Bayne, C. K. 1976. Early medial Pleistocene faunas 
of Meade County, Kansas. Kansas Geological Sur- 
vey, Guidebook Series, 1:1-25. 

BEHRENSMEYER, A. K., AND A. P. HILL (EpDs.). 1980. 
Fossils in the making: vertebrate taphonomy and 
paleoecology. The University of Chicago Press, Chi- 
cago, 338 pp. 

BENTON, M. J. (ED.) 1988. The phylogeny and clas- 
sification of the tetrapods. Volume 2: Mammals. The 
Systematics Association Special Volume, 35B:1-329. 

BLACKBURN, D. G., J. M. TAYLor, AND H. A. 
PADYKULA. 1988. Trophoblast concept as applied 
to therian mammals. Journal of Morphology, 196: 
127-136. 

Bown, T. M., AND K. D. Rose. 1987. Patterns of 
dental evolution in early Eocene anaptomorphine 
primates (Omomyidae) from the Bighorn Basin, 
Wyoming. Paleontological Society Memoir, 23:1- 
162. 

CARLETON, M. D., AND R. E. EsHELMAN. 1979. A 
synopsis of fossil grasshopper mice, genus Onycho- 
mys, and their relationships to recent species. The 


University of Michigan, Museum of Paleontology, 
Papers on Paleontology, 21(C. W. Hibbard Memo- 
rial Volume 7):1-63. 

CIFELLI, R. L. 1990. Cretaceous mammals of south- 
ern Utah. IV. Eutherian mammals from the Wah- 
weap (Aquilan) and Kaiparowits (Judithian) for- 
mations. Journal of Vertebrate Paleontology, 10:346- 
360. 

Ciemens, W. A., JR. 1963. Fossil mammals of the 
type Lance formation Wyoming. Part I. Introduction 
and Multituberculata. University of California Pub- 
lications in Geological Sciences, 48:1-105. 

. 1966. Fossil mammals of the type Lance for- 

mation Wyoming. Part II. Marsupialia. University 

of California Publications in Geological Sciences, 62: 

1-122. 

. 1973. Fossil mammals of the type Lance for- 
mation, Wyoming. Part III. Eutheria and summary. 
University of California Publications in Geological 
Sciences, 94:1-102. 

Cope, E. D. 1871. Preliminary report on the Verte- 
brata discovered in the Port Kennedy bone cave. 
Proceedings of the American Philosophical Society 
of Philadelphia, 12:73-102. 

. 1899. Vertebrate remains from Port Kennedy 
bone deposit. Journal of the Academy of Natural 
Sciences of Philadelphia, Second Series, | 1:193-267. 

Crompton, A. W. 1971. The origin of the tribosphe- 
nic molar. Pp. 65-87, in Early mammals (D. M. 
Kermack and K. A. Kermack, eds.). Zoological Jour- 
nal of the Linnean Society, 50, suppl., 1:1-203. 

Crompton, A. W., AND A. L. Sun. 1985. Cranial 
structure and relationships of the Liassic mammal 
Sinoconodon. Zoological Journal of the Linnean So- 
ciety, 85:99-119. 

DAMUTH, J., AND B. J. MACFADDEN (EDS.). 1990. Body 
size in mammalian paleobiology: estimation and bi- 
ological implications. Cambridge University Press, 
Cambridge, Great Britain, 397 pp. 

Darwin, C. 1859. On the origin of species by means 
of natural selection. .. Murray, London, 490 pp. 
Dopson, P., AND D. WEXxLAR. 1979. Taphonomic 
investigations of owl pellets. Paleobiology, 5:275- 

284. 

Domnina, D. P., C. E. RAy, AND M. C. MCKENNA. 
1986. Two new Oligocene desmostylians and a dis- 
cussion of Tethytherian systematics. Smithsonian 
Contributions to Paleobiology, 59, 56 pp. 

Dorr, J. A., JR., D. R. SPEARING, AND J. R. STEIDT- 
MANN. 1977. Deformation and deposition between 
a foreland uplift and an impinging thrust belt: Ho- 
back Basin, Wyoming. Geological Society of Amer- 
ica, Special Paper, 177, 82 pp. 

Dott, R. H., Jr., AND R. L. BATTEN. 1971. Evolution 
of the Earth. McGraw-Hill Book Company, New 
York, 649 pp. 

EDINGER, T. 1948. Evolution of the horse brain. Geo- 
logical Society of America, Memoir 25:1-174. 

ELDREDGE, N., AND S. J. GouLp. 1972. Punctuated 
equilibria: an alternative to phyletic gradualism. Pp. 
82-115, in Models in Paleobiology (T. J. M. Schopf, 
ed.). Freeman, Cooper & Company, San Francisco, 
250 pp. 

Emry,R.J. 1970. A North American Oligocene pan- 
golin and other additions to the Pholidota. Bulletin 


ZA2 ZAKRZEWSKI AND LILLEGRAVEN 


of the American Museum of Natural History, 142: 
455-510. 

ENGsTRoM, M. D., J. R. CHOATE, AND H. H. GENoways. 
Taxonomy. Pp. 179-199, in Seventy-five years of 
mammalogy (1919-1994) (E. C. Birney and J. R. 
Choate, eds.). Special Publication, The American 
Society of Mammalogists, 1 1:1-433. 

EVERNDEN, J. F., D. E. SAvAGE, G. H. CurTIs, AND G. 
T. James. 1964. Potassium-argon dates and the 
Cenozoic mammal chronology of North America. 
American Journal of Science, 262:145-198. 

FLYNN, J. J., B. J. MACFADDEN, AND M. C. MCKENNA. 
1984. Land-mammal ages, faunal heterochrony, and 
temporal resolution in Cenozoic terrestrial sequenc- 
es. Journal of Geology, 92:687-705. 

Fox, R. C. 1989. The Wounded Knee local fauna 
and mammalian evolution near the Cretaceous-Ter- 
tiary boundary, Saskatchewan, Canada. Paleonto- 
graphica, Abstract A, 208:11-59. 

GALUSHA, T., AND J. C. Buick. 1971. Stratigraphy of 
the Santa Fe Group, New Mexico. Bulletin of the 
American Museum of Natural History, 144:1-127. 

Gazin, C. L. 1953. The Tillodontia: an early order 
of mammals. Smithsonian Miscellaneous Collec- 
tions, 121:1-110. 

GINGERICH, P. D. 1985. Species in the fossil record: 
concepts, trends, and transitions. Paleobiology, 11: 
27-41. 


1986. George Gaylord Simpson: empirical 
theoretician. Pp. 3-9, in Vertebrates, phylogeny, and 
philosophy (K. M. Flanagan and J. A. Lillegraven, 
eds.). Contributions to Geology, University of Wy- 
oming, Laramie, Special Paper, 3:1-361. 

GouLp, S. J. 1985. The paradox of the first tier: an 
agenda for paleobiology. Paleobiology, 11:2-12. 

GOULD, S. J., AND N. ELDREDGE. 1977. Punctuated 
equilibria: the tempo and mode of evolution recon- 
sidered. Paleobiology, 3:115-151. 

GRAHAM, R. W., AND E. L. LUNDELIus, JR. 1984. 
Coevolutionary disequilibrium and Pleistocene ex- 
tinctions. Pp. 223-249, in Quaternary extinctions: a 
prehistoric revolution (P. S. Martin and R. G. Klein, 
eds.). University of Arizona Press, Tucson, 891 pp. 

Grecory, J. T., J. A. BAcsKAI, G. V. SHKURKIN, M. 
C. WINANS, AND B. H. RAUSCHER. 1992. Buibliog- 
raphy of fossil vertebrates 1989. Society of Verte- 
brate Paleontology, Lincoln, Nebraska, 495 pp. 

Guitpay, J. E. 1971. The Pleistocene history of the 
Appalachian mammal fauna. Pp. 233-262, in The 
distributional history of the biota of the southern 
Appalachians. Part III: Vertebrates (P. C. Holt, ed.). 
Virginia Polytechnic Institute and State University 
Research Division Monograph, Blacksburg, 4:1-306. 

GutTurig, R. D. 1984. Mosaics, allelochemics and 
nutrients: an ecological theory of late Pleistocene 
megafaunal extinctions. Pp. 259-298, in Quaternary 
extinctions: a prehistoric revolution (P. S. Martin 
and R. G. Klein, eds.). University of Arizona Press, 
Tucson, 891 pp. 

Hai, E.R. 1946. [Review of] Simpson, George Gay- 
lord. The principles of classification and a classifi- 
cation of mammals. Journal of Mammalogy, 27:287- 
288.. 

HENNIG, W. 1966. Phylogenetic systematics. Uni- 
versity of Illinois Press, Urbana, 263 pp. 

Hesse, C. 1941. [Review of] Nomenclature and cor- 


relation of the North American continental Tertiary 
(Report by H. E. Wood, I], et al.). Journal of Mam- 
malogy, 22:458. 

HipBarp, C. W. 1941. The Borchers fauna, a new 
Pleistocene interglacial fauna from Meade County, 
Kansas. Kansas Geological Survey Bulletin, 38:197- 
220. 


1947. The status of the genera Anaptogonia 

and Sycium Cope. Journal of Mammalogy, 28:67. 

1949. Techniques of collecting microverte- 
brate fossils. Contributions from the Museum of Pa- 
leontology, University of Michigan, 8:7-19. 

Hopson, J. A., AND H. R. BARGHUSEN. 1986. An 
analysis of therapsid relationships. Pp. 83-106, in 
The ecology and biology of mammal-like reptiles (N. 
Hotton, III, P. D. MacLean, J. J. Roth, and E. C. 
Roth, eds.). Smithsonian Institution Press, Wash- 
ington, D.C., 326 pp. 

Hux.ey, T. H. 1870. The anniversary address of the 
president. Quarterly Journal of the Geological So- 
ciety of London, 26:xxix-/x1v. 

IzetT, G. A. 1990. The Cretaceous/Tertiary bound- 
ary interval, Raton Basin, Colorado and New Mex- 
ico, and its content of shock-metamorphosed 
minerals; evidence relevant to the K/T boundary 
impact-extinction theory. Geological Society of 
America, Special Paper 249:1-100. 

James, H. F., ET AL. 1987. Radiocarbon dates on 
bones of extinct birds from Hawaii. Proceedings of 
the National Academy of Sciences, USA, 84:2350- 
2354. 

JENKINS, F. A., JR. 1974. Tree shrew locomotion and 
the origin of primate arborealism. Pp. 85-115, in 
Primate locomotion (F. A. Jenkins, Jr., ed.). Aca- 
demic Press, New York, 390 pp. 

JEPSEN, G. L. 1970. Bat origins and evolution. Pp. 
1-64, in Biology of bats (W. A. Wimsatt, ed.). Ac- 
ademic Press, New York, 1:1-404. 

JERISON, H. J. 1973. Evolution of the brain and in- 
telligence. Academic Press, New York, 482 pp. 

KELLOGG, R. 1936. A review of the Archaeoceti. Car- 
negie Institution of Washington Publication, 482:1- 
366. 

Kortn, W. W. 1979. Taphonomy of microvertebrate 
fossil assemblages. Annals of Carnegie Museum, 48: 
235-285. 

Krause, D. W. 1982. Jaw movement, dental func- 
tion, and diet in the Paleocene multituberculate Pri- 
lodus. Paleobiology, 8:265-281. 

LILLEGRAVEN, J. A. 1969. Latest Cretaceous mam- 
mals of upper part of Edmonton formation of Al- 
berta, Canada, and review of marsupial-placental di- 
chotomy in mammalian evolution. University of 
Kansas Paleontological Contributions, 50:1-122. 

Linpsay, E. H., R. F. BUTLER, AND N. M. JOHNSON. 
1981. Magnetic polarity zonation and biostratig- 
raphy of late Cretaceous and Paleocene continental 
deposits, San Juan Basin, New Mexico. American 
Journal of Science, 281:390-435. 

MartIn, P.S. 1967. Pleistocene overkill. Pp. 75-120, 
in Pleistocene extinctions, the search for a cause (P. 
S. Martin and H. E. Wright, Jr., eds.). Yale Univer- 
sity Press, New Haven, Connecticut, 453 pp. 

. 1984. Prehistoric overkill: the global model. 

Pp. 354-403, in Quaternary extinctions: a prehis- 


PALEOMAMMALOGY 215 


toric revolution (P. S. Martin and R. G. Klein, eds.). 
The University of Arizona Press, Tucson, 891 pp. 

MATTHEW, W. D. 1915. Climate and evolution. An- 
nals of the New York Academy of Sciences, 24:171- 
318. 


. 1939. Climate and evolution. Second ed. Spe- 
cial Publication of the New York Academy of Sci- 
ences, 1:1-223. 

McKenna, M. C. 1962. Collecting small fossils by 
washing and screening. Curator, 3:221-235. 

1973. Sweepstakes, filters, corridors, Noah’s 

arks, and beached Viking funeral ships in palaeo- 

geography. Pp. 295-308, in Implications of conti- 
nental drift to the Earth sciences (D. H. Tarling and 

S. K. Runcorn, eds.). Academic Press, London, 1:1- 

622. 


. 1975. Toward a phylogenetic classification 
of the Mammalia. Pp. 21-46, in Phylogeny of the 
Primates: a multidisciplinary approach (W. P. Luck- 
ett and F. S. Szalay, eds.). Plenum Press, New York, 

483 pp. 

. 1983. Holarctic rearrangement, cosmic events, 
and Cenozoic terrestrial organisms. Annals of the 
Missouri Botanical Garden, 70:459-489. 

McNas, B. K. 1978. The evolution of endothermy 
in the phylogeny of mammals. The American Nat- 
uralist, 112:1-21. 

Me.tetTtT, J. 1974. Scatological origin of microver- 
tebrate fossil accumulations. Science, 185:349-350. 

Miao, D. 1991. On the origins of mammals. Pp. 579- 
597, in Origins of the higher groups of tetrapods: 
controversies and consensus (H.-P. Schultze and L. 
Trueb, eds.). Cornell University Press, Ithaca, 724 
pp. 

MILter, G. S, AND J. W. GipLey. 1918. Synopsis of 
the supergeneric groups of rodents. Journal of the 
Washington Academy of Science, 8:431-448. 

Newman, C. M., J. E. COHEN, AND C. Kipnis. 1985. 
Neo-Darwinian evolution implies punctuated equi- 
libria. Nature, 315:400-401. 

Novacek, M. J. 1987. Auditory features and affinities 
of the Eocene bats Jcaronycteris and Palaeochirop- 
teryx (Microchiroptera, incertae sedis). American 
Museum Novitates, 2877:1-18. 

. 1990. Morphology, paleontology, and the 
higher clades of mammals. Current Mammalogy, 
2:507-543. 

Otson, E. C. 1980. Taphonomy;: its history and role 
in community evolution. Pp. 5-19, in Fossils in the 
making: vertebrate taphonomy and paleoecology (A. 
K. Behrensmeyer and A. P. Hill, eds.). The Univer- 
sity of Chicago Press, Chicago, 338 pp. 

Ossorn, H. F. 1921. Resemblances and contrasts 
between zoologic and paleontologic research in 
mammalogy. Desirability of uniform standards and 
systems in classification, in description, in measure- 
ments, in reasoning. Journal of Mammalogy, 2:1- 
11. 

PROTHERO, D. R., AND R. M. SCHOCH (EDs.). 1989. 
The evolution of perissodactyls. Oxford University 
Press, London, 537 pp. 

Rapinsky, L. 1981. Brain evolution in extinct South 
American ungulates. Brain, Behavior and Evolution, 
18:169-187. 

REPENNING, C. A., C. E. RAY, AND D. GRIGORESCU. 


1979. Pinniped biogeography. Pp. 357-369, in His- 
torical biogeography, plate tectonics, and the chang- 
ing environment (J. Gray and A. J. Boucot, eds.). 
Oregon State University Press, Corvallis, 500 pp. 

Rowe, T. 1988. Definition, diagnosis, and origin of 
the Mammalia. Journal of Vertebrate Paleontology, 
8:241-264. 

SAVAGE, D. E. 1951. Late Cenozoic vertebrates of the 
San Francisco Bay region. University of California 
Publications in Geological Sciences, 28:215-314. 

SAVAGE, D. E., AND D. E. RussELL. 1983. Mamma- 
lian paleofaunas of the world. Addison-Wesley Pub- 
lishing Company, London, 432 pp. 

ScHocH, R. M. 1986. Systematics, functional mor- 
phology and macroevolution of the extinct mam- 
malian order Taeniodontia. Bulletin of the Peabody 
Museum of Natural History, Yale University, New 
Haven, Connecticut, 42:1-307. 

Simons, E. L., ANDR. F. KAy. 1983. Oatrania, a new 
basal anthropoid primate from Fayum, Oligocene of 
Egypt. Nature, 304:624-626. 

Simpson, G. G. 1928. A catalogue of the Mesozoic 
Mammalia in the Geological Department of the Brit- 
ish Museum. Oxford University Press, London, 215 
pp. 

. 1929. American Mesozoic Mammalia. 

Memoir of the Peabody Museum of Yale University, 

3:1-171. 

1944. Tempo and mode in evolution. Co- 

lumbia University Press, New York, 237 pp. 

1945. The principles of classification and a 

classification of mammals. Bulletin of the American 

Museum of Natural History, 85:1-350. 

. 1952. Probabilities of dispersal in geologic 

time. Pp. 163-176, in The problem of land connec- 

tions across the South Atlantic, with special reference 
to the Mesozoic (E. Mayr, et al., eds.). Bulletin of 

the American Museum of Natural History, 99:79- 

258. 


1953a. Evolution and geography. Oregon 

State System of Higher Education, Condon Lectures, 

Corvallis, 64 pp. 

1953b. The major features of evolution. Co- 

lumbia University Press, New York, 434 pp. 

1971. Concluding remarks: Mesozoic mam- 
mals revisited. Pp. 181-198, in Early mammals (D. 
M. Kermack and K. A. Kermack, eds.). Zoological 
Journal of the Linnean Society, 50, suppl., 1:1-203. 

SKINNER, M. F., S. M. SKINNER, AND R. J. Gooris. 
1977. Stratigraphy and biostratigraphy of late Ce- 
nozoic deposits in central Sioux County, western Ne- 
braska. Bulletin of the American Museum of Natural 
History, 158:263-371. 

SwIsHER, C. C., III, AND D. R. PROTHERO. 1990. Sin- 
gle-crystal *°?Ar/**Ar dating of the Eocene-Oligocene 
transition in North America. Science, 249:760-762. 

SZALAY, F. S., M. J. NovACEK, AND M. C. MCKENNA 
(EDs.). 1993. Mammal phylogeny. Springer-Verlag, 
New York, 1:1-249, 2:1-321. 

TEDFORD, R. H. 1974. Marsupials and the new pa- 
leogeography. Pp. 109-126, in Paleogeographic 
provinces and provinciality (C. A. Ross, ed.). Society 
of Economic Paleontologists and Mineralogists, Spe- 
cial Publication, 21, 1-233. 

Vooruigs, M. R. 1969. Taphonomy and population 


214 ZAKRZEWSKI AND LILLEGRAVEN 


dynamics of an early Pliocene vertebrate fauna, Knox 
County, Nebraska. Contributions to Geology, Uni- 
versity of Wyoming, Laramie, Special Paper, 1, 1- 
69. 

Wess, S. D. 1983. The rise and fall of the late Mio- 
cene ungulate fauna in North America. Pp. 267-306, 
in Coevolution (M. H. Nitecki, ed.). The University 
of Chicago Press, Chicago, 392 pp. 

. 1985. Late Cenozoic mammal dispersals be- 
tween the Americas. Pp. 357-386, in The great biotic 
interchange (F. G. Stehli and S. D. Webb, eds.). Ple- 
num Press, New York, 532 pp. 

Wuite, J. A., AND B. L. KELLER. 1984. Evolutionary 
stability and ecological relationships of morphology 
in North American Lagomorpha. Pp. 58-66, in Pa- 
pers in vertebrate paleontology honoring Robert 
Warren Wilson (R. M. Mengel, ed.). Special Publi- 
cation, Carnegie Museum of Natural History, 9, 1- 
186. 

WIBLE, J. R., AND J. A. Hopson. 1993. Basicranial 
evidence for early mammal phylogeny. Pp. 45-62, 
in Mammal phylogeny: Mesozoic differentiation, 
multituberculates, monotremes, early therians, and 
marsupials (F. S. Szalay, M. J. Novacek, and M. C. 
McKenna, eds.). Springer-Verlag, New York, 249 


pp. 

Witson, J. A. 1978. Stratigraphic occurrence and cor- 
relation of early Tertiary vertebrate faunas, Trans- 
Pecos Texas, Part I: Vieja area. Texas Memorial Mu- 
seum Bulletin, 25:1—42. 


Witson, R. W. 1986. The Paleogene record of the 
rodents: fact and interpretation. Pp. 163-175, in 
Vertebrates, phylogeny, and philosophy (K. M. Flan- 
agan and J. A. Lillegraven, eds.). Contributions to 
Geology, University of Wyoming, Laramie, Special 
Paper, 3:1-361. 

Woop, A. E. 1955. A revised classification of the 
rodents. Journal of Mammalogy, 36:165-187. 

Woop, H. E., II, er at. 1941. Nomenclature and 
correlation of the North American continental Ter- 
tiary. Bulletin of the Geological Society of America, 
52:1-48. 

WoobDBuRNE, M. O. (ED.). 1987. Cenozoic mammals 
of North America: geochronology and biostratigra- 
phy. University of California Press, Berkeley, 336 
pp. 

WoopsurRng, M. O., AND W. J. ZINSMEISTER. 1984. 
The first land mammal from Antarctica and its bio- 
geographic implications. Journal of Paleontology, 58: 
913-948. 

ZAKRZEWSKI, R. J. 1975. Pleistocene stratigraphy and 
paleontology in western Kansas: the state of the art, 
1974. Pp. 121-128, in Studies on Cenozoic pale- 
ontology and stratigraphy in honor of Claude W. 
Hibbard (G. R. Smith and N. E. Frieland, eds.). Uni- 
versity of Michigan, Papers on Paleontology, 12 (C. 
W. Hibbard Memorial Volume 3), 143 pp. 


BIOGEOGRAPHY 


SYDNEY ANDERSON AND BRUCE D. PATTERSON 


Introduction 


| Beary the study of the distribu- 
tion of life, seeks to comprehend an 
immense range and diversity of phenome- 
na. Most broadly conceived, the field ranges 
from the domain of astrophysics (the dis- 
tribution of matter in the universe and the 
physical laws of radiation, gravitation, and 
others which affect this) to ecological and 
behavioral interactions that govern the spa- 
tial distribution of individuals within local 
demes. For the purposes of this brief his- 
torical review, and in parallel with editorial 
opinion in biogeographic journals (see 
Blondel, 1987), however, we consider bio- 
geographic patterns and processes ranging 
from global climates and drifting continents 
on the one hand to local communities and 
species responses at distributional limits on 
the other (e.g., Reichman, 1984). Concep- 
tually, at least, our coverage transcends the 
fields now known as historical biogeography 
and landscape and geographical ecology, al- 
though space limitations preclude detailed 
treatment of all aspects. 

Biogeographic patterns and processes are 
sensitive to variations in time, space, and 
biological organization. These patterns and 
processes may be categorized according to 
the scales of these crucial factors. ““Ecolog- 
ical’’ time-periods may be contrasted with 


215 


a 


x 


on : ” 
an oS 
At 
a 
Bee ar @ 4 


ie 


“evolutionary” ones—the former denotes 
the years, decades, and centuries over which 
ecological processes, such as dispersal, suc- 
cession, altered resource-use patterns, and 
others take place. Evolutionary time peri- 
ods may involve different types of organ- 
ismal responses, including changing gene 
frequencies, local adaptation and genetic 
drift, and speciation. In similar fashion, 
processes that operate over local spatial 
scales differ from those involved in larger 
(regional or continental) patterns. Finally, 
the processes of environmental stimulus and 
organismal response are mediated in fun- 
damentally different ways by species pop- 
ulations (through the genetics of reproduc- 
tion and adaptation) and biotas (through 
competition, predation, and other com- 
munity-level processes). 

We attempt here to chronicle the devel- 
opment of mammalian biogeography over 
the last 75 years. After considering some 
trends that generally apply to all biogeo- 
graphic subdisciplines, we use the different 
scales of time, space, and biological orga- 
nization to organize our discussion of var- 
ious topics. We first treat studies of species 
over ecological time, proceeding then to 
those on biotas (but emphasizing mam- 
malian faunas) over ecological time, species 


216 ANDERSON AND PATTERSON 


over evolutionary time, and faunas over 
evolutionary time. Within each unit, we 
have attempted to arrange patterns and pro- 
cesses in order of increasing spatial scope. 


Historical Trends 


By 1919 the discipline of biogeography 
was already vigorous and well established. 
That different plants and animals lived in 
different places was known in classical times, 
but elaboration and formalization of that 
recognition increased dramatically in the 
three centuries before the founding of the 
American Society of Mammalogists. Im- 
portant contributions were made by early 
19th Century workers, including von Hum- 
boldt and Bonpland, de Candolle, and Lyell 
(see Nelson, 1978). Sclater’s (1858) classi- 
fication of the world’s avifauna into bio- 
geographic regions and subregions ranks as 
a major biogeographic development of the 
19th Century. This work, and the burgeon- 
ing inventories of worldwide faunas assem- 
bled by imperial Europe, permitted Lydek- 
ker (1986) to develop a strikingly accurate 
discussion of regionalism in mammalian 
faunas. The life-zone concept of Merriam 
(1890) was another major event, at least for 
understanding the ecological factors that af- 
fect the distribution of North American 
mammals. In addition, under Merriam’s 
leadership, the U.S. Biological Survey had 
produced a growing series of taxonomic re- 
visions and regional faunal accounts that 
were published mainly in the North Amer- 
ican Fauna series—43 numbers had been 
published by 1919. Based on copious col- 
lections, many of these accounts included 
detailed distribution maps of species, veg- 
etation types, and finally formal “life zones.” 
Moreover, they contributed importantly to 
the developing polytypic species concept, 
with its explicit recognition of geographic 
variation, and thence eventually to the 
*““modern synthesis” of evolutionary theory. 
Just before the founding of the society, Mat- 
thew’s (1915) Climate and Evolution was 
published, postulating the northern origin and 


southward dispersal of many mammalian 
groups; Matthew’s thesis emphasized the im- 
portance of history in interpreting biogeo- 
graphic patterns. This paper had consider- 
able influence on mammalian biogeography, 
in significant part through the work of his 
student G. G. Simpson. However, some of 
the details and major assumptions in Mat- 
thew’s five-point thesis have required mod- 
ification. 

A more enduring contribution to this field 
was Wegener’s hypothesis of drifting con- 
tinents, originally presented in 1912. This 
revolutionary hypothesis was soundly re- 
jected by Matthew and many other mam- 
malogists, and its revival required the pas- 
sage of half a century and the discovery of 
a geophysical mechanism, plate tectonics, 
to allow drift. 

Since the founding of the ASM, a number 
of trends are discernable in the development 
of biogeography. Simple, general patterns 
were dissected to reveal more complex ones. 
Emphases shifted from purely descriptive 
accounts to increasingly quantitative and 
predictive ones. Models of biogeographic 
processes were developed, initially static and 
then increasingly more dynamic in char- 
acter. Some of these trends have long been 
evident—in a mid-century appraisal, Hubbs 
(1958:470) noted “‘a shift from the classical, 
purely descriptive biogeography to a kinetic 
approach, which is more concerned with 
processes and explanations than with the 
classification of the earth into a hierarchy 
of biogeographical regions.” 

As biogeography matured, gains in ana- 
lytical rigor have been achieved, sometimes 
at the expense of flexibility and breadth. In 
its infancy, biogeography had spanned all 
or most of natural history, but as it matured, 
rival schools developed around narrower 
concepts (e.g., island biogeography) or ap- 
proaches (e.g., numerical biogeography). 
Biogeographers were themselves classified 
as champions of dispersal or vicariance, or 
devotees of equilibrium or historical schools. 
Even finer distinctions were thought nec- 
essary to reflect philosophical differences 


BIOGEOGRAPHY 217, 


within these categories (e.g., vicariance bio- 
geography, phylogenetic biogeography, and 
panbiogeography). For the most part, bio- 
geographic discords reflected parallel acri- 
mony in the sister disciplines of systematics 
and ecology, which also experienced philo- 
sophical and technological revolutions dur- 
ing the 1960s and 1970s (Hull, 1988; Mc- 
Intosh, 1985). Perhaps because extended 
critical discussion has exposed the short- 
comings of each approach, biogeography to- 
day is best carried out under a banner of 
pluralism (McIntosh, 1987). 


Species Over Ecological Time 
Periods 


How is the distribution of a species re- 
lated to its abundance? This relationship was 
explored, mostly from an ecological per- 
spective, by Andrewartha and Birch (1954). 
The thesis of the book is that “distribution 
and abundance are but two aspects of one 
phenomenon.” The book is replete with 
biogeographical implications. They showed 
that common species may be rare in mar- 
ginal parts of their ranges and that there is 
no fundamental distinction between the ex- 
tinction of a local population and the ex- 
tinction of a species except that, in the latter 
case, the population becoming extinct hap- 
pens to be the last one of the species. 

How is the distribution of a species lim- 
ited? Trying to understand those limits be- 
gan simply enough with concepts such as 
Liebig’s “‘law of the minimum” proposed 
for the limits to growth in plants (discussed 
by Hesse et al., 1937:21). This interesting 
question continues to attract speculation and 
investigation. A wholesale shift in distri- 
bution of a local fauna of mammals accom- 
panying changes in local climates in the 
Pleistocene was described by Guilday et al. 
(1964). 

The dynamic nature of species limits on 
a shorter time scale is indicated by numer- 
ous documented cases in which boundaries 
of individual species have expanded or con- 


tracted recently in North America over a 
relatively few years of time. For example, 
Dasypus novemcinctus (Fitch et al., 1952; 
Smith and Lawlor, 1964), Baiomys taylori 
(Baccus et al., 1971), and Sigmodon hispi- 
dus (Genoways and Schlitter, 1967), ex- 
tended their ranges northward; Marmota 
monax extended westward in Kansas 
(Choate and Reed, 1986); Lepus californicus 
extended eastward in Texas (Packard, 1963); 
Spermophilus richardsoni (Hansen, 1962) 
extended southward in Colorado; and since 
1960 Sorex cinereus, Microtus pennsylvani- 
cus, Mustela nivalis, and Zapus hudsonius 
have extended southward in Kansas (Frey, 
1992). It is more difficult to demonstrate 
retractions in ranges, but surely these have 
been occurring as well. The retractions in 
ranges of many larger mammals, such as 
grizzly bears, mountain lions, gray wolves, 
and wapiti in North America, need no fur- 
ther documentation here. Another recent 
mammalian example is the correlation of 
hours of darkness (about 7.3 hours in this 
case) needed for feeding with the northern 
limit of an Asian porcupine (Alkon and 
Saltz, 1988). 

In 1957, Darlington’s book Zoogeogra- 
phy: the Geographical Distribution of Ani- 
mals summarized distribution of the major 
groups of terrestrial vertebrates. Questions 
posed (p. vii) were: (1) What is the main 
pattern of animal distributions? (2) How has 
the pattern been formed? (3) Why has the 
pattern been formed? and, (4) What does 
animal distribution tell about ancient lands 
and climates? 

The answers (Darlington, 1957:618) were: 
(1) The main pattern is a “‘concentration of 
the largest, most diverse, least-limited fau- 
nas in the main tropical regions of the Old 
World; limitation caused by climate north 
of the tropics; and limitation and differen- 
tiation caused by barriers in South America 
and Australia.” (2) The pattern has been 
formed by spread of successive dominant 
groups from the Old World tropics over 
much or all of the world, followed by zo- 
nation and differentiation according to cli- 


218 ANDERSON AND PATTERSON 


mate and ocean barriers, and by retreat and 
replacement of old groups as new ones 
spread.”’ (3) The pattern has been formed 
‘‘because evolution has tended to produce 
the most dominant animals in the largest 
and most favorable areas, which for most 
vertebrates are in the main regions of the 
Old World tropics” (see Darlington, 1957: 
569 for brief comments on probabilities and 
dominance). (4) Animal distribution tells us 
that ‘‘as far back as can be seen clearly, the 
main pattern of continents and climates 
seems to have been the same as now.”’ From 
a slightly skeptical point of view, we may 
now judge that the compilation of sum- 
maries of distributions of different groups 
may have been a greater contribution than 
the set of answers or conclusions. 

A hypothesis that dominant animals usu- 
ally move to gain advantages rather than to 
escape disadvantages is repeatedly asserted 
in various contexts (e.g., Darlington, 1957: 
620, 637, and ranging from major groups 
of vertebrates to races of humans). The con- 
cept lacks clear definition and has a teleo- 
logical implication that is, at best, mislead- 
ing. An interesting exchange on the 
application of the concept to human races 
was published in the Journal of Mammal- 
ogy (Hall, 1946; Hill, 1947). 

In The Mammals of North America (Hall 
and Kelson, 1959) was a chapter (of 8 pages) 
on zoogeography (by Hall). The questions 
posed were: ““What patterns emerge from 
the 500 maps showing the geographic dis- 
tribution of North American mammals? 
What factors account for these patterns?” 
and, ‘““Why are there fewer kinds of mam- 
mals in one area than in another?” The ma- 
jor patterns discussed are: (1) the distinction 
of three major regions with largely different 
faunas, namely boreal, temperate, and trop- 
ical; (2) the presence of more temperate than 
boreal species, and more tropical than tem- 
perate; (3) the presence of zonation within 
each of the major regions; (4) the presence 
in North America of more species thought 
to have come from Asia than vice versa, 
and the presence in South America of more 


species from North America than the re- 
verse; and (5) the presence of an unusually 
large number of subspecies in the South- 
west. 

The major factors said to account for these 
patterns are: (1) temperature was regarded 
as a major factor in determining mamma- 
lian distributions from north to south; (2) 
the number of different habitats that are 
available is positively correlated with the 
number of species, both on the large spatial 
scale of regions and on the smaller scale of 
zones and local areas within zones; (3) the 
greater vigor of ““mammals of a large land 
area [which] more often than not prevail 
over their counterparts of a small land area 
when the two are brought into competition” 
(p. xx1x); and (4) advances and recessions 
of glaciation and accompanying aridity in 
areas from west to east within the temperate 
zone. The relevance of paleontological his- 
tory was mentioned briefly. Hall (1981) 
shortened the original eight-page discussion 
of zoogeography to one page and included 
no basically different interpretations. Nei- 
ther the patterns nor their explanations dif- 
fered greatly from what could be found in 
earlier literature. 

Returning to the hypothesis that animals 
from a greater land mass are more vigorous, 
we note that Hall in Hall and Kelson (1959) 
incidentally presented two other and con- 
trary hypotheses. A probabilistic explana- 
tion appears in a footnote on p. xxvi, relat- 
ing to the relative contributions of the South 
American and Central American tropics. 
Elsewhere (p. xxv) he noted that ‘North 
America and Eurasia might properly be 
thought of as one continuous region—the 
Holarctic region,”’ which has only recently 
been broken by the barrier of the Bering Sea. 
In current terminology this is simply a vi- 
cariant event and the original hypothesis 
about different areas of different sizes and 
about vigor seems irrelevant, at least as it 
relates to Asia and North America. A prob- 
abilistic model was discussed by Horton 
(1974), basically as a null hypothesis, and 
the conclusion was reached that it is not 


BIOGEOGRAPHY 219. 


necessary to invoke the concept of relative 
species dominance as a determinant of the 
direction of species movement in many 
cases. Frequently the term “dominance” has 
been used in the literature somewhat in- 
consistently and without careful definition, 
with resulting confusion. 

A few years after the publication of Hall 
and Kelson (1959), Eduardo Rapoport came 
to the Department of Mammalogy at the 
American Museum of Natural History and 
asked one of us (Anderson) what similar 
works might exist for the mammals of other 
continents. Unfortunately, the answer was 
none. Since then a set of maps for Australian 
species has become available (Strahan, 1983) 
and has been the subject for biogeographical 
analysis from the standpoint of areography 
(Anderson and Marcus, 1992). A three vol- 
ume work on South American mammals 
when completed will provide maps (two 
volumes have been published, Eisenberg, 
1989, and Redford and Eisenberg, 1992). 
Another three-volume work with maps for 
South American mammals has been in 
preparation for many years (to be edited by 
S. Anderson, A. L. Gardner, and J. L. Pat- 
ton). There is no comparable compilation 
with maps for Africa. Most of Eurasia lies 
in the Palaearctic Region, for which a set of 
maps was published by Corbet (1978), and 
the remainder lies within the Oriental or 
Indomalayan Region, recently treated by 
Corbet and Hill (1992) and including a set 
of maps. No subsequent biogeographical or 
areographic analyses based on these two sets 
of maps has been published yet. Inciden- 
tally, faunal lists, whether regional or on 
some more local scale, even when not ac- 
companied by maps, have traditionally been 
basic sources for biogeographic data and 
their importance needs to be acknowledged 
here. 

The set of maps for North American 
mammals in Hall and Kelson (1959) was 
used as the source of data in subsequent 
analyses by several authors. The question 
of how many species occur in different areas 
was addressed by Simpson (1964), who tal- 


lied numbers of species postulated (on the 
basis of the published range maps) to occur 
in each of the squares of a 150-mile grid. 
These sources were used in a more detailed 
examination of the relative contributions of 
different groups of mammals to the latitu- 
dinal gradient in species numbers by Wilson 
(1974). He noted ‘“‘the lack of increase in 
species density” toward the tropics when 
quadrupedal mammals are considered alone, 
the major contributors to the latitudinal ef- 
fect being the bats. He considered also the 
possible effect of the lesser amount of space 
available in Central America. Wilson’s 
studies provided a much “finer grained” 
look than the tallies by three major regions 
in Halls’ (1981) analysis. Even finer detail 
is worthy of analysis (but there is a limit to 
the ability of progressively smaller units of 
space to yield meaningful geographic infor- 
mation, as was discussed by Anderson, 
1972). Willig and Sandlin (1991) compared 
the effects of quadrat and latitudinal band 
methodologies on detection of latitudinal 
gradients in species richness. 

What is the frequency distribution of sizes 
of geographic ranges among all possible 
ranges for species of North American mam- 
mals? This question was addressed by An- 
derson (1977), using the same set of maps, 
and he noted that “‘it is clear that the species 
are not spread evenly, but that they are about 
an order of magnitude (10 times) less ‘con- 
centrated’ in each successively larger order 
of magnitude of range” (Fig. 1). 

Various analyses have focused on areas 
of distributions. For example, Armstrong 
(1972:354) noted that ‘““Areographic anal- 
ysis is of interest because it enables the pro- 
visional segregation of faunal elements of 
possible historical integrity from assem- 
blages with compatible and complementa- 
ry, yet coincidental ecology.” The areo- 
graphic analysis referred to was the sorting 
of species into groups with respect to the 
locations of their geographic ranges. Thus, 
in Colorado, Armstrong recognized nine 
‘“‘faunal elements” such as Cordilleran, Chi- 
huahuan, Neotropical, and Great Basin. 


220 ANDERSON AND PATTERSON 


10 +1 


10-4 


NUMBER OF SPECIES PER 100 KM? INCREMENT IN SIZE OF AREA 


10' 2 3 


4 5 6 10’ 


AREA IN KM? 


Fic. 1.—Graph for North American terrestrial mammals showing the number of species (averaged 
for each succeeding order of magnitude) having ranges of any given size. Counts are grouped in 100 
km? increments. The negative values on the ordinate are powers of 10, thus 10~* or 0.0001 species 
per 100 km? increment for a range of 10° (1,000,000) km? means that there are so few species with 
ranges of this size that most increments or size-classes of 100 km? are unoccupied and, and on the 
average, there is about one species for each 10,000 size-classes (Anderson, 1977:12). It may be 
reasonably inferred from these data that at any range size a species has a greater probability of losing 


range than of increasing its range. 


Most publications in mammalogy, or in 
biogeography, fit an existing mold. Al- 
though they contain new information, test 
an existing theory, or otherwise contribute 
to knowledge, they seem basically familiar 
as to topic, concept, assumptions, empha- 
sis, and methodology. Occasionally a pub- 
lication breaks new intellectual ground. Ar- 
eography (Rapoport, 1982; an earlier 
Spanish edition, published in 1975, was not 
widely distributed) was such a publication. 


Clearly the author was thinking along new 
lines, developing new methods, and asking 
new questions. Let us briefly consider some 
of these. 

Do North American species of mammals 
belonging to different orders and families 
have geographic ranges of different sizes 
(Rapoport, 1982:7)? Mean ranges were giv- 
en for 9 orders and 14 families. Arithmetic 
means were used and differences noted. For 
example, the mean for Carnivora, the order 


BIOGEOGRAPHY 224 


with the largest ranges, was about eight times 
that of the Rodentia. Graphs of range size 
distributions for six orders were published 
by Anderson (1977:10). Rapoport used 
‘“‘square megametres”’ as his unit of mea- 
surement and defined a megametre (Mm) 
as 100 km. The prefix mega is usually used 
for million rather than 100 thousand, so 
these discrepant usages need to be taken into 
account when comparing data in Ander- 
son’s paper with those in Rapoport’s. One 
square megameter as used by Rapoport is 
equal to 1 x 104 km? as used by Anderson, 
and Anderson used geometric means in- 
stead of arithmetic means. 

What are the mean geographic ranges of 
bats with different feeding habits (Rapoport, 
1982:9)? Those that eat animal food are 
more widespread than those that eat plant 
food. Whether this would remain true if their 
entire ranges are included, rather than just 
the parts of ranges within North America, 
remains to be tested. 

What is the frequency distribution of the 
sizes of ranges of species among all possible 
sizes (Rapoport, 1982:13)? This question 
was investigated at about the same time, but 
independently of Anderson’s work, which 
was published in 1972. Both authors point- 
ed out the logarithmic or “hollow curve” 
distribution. 

How are the ranges of subspecies distrib- 
uted in space and in size relative to each 
other (Rapoport, 1982:27)? Various aspects 
of this were discussed and it was noted that 
“There is a tendency to increase the perim- 
eter of the irregularity of the species’ exter- 
nal frontiers when the number of subspecies 
increases.” 

How are numbers of subspecies with 
ranges surrounded by the ranges of other 
subspecies correlated with the total number 
of subspecies recognized within the species 
(Rapoport, 1982:31)? The correlation of in- 
ternal subspecies and total number of sub- 
species in the species is +0.979. 

Do the ranges of subspecies relative to 
each other differ among taxonomic groups 
(Rapoport, 1982:35)? The relative numbers 


that are considered to be contiguous, in- 
cluded, disjunct, and superimposed, differ 
some among the species of different orders, 
but the significance, both statistically and 
biologically, is unclear. 

Does the size of the range of the most 
widespread subspecies agree with an equi- 
table model or a random model (Rapoport, 
1982:41)? A broken stick model was dis- 
cussed and it was concluded that the divi- 
sion of lands among subspecies “‘seems to 
be a stochastic process” rather than an eq- 
uitable one. This question was considered 
in some detail and with the same conclusion 
by Anderson and Evensen (1978). 

Does the total size of the range ofa species 
affect the way it is divided into subspecies 
(Rapoport, 1982:42)? “It seems that in the 
very widespread species the bigger land- 
owners (sspp.) have a better chance of de- 
veloping into very big landowners,” and that 
as the size of a subspecies’ range decreases 
it becomes less likely to fragment into two 
parts. As a result there is greater equitability 
among the ranges of smaller subspecies. The 
author noted that this poses more questions 
than it answers. [In a way this is more stim- 
ulating than the common procedure in which 
an author concludes that we now have “‘ex- 
plained”’ something or other. ] 

The focus of areography on the areas of 
distribution or ranges of species, and on the 
sizes, shapes, and locations in space and 
time of these ranges, leads to other types of 
questions and answers. For example, An- 
derson (1977:11) was led to conclude that 
species “‘are about an order of magnitude 
(10 times) less ‘concentrated’ in each suc- 
cessively larger order of magnitude of range”’ 
(Fig. 1). This led Anderson (1985) to the 
conclusion that ‘“‘the geographic range of a 
species, regardless of its size, is more likely 
to decrease than to increase.”” The former 
‘““conclusion’”’ summarizes an observed pat- 
tern at one time, whereas the latter is an 
inference from that pattern and from the- 
oretical assumptions and considerations and 
may well be true over time spans of different 
duration. These two conclusions seem to 


ype ANDERSON AND PATTERSON 


180° 120° 60° 60° 0° 0° 120° 180° 
LLL TAS SS <6 TOA SEE 
AEE ag E77 LT La SS 
° Pet AN ag Turcee sae aaeae 
60° RES om 1 = coy 
ie / a = rapetai area 
Z s tp HA mma 
+ [AEH Ae | 
SR4abaN®): 6Aw) 
° ! i ag, Ug aenet 
se La RGN TY 


AG. ROT 


. Nearctic 

. Palearctic 
. Neotropical 
. Argentine 


<b ab cb cpap 


' 
H 
Ox 
ERS 
ererezey. 


— 
Aw 
aq 


. Mediterranean 
. Ethiopian 

. Oriental 

. Australian 

. West Indian 


0 1060 2000 3000 
Caen! Veet aE 


. Madagascan 


Fic. 2.—Four mammal faunal regions; (1) Holarctic, (2) Latin America, (3) Afro-Tethyan, and (4) 


Island, and ten subregions listed on map (Smith, 1983:462). 


have escaped the attention of Pagel et al. 
(1991:796) when they wrote that ‘“‘species 
with very small, and species with very large, 
geographic ranges are scarce.’’ Only half of 
this is true. Large ranges are scarce, small 
ranges are not. The inference (discussed by 
Anderson, 1985) about the relative proba- 
bilities of increases and decreases of range 
sizes in a dynamic Markovian system over 
time was not mentioned in their discussion 
of ecological aspects of the distribution of 
range sizes of North American mammals. 


Biotas Over Ecological Time 
Periods 


Regions and subregions. —Coincidence in 
the range limits of organisms points to broad 
biogeographic similarities among some lo- 
cal biotas and fundamental differences be- 
tween others. Lydekker’s (1896) account A 
Geographical History of Mammals served 
as a principal reference for mammalian geo- 
graphical classifications. Lydekker (1896) 
divided the world into three realms: the No- 


togaeic (including Australian, Polynesian, 
Hawaiian, and Austro—Malayan regions), the 
Neogaeic (limited to the Neotropical re- 
gion), and the Arctogaeic (including Mala- 
gasy, Ethiopian, Oriental, Holarctic, and 
Sonoran regions). Given the careful work 
represented in these early accounts, more 
recent contributions to the subject have in- 
volved either detailed analyses of biotic 
limits at the interface of such regions, or 
more quantitative approaches that create a 
fuller hierarchy of biogeographic regional- 
ism. Additions to this hierarchy awaited 
better inventories, but simple classification 
is merely a first step in attempting to un- 
derstand biogeographic patterns and pro- 
cesses. 

On a world scale an analysis of terrestrial 
mammal faunal regions using multidimen- 
sional scaling produces a classification (Fig. 
2) with four regions and ten subregions that 
is ‘more efficient and more internally con- 
sistent”? than the classic Sclater-Wallace 
classification (Smith, 1983). Smith’s anal- 
ysis was based on maps for families of living 
mammals (Anderson and Jones, 1979). 


BIOGEOGRAPHY Le 


How many faunal areas (whatever they 
may be named) should be recognized and 
where are their boundaries? This questions 
as it relates to mammals was addressed by 
Hagmeier and Stults (1964) and Hagmeier 
(1966), using squares in a 50-mile grid to 
tally boundaries of species from the set of 
North American distribution maps for 
mammals (from Hall and Kelson, 1959). A 
thoughtful critique of the analyses of Hag- 
meier and Simpson was written by Murray 
(1968). Many other detailed analyses at- 
tempting to define or recognize faunal 
regions, provinces, or similar areas have 
been conducted (for example Matson, 1982). 

Life zones.—Life-zone concepts mostly 
date from Merriam’s (1890) description of 
biotic associations on the San Francisco 
Mountains in northern Arizona. Merriam 
noted the resemblance between the replace- 
ment of communities as elevation de- 
creased along a transect with the replace- 
ment observed as latitude decreased on a 
continental scale. Attributing the underly- 
ing causation to physical effects of temper- 
ature and precipitation gradients, Merriam 
(1894) proposed a large-scale classification 
of habitats that proved ultimately unsatis- 
factory when applied on a larger continental 
or global scale. Reasons for this were ob- 
vious to Lydekker (1896), who recognized 
that historical opportunity was also a fun- 
damental factor. A clear correlation of fau- 
nas or biotas with elevation and latitude 
would only exist, as in North America, where 
mountain ranges tended to be north-south 
in orientation. East—west chains, such as the 
Pyrenees, Alps, and Himalayas, do not per- 
mit organisms that are restricted to the arc- 
tic or temperate zones of higher elevations 
to have direct access to corresponding belts 
at higher latitudes. 

As early as 1923, Dice criticized the con- 
cept of life zones on the grounds that in- 
dividual limitations of species ranges, and 
their common correlation with habitat, ren- 
dered this concept inaccurate and prone to 
error. Two other systems for delineating and 
naming biotic associations in North Amer- 


ica were the biomes of Clements and Shel- 
ford (1939) and the biotic provinces of Blair 
and Hubbell (1938) and Dice (1943). 

None of these specific systems is now in 
widespread use. Nevertheless, questions 
such as the following are fundamentally un- 
resolved and continue to attract attention. 
In what ways is it useful to delineate and 
designate biotic or faunal areas? How should 
this be done, on ecological grounds, on spa- 
tial or areal grounds, or on some combined 
ecogeographical basis? What exactly do we 
represent by these schemes? What conclu- 
sions or what predictions can be drawn 
therefrom? 

Diversity patterns and their correlates. — 
Early naturalists recognized that biotic di- 
versity 1S unequally distributed over the 
planet, being greater in tropical regions. 
Hershkovitz (1987) has shown that 26% of 
the world’s mammal fauna known to Lin- 
neaus in 1758, and 31% known to Buffon 
in his 1753-1789 compilations, came from 
the neotropics; corresponding figures for 
North America are 13% and 19%, respec- 
tively. Tropical “‘hotspots”’ of diversity at- 
tracted the attention of naturalist-explorers, 
whose collections continue to serve in the 
description and reappraisal of biotic diver- 
sity. It is a sad statement of modern science 
that, after 200 years, we still cannot esti- 
mate, to the nearest order of magnitude, the 
number of species coexisting on the planet 
(May, 1988). 

Diversity has been related to a host of 
abiotic and biotic factors. A complete listing 
and discussion is beyond the scope of this 
work but includes spatial heterogeneity, 
competition, predation, stability, produc- 
tivity, predictability, and seasonality (Em- 
len, 1973). Abiotic variables commonly 
correlated with these include area, latitude, 
longitude, elevation, precipitation, soils, and 
many others. Wright (1983) offered the 
““species-energy” relationship as an ulti- 
mate explanation for many of these more 
proximate factors. 

Among numerous mammalian examples, 
several recent papers illustrate diverse ap- 


I) 
we) 
aN 


CURACAO 
BONAIRE 


ARUBA 


iy 


ANDERSON AND PATTERSON 


GRENADA 


fom 


TRINIDAD 


VE NSE Zo ew 


Fic. 3.—Diagrammatic comparison of islands off the north coast of South America. The size of 
each square is proportional to the size of the indicated island. The shading of each square indicates 
the vegetation type of the island (black—rain forest, diagonal lines—traces of rain forest, white— 
xerophytic vegetation only). The numbers refer to the number of bat species recorded from the island. 
The straight line near the bottom represents the Venezuelan coastline, the zigzag line above it the 
100-fathom line (Koopman, 1958:432). 


proaches to characterizing diversity and use 
specific analytical features to heighten their 
resolution, including Flessa (1975), McCoy 
and Connor (1980), and Willig and Sandlin 
(1991). References cited in these works il- 
lustrate the breadth of the field. 

Island biogeography. —Knowledge of 
mammals on Mediterranean islands existed 
in classical Greek and Roman times. These 
studies were surprisingly contemporary in 
scope, identifying two of the most salient 
and prevalent properties of island life: (1) 
insular species often occur nowhere else and 
sometimes differ dramatically from main- 
land taxa; and (2) many insular forms are 
now extinct or endangered, illustrating their 
vulnerability to environmental and climatic 
changes. Such classical studies were known 
to both Darwin and Wallace in their pio- 
neering works on evolution and biogeog- 
raphy. At the founding of the ASM, mam- 
malian studies of island biogeography were 
relatively well integrated. One has only to 
consider the arguments of Grinnell and 
Swarth (1913), who published a truly mod- 
ern analysis of the disjunct bird and mam- 
mal faunas inhabiting the isolated San Ja- 
cinto Mountains in southern California (see 
below). 

Koopman (1958) examined the effects of 
island size, island isolation, former land 
connections, and ecological habitats on bats 


inhabiting islands off the north coast of 
South America. This multi-factorial ap- 
proach is informative (Fig. 3), because it 
demonstrates that analyses at many differ- 
ent scales of space, time, and organic di- 
versity are necessary. An analysis of non- 
volant mammals in southeast Asia enabled 
Heaney (1986) to postulate changing ex- 
tinction rates (Fig. 4) over time on a hy- 
pothetical island of 10,000 km7?. 

Studies of island biogeography changed 
most dramatically with the publication of 
the “equilibrium theory” of island bioge- 
ography by MacArthur and Wilson (1963, 
1967). This theory explained the species 
richness of any area as the product of two 
opposing rates: the migration of new species 
(J) and the extinction of existing ones (£). 
Ata certain species richness, determined by 
an island’s distance from colonization 
sources (affecting J) and its area (affecting 
E), the number of species tends toward a 
dynamic equilibrium —at this point, species 
composition changes (“‘turnover’’), but spe- 
cies number is maintained at equilibrium. 
Mostly forgotten in the stormy debates en- 
gendered by the equilibrium theory was its 
authors’ qualification (MacArthur and Wil- 
son, 1967:20-21): ‘‘a perfect balance be- 
tween immigration and extinction might 
never be reached .. . but to the extent that 
the assumption of a balance has enabled us 


BIOGEOGRAPHY 225 


50 


Lf 


40 


30 


20 


Extinction rate per 10000 years (%) 


ce) 2 4 6 8 


10 12 14 16 18 


Time after isolation (10 OOO years) 


Fic. 4.—Extinction rate for non-volant mammal faunas in SE Asia. The two filled circles are data 
points discussed in Heaney’s text; the curved line is an estimate of actual extinction rates for an island 
of 10,000 km? following isolation from the mainland. The exact shape of the curve is problematical 


(Heaney, 1986:155). 


to make valid new predictions, the equilib- 
rium concept is useful as a step... .” 

The equilibrium model stimulated much 
research activity, including a substantial 
body focused on mammals (reviewed by 
Brown, 1986). Despite their limited number 
of species compared to some other animal 
groups, mammals exhibit an enormous 
range of vagilities, from volant migrators to 
sessile burrowers. Variation in body size and 
generation time is also immense. Thus, 
mammals show a broad spectrum of dis- 
persal abilities and extinction proneness, and 
a correspondingly wide range of insular dis- 
tributional patterns, all of which make this 
group well suited for tests of biogeographic 
theory. 

Mammalian studies in island biogeogra- 
phy have critically altered basic paradigms 
within the field. Lomolino (1984, 1986) 
studied various aspects of mammalian dis- 
persal as they relate to colonization of near- 
shore islands in the St. Lawrence Seaway, 
finding that the traits that lead to initial col- 
onization of islands are not necessarily those 
promoting stable persistence there. Using 
this system and others, Lomolino (1986) 


showed that two principal variables of equi- 
librium theory, area and isolation, are not 
always independent in their effects on mam- 
malian species richness but may interact to 
produce “compensatory effects.”” For ex- 
ample, small near-shore islands may have 
a surfeit of species, because high rates of 
colonization from the mainland sometimes 
overwhelm high extinction rates expected 
from their limited areas (see also Hanski, 
1986). 

The greatest revision of island biogeog- 
raphy since MacArthur and Wilson has been 
the recognition that history plays a critical 
role in the derivation and status of insular 
biotas, even over “ecological time.’”? Em- 
pirical studies on many groups were to show 
that few archipelagos were actually at equi- 
librium (Gilbert, 1980); most appeared to 
be approaching a theoretical equilibrium, 
either from above (“biotic relaxation” via 
an excess of extinctions) or from below (un- 
der-saturation via an excess of coloniza- 
tions). Brown (1971) was the first to place 
such “‘nonequilibrial” island systems in the 
theoretical context of the MacArthur-Wil- 
son model. His now-classic study of mam- 


226 ANDERSON AND PATTERSON 


mals inhabiting Great Basin mountaintops 
presented in modern terms the arguments 
advanced by Grinnell and Swarth (1913) 58 
years earlier, that isolated mountain ranges 
were freely populated during favorable 
Pleistocene episodes and, following their 
disjunction, have suffered an excess of area- 
dependent extinctions. Brown’s (1971) in- 
terpretation of Great Basin faunal dynamics 
later received empirical support from Gray- 
son’s (1987) study of the Pleistocene fossils. 
Other studies have utilized Pleistocene rec- 
ords to substantiate inferences of historical 
derivation and dynamics of modern species 
richness (Ayer, 1936; Harris, 1990; Hope, 
1973; Heaney, 1984, 1986, 1991; Morgan 
and Woods, 1986; Patterson, 1984). 

The role of history in island biogeography 
is an area of considerable research activity, 
much of it involving mammals. Lawlor 
(1986) found significant differences in spe- 
cies-area slopes for mammalian faunas in- 
habiting “oceanic” archipelagos (populated 
de novo via overwater colonization) and 
““landbridge” archipelagos (fragments of 
formerly continuous areas subjected to local 
extinctions). Species-area slopes had pre- 
viously been considered “‘devoid of biolog- 
ical meaning” (Connor and McCoy, 1979) 
because earlier analyses failed to take ac- 
count of history. History also appears to 
influence patterns of insular species com- 
position in predictable ways. Patterson and 
Atmar (1986) showed that, in landbridge 
island archipelagos, species composition 
shows a nested subset pattern in which 
smaller islands support nested subsets of the 
species present on larger islands. Oceanic 
islands rarely show this highly non-random 
pattern (see also Patterson, 1990). 


Species Over Evolutionary Time 
Periods 


How should variability among popula- 
tions on a geographic scale and within 
species be analyzed and expressed or rep- 


resented? How should it be treated taxo- 
nomically? These are long-standing ques- 
tions. The quality and quantity of basic data 
on variation have been increasing through- 
out this century. This has made it both pos- 
sible and desirable to consider whether the 
traditional use of subspecies in taxonomy 
should be changed, whether by abandon- 
ment, modification, or supplementation. 
New methods of computation became read- 
ily available after 1950, and with the pro- 
liferation of electronic computers. New con- 
cepts, including phenetic analysis (in the 
form of numerical taxonomy and subse- 
quently in clustering procedures in biogeo- 
graphical analysis), phylogenetic systemat- 
ics (growing chiefly from Hennig, 1966), and 
vicariance biogeography (rooted in the work 
of Croizat, 1976, and other papers as early 
as 1958, but rapidly mutating or splintering 
in various ways), have stimulated work also. 

In the 1950s, concepts of subspecies and 
their nomenclature were discussed in a se- 
ries of articles, mostly in the pages of Sys- 
tematic Zoology. These included, in order 
of increasing perceived utility: Burt (1954), 
Hagmeier (1958), Doutt (1955), Lidicker 
(1962), Anderson (1966), Dillon (1961), and 
Durrant (1955). Today, subspecies have not 
been abandoned completely but are still in 
use at least in mammalogy and ornithology. 
There seem to have been some modifica- 
tions in the concept in that subspecies may 
be used on the average a little more cau- 
tiously and at a bit higher level (in the con- 
tinuum of degrees of difference between 
populations) (see also Engstrom et al., 1994). 
We feel that subspecies still have some use 
as a matter of convenience, but that the level 
in the continuum selected for subspecific 
recognition in any particular case is not in- 
herently more interesting biologically than 
other possible levels. Supplementary meth- 
ods are now in common use to deal with 
continuously varying degrees of difference, 
in both taxonomy and biogeography. 

The role of vicariance, currently defined 
as the splitting of a formerly continuous 
range or area into two or more parts, has 


BIOGEOGRAPHY 22d 


received considerable attention in recent 
decades. In a frequently cited paper on Ca- 
ribbean biogeography, Rosen (1975) pre- 
sented a vicariance model. It was said to 
have used data from mammals, but what 
data and how they were in fact used, or what 
the conclusions have to do with mammalian 
distribution is not clear; we are simply left 
with the implication that mammals con- 
form to the vicariance model. The descrip- 
tion of the method indicated that only 
monophyletic groups or individual species 
(which are regarded as monophyletic groups 
of populations) should be used. The author 
did not indicate which specific groups with- 
in the Mammalia were used in the analysis, 
nor which were hypothesized to be or dem- 
onstrated to be monophyletic. None of the 
five sources cited for mammalian data ex- 
plicitly indicated which groups may be 
monophyletic. 

Later studies of mammals in the Carib- 
bean region have provided little to docu- 
ment the relevance of the vicariance model 
to the distribution of modern mammals. 
Based on their study of bats, Baker and Gen- 
oways (1978) reported that dispersal by flight 
seems to be the most logical explanation for 
the present Antillean bat fauna, but also 
noted that “the fact that the vicariance 
model is not the best one to explain the 
origin of the bat fauna should not be taken 
as an indictment against the model” (Baker 
and Genoways, 1978:72). MacFadden 
(1980) suggested that the insectivores Ne- 
sophontes and Solenodon may be relicts of 
an early continental fauna. MacPhee and 
Woods (1982) concluded in stronger terms 
that “‘on the whole, long-distance, over-wa- 
ter rafting from the Americas remains the 
most likely mechanism for past land ver- 
tebrate immigration into the Caribbean.” 
Ernest Williams (in Woods, 1990) sum- 
marized the history of West Indian bioge- 
ography and the relative contributions of 
dispersalist and vicariance models. Because 
organisms do disperse and barriers do arise 
any model to be adequate in comprehen- 
siveness must include both dispersal and 


vicariance. The problem is to evaluate the 
roles of both processes in particular situa- 
tions as well as in general. In the same sym- 
posium volume, Karl Koopman (in Woods, 
1990:639) was “in full agreement with Ba- 
ker and Genoways (1978),” J. Knox Jones, 
Jr. (Gn Woods, 1990:653) also agreed “‘that 
overwater dispersal best explains present 
chiropteran distribution on Caribbean is- 
lands,” and Charles Woods (1990:741) also 
postulated overwater dispersal and evolu- 
tionary radiation on the islands as the prin- 
cipal factors in the origin of the rodent fau- 
na. 

In the 1970s, interest in developing meth- 
ods and actual applications of cladistic sys- 
tematics to biogeography were increasing. 
This decade of activity culminated ina 1979 
symposium at the American Museum of 
Natural History (Nelson and Rosen, 1981). 
In following years several text or reference 
books on this approach to biogeography were 
published (e.g., Humphries and Parenti, 
1986; Nelson and Platnick, 1981; Wiley, 
1988). Two special issues of Systematic Zo- 
ology (1988, nos. 3 and 4) included papers 
given at a later symposium on vicariance 
biogeography. There have not been many 
applications, successful or unsuccessful, 
specifically to mammalian biogeography. 
The magnitude and relative importance of 
vicariance biogeography to mammalian 
biogeography in the long term remains to 
be seen. 


Biotas Over Evolutionary Time 
Periods 


Biotic processes involving mammalian 
faunas of sizeable areas over relatively long 
time periods have attracted interest from 
several quarters between 1919 and 1994. 
Among these processes (and conceptual 
schemes for dealing with them) are the fol- 
lowing: 

Distributional “‘tracks.’’—In his famous 
three-volume opus, Croizat (1958:74) 


228 ANDERSON AND PATTERSON 


opined ““When we mention zoogeography, 
we most likely imagine a science of dis- 
persal.... In reality, what we get today as 
‘zoogeography,’ regardless of beauty of 
package and sound of label, is the lore orig- 
inally broadcast by Darwin and later on re- 
furbished by Matthew and his successors, 
Simpson, Mayr, etc.” Croizat’s solution to 
this was the formulation of “‘panbiogeog- 
raphy.” Relying on graphical analyses of the 
geographic ranges of taxa (called tracks), 
panbiogeography seeks to identify ancestral 
patterns of spatial and temporal distribu- 
tion, of which modern distributions are but 
relict fragments. Many of the 2,750 pages 
of Panbiogeography are devoted to mam- 
malian examples and their interpretation by 
Croizat. There are many other ways of using 
spatial patterns to gain insight into faunal 
development and composition. For exam- 
ple, the simple and empirical superimpo- 
sition of species boundaries on one map, 
whether showing a group with similar dis- 
tributions such as those centering on the 
Chihuahuan desert or all of the species of a 
larger group such as mammals, can be in- 
formative (e.g., Anderson, 1972; Anderson 
and Marcus, 1992; Armstrong, 1972; Find- 
ley, 1969; and Jones et al., 1985). 

Continental stability.—Simpson (1953) 
commented that “It remains possible that 
there were transoceanic continents or bridg- 
es or that continents drifted in the Triassic 
or earlier, but there is little good evidence 
that such was the fact. In any case such re- 
mote events would have little or no bearing 
on the present distribution of living things.” 
Within a few years there was plenty of ev- 
idence that continents were drifting, not only 
in the Triassic but at present, and there was 
serious consideration of the bearing of con- 
tinental drift on the present distribution of 
living things. Serious disagreements remain 
on the relative importance of that process 
compared to others. 

Centers of endemism. —This term has ap- 
peared in the literature in recent years and 
has been related to the concept of refugia 


noted below. It is not always clear what an 
author may mean by a “center of ende- 
mism.” In some cases it refers to a clustering 
of the centers of the geographic ranges of a 
number of species, especially species with 
rather small ranges. The occurrence of such 
a cluster is interpreted as evidence for a for- 
mer refugium. In other cases a center of 
endemism refers to an area in which a rel- 
atively high percentage of the species are 
endemic thereto. 

Ecological complementarity of different 
regions. —The extent to which (and the ways 
in which) faunas of similar major habitats, 
such as forests, deserts, or grasslands, are 
comparable has attracted recent interest and 
deserves more exploration. Such compari- 
sons involve both ecological and evolution- 
ary time scales and have as much to con- 
tribute to ecology, systematics, and 
functional morphology as to biogeography. 
The comparison of vertebrates in North 
American and South American deserts (the 
Sonoran and Monte, respectively) by Blair 
et al. (1976) is one example. 

Refugia. —The idea that certain geo- 
graphic regions have served as refugia for 
biotas against the vicissitudes of climate or 
competitors has been employed in various 
contexts. Over shorter Quaternary time 
scales, the somewhat controversial notion 
that warmer wet-dependent biotas endured 
Pleistocene cold or arid fluxes in isolated 
refugia has gained great application and ac- 
ceptance throughout the world. 

Findley (1969) applied this concept to in- 
terpreting distributions of montane and 
desert mammals in the southwestern part 
of the United States. The idea that refugia 
may have been important in tropical as well 
as temperate regions developed more re- 
cently. Cerquiera (1982) and Kinzey (1982) 
suggested that refugia applied to primate 
distributions in South America. Detailed 
map data for other mammalian groups have 
not been available to test this model. Most 
discussions and examples have centered on 
plants, birds, and butterflies for which map 


BIOGEOGRAPHY 229 


data were available (Whitmore and Prantz, 
1987). 

Over vast geologic time scales, the island 
continents of Australia and South America 
served as Tertiary and Quaternary refuges 
for various archaic groups of mammals, 
some still living, that were replaced by later 
lineages on other continents (e.g., Simpson, 
1980). 

Interchange. —Studies of faunal inter- 
change analyze the patterns and processes 
involved when historically differentiated bi- 
otas intermingle. Different geographic the- 
aters and faunas provide insights into var- 
ious levels of this dynamic biogeography. 
Musser’s continuing studies of mammals, 
especially murid rodents, from Sulawesi 
probe the limits of Wallace’s line and its 
effects on biological evolution. Hoffmann, 
Vorontsov, and their coworkers focused 
more than a decade of work on the dynamic 
character of Beringia, a trans—oceanic land- 
bridge that opened and closed repeatedly 
through the Pleistocene with the waxing and 
waning of continental glaciers, permitting 
interchange of Palearctic and Nearctic ele- 
ments of the Holarctic biota. Surely the best 
studied example of biotic interchange in- 
volves the Nearctic and Neotropical faunas 
juxtaposed by the emergence of a Pana- 
manian land bridge roughly 3 million years 
ago (e.g., Stehli and Webb, 1985). The strong 
differentiation of faunas isolated through- 
out the Cenozoic, their biotic diversity, and 
a detailed chronology of events derived from 
abundant fossil remains combine to pro- 
duce this unparalleled record of biogeo- 
graphic dynamics. 

Conservation biogeography. —Of course, 
mammalogists were leading figures in the 
nascent conservation movement in North 
America, but much of their activity was in 
fields other than biogeography. ‘Applied 
biogeography” took root soon after the for- 
mulation of the Equilibrium Theory of Is- 
land Biogeography (MacArthur and Wilson, 
1967). This theory was quickly applied to 
the conservation of species in habitat frag- 


ments, distilling a series of “‘geometrical 
rules” of reserve design (summarized by Di- 
amond, 1976) that were at once the subjects 
of both acclaim and criticism. Mammalo- 
gists have contributed significantly to con- 
tinued refinements of this field. 

East (1981), Heaney (1986), and New- 
mark (1987) drew a series of conclusions 
about conservation of African, Philippine, 
and North American mammals, respective- 
ly, based on correlations between extinction 
rates and reserve or island area. Patterson 
and Atmar (1986) also argued that parks 
need to be large to fulfill their basic function, 
basing their conclusions on analyses of spe- 
cies composition. In a nested subset pattern, 
many small fragments each tend to support 
the same set of species; rare or narrowly 
restricted endemics most in need of protec- 
tion are found only in the largest, richest 
fragments. Kitchener et al. (1980) devel- 
oped a point-by-point assessment of the 
conservation value of heath fragments to 
small mammals of western Australia. The 
role of corridors between fragments in help- 
ing to sustain isolated populations of rain- 
forest possums was examined by Laurance 
(1990), substantiating a “‘rescue effect”’ pre- 
viously hypothesized by Brown and Kodric- 
Brown (1977); this subdiscipline is attract- 
ing much current attention. By showing that 
the majority of species in all higher taxa 
have small geographic ranges, Anderson 
(1985) underscored the vulnerability of most 
taxa to localized environmental changes. 
Literature on the role of biogeographic the- 
ory in conserving diverse tropical com- 
munities was recently reviewed by Patter- 
son (1991). 

Readers familiar with the biogeographic 
literature will readily appreciate how cur- 
sory this review has been. In fact, there are 
few aspects of biology that do not have bio- 
geographic consequences, either by affecting 
geographic range limits, abundance within 
the range, or patterns of species coexistence. 
However, we hope that our survey succeeds 
in indicating the breadth and pluralistic na- 


230 


ture of biogeographic research and its sub- 
stantial role within mammalogy, past, pres- 
ent, and future. 


Literature Cited 


ALKon, P. U., AND D. SALTz. 1988. Foraging time 
and the northern range limits of Indian crested por- 
cupines (Hystrix indica Kerr). Journal of Biogeog- 
raphy, 15:403-408. 

ANDERSON, S. 1966. Taxonomy of gophers, especially 
Thomomys in Chihuahua. Systematic Zoology, 15: 
189-198. 

1972. Mammals of Chihuahua, taxonomy 

and distribution. Bulletin of the American Museum 

of Natural History, 148:149-410. 

. 1977. Geographic ranges of North American 

terrestrial mammals. American Museum Novitates, 

2629:1-15. 

1985. The theory of range-size (RS) distri- 
butions. American Museum Novitates, 2833:1-20. 

ANDERSON, S., AND M. K. EvENSEN. 1978. Random- 
ness in allopatric speciation. Systematic Zoology, 27: 
421-430. 

ANDERSON, S., AND J. K. Jones, JR. 1979. Recent 
mammals of the world: a synopsis of families. Ron- 
ald Press, New York, 453 pp. 

ANDERSON, S., AND L. F. MARcus. 1992. Aerography 
[sic] of Australian tetrapods. Australian Journal of 
Zoology, 40:627-651. 

ANDREWARTHA, H. G., AND L. C. Brrcu. 1954. The 
distribution and abundance of animals. University 
Chicago Press, Chicago, 782 pp. 

ARMSTRONG, D. M. 1972. Distribution of mammals 
in Colorado. Monograph of the Museum of Natural 
History, The University of Kansas, 3:x + 415 pp. 

Ayer, M. Y. 1936. The archeological and faunal ma- 
terial from Williams Cave, Guadalupe Mountains, 
Texas. Proceedings of the Academy of Natural Sci- 
ence, Philadelphia, 88:599-618. 

Baccus, J. T., R. E. GREER, AND G. G. RAuN. 1971. 
Additional records of Baiomys taylori (Rodentia: 
Cricetidae) for northern Texas. The Texas Journal 
of Science, 23:148-149. 

BAKER, R. J., AND H. H. GeENoways. 1978. Zooge- 
ography of Antillean bats. Academy of Natural Sci- 
ences of Philadelphia, Special Publication, 13:53- 
97. 

Biarr, W. F., AND T. H. Hussey. 1938. The biotic 
districts of Oklahoma. The American Midland Nat- 
uralist, 20:425-454. 

Biair, W. F., A. C. HULSE, AND M. A. Mares. 1976. 
Origins and affinities of vertebrates of the North 
American Sonoran Desert and the Monte Desert of 
northwestern Argentina. Journal of Biogeography, 
3:1-18. 

BLONDEL, J. 1987. From biogeography to life history 
theory: a multithematic approach illustrated by the 
biogeography of vertebrates. Journal of Biogeogra- 
phy, 14:405-422. 

Brown, J. H. 1971. Mammals on mountaintops: 
nonequilibrium insular biogeography. The Ameri- 
can Naturalist, 105:467-478. 


ANDERSON AND PATTERSON 


. 1986. Two decades of interactions between 
the MacArthur—Wilson model and the complexities 
of mammalian distributions. Pp. 231-251, in Island 
biogeography of mammals (L. R. Heaney and B. D. 
Patterson, eds.). Biological Journal of the Linnean 
Society 28 (1&2) and Academic Press, London, 271 
pp. 

Brown, J. H., AND A. Kopric-Brown. 1977. Turn- 
over rates in insular biogeography: effect of immi- 
gration on extinction. Ecology, 58:445-449. 

Burt, W.H. 1954. The subspecies category in mam- 
mals. Systematic Zoology, 3:99-104. 

CERQUIERA, R. 1982. South American landscapes and 
their mammals. Pp. 53-75, in Mammalian biology 
in South America (M. A. Mares and H. H. Geno- 
ways, eds.). Special Publication, Pymatuning Lab- 
oratory of Ecology, University of Pittsburgh, 6:1- 
539. 

CHOATE, J. R., AND K. M. REED. 1986. Historical 
biogeography of the woodchuck in Kansas. Prairie 
Naturalist, 18:37-42. 

CLEMENTS, F. E., AND V. E. SHELFORD. 1939. Bio- 
ecology. J. Wiley & Sons, New York, vi + 425 pp. 

Cockrum, E. L. 1948. The distribution of the hispid 
cotton rat in Kansas. Transactions of the Kansas 
Academy of Science, 51:306-312. 

Connor, E. F., AND E. D. McCoy. 1979. The statis- 
tics and biology of the species—area relationship. The 
American Naturalist, 113:791-833. 

CorseT, G. B. 1978. The mammals of the Palaearctic 
Region: a taxonomic review. British Museum (Nat- 
ural History), London, and Cornell University Press, 
Ithaca, 314 pp. 

CorseT, G. B., AND J. E. Hitt. 1992. The mammals 
of the Indomalayan Region: a systematic review. 
Natural History Museum Publications, Oxford Uni- 
versity Press, Oxford, 6 + 488 pp. 

CroizaT, L. 1958. Panbiogeography, or an introduc- 
tory synthesis of zoogeography, phytogeography, and 
geology; with notes on evolution, systematics, ecol- 
ogy, anthropology, etc. Vol. 1—The New World, 
1,018 pp. Vols. Ila and IIb— The Old World, by the 
author, Caracas, Venezuela, 1,731 pp. 

CROIZAT-CHALEN, L. C. M. 1976. Buiogeografia an- 
alytica y sintetica (““Panbiogeografia”’) de los Amer- 
icas. Biblioteca de la Academia de Ciencias Fisicas, 
Matematicas y Naturales (Caracas), vol. 15, 16, 1- 
454, 455-890. 

DARLINGTON, P. J., JR. 1957. Zoogeography: the geo- 
graphical distribution of animals. J. Wiley & Sons, 
New York, 675 pp. 

DIAMOND, J. M. 1976. Relaxation and differential 
extinction on land—bridge islands: applications to 
natural preserves. Pp. 616-628, in Proceedings of 
the 16th International Ornithological Congress, Can- 
berra, Australia. 

Dice, L. R. 1923. Life zones and mammalian distri- 
bution. Journal of Mammalogy, 4:39-47. 

1943. The biotic provinces of North Amer- 
ica. University of Michigan Press, Ann Arbor, 78 
pp. 

Ditton, L. S. 1961. Historical subspeciation in the 
North American marten. Systematic Zoology, 10: 
49-64. 

Doutrt, J. K. 1955. Microgeographic races in mam- 
mals. Systematic Zoology, 4:179-185. 


BIOGEOGRAPHY Zon 


DuRRANT, S. D. 1955. In defense of subspecies. Sys- 
tematic Zoology, 4:186-190. 

East, R. 1981. Species—area curves and populations 
of large mammals in African savanna reserves. Bi- 
ological Conservation, 21:111-126. 

EISENBERG, J. F. 1989. Mammals of the Neotropics, 
the northern Neotropics, vol. 1. University of Chi- 
cago Press, Chicago, 449 pp. 

EMLEN, J. M. 1973. Ecology: an evolutionary ap- 
proach. Addison-Wesley, Reading, Massachusetts, 
493 pp. 

ENGsTrRoM, M. D., J. R. CHOATE, AND H. H. GENowaAys. 
1994. Taxonomy. Pp. 179-199, in Seventy-five years 
of mammalogy (1919-1994) (E. C. Birney and J. R. 
Choate, eds.). Special Publication, The American 
Society of Mammalogists, 1 1:1-433. 

FINDLEY, J. S. 1969. Biogeography of southwestern 
boreal and desert mammals. University of Kansas, 
Museum of Natural History Miscellaneous Publi- 
cation, 51:113-128. 

Fitcu, H. S., P. GoopRUM, AND C. NEwMaAN. 1952. 
The armadillo in the southeastern United States. 
Journal of Mammalogy, 33:21-37. 

FiessA, K. W. 1975. Area, continental drift, and 
mammalian diversity, Paleobiology, 1:189-194. 
Frey, J. K. 1992. Response of a mammalian faunal 
element to climatic changes. Journal of Mammalogy, 

73:43-S0. 

Genoways, H. H., AND D. A. ScHLITTER. 1967. 
Northward dispersal of the hispid cotton rat in Ne- 
braska and Missouri. Transactions of the Kansas 
Academy of Science, 69:3-4. 

GILBERT, F.S. 1980. The equilibrium theory of island 
biogeography: fact or fiction? Journal of Biogeogra- 
phy, 7:209-235. 

Grayson, D. K. 1987. The biogeographic history of 
small mammals in the Great Basin: observations on 
the last 20,000 years. Journal of Mammalogy, 68: 
359-375. 

GRINNELL, J., AND H. S. SwartH. 1913. An account 
of the birds and mammals of the San Jacinto area 
of southern California [with remarks upon the be- 
havior of geographic races on the margins of their 
habitats]. University of California, Publications in 
Zoology, 10:197-406. 

GuILbAyY, J. E., P. S. MARTIN, AND A. D. McCrapy. 
1964. New Paris No. 4: a late Pleistocene cave de- 
posit in Bedford County, Pennsylvania. Bulletin of 
the National Speleological Society, 26:121-194 + 2 
color pls. 

HAGMEIER, E. M. 1958. Inapplicability of the sub- 
species concept to North American marten. System- 
atic Zoology, 7:1-7. 

1966. A numerical analysis of the distribu- 
tional patterns of North American mammals. II. Re- 
evaluation of the provinces. Systematic Zoology, 15: 
279-299. 

HAGMEIER, E. M., AND C. D. Stutz. 1964. A nu- 
merical analysis of the distributional patterns of North 
American mammals. Systematic Zoology, 13:125- 
155. 

HAtt, E. R. 1946. Subspecies of man at the peace 
table. Journal of Mammalogy, 27:358-364. 

. 1981. The mammals of North America. Sec- 

ond ed. John Wiley & Sons, New York, 1:1-600 + 

90, 2:601-1181 + 90. 


HALL, E. R., AND K. R. KEtson. 1959. The mammals 
of North America. Ronald Press, New York, 2 vols, 
1,083 pp + 79 pp. index. 

Hansen, R. M. 1962. Dispersal of Richardson ground 
squirrel in Colorado. The American Midland Nat- 
uralist, 68:58-66. 

HanskI, I. 1986. Population dynamics of shrews on 
small islands accord with the equilibrium model. Pp. 
23-36, in Island biogeography of mammals (L. R. 
Heaney and B. D. Patterson, eds.). Biological Journal 
of the Linnean Society 28 (1&2) and Academic Press, 
London, 271 pp. 

Harris, A. H. 1990. Fossil evidence bearing on 
Southwestern mammalian biogeography. Journal of 
Mammalogy, 71:219-229. 

HEANEY, L.R. 1984. Mammalian species richness on 
islands on the Sunda Shelf, Southeast Asia. Oecolo- 
gia, 61:11-17. 

1986. Biogeography of mammals in Asia: 

estimates of rates of colonization, extinction and spe- 

ciation. Pp. 127-165, in Island biogeography of 
mammals (L. R. Heaney and B. D. Patterson, eds.). 

Biological Journal of the Linnean Society, 28 (1&2) 

and Academic Press, London, 271 pp. 

1991. An analysis of patterns of distribution 
and species richness among Philippine fruit bats 
(Pteropodidae). Bulletin of the American Museum 
of Natural History, 206:145-167. 

HENNIG, W. 1966. Phylogenetic systematics. Uni- 
versity of Illinois Press, Urbana, 263 pp. 

HERSHKOvITZ, P. 1987. A history of the recent mam- 
malogy of the Neotropical Region from 1492 to 1850. 
Pp. 11-98, in Studies in neotropical mammalogy: 
essays in honor of Philip Hershkovitz (B. D. Patter- 
son and R. M. Timm, eds.). Fieldiana: Zoology, new 
series, number 39, 506 pp. 

HEssE, R., W. C. ALLEE, AND K. P. ScHmipt. 1937. 
Ecological animal geography. John Wiley & Sons, 
New York, xiv + 597 pp. 

Hit, J. E. 1947. A zoologist looks at raciology: a 
reply to Dr. E. Raymond Hall’s “‘Zoological subspe- 
cies of man at the Peace Table.” Journal of Mam- 
malogy, 28:87-89. 

Hore, J. H. 1973. Mammals of the Bass Straits Is- 
lands. Proceedings of the Royal Society of Victoria, 
85:163-195. 

Horton, D. R. 1974. Species movement in zooge- 
ography. Journal of Biogeography, 1:155-158. 

Husss, C. L. (Ep.). 1958. Zoogeography. American 
Association for the Advancement of Science, Pub- 
lication 51, 509 pp. 

Hu.t, D. 1988. Science as a process: an evolutionary 
account of the social and conceptual development 
of science. University of Chicago Press, Chicago, 586 
pp. 

HuMpPurIes, C. J., AND L. PARENTI. 1986. Cladistic 
biogeography. Oxford Monographs on Biogeogra- 
phy. Clarendon Press, Oxford, 98 pp. 

Jones, J. K., Jk., D. M. ARMSTRONG, AND J. R. CHOATE. 
1985. Guide to mammals of the plains states. Uni- 
versity of Nebraska Press, Lincoln, 371 pp. 

Kinzey, W. G. 1982. Distribution of primates and 
forest refuges. Pp. 445-482, in Biological diversifi- 
cation in the tropics (G. T. Prance, ed.). Columbia 
University Press, New York, 714 pp. 

KITCHENER, D. J., A. CHAPMAN, B. G. Muir, AND M. 


ios ANDERSON AND PATTERSON 


PALMER. 1980. The conservation value for mam- 
mals of reserves in the western Australian wheatbelt. 
Biological Conservation, 18:179-207. 

Koopman, K. F. 1958. Land bridges and ecology in 
bat distribution on islands off the northern coast of 
South America. Evolution, 12:429-439. 

LAuRANCE, W. F. 1990. Comparative responses of 
five arboreal marsupials to tropical forest fragmen- 
tation. Journal of Mammalogy, 71:641-653. 

Lawior, T. E. 1986. Comparative biogeography of 
mammals on islands. Pp. 99-125, in Island bioge- 
ography of mammals (L. R. Heaney and B. D. Pat- 
terson, eds.). Biological Journal of the Linnean So- 
ciety, 28 (1&2) and Academic Press, London, 271 
pp. 

Lipicker, W. Z., JR. 1962. The nature of subspecies 
boundaries in a desert rodent and its implications 
for subspecies taxonomy. Systematic Zoology, 11: 
160-171. 

Lomo.ino, M. V. 1984. Immigrant selection, pre- 
dation and the distributions of Microtus pennsyl- 
vanicus and Blarina brevicauda on islands. The 
American Naturalist, 123:468-483. 

1986. Mammalian community structure on 

islands: the importance of immigration, extinction, 

and interactive effects. Pp. 1-21, in Island bioge- 
ography of mammals (L. R. Heaney and B. D. Pat- 
terson, eds.). Biological Journal of the Linnean So- 

ciety, 28 (1&2) and Academic Press, London, 271 


pp. 

Lona, C. A. 1963. Mathematical formulas expressing 
faunal resemblance. Transactions of the Kansas 
Academy of Science, 66:138-140. 

LyDEKKER, R. 1896. A geographical history of mam- 
mals. Cambridge University Press, Cambridge, 400 
pp. 

MacArTHur, R. H., AND E. O. WiLson. 1963. An 
equilibrium theory of island biogeography. Evolu- 
tion, 17:373-387. 

1967. The theory of island biogeography. 
Monographs in Population Biology, Princeton Uni- 
versity Press, Princeton, 1:1—203. 

MacFappen, B. J. 1980. Rafting mammals or drift- 
ing islands?: biogeography of the Greater Antillean 
insectivores Nesophontes and Solenodon. Journal of 
Biogeography, 7:11-—22. 

MaAcPHEE, R. D. E., AND C. A. Woops. 1982. A new 
fossil cebine from Hispaniola. American Journal of 
Physical Anthropology, 58:419-436. 

Matson, J. O. 1982. Numerical analysis of rodent 
distributional patterns in Zacatecas, México. Journal 
of Mammalogy, 63:73-84. 

MaTTHEw, W. D. 1915. Climate and evolution. An- 
nals of the New York Academy of Science, 24:171- 
318. 

May, R. M. 1988. How many species are there on 
Earth? Science, 241:1441-1449. 

McCoy, E. D., AND E. F. Connor. 1980. Latitudinal 
gradients in the species diversity of North American 
mammals. Evolution, 34:193-203. 

McIntTosu, R. P. 1985. The background of ecology. 
Cambridge studies in ecology. Cambridge Univer- 
sity Press, Cambridge, 383 pp. 

. 1987. Pluralism in ecology. Annual Review 

of Ecology and Systematics, 18:321-342. 


MERRIAM, C. H. 1890. Results of a biological survey 
of the San Francisco Mountain region and desert of 
the Little Colorado, Arizona. North American Fau- 
na, 3:136 pp. + frontisp., 14 pls., and 5 maps [in- 
cluding one section by L. Stejneger]. 

1894. Laws of temperature control of the 
geographic distribution of terrestrial animals and 
plants. National Geographic Magazine, 6:229-238. 

MoraGan, G. S., AND C. A. Woops. 1986. Extinction 
and the zoogeography of West Indian land mam- 
mals. Pp. 167-203, in Island biogeography of mam- 
mals (L. R. Heaney and B. D. Patterson, eds.). Bi- 
ological Journal of the Linnean Society, 28 (1&2) 
and Academic Press, London, 271 pp. 

Murray, K. F. 1968. Distribution of North Amer- 
ican mammals. Systematic Zoology, 17:89-102. 
NEtson, G. 1978. From Candolle to Croizat: com- 
ments on the history of biogeography. Journal of the 

History of Biology, 11:269-305. 

NELSON, G., AND N. PLATNICK. 1981. Systematics 
and biogeography: cladistics and vicariance. Colum- 
bia University Press, New York, 567 pp. 

NELSON, G., AND D. E. ROSEN (EDs.). 1981. Vicari- 
ance biogeography, a critique. Columbia University 
Press, New York, 593 pp. 

Newmark, W. D. 1987. A land-bridge island per- 
spective on mammalian extinctions in western North 
American parks. Nature, 325:430—-432. 

PACKARD, R.L. 1963. Distribution of the black-tailed 
jackrabbit in eastern Texas. The Texas Journal of 
Science, 15:107-110. 

PAGEL, M. D., R. M. May, AND A. R. Coie. 1991. 
Ecological aspects of the geographical distribution 
and diversity of mammalian species. The American 
Naturalist, 137:791-815. 

Patterson, B. D. 1984. Mammalian extinction and 
biogeography in the Southern Rocky Mountains. Pp. 
247-293, in Extinctions (M. H. Nitecki, ed.). Uni- 
versity of Chicago Press, Chicago, 354 pp. 

. 1990. On the temporal development of nest- 

ed subsets patterns of species composition. Oikos, 

59:330-342. 

. 1991. The integral role of biogeographic the- 
ory in the conservation of tropical forest diversity. 
Pp. 124-149, in Latin American mammalogy: his- 
tory, biodiversity, and conservation (M. A. Mares 
and D. J. Schmidly, eds.). University of Oklahoma 
Press, Norman, xviii + 468 pp. 

PATTERSON, B. D., AND W. ATMAR. 1986. Nested 
subsets and the structure of insular mammalian fau- 
nas and archipelagos. Pp. 65-82, in Island bioge- 
ography of mammals (L. R. Heaney and B. D. Pat- 
terson, eds.). Biological Journal of the Linnean 
Society, 28 (1&2) and Academic Press, London, 271 
pp. 

PATTERSON, B. D., AND J. H. BRown. 1986. Region- 
ally nested patterns of species composition in gra- 
nivorous rodent assemblages. Journal of Biogeog- 
raphy, 18:395-402. 

Rapoport, E. H. 1982. Areography, geographical 
strategies of species. Pergamon Press, Oxford, xvi + 
269 pp. 

REDFORD, K. H., & J. F. EISENBERG. 1992. Mammals 
of the Neotropics, the southern cone, vol. 2., Chile, 


BIOGEOGRAPHY 233 


Argentina, Uruguay, and Paraguay. University of 
Chicago Press, 430 pp. 

REICHMAN, O. J. 1984. Spatial and temporal varia- 
tion of seed distributions in Sonoran desert soils. 
Journal of Biogeography, 11:1-11. 

Rosen, D.E. 1975. A vicariance model of Caribbean 
biogeography. Systematic Zoology, 24:431-464. 

ScLATER, P. L. 1858. On the general geographical 
distribution of the members of the Class Aves. Pro- 
ceedings of the Linnaean Society, London, 2:130- 
146. 

Smpson, G. G. 1953. Evolution and geography, an 
essay on historical biogeography, with special ref- 
erence to mammals. Condon Lectures, Oregon State 
System of Higher Education, 64 pp. 

. 1964. Species density of North American Re- 

cent mammals. Systematic Zoology, 13:57-73. 

1980. Splendid isolation: the curious history 
of South American mammals. Yale University Press, 
New Haven, Connecticut, 266 pp. 

Smitu, C.H. 1983. A system of world mammal fau- 
nal regions. I. Logical and statistical derivation of 
the regions. Journal of Biogeography, 10:455-466. 

1983. A system of world mammal faunal 
regions. II. The distance decay effect upon inter-re- 
gional affinities. Journal of Biogeography, 10:467- 
482. 

SmiTH, J. D., AND T. E. LAwtor. 1964. Additional 
records of the armadillo in Kansas. The Southwest- 
ern Naturalist, 9:48-49. 


STEHLI, F. G., AND S. D. WEBB (EDS.). 1985. The great 
American biotic interchange. Plenum Press, New 
York, 532 pp. 

STRAHAN, R. 1983. The Australian Museum com- 
plete book of Australian mammals. Angus & Rob- 
ertson Publishers, Sydney, xvi + 530 pp. 

WEGENER, A. 1912. ‘Die Entstehung der Konti- 
nente.”’ Dr. A. Petermanns Mitteilungen aus Justus 
Perthes’ Geographischer Anstart, year 58 (April), pp. 
185-195; (May), pp. 253-256; (June), pp. 305-309. 

WHITMORE, T. C., AND G. T. PRANTZ (EDS.). 1987. 
Biogeography and Quaternary history in tropical 
America. Clarendon Press, New York, x + 214 pp. 

Witey,E.O. 1988. Vicariance biogeography. Annual 
Review of Ecology and Systematics, 19:513-542. 

WiLic, M. R., AND E. A. SANDLIN. 1991. Gradients 
of species density and species turnover in New World 
bats: a comparison of quadrat and band methodol- 
ogies. Pp. 81-96, in Latin American mammalogy: 
history, biodiversity, and conservation (M. A. Mares 
and D. J. Schmidly, eds.). University of Oklahoma 
Press, Norman, Oklahoma, 468 pp. 

WILson, J. W., III. 1974. Analytical zoogeography of 
North American mammals. Evolution, 28:124—-140. 

Woops, C. A. (ED.). 1990. Biogeography of the West 
Indies, past, present, and future. Sandhill Crane Press, 
Gainesville, Florida, 878 pp. 

WRIGHT, D. H. 1983. Species-energy theory: an ex- 
tension of species—area theory. Oikos, 41:496-506. 


ANATOMY 


CARLETON J. PHILLIPS 


Introduction 


he Editors of this volume asked me to 
write about “anatomy” as it relates to 
mammalogy, using 1919 as a starting point. 
This seemed like a straight-forward task for 
a mammalogist interested in anatomy. Af- 
ter all, it is perfectly rational to believe that 
knowledge of structure-function complexes 
is an underpinning for understanding eco- 
logical, physiological, developmental, and 
behavioral patterns in mammals. Structure 
in static form is the cornerstone of system- 
atics and structure in dynamic form is the 
cornerstone of understanding adaptive ra- 
diation. It also is reasonable to think that 
an historical perspective on mammalian 
anatomy could be developed with 1919 as 
the starting point. Indeed, D’Arcy Went- 
worth Thompson’s treatise, On Growth and 
Form, had been published only a few years 
earlier, in 1917. This remarkable example 
of early 20th-Century scholarship still is a 
seminal source of ideas and hypotheses in- 
tegrating anatomy, physiology, and devel- 
opment in an evolutionary setting (Bonner, 
1966). 

Regardless of the expectations of the Ed- 
itors, the assigned task far exceeded what 
could be reviewed in a brief essay. After all, 
what is anatomy? In a broad sense it is an 


234 


academic discipline that presently encom- 
passes a stunning array of topics. At one 
extreme there is ultrastructure of mamma- 
lian cells and organelles, cell cytoskeletal 
features, intracellular localizations of secre- 
tory products and mRNAs, and three-di- 
mensional configurations of basal laminae 
and neural networks; at the other extreme 
there is functional gross anatomy in the con- 
text of feeding, locomotion, physiology, or 
ecology. 

Unlike some other scientific disciplines, 
the field of anatomy has expanded dramat- 
ically since 1919. So, while some research 
still retains the basic descriptive elements 
of earlier gross and microscopic anatomy 
(e.g., Brylski, 1933; Carleton, 1985; Forman 
and Phillips, 1993; Lay, 1993), technolog- 
ical advances in microscopy, cytochemis- 
try, and immunohistochemistry, and foun- 
dation research in molecular biology and 
biochemistry have pushed the frontier of 
the discipline to the subcellular level (Phil- 
lips and Tandler, 1987). Today, a com- 
parative anatomical study might involve 
immunohistochemical localization of neu- 
rotransmitters in the retina (e.g., Studholme 
et al., 1987), regulatory peptides in the di- 
gestive tract (Mennone et al., 1986), the ge- 


ANATOMY 2905 


ometry of neurons (Purves and Lichtman, 
1985), somatotrophic mapping of the brain 
(Calford et al., 1985), comparative ultra- 
structure (Phillips, 19855; Tandler et al., 
1990) or morphological modifications to 
cellular organelles such as mitochondria 
(Tandler and Phillips, 19935), or Golgi 
complexes in secretory cells (Tandler et al., 
in press a). Moreover, technology has con- 
verted the discipline from a largely descrip- 
tive enterprise to an experimental, function- 
oriented science. The latter being true at 
both gross and microscopic levels of anat- 
omy, one finds functional analyses of gaits, 
climbing, swimming, and dentition (e.g., 
Cartmill, 1985; Fish, 1982; Hildebrand, 
1985a, 1985b; Kiltie, 1981; Van de Graaff 
et al., 1982), integrations of gross and mi- 
croscopic anatomy with physiology (Dia- 
mond, 1992; Rodriguez-Colunga et al., 
1992; Sands et al., 1977), as well as analyses 
of the genetic basis of proteins composing 
the eye lens (Piatigorsky and Wistow, 1989), 
or genes responsible for the microanatom- 
ical structure of mammalian tooth enamel 
(Greenberg et al., 1984). 

Collectively, a staggering amount of sci- 
entific and quasi-scientific information on 
mammalian anatomy has been accumulat- 
ed since 1919. Thousands of journal arti- 
cles, hundreds of institutional publications 
and monographs, and hundreds of books 
would qualify for consideration. Currently, 
Kent Van de Graaff has estimated that more 
than 80 journals carry anatomical articles 
about mammals (Van de Graaff, in litt.). 
Some of these are obvious: e.g., Anatomy 
and Embryology; Journal of Morphology; 
Journal of Anatomy; and Anatomical Rec- 
ord. However, most would be overlooked 
by an anatomist of 1919 vintage. For ex- 
ample, there are many with such unlikely 
titles as The Journal of Wildlife Manage- 
ment; Journal of Ultrastructure Research; 
European Journal of Cell Biology, Micros- 
copy Research and Techniques; Structural 
Biology, Differentiation; Cell and Tissue 
Research, Brain; Behavior and Evolution; 
or Growth. All of these regularly publish 


articles with information that includes some 
aspect of mammalian anatomy, particularly 
microscopic and subcellular anatomy. So, 
it is apparent from the foregoing synopsis 
that only a very trimmed version of the sub- 
ject proposed by the Editors would be man- 
ageable for the present essay. 

Ultimately, I decided to focus on anato- 
my in the context of American mammalogy. 
Thus, I offer apologies to the large number 
of potential readers, especially my Euro- 
pean colleagues and specialists in functional 
anatomy or cell structure, whose excellent 
work or specific interests in anatomy either 
are not mentioned or not cited extensively 
in this essay. Moreover, I trust that all read- 
ers will appreciate the fact that I have been 
very judicious in selecting citations: my in- 
tention has been to provide easily accessible 
examples, many taken from the Journal of 
Mammalogy, that could serve to illustrate 
a particular point. I should mention, how- 
ever, that even with the many citations taken 
from the Journal of Mammalogy, there are 
at least three times more articles (including 
many of excellent quality) that might have 
been cited from this journal alone! I also 
have used the opportunity to introduce some 
literature from historians of science and 
philosophers of science, whose perspectives 
are welcome in any attempt to understand 
our present state as mammalogists. How- 
ever, the fact remains that only a small frac- 
tion of the huge body of anatomical litera- 
ture is mentioned herein. 

Aside from narrowing the overall topic of 
anatomy to manageable proportions, a fo- 
cus on anatomy in American mammalogy 
allowed me to explore, from a personal per- 
spective, two observations about anatomy 
in terms of our discipline. First, while 
“mammalian anatomy” is a useful umbrella 
term, there appear to be at least three dif- 
ferent kinds of anatomical research on 
mammals. One of these is conducted in the 
American medical school environment (and 
in dental and veterinary medical schools), 
another is conducted by persons who regard 
themselves as morphologists or experimen- 


236 PAT LETS 


tal zoologists rather than anatomists or 
mammalogists, and yet another is conduct- 
ed by persons who primarily were trained 
as museum-based systematic mammalo- 
gists. Writing about different kinds of mam- 
malian anatomy may strike the reader as an 
exercise in semantic hair-splitting, but these 
categories of anatomy actually represent dif- 
ferent scientific endeavors. I reached my 
conclusions about this subject in part from 
personal experience. For example, I have 
had opportunities to collaborate with anat- 
omists such as Bernard Tandler and Carlin 
Pinkstaff who were based in medical or den- 
tal schools or trained primarily in zoology, 
and anatomists such as G. Lawrence For- 
man whose background, like mine, was fun- 
damentally shaped by museum-based train- 
ing in systematic mammalogy. I also have 
presented papers on mammalian anatomy 
at meetings of both the American Society 
of Mammalogists and the American Asso- 
ciation of Anatomists. Finally, over the years 
(especially early in my career), I had nu- 
merous opportunities to visit with mam- 
malogists such as E. Raymond Hall (The 
University of Kansas), Rollin H. Baker 
(Michigan State University), William H. 
Burt (University of Michigan), and David 
H. Johnson (United States National Mu- 
seum), and zoologists such as Karl Stiles and 
H.R. Hunt (Michigan State University), and 
Tracy Storer (University of California, Da- 
vis), whose careers (excepting Baker) ex- 
tended back nearly to the starting point of 
the present essay. My assertion, however, is 
not uniquely derived just from personal ex- 
perience. Indeed, a number of historians of 
science who gathered in preparation for the 
Centennial Celebration of the American So- 
ciety of Zoologists (held in 1989) more-or- 
less came to the same conclusion (Rainger 
et al., 1988). So, one objective of my essay 
will be to explore the history and intellectual 
or conceptual frameworks of what I term 
medical school anatomy, zoological mor- 
phology, and mammalogical anatomy. 

My second observation is that there is 
some contradictory evidence about the role 
of anatomical research as it is conducted 


within the field of mammalogy. At one ex- 
treme, there is the obvious fact that nearly 
every taxonomic, or systematic, article on 
mammals has anatomical illustrations and, 
sometimes, novel anatomical descriptions. 
At the other extreme, anatomical research 
papers are cited only rarely in most general 
books or faunal accounts. Typical faunal ac- 
counts do not draw upon anatomical infor- 
mation and anatomical data are rarely in- 
cluded in geographic overviews. Standard 
reference works in mammalogy—say John 
Eisenberg’s overview of the mammalian ra- 
diations (Eisenberg, 1981) or Terry 
Vaughan’s classic mammalogy text 
(Vaughan, 1978) have attractive anatomical 
line drawings, but virtually none of the 
modern anatomical literature from non- 
mammalogical sources is cited in either book 
and little of the data from the various mod- 
ern kinds of anatomical research, which 
range from cinematographic analyses of 
feeding or climbing to patterns of innerva- 
tion, brain structure, and cell ultrastructure, 
have been integrated into the texts. Draw- 
ings of skulls and lower jaws, joint articu- 
lations, and dental cusp patterns are appli- 
cations of anatomical illustration, but are 
not representative of ““anatomy” in a mod- 
ern scientific sense. 

In contrast to the limited integration of 
anatomy into some seminal texts and faunal 
accounts, many high quality anatomical pa- 
pers have been published in the Journal of 
Mammalogy. Authorship of these contri- 
butions has been international in scope and 
many of these papers, particularly those 
dealing with locomotion, dentition, glands, 
tongues and jaw systems and digestive sys- 
tems, collectively, comprise an important, 
fundamental contribution to knowledge 
about mammals. In fact, the Journal of 
Mammalogy may be the best single source 
of information about integumentary glands 
in mammals (e.g., Atkeson and Marchin- 
ton, 1982: Dapson et al., 1977; Eadie, 1938; 
Estes et al., 1982; Jones and Plakke, 1981; 
Quay, 1965, 1968). Anatomical articles 
published in the Journal of Mammalogy are 
distinctive in two ways: most are compar- 


ANATOMY 207 


ative (i.e., more than one species is consid- 
ered); and most are integrative (i.e., the an- 
atomical data are in some way integrated 
with ecology or physiology or some other 
aspect of mammalian biology). 

The ASM also has strongly supported the 
publication of anatomical contributions in 
its special publication series, sometimes as 
morphological monographs (Altenbach, 
1979) and sometimes within books focused 
on the biology of particular taxa (e.g., Gen- 
oways and Brown, 1993; Tamarin, 1985). 
Museum-based publications constitute an- 
other significant portion of the North Amer- 
ican literature in mammalogy. Historically, 
such publications frequently have con- 
tained specialized types of anatomical in- 
formation about mammals: for example, in 
such publications one might find basic de- 
scriptive histology (Miller, 1895); discus- 
sion of anatomical adaptations in marine 
mammals (Howell, 1929); explanations of 
flight anatomy in bats (Vaughan, 1959); in- 
terspecific comparisons of the structure of 
the baculum (Burt, 1960); dental morphol- 
ogy and development (Phillips, 1971); the 
tragus in the ears of bats (Smith, 1972); bat 
skulls, dentitions, and skeletal features in 
context of ecology and evolution (Freeman, 
1981; Freeman and Lemen, 1991); histo- 
morphological comparisons of female re- 
productive tracts in bats (Hood and Smith, 
1983); comparative histology, histochem- 
istry, and ultrastructure of gastric mucosae 
in correlation with dietary habits (Forman, 
1972; Phillips et al., 1984); ultrastructure of 
secretory cell products (Phillips et al., 
1987a); comparative anatomy of hyoid 
musculature (Griffiths and Smith, 1991); 
comparative morphology of the cochlea in 
microchiropteran bats (Novacek, 1991); and 
comparative morphology of the glans penis 
in three genera of bats (Ryan, 1991). 


Paradigms and Conceptual 
Frameworks 


Between 1890 and 1915, American aca- 
demic biologists diverged into several dis- 


tinctive professional subsets characterized, 
in part, by differences of opinion about what 
constituted “‘science.”” At one extreme the 
process of science was strictly descriptive, 
whereas at the other extreme it was strictly 
experimental (Benson, 1988; Rainger et al., 
1988). By 1919, when the newly formed 
ASM first published the fledgling Journal of 
Mammalogy, this divergence was reflected 
among “‘anatomists” and ““morphologists,”’ 
both of whom investigated mammalian 
anatomy. The anatomists and morpholo- 
gists of 1919 represented two very different 
academic camps. Anatomists largely fa- 
vored descriptive work and typically were 
employed by medical schools where their 
academic function was to train young phy- 
sicians (Appel, 1988). By way of contrast, 
the morphologists, whose studies were be- 
coming more and more experimental, were 
employed by college and university aca- 
demic departments of zoology or biology 
(Rainger et al., 1988). 

Between the two academic camps, it was 
the anatomists rather than the morpholo- 
gists per se who had the most influence on 
the early ASM and on the field of mam- 
malogy before it was codified into an aca- 
demic discipline. The reasons for this are 
twofold. First, some ‘“‘founding fathers” of 
mammalogy were anatomists either by ac- 
ademic experience or by virtue of the med- 
ical profession. Harrison Allen and Gerrit 
S. Miller, Jr., are but two examples of early 
American mammalogists who could be 
identified as anatomists and both published 
excellent anatomical and microanatomi- 
cal papers (e.g., Allen, 1880, 1885; Miller, 
1895). Second, as an academic endeavor, 
anatomy was a largely descriptive mammal- 
oriented activity. Indeed, anatomists in 1919 
did not necessarily focus on human beings 
or medicine as we know it today, and “‘lab- 
oratory” species of mammals still needed 
to be investigated in fundamental ways. Ac- 
cordingly, descriptions of mammalian an- 
atomical characteristics as provided by 
medical school faculty members were 
prominent features of the Journal of Mam- 
malogy between 1925 and 1950. 


238 PHILLIPS 


In 1919, morphologists worked in the 
context of zoology and their research inter- 
ests and teaching differed considerably from 
those of the medical school anatomists 
(Benson, 1988). Insofar as research is con- 
cerned, the morphologists investigated both 
invertebrates and vertebrates. Their work 
was somewhat comparative, but species of 
animals usually were valued for their utility 
as models for testing hypotheses rather than 
because of their intrinsic value or because 
of a curiosity about the species themselves. 
In the academic arena, the morphologist’s 
pedagogic goals focused on graduate stu- 
dents and the challenge of research training 
rather than on teaching medical students the 
art of practicing medicine (Benson, 1988). 
Today, 75 years after the birth of the ASM, 
the fields of medical anatomy and zoolog- 
ical morphology still differ dramatically 
from each other and both are surprisingly 
different from mammalogical anatomy. 
Mammalogical anatomy developed as a 
unique form of scholarship within the broad 
context of “anatomy.” This uniqueness is 
partly due to the fact that mammalogical 
anatomy was created within museum-based 
American mammalogy rather than within 
either medical school anatomy or zoology 
department morphology. 

If the assertion that the subject matter of 
mammalian anatomy is shared by three sep- 
arate academic groups seems remarkable, 
maybe even preposterous, one could sub- 
stantiate it fairly easily by comparing con- 
tents and citation sources in the American 
Journal of Anatomy, Anatomical Record, 
Journal of Morphology, and American Zo- 
ologist with the contents and citation sources 
in the Journal of Mammalogy. There is re- 
markably little overlap among articles and 
sources of information between or among 
these journals. It is true, of course, that all 
of these journals publish articles about 
mammals. What impresses me, however, 1s 
the extent to which the articles reflect dif- 
ferent scientific perspectives. These differ- 
ences might suggest a lack of interchange or 
cognizance of one discipline for another, but 


perhaps they simply reflect the fact that the 
practitioners do not share a common schol- 
arly heritage. My thesis is that modern 
mammalogical anatomy, medical school 
anatomy, and zoological morphology, as ac- 
ademic endeavors, differ in ways that are 
important to appreciate because these dif- 
ferences have served to influence, perhaps 
even channel, research over the past 75 
years. 

To the non-scientist, “science” often is 
regarded as a single enterprise conducted 
under a common set of rules referred to as 
the Scientific Method. Scientists generally 
understand, however, that their own work 
can differ in many ways from the scholar- 
ship of another scientific discipline. When 
confronted with the task of describing or 
explaining differences between their own and 
other scientific disciplines, many scientists 
find it difficult to articulate their perception 
of the difference. Indeed, sometimes there 
is little more than a vague sense that, “we 
do things differently.”” Even so, the sum of 
the differences just within the biological sci- 
ences is real enough to cause conflict, intra- 
departmental battles over college and uni- 
versity science curricula, and severe 
competition for funding. A philosopher of 
science, Thomas Kuhn, recognized the sig- 
nificance of these subtle non-uniformities 
within scientific disciplines. To write about 
this phenomenon, he (Kuhn, 1962) used the 
term “paradigm” to describe subunits of 
scholarship within broad fields of science; 
specifically he defined a paradigm as a co- 
herent research tradition, including the rules 
and standards under which research is con- 
ducted. The components of a paradigm are 
varied, but could be expected to include an 
ethical perspective, cultural and academic 
origins, historical context, oral and written 
traditions, and the flavor of the personalities 
of the founders. Scholarly paradigms would 
be expected to incorporate a set of theories 
or assumptions and, although it may be dif- 
ficult to define, one might expect a biolog- 
ical paradigm to espouse a particular con- 
cept of the nature of the world (Kuhn, 1962). 


ANATOMY 2o9 


Finally, paradigms are defined implicitly 
rather than explicitly, so boundaries and 
membership often make more sense in ret- 
rospect than at a particular moment in time. 
The idea of scholarly paradigms can be ap- 
plied readily to the subdivisions described 
within the broad subject of anatomical re- 
search on mammals. In the United States, 
medical school anatomy, zoological mor- 
phology, and mammalogical anatomy are 
different paradigms. 

In the present essay, I explore some of the 
many components of a paradigm. However, 
one of the most important is what I term 
‘“conceptual framework.’’ A conceptual 
framework grows from the favorite theories 
and assumptions that underlie a scholarly 
paradigm. However, a conceptual frame- 
work to a large extent is the summation of 
how a paradigm deals with particular the- 
ories and assumptions and, therefore, the 
conceptual framework of one paradigm 
might differ from that of another, even 
though they are based on a single, common 
theory. 

Conceptual frameworks are important 
because they root a scientist’s research, link 
the results into some broader context, and 
influence the pathways of future research. 
Differences among conceptual frameworks 
unquestionably produce the most signifi- 
cant intellectual distinctions that can be 
made between scientists and, ultimately, 
paradigms. To many readers, Darwinian 
evolution is the single most obvious theory 
in all of biology, so it is constantly surprising 
to discover that evolution is not at the core 
of all scholarly paradigms in biology. In- 
deed, historically, Darwinian evolution was 
the central theoretical feature of zoological 
morphology, but was not central to either 
medical school anatomy or mammalogical 
anatomy. 

The conceptual framework underlying the 
research of medical school anatomists could 
be described as a linear pathway within 
which the scientific method is applied to 
“questions” that unfold one after the other, 
often on the basis of technological advance- 


ment. By way of contrast, traditional tax- 
onomy (ultimately systematics and cladis- 
tics) has provided the paradigm of 
mammalogical anatomy with a distinctive, 
non-linear type of conceptual framework. 
Taxonomic arrangements were the bases 
upon which interspecific anatomical com- 
parisons were made at the time when mu- 
seum-based mammalogical anatomy de- 
veloped. 

My assertion that evolutionary theory 
does not serve as the conceptual framework 
in medical school anatomy may not surprise 
many readers, but the same might not be 
true of my assertions about mammalogical 
anatomy and zoological morphology. Giv- 
en the number of mammalogists presently 
interested in evolutionary biology, why were 
classification processes rather than evolu- 
tionary theory the original bases for the con- 
ceptual framework of mammalogical anat- 
omy? The answer lies at least partly within 
the basic divergence between museum-based 
and laboratory-based natural science of the 
19th Century (Benson, 1988; Kohlstedt, 
1988). The museum-based branch, which 
strongly influenced many founders of mod- 
ern mammalogy and mammalogical anat- 
omy, was dominated by Louis Agassiz. This 
was Significant because, as Kohlstedt (1988) 
has pointed out, Agassiz’s 1848 textbook, 
Principles of Zoology, was used by his dis- 
ciples in the museum community until late 
in the century. Agassiz wrote that the di- 
versity of animal life was an “exhibition of 
the divine thought” and that human beings 
*“*... being made in the spiritual image of 
God ... [are] competent to rise to the con- 
ception of His plan and purpose in the works 
of the Creation.” Agassiz felt that the tax- 
onomic activities that characterized muse- 
um-based research should include study of 
the “plan and purpose of God in His cre- 
ation” (Kohlstedt, 1988). I do not mean to 
argue that early museum-based mammal- 
ogy was “creationist” in the modern sense. 
I simply wish to explain how as a museum- 
based science, mammalogical anatomy in- 
herited an intellectual perspective and con- 


240 PHILLIPS: 


ceptual framework strikingly different from 
that of the morphologists employed in zo- 
ology departments at the turn of the century. 
The morphologists were not disciples of Ag- 
assiz: in the late 19th Century they actively 
avoided classification and focused instead 
on testing evolutionary theory, primarily 
through experimental research in embry- 
ology (Benson, 1988). 


The Early History 


In order to appreciate fully mammalogi- 
cal anatomy and its academic cousins, one 
must survey the foundations of anatomy, 
mammalogy, and morphology before 1919. 
It is difficult to imagine college and univer- 
sity life and academic structure in the 1880s, 
but it bore little resemblance to the present 
time. The subdisciplines within biology were 
as yet undefined. Aside from the need to 
educate another generation of teachers, the 
raison détre of the professorate was unclear 
except at a few august institutions with 
philosophical scholars in the European tra- 
dition. Research and science as we know 
it—or as we use these terms—were very dif- 
ferent from today and do not appear to have 
been the major foci of academic activity 
that they eventually became. The profes- 
sorate was not yet a national cadre of re- 
searchers (Rainger et al., 1988). Instead of 
research, curriculum and curricular issues 
had priority. 

In seeking individuals who affected the 
paradigms of mammalogical anatomy, 
medical school anatomy, and zoological 
morphology, we would find that many of 
the founders of medical school anatomy 
spent most of their time teaching young 
physicians (Appel, 1988). The founders of 
zoological morphology would be found 
among the self-described “‘naturalists’’ who 
taught various life science subjects in col- 
leges and universities from the 1860s 
through the 1880s (Benson, 1988). By way 
of contrast, most of the academic grandfa- 
thers of North American mammalogy were 


among the “natural historians” of the 1870s 
through the 1880s. These forebearers of our 
discipline in North America concentrated 
on taxonomic studies of museum collec- 
tions and used museum collections as the 
central pedagogic tool for training future 
teachers of natural science (Kohlstedt, 1988). 
When not at the museum, they were some- 
where afield, gun and traps in hand (Ster- 
ling, 1991). The cultural component of 
mammalogical anatomy traces to the fact 
that many prominent early mammalogists 
spent as much time exploring and collecting 
as with college students, and more time 
skinning, cataloging, and identifying speci- 
mens than studying the revolutionary con- 
cepts of embryology, cytology, and evolu- 
tion that occupied the thoughts of the 
zoological morphologists on the campuses 
of the Northeast and Midwest. 

The medical school anatomists.—Gen- 
erally speaking, the American anatomists 
did not spend time afield; they either were 
too busy with the medical arts or, more like- 
ly, did not regard the elemental activities of 
natural history as worthy of their time. They 
formally organized into an academic asso- 
ciation in 1888 at the Congress of American 
Physicians and Surgeons held in Philadel- 
phia. Throughout the 19th Century there 
was a certain elitism about natural science 
as conducted in the vicinity of Philadelphia 
so that the anatomists’ selection of a city in 
which to organize themselves is telling. In- 
deed, the year before, in 1887, George B. 
Goode had addressed the Biological Society 
of Washington and paraphrased from Pick- 
ard’s text on the History of Zoology. Pickard 
had asserted, “.. . zoology the world over, 
has sprung from the study of human anat- 
omy, and. .. American zoology took its rise 
and was fostered chiefly in Philadelphia by 
the professors in the medical schools.” 
Goode did not buy Pickard’s idea, and went 


on to remark, “... there were good zoolo- 
gists in America long before there were 
medical schools, and ... Philadelphia was 


not the cradle of American natural history” 
(Kohlstedt, 1991). This quotation 1s telling 


ANATOMY 241 


because it not only illustrates the signifi- 
cance of Philadelphia in American science, 
but also because it underscores the sense of 
competition that forced the divergence be- 
tween medical school anatomists and zoo- 
logical morphologists. 

The organizers of the first society of anat- 
omists described themselves as the Asso- 
ciation of American Anatomists, but 
changed their name to the American As- 
sociation of Anatomists in 1909 (Appel, 
1988). It is worth recalling that one of the 
prominent leaders of this founding group 
was Harrison Allen, who at the time was 
writing papers on the anatomy of bats and 
rodents and qualifies as an early figure in 
mammalogy. But, aside from Allen, who 
were these people and why did they formally 
organize themselves? Essentially, the anat- 
omists may have been motivated by a desire 
to separate themselves from “‘physicians”’ 
in the sense of distinguishing between a 
scholarly pursuit—studying anatomy—and 
non-scholarly professional practice (see Ap- 
pel, 1988). Strictly speaking, many of these 
anatomists were not just medical practi- 
tioners. Instead, many were faculty mem- 
bers at such medical schools as Harvard 
College, Yale, or Johns Hopkins. In this ca- 
pacity they were charged with responsibility 
for educating young physicians. In retro- 
spect they also seem to have been struggling 
to define their roles in ways that matched 
the prevailing ideas of scholarship. So, in 
modern terms they may have been seeking 
a vehicle by which their avocation—de- 
scriptive gross anatomy—could be incor- 
porated into their job description as pro- 
fessors of anatomy. 

The zoological morphologists. —Regard- 
less of how one interprets the origins of the 
American Association of Anatomists, the 
most salient fact is extremely clear: the 
College of Physicians and Surgeons and its 
offspring association diverged from the 
academic milieu of the zoological mor- 
phologists, or naturalists. The relationship 
between the morphologists and medical 
school anatomists was not neutral; the mor- 


phologists openly regarded anatomy as a 
““dead”’ discipline. In the view of the mor- 
phologists, the basic structure of mammals, 
birds, and other vertebrates already had been 
described, so the principal task of anatomy 
had been completed (Appel, 1988; Benson, 
1988). Why then would one wish to contin- 
ue to pursue the subject? Indeed, the zoo- 
logical morphologists and their colleagues 
were caught up in the sweeping philosoph- 
ical issue of evolution and the practice of 
experimental “science.” Additional gross 
anatomical descriptions probably seemed 
irrelevant. In terms of heritage, the Amer- 
ican anatomists thus were excluded from 
the scholarship and ambiance of the life sci- 
ences as practiced in college and university 
zoology departments (Appel, 1988). 

Ironically, however, it was microanato- 
my, embryology, histology, and cytology— 
all topics that eventually were studied by 
medical school anatomists—that attracted 
the attention of naturalists and served as 
early exemplars of 19th Century zoological 
morphology. The zoological morphologists 
pursued research projects that combined 
theoretical interests in evolution with new 
technology. In particular, they used the ever- 
improving technical skills of Germans who 
manufactured high quality lenses and pro- 
vided rotary microtomes that could be used 
to slice tissues into thin, translucent sections 
suitable for the optical microscope (Benson, 
1988). The cell theory of Schleiden and 
Schwann and the hypothesized relationship 
between ontogeny and evolution provided 
the zoological morphologists with concep- 
tual frameworks for their research. This then 
was the academic heritage of the zoological 
morphologist: like their medical school 
counterparts they were laboratory-based, but 
unlike their medical school counterparts, 
their studies were rooted in evolutionary 
theory. 

The zoological morphologists dominated 
development of college and university ac- 
ademic departments from the 1880s on- 
ward (Benson, 1988; Maienschein, 1988). 
Meanwhile, the traditional gross anatomists 


242 Jed ie i Ed OF eas 


dominated the medical schools until Wat- 
son and Crick elucidated the structure of 
the DNA molecule, a feat that gave birth to 
cell and molecular biology and revolution- 
ized the life sciences. 

The mammalogical anatomists. — Strictly 
speaking, mammalogical anatomists—or the 
foundations of the science of mammalogy — 
were not included in the academic milieu 
of either the zoological morphologists or the 
anatomists. The academic origins of North 
American mammalogy and, ultimately, the 
paradigm of mammalogical anatomy, can 
be found within the culture ofa third group— 
the museum-based natural historians. Su- 
perficially, it might seem that the natural 
historians were the forerunners of the nat- 
uralists and the naturalists were forerunners 
of all modern biologists. However, as Ben- 
son (1988) has explained, there was a strik- 
ing divergence between natural historians 
and naturalists just as there was between the 
naturalists and anatomists. The natural his- 
torians collected and stored specimens of 
animals, which they used as the basis for 
instruction, exhibition, and personal taxo- 
nomic study. Because specimens were stored 
in museums, the museum environment was 
home to the natural historians who were 
founders of North American mammalogy. 
The naturalists deliberately diverged from 
the museum-based natural historians: in- 
stead of specimen-based instruction for fu- 
ture school teachers, public exhibitions, and 
taxonomy, the naturalists focused on re- 
search and research training and graduate 
education (Kohlstedt, 1988). The zoological 
morphologists who were derived from 
among these original naturalists thus held 
little in common with museum-based nat- 
ural historians (Benson, 1988; Kohlstedt, 
1988). Indeed, in the biology department at 
Johns Hopkins of the 1880s, the courses 
included histology, mammalian anatomy, 
comparative osteology, and embryology, but 
no taxonomy or classification (Benson, 
1988). 

In the first paragraph of this section of my 
essay, and elsewhere, I have used the term 


“culture” in reference to the origins of 
mammalogical anatomy. I selected this word 
because mammalogical anatomy was heavi- 
ly influenced by factors other than tradi- 
tional academic scholarship. In particular, 
hunting and trapping, exploration, collect- 
ing, and general “outdoorsmanship” un- 
derlie the origins of museum-based mam- 
malogy and are some of the most 
fundamental reasons why anatomy in the 
context of mammalogy is totally different 
from the types of anatomy practiced by typ- 
ical zoology department morphologists or 
by medical school faculty members. The 
field activites that characterized early mam- 
malogy meant that future mammalogists 
would select wild mammals as research ob- 
jects instead of laboratory species. The cul- 
ture of exploration meant that mammalo- 
gists would be just as likely (maybe more 
likely) to investigate exotic species in the 
most remote places as they would be to in- 
vestigate species that could be obtained near 
campus. Thus, in many ways the culture 
from which mammalogy grew Is responsible 
for the broad perspectives of mammalogists 
and mammalogical anatomists. However, 
the primitive style in which field work was 
conducted also placed technical and intel- 
lectual limitations on what could or could 
not be investigated. By growing from, and 
then embracing, the culture of natural his- 
tory, mammalogy both gained and lost. The 
discipline clearly was committed to a path- 
way that would diverge from whatever form 
of mammalogical science might be con- 
ducted by medical school anatomists and 
morphologists in zoology departments. 
The cultural roots of North American 
natural history—and ultimately mammal- 
ogy—trace to Thomas Jefferson (Wilson and 
Eisenberg, 1990). In 1804, while serving as 
President of the United States, Jefferson sent 
Meriwether Lewis and William Clark on a 
lengthy collecting survey west of the Mis- 
sissipp1 River. The motives behind the ex- 
pedition have been debated by historians 
and some think that Jefferson’s scientific in- 
terests merely covered his real intentions, 


ANATOMY 243 


which were territorial and political (Brodie, 
1974). Considering Jefferson’s personality, 
however, it seems obvious that his interest 
in natural science was a significant part of 
the story. Indeed, Jefferson was very inter- 
ested in the possibility that unusual animals 
inhabited the continent. More importantly, 
perhaps, Jefferson made clear his intent that 
faunal survey was a goal of the expedition. 
Thus, the Lewis and Clark expedition was 
not merely an effort to carry forth the flag; 
it represented the beginning of the concept 
of government-sponsored science and served 
as the prototype for later major expeditions 
including J. W. Powell’s extensive surveys 
(Kohlstedt, 1991; Powell, 1925). Under Jef- 
ferson’s guidance, specimen collecting was 
planned as a major component of the Lewis 
and Clark expedition and consideration was 
given to the return and deposition of spec- 
imens. 

Many of the progenitors of North Amer- 
ican mammalogy preferred the gun and trap 
and the bedroll and camp fire to virtually 
anything else. An outdoor perspective and 
an emphasis on purposeful collecting were 
their twin legacies to mammalogy. Men such 
as E. W. Nelson and E. A. Goldman are part 
of the breeding stock of mammalogy. To be 
frank, they were neither scholars nor sci- 
entists. Given their limited educational ex- 
perience and their interests (see Sterling, 
1991), it is unlikely that they were aware of 
the emerging cell theory and the controversy 
about ontogeny and evolution that were be- 
ing hotly debated in the hallways of aca- 
demé. 

In the late 19th Century and early 1900s, 
C. Hart Merriam sent Nelson and Goldman 
deep into Mexico, where they perfected the 
art of travel under adverse conditions, the 
strategy for collecting, and the techniques 
of field preservation. They developed ways 
of shipping specimens safely, habitually took 
notes, and became highly efficient at making 
camps (Sterling, 1991). 

To appreciate fully the cultural impor- 
tance of men like Nelson and Goldman, one 
must understand that they did more than 


set a tone for mammalogy. Aspects of their 
lives have been recapitulated to a remark- 
able degree by succeeding generations of 
mammalogists, including some of those who 
principally are mammalogical anatomists. 
Many living North American mammalo- 
gists have explored and collected mammals 
in every corner of the planet and, like Nel- 
son and Goldman, have traveled on foot or 
horseback, lived in the rudest of conditions, 
and have endured the persistent assault of 
extremes in weather and countless insect 
and acarine pests. The pursuit of field work 
in the tradition of Nelson and Goldman has 
produced a certain outlook, a certain breed 
of scientist who, as Michael Mares put it, is 
**’.. accustomed to the hardships ... in- 
cluding disrupted home lives, unsympa- 
thetic administrators, and frequent health 
problems [and therefore do] not suffer fools 
gladly” (Mares, 1991:63). Many modern 
mammalogists could have written the same 
lines as Nelson, who said, “... I often get 
thoroughly disgusted with [field work] and 
yet there is a fascination about the life I am 
leading that keeps me going despite myself” 
(from Sterling, 1991:40). 

For many years, the technology of mam- 
malogical anatomy was purely an extension 
of the art created by the 19th Century nat- 
ural historians. Thus, even in the early 1960s 
the practice of field work and field collection 
differed little from the process as practiced 
in 1895 by Nelson and Goldman. So, in- 
sofar as anatomical studies are concerned, 
the museum laboratory portion of any re- 
search was destined to be archaic in com- 
parison to what could be accomplished in 
the laboratories of a medical school anat- 
omist. Although the adherence of mam- 
malogists to their field and collecting 
traditions clearly limited the types of ana- 
tomical research that could be conducted, 
it also provided a remarkable platform for 
access to new data and the scientific future 
is bright with prospects for mammalogical 
contributions to cell and molecular biology 
and biochemistry. 

By the late 1960s young mammalogists 


244 PHILLIPS 


began to pack hand-powered centrifuges and 
small microscopes in their baggage and by 
1972, some 80 years after Nelson and Gold- 
man explored and collected along the Pa- 
cific coast of Jalisco, Mexico, a new method 
of fixation of tissues for transmission elec- 
tron microscopy was field-tested for the first 
time. In many ways nothing had changed: 
the fixative was formulated over a kerosene 
burner in a rude camp that resembled the 
one used by Nelson and Goldman in ap- 
pearance, atmosphere, and geographic lo- 
cation. The fixative failed its first test, but 
was perfected and field-tested in Suriname 
by 1981 (Phillips, 1985a). Subsequently it 
has made possible the exploration of com- 
parative cell ultrastructure and cytochem- 
istry with virtually any species of mammal 
collected anywhere (e.g., Nagato et al., 1984; 
Phillips, 19855; Tandler et al., 1986; Tand- 
ler and Phillips, 1993a). A process once re- 
served for the laboratories of medical school 
anatomists now can be applied in the con- 
ceptual framework of mammalogical anat- 
omy. Thus, mammalogical anatomists have 
slowly acquired the technologies necessary 
for modernizing mammalogical field work 
(e.g., Forman and Phillips, 1988). 

The schism between natural historians and 
naturalists. —Away from the field, the early 
natural historians mostly were associated 
with museums rather than college academic 
departments, and this distinction was more 
than administrative. Indeed, a deep philo- 
sophical schism separated the “‘laboratory- 
based”’ naturalists from the ‘“‘museum- 
based”’ natural historians in the late 19th 
Century (Benson, 1988). While the natu- 
ralists debated ontogeny and evolution, the 
natural historians were captivated by di- 
versity and focused their energy on collect- 
ing, cataloging, housing, and describing 
specimens. 

As we look backward in time, it is ap- 
parent that these two camps were deliberate 
in their divergence. The laboratory-based 
naturalists controlled the college curricula 
and regarded the natural historians as non- 
scientific amateurs (Benson, 1988). The nat- 


ural historians intellectually barricaded 
themselves in their museums and expressed 
concern that students were not receiving ad- 
equate training in taxonomy. To appreciate 
fully the significance of this schism, one need 
only examine an example of the academic 
pathways that ultimately led to the disci- 
pline of Zoology at the University of Chi- 
cago, and the development of the Museum 
of Vertebrate Zoology at the University of 
California, Berkeley. 

When the prominent natural historian 
David Starr Jordan organized a new college 
at Palo Alto, California (now Stanford Uni- 
versity), he received very specific advice 
from George W. Peckham. In 1881, in a 
letter to Jordan, Peckham wrote: 


**...1t seems to me that Morphology and 
Embryology have usurped too much of 
the attention of the workers in the un1- 
versities of America. I really believe that 
there has been more bad cell-making than 
bad species-making. The new Clark Uni- 
versity under my friend Dr. Whitman 
[Charles Otis Whitman] will turn out nu- 
merous young morphologists, but not a 
man with any sympathy for general Nat- 
ural History work” (Benson, 1988). 


Jordan seems to have taken to heart the 
idea that natural history and cell-making 
could never co-exist, and this was reflect- 
ed—reinforced—in his academic descen- 
dants who established their own institutions 
devoted to natural history and taxonomy. 
In particular, Jordan had a strong influence 
on Joseph Grinnell, who was to become the 
academic grandfather of mammalogy (Jones, 
1991). Grinnell was perfect for the task. Al- 
though small in stature and reportedly shy 
by nature, he had a scholar’s demeanor and 
was a demanding task master. E. Raymond 
Hall, one of Grinnell’s many successful stu- 
dents, frequently told me of Grinnell’s dog- 
matic, pedantic nature, which Hall himself 
had inherited. In keeping with the culture 
of mammalogy, Grinnell was born, in 1877, 
some 40 miles from Fort Sill in the “Indian 
Territory” now called Oklahoma. As a 


ANATOMY 245 


youngster, Grinnell went to the wilderness 
of Alaska where he developed a reputation 
as a collector, which ultimately served as 
one of his major credentials in securing his 
positions with Jordan and the Museum of 
Vertebrate Zoology at Berkeley (Dunlap, 
1988: Jones, 1991). 

While Jordan was following Peckham’s 
advice, Charles Whitman retained his focus 
on evolutionary theory, morphology, and 
cytology. After leaving Clark University, 
Whitman essentially fathered zoology at the 
University of Chicago and his department 
became home to such luminaries as W. C. 
Allee, Sewall Wright, and zoological mor- 
phologists such as Libby H. Hyman (Maien- 
schein, 1988). 

The impact of Joseph Grinnell on the field 
of mammalogy and the paradigm of mam- 
malogical anatomy hardly can be exagger- 
ated; he and his academic descendants have 
published more than 5,000 scientific papers 
and books (Jones, 1991). Because he played 
so important a role in establishing and cod- 
ifying our academic discipline in North 
America, it is noteworthy that Grinnell was 
remarkably narrow in academic ideology. 
Indeed, while college departments were di- 
versifying and the naturalists of old were 
reorganizing into zoological morphologists, 
cytologists, embryologists, and geneticists, 
it almost seems as though Grinnell and oth- 
er natural historians retrenched even further 
by actively restricting themselves and their 
students to more and more narrowly de- 
fined forms of taxonomy and zoogeography. 
Grinnell prohibited his students in the Mu- 
seum of Vertebrate Zoology from taking 
courses or pursuing projects outside of the 
narrow confines of the museum environ- 
ment. This suited some of his students just 
fine, but those possessed of broader interests 
in biology may have found the restraints an 
impediment to their personal intellectual 
development. For example, when Tracy 
Storer—a zoologist by any measure — wished 
to investigate ecological principles, he was 
pressured by Grinnell to refocus on muse- 


um-based taxonomic research with dead 
rather than living animals (Dunlap, 1988). 
The gap between the descendants of the 
original naturalists and the natural histori- 
ans who gathered together in the natural 
history museums was further enforced by 
the fact that museums often were physically 
separated from academic departments and 
went so far as to create their own scientific 
publications to provide unspoiled outlets for 
the products of their research. 

One of the interesting and historically rel- 
evant side-lights to the isolationism of the 
museum-based mammalogists occurred un- 
der E. Raymond Hall’s directorship of the 
Museum of Natural History at The Uni- 
versity of Kansas. Hall (E. R. Hall, pers. 
comm.) recognized the development of 
mammalogical anatomy as a distinctive 
paradigm and was convinced that medical 
school anatomy could profit from the intro- 
duction of mammalogical anatomists. Con- 
sequently, he fairly frequently suggested that 
students develop skills that would qualify 
them for employment on medical school 
faculties and one of his students, Phillip H. 
Krutzsch, became the first Chairman of 
Anatomy at the University of Arizona 
School of Medicine. 

Ethics, codification, and splintering of 
mammalogical anatomy.—We have ex- 
amined a variety of elements that contrib- 
uted to the paradigm of mammalogical 
anatomy: the divergence of natural histo- 
rians from the academic naturalists; the im- 
pact of outdoorsmanship and the tradition 
of exploration; the museum-based collec- 
tion as a research and pedagogic resource; 
and the pervasive impact of taxonomy-sys- 
tematics-cladistics as a conceptual frame- 
work. Before addressing the influences of 
taxonomy and natural history in more de- 
tail, we should briefly consider how ethical 
perspectives and codification processes oc- 
cur in scholarly paradigms. In the case of 
mammalogical anatomy, codification of a 
coherent research process occurred over a 
lengthy period of time, but principally was 


246 PHTELIPS 


in place by the time that the first generation 
of Grinnell’s students departed from Berke- 
ley to establish their own programs and mu- 
seums. 

There are many differences among the 
paradigms of mammalogical anatomy, 
medical anatomy, and zoological morphol- 
ogy, and one might presume that these dif- 
ferences tend to limit the direct competition 
that otherwise could occur. One of the more 
interesting examples is the extent to which 
the paradigms have subdivided the research 
subjects. Mammalogical anatomists are in- 
terested in most species of mammals, but 
tend not to be interested in the anatomy of 
either laboratory species or human beings. 
This characteristic of mammalogical anat- 
omy has been codified, at least in part, 
through the Journal of Mammalogy. A re- 
view of articles published since 1919 re- 
veals that articles on anatomy of laboratory 
mammals had become scarce by 1950 and 
that over the last 40 years they are essen- 
tially nonexistent. In effect, editorial policy 
(perhaps de facto) has restricted, or helped 
codify, the species that are suitable for an- 
atomical studies in mammalogy. This cod- 
ification did not originate within the ASM; 
it is reflected also in the museum-based an- 
atomical publications from the time of Jo- 
seph Grinnell. As a comparison, the codi- 
fication of using laboratory species or human 
tissues in medical school anatomy seem- 
ingly has been driven by a) restrictions on 
funding support for research, and b) the idea 
that a single example (possibly equivalent 
to a single species) is adequately represen- 
tative of most mammals. In the paradigm 
of mammalogical anatomy, the selection of 
a few species appears equivalent to reducing 
mammalian diversity to a world of “‘the 
mouse,” “the rat,” and “the dog.” It is far 
easier to describe paradigms than to allocate 
individuals to particular paradigms, and, in 
fact, some individuals very likely are able 
to shift from one paradigm to another. Thus, 
it should not be surprising that some of the 
prominent members of the medical school 


anatomy paradigm, for example Don Faw- 
cett of Harvard Medical School, Carlin 
Pinkstaff of West Virginia University School 
of Medicine, and Frank Kallen of SUNY- 
Buffalo, are interested in wild species. Other 
scientists are more paradigm-bound, and it 
is their work that actually helps to define a 
paradigm. Moreover, in contrast to the 
Journal of Mammalogy, most anatomical 
journals welcome articles on the anatomy 
of non-laboratory species, so long as authors 
clearly explain why anatomical data from a 
“new” species might not be redundant to 
data from the mouse or rat. 

The formation of paradigms is not pre- 
ordained, does not appear to follow a set of 
rules, and generally is understandable only 
in historical terms. One important excep- 
tion to this generalization may have oc- 
curred during the time that mammalogical 
anatomy was being codified. Not long after 
1919, within mammalogy there was an in- 
ternal clash over research and, especially, 
agreed-upon scientific ethics. [Although, the 
point should be made that “‘ethics”’ was not 
recognized as the issue at that time.] It is 
particularly interesting that this clash in- 
volved several of the original field collec- 
tors, especially E. A. Goldman, on the one 
hand and Grinnellian scholars on the other. 
The battle, which has been discussed and 
analyzed in historical detail by Thomas 
Dunlap (1988), centered on predator con- 
trol policies and involved Goldman in his 
post-Mexico career as a Washington bu- 
reaucrat. In an ethical sense, Goldman and 
his supporters took the position that some 
species of mammals are more valuable than 
others. Value was dictated by human econ- 
omy. The Grinnellian scholars, led primar- 
ily by E. Raymond Hall, essentially took the 
ethical position that all mammals have equal 
intrinsic value. As part of their strategy, Hall 
and his colleagues seized the academic high 
ground and, recalling Goldman’s relatively 
weak academic credentials, they attacked 
Goldman’s understanding of science and 
ability to interpret data (Dunlap, 1988; E. 


ANATOMY 247 


R. Hall, pers. comm.). A successful effort 
was made to use the ASM and Journal of 
Mammalogy in the fight against predator 
control policies of the federal government 
(Dunlap, 1988), which ultimately was a fight 
that helped further define one paradigm and 
nearly created another. 

The predator control conflict is relevant 
to our discussion because it illustrates how 
a paradigm can struggle for self-definition. 
In this case, the museum-based academic 
mammalogists codified their ethical posi- 
tion on the intrinsic value of all mammals 
and set their own standards for research. 
Moreover, the resulting split almost created 
a new paradigm for anatomical research on 
mammals—a paradigm based on wildlife 
biology or management. So it is that one 
can find a specialized type of anatomical 
information on selected species of mam- 
mals in The Journal of Wildlife Manage- 
ment. Studies of mammalian anatomy are 
a relatively small component of wildlife bi- 
ology, but are distinctive enough to be con- 
trasted with museum-based mammalogical 
anatomy. There are three major differences: 
absence of a taxonomic or systematic com- 
ponent; emphasis on application of data to 
management issues or biological founda- 
tions of management; and restriction of re- 
search to species judged to be of suitable 
economic (game) value. In terms of appli- 
cation, one finds articles in both The Jour- 
nal of Wildlife Management and the Journal 
of Mammalogy on topics such as the fol- 
lowing: use of anatomical features of teeth 
and skulls in age determination (e.g., Kirk- 
patrick and Sowls, 1962; Marks and Erick- 
son, 1966); use of thymus, other glands, and 
the kidney as indicators of nutritional and 
developmental status (Hoffman and Rob- 
inson, 1966; Ozoga and Verme, 1978; Ran- 
som, 1965); basic anatomy of game species 
(Short et al., 1965); the effects of environ- 
mental conditions and diet on growth and 
fat accretion in game species (Abbott et al., 
1984; Holter and Hayes, 1977; Klein et al., 
1987; Verme, 1979); and descriptive and 


morphometric data on physical morphol- 
ogy of different age classes (e.g., Lochmiller 
et al., 1987). 


The Influence of Taxonomy 


Taxonomy has influenced mammalogical 
anatomy in three ways: 1) as mammalian 
taxonomy became codified into a predict- 
able process, certain types of anatomical data 
were gathered and described; 2) certain types 
of descriptive information became “‘accept- 
able” matters for publication; and 3) any 
comparative anatomy undertaken by prac- 
titioners was conducted in terms of a tax- 
onomic hierarchy. Because taxonomy was 
the mainstay of early 20th-century mam- 
malogy, much of the early anatomical “‘re- 
search” by mammalogists resembled a se- 
ries of practical exercises. Descriptions of 
certain structural elements, most notably the 
skeleton and dentition, were essential to 
taxonomy. Thus, early mammalogists spent 
much of their working time describing skulls, 
jaws, and teeth in careful detail. This pro- 
cedure has not changed; the current version 
of this type of descriptive anatomy is vir- 
tually indistinguishable from that of 100 
years ago. 

It is important to understand that anat- 
omy in the context of taxonomy was never 
intended to solve anatomical puzzles and 
certainly not intended to shed light on the 
sweeping theoretical concepts that attracted 
the zoological morphologists. When new in- 
formation about structure was obtained by 
mammalogical anatomists, it was almost by 
accident and usually was treated as inci- 
dental to the main theme of the research. 
Taxonomy codified the pattern of obser- 
vation so that the anatomical descriptions 
were tailored into a suitable format. In other 
words, the observations used for the written 
anatomical descriptions were predeter- 
mined by what was needed for comparisons 
to related species. An example of how this 
influenced anatomical descriptions by 


248 PHILLIPS 


mammalogists can be seen in a paper by E. 
Raymond Hall. When Hall had an oppor- 
tunity to examine the post-cranial skeleton 
from a rare species of bat, he did so only in 
the context of its comparison to a species 
in a related genus (Hall, 1935). 
Observations born of taxonomy tend to 
deflect other anatomical issues. So, most 
mammalogical anatomy in the context of 
taxonomy is focused on pure comparisons 
and contrasts between species rather than 
functional concepts. The ““channeled”’ prac- 
tical anatomy derived from the taxonomic 
framework of North American mammalogy 
has continued on to this day, passsed down 
primarily through the Grinnellian academic 
lineage. An excellent example of this phe- 
nomenon may be seen in a summary paper 
by Jones and Genoways (1970), who re- 
viewed new (ca. 1970) aspects of bat anat- 
omy that could be used in modern types of 
systematic studies. Application of this ap- 
proach, at the level of the light microscope. 
may be seen in an article in which Hood 
and Smith (1982) used histomorphological 
features in a cladistic analysis. One thus finds 
many examples of modern investigations in 
mammalogical anatomy, even some con- 
ducted with histochemical and ultrastruc- 
tural methods, presented in a context and 
style familiar to taxonomists since 1885, but 
virtually unrecognizable to modern medical 
anatomists or zoological morphologists. One 
of the most striking recent examples can be 
seen in G. Lawrence Forman’s Ph.D. dis- 
sertation at the University of Kansas. The 
histological and histochemical comparisons 
of gastric mucosa in species of microchi- 
ropteran bats duplicated the telegraphic style 
of taxonomic papers (Forman, 1972). 
Moreover, his research “laboratory” was 
housed in the museum penthouse so his 
slides were warmed by being placed on an 
empty tin of pipe tobacco that had been 
lined with aluminum foil. The slides then 
were warmed by a goose-neck lamp rather 
than by means of the slide warmers across 
campus in the departmental histology fa- 


cility. Slides prepared by this means were 
then cleared, dehydrated, and stained in 
chemical solutions kept in empty baby-food 
jars that were stored on a nearby shelf. 

A recognition of the importance of quan- 
titation was another major impact of tax- 
onomy on mammalogical anatomy. The 
correct way of taking and recording mea- 
surements of skeletal materials and teeth 
concerned everyone in taxonomy. One of 
the first papers published in the new Journal 
of Mammalogy was John C. Phillips’ de- 
scription of how to measure deer skulls 
(Phillips, 1919). The introduction of “‘dial”’ 
calipers was seen as a means of improving 
repeatability. This initiated a trend in which 
more and more measurable anatomical 
characters were sought. Both B. Elizabeth 
Horner (Horner, 1944) and Sydney Ander- 
son (Anderson, 1968) described new types 
of craniometers that facilitated the process 
of taking skull and dental measurements. 
The use of morphometry and statistics to 
investigate geographic, populational, onto- 
genetic, and interspecific variation in skel- 
etal elements evolved from this technology 
and from the availability of skeletal mate- 
rials in museum collections. The measure- 
ment of cranial and dental characters used 
in taxonomy eventually resulted in efforts 
to separate variation due to inheritance from 
variation due to other factors (Bader, 1965; 
Strandskov, 1942): and to use allometric 
techniques for comparing skeletal anatomy 
(e.g., Goldstein, 1972; Nelson and Shump, 
1978). 

The value of dentition to taxonomy (and 
to paleontological mammalogy) 1s obvious 
and it is not surprising, therefore, to find 
that large numbers of papers on dental anat- 
omy have been published by mammalogical 
anatomists. In addition to numerous de- 
scriptions of particular teeth in certain spe- 
cies or groups of mammals, the Journal of 
Mammalogy is an exceptionally rich re- 
source of information about such disparate 
topics as genetics of tooth development (Gill 
and Bolles, 1982), dental homologies (Zieg- 


ANATOMY 249 


ler, 1971), eruption of teeth (Shadle, 1936; 
Slaughter et al., 1974), dental ontogeny (Bir- 
ney and Timm, 1975), dental functions and 
coronal morphology (Chiasson, 1957; Kil- 
tie, 1981), enamel structure (Flynn et al., 
1987: Krause and Carlson, 1987), age de- 
termination and parturition (Klevezal and 
Myrick, 1984; Phillips et al., 1982), dental 
evolution (Gingerich and Rose, 1979; Klin- 
gener, 1963; Phillips and Oxberry, 1972), 
and quantitative variation (Gingerich and 
Winkler, 1979). Moreover, this interest in 
dentition has carried over to an interest in 
mastication (Herring, 1985; Reduker, 1983; 
Riley, 1985; Wilkins and Woods, 1983). 

Museum-based mammalogical anatomy 
has appeared in investigations of tooth 
structure in many ways. One of the more 
unusual twists in the taxonomic trail led to 
the mouths of phyllostomid bats of the ge- 
nus Leptonycteris. Periodontal disease and 
dissolution of mineralized tissue was ob- 
served in L. nivalis, but not in a broadly 
distributed relative, L. sanborni. This “‘tax- 
onomic”’ character was traced to species- 
specific infestations of macronyssid mites 
(Phillips et al., 1969). 

The search for new taxonomic characters 
also has led to one of the most distinctive 
topics in mammalogical anatomy seen 
within the covers of the Journal of Mam- 
malogy and in museum publications. Be- 
ginning in 1940, the Journal of Mammalogy 
started publishing articles describing the os 
penis (baculum) and os clitorides of rodents 
and bats. The first article, on sciurid bacula 
(Wade and Gilbert, 1940), set the stage for 
a series of papers that described, compared, 
and, occasionally, offered functional hy- 
potheses (e.g., Blair, 1942; Burt and Bar- 
kalow, 1942; Layne, 1952; Krutzsch, 1959, 
1962; Patterson and Thaeler, 1982). Inter- 
est in the baculum and reproductive sys- 
tems in general appears to have led to an 
interest in using the soft anatomy of the 
phallus in taxonomic studies (Lidicker, 
1968). One such paper, based upon micro- 
scopic observations of the penis in species 


of bats and primates by a Grinnellian aca- 
demic grandson, James D. Smith (Smith and 
Madkour, 1980), helped to touch off a 
sometimes bitter, sometimes anachronistic, 
international debate about the origin of bats 
(see Goodman, 1991, for an overview). 

The use of microscopic data in mam- 
malian taxonomy offers yet another set of 
examples of the unique nature of mam- 
malogical anatomy. For instance, in keeping 
with the established pattern of mammalog- 
ical anatomy, Smith and Madkour (1980) 
did not publish their histological observa- 
tions on the penis in bats in an anatomical 
journal. Instead, their findings were written 
in taxonomic style and published without 
any photomicrographs in the “proceedings” 
of a meeting hosted by The Museum, Texas 
Tech University. To a traditionally-trained 
microanatomist, photomicrographs are 
taken as “hard” data, so while Smith and 
Madkour’s paper helped create a furor in 
taxonomist circles, it might have been un- 
publishable in more traditional anatomic 
circles. Comparative microanatomy of 
mammalian spermatozoa 1s another area in 
which mammalogical anatomists have in- 
fluenced taxonomy. In turn, many of the 
articles on sperm morphology have been in- 
fluenced more by the traditions of taxono- 
my than by the style of histological and his- 
tochemical data in other journals. 

The taxonomic format gradually is being 
dropped by mammalogical anatomists in 
favor of formats more in keeping with the 
style in traditional anatomical journals. Ex- 
amples of this conversion in mammalogical 
anatomy in the Journal of Mammalogy in- 
clude articles based on scanning electron 
microscopy of hair structure (Brian et al., 
1987; Homan and Genoways, 1978; Short, 
1978) and transmission electron micros- 
copy of the retina in rodents (Feldman and 
Phillips, 1984). Moreover, even the inte- 
gration of mammalogical anatomy, system- 
atics, and molecular evolution can be ex- 
pected to occur in the coming decades 
(Phillips et al., 1993). 


250 PHILLIPS 


The Influence of Natural History 


Natural history, the linchpin of North 
American mammalogy, has influenced 
mammalogical anatomy in three ways. First, 
there is the fact that natural history involves 
field work and wild species in a natural set- 
ting. There is a tradition of exploration, col- 
lection, and faunal survey in mammalogy. 
Second, there is the semiformal codification 
of natural history (what to look at, what 
information to record, what to share with 
others). Third, within natural history it is 
acceptable to note bits and pieces of infor- 
mation—oddities and abnormalities. 

Natural historians were regarded as am- 
ateurs by the early zoologist-naturalists who 
formed the nuclei of university and college 
zoology departments (Benson, 1988). In part 
this was due to the fact that many of the 
early survey personnel lacked advanced for- 
mal education. However, another aspect of 
this attitude was the hearsay aspect of the 
information disseminated by the natural 
historians. Indeed, at the end of the 19th 
Century and into the early 20th Century, 
writing in natural history was a curious 
amalgamation of fact and fiction, keen ob- 
servation and anthropomorphism (see 
Dunlap, 1988). In effect, the Grinnellian era 
was devoted to a process that I have termed 
“codification.” That is, the ground rules and 
style of research in natural history were 
gradually organized into a format resem- 
bling what was accepted as “‘science”’ in bi- 
ology. This process unfolds dramatically if 
one reads the Journal of Mammalogy from 
1919 to 1945. Likewise, museum-based 
mammalogical anatomy inherited a natural 
history component that influenced the par- 
adigm. 

By tradition, the natural historians an- 
cestral to mammalogical anatomists were 
keen observers who noticed virtually any- 
thing that was unusual in the field or in 
features of the specimens that they exam- 
ined after their collecting trips. The idea of 
specifically recording observations of such 


features as integumentary glands, coats, spe- 
cial sensory structures, and abnormalities 
was codified early in the history of the ASM 
by Ernest Thompson Seton (Seton, 1919). 
In 1927, Joseph Grinnell presented a ver- 
sion of what should be acceptable in natural 
history studies, and this in turn formed the 
basis of E. Raymond Hall’s version (Hall, 
1955). 

After a review of the Journal of Mam- 
malogy, one is struck with the sensation that 
many of the mammalogical anatomists took 
seriously Seton’s (1919) suggestion about 
noting the presence or absence and char- 
acteristics of glands. It almost seems as 
though Seton’s paper, published in the first 
volume of the Journal, initiated an entire 
series of investigations of glands and gland 
structure. The Journal thus contains what 
may be the most extensive set of articles on 
this aspect of mammalian anatomy ever 
published. Indeed, the Journal of Mam- 
malogy is by far the best single source of 
basic information about cutaneous and oth- 
er integumentary glands in insectivores 
(Dryden and Conaway, 1967; Eadie, 1938), 
pikas (Harvey and Rosenberg, 1960), bats 
(Hood and Smith, 1984; Phillips et al., 
1987b; Werner and Lay, 1963; Valdivieso 
and Tamsitt, 1964), rodents (Eriksson, 1981; 
Quay, 1962, 1965, 1968; Quay and Tomich, 
1963), and ungulates (Quay and Miller- 
Schwarze, 1970). 

Integumentary glands have attracted so 
much attention in part because they are of- 
ten obvious to an observer. The most no- 
table scientific reason, however, is that (as 
was apparent even to early natural histori- 
ans) skin glands are important to mam- 
malian biology. Observers in the field noted 
that individuals in some species seemed to 
react to each other based on olfactory cues; 
individuals frequently appeared to sniff cer- 
tain areas of the skin on conspecifics. Ob- 
servers also noted that glands were often 
more prominent in males than in females. 
Subsequently it was shown that mammalian 
skin glands typically are responsive to an- 
drogenic stimulation (Jannett, 1975; Quay, 


ANATOMY 204 


1968), so secretions have the potential of 
being sex-specific. In addition to sex-spe- 
cific secretions, integumentary glands some- 
times harbor symbiotic bacteria that ap- 
parently are involved in scent production 
(Studier and Lavoie, 1984; Tandler et al., 
in press b). Although the histology and his- 
tochemistry of integumentary glands are not 
often discussed in detail in general texts, at 
least their roles in behavior now are widely 
appreciated (Miiller-Schwarze, 1983). 

Although many individuals have contrib- 
uted to the knowledge of mammalian skin 
glands, William B. Quay heads the list. Ref- 
erences to his extensive comparative anal- 
yses can be seen in articles ranging from 
modern reviews of knowledge about the in- 
tegument and lipid-secretion in mammals 
to microanatomy of microtine rodents (see 
Quay, 1965, for example). It may not be 
surprising to discover that Quay’s academic 
roots in part trace to the Museum of Zo- 
ology at the University of Michigan. The 
Museum of Zoology in Quay’s time (late 
1950s) was blessed with two of Joseph Grin- 
nell’s students, William H. Burt and Em- 
mett T. Hooper. Quay’s research, right from 
the beginning, exemplified both the taxo- 
nomic and the natural history components 
of mammalogical anatomy. Keeping in the 
tradition, some of his histological and his- 
tochemical articles appeared in Museum of 
Zoology publications. However, Quay was 
able to expand the border of his research 
beyond the traditions set by taxonomy and 
natural history. He proceeded to the exper- 
imental type of research favored by both 
medical anatomists and zoological mor- 
phologists; he was not satisfied with descrip- 
tion and was willing to pursue his subject 
at a chemical level. 

Feeding habits, diet, and feeding adap- 
tations comprise another area that attracted 
the attention of the early natural historians 
and their observations help set that stage 
for detailed anatomical investigations of 
tongues, salivary glands, and digestive tracts 
in mammals (Doran and Allbrook, 1973; 
Golley, 1960; Greenbaum and Phillips, 


1974; Horner et al., 1964; Kubota and Hor- 
luchi, 1963; Phillips et al., 19875). 

In closing this section, it is worth men- 
tioning briefly another aspect of the para- 
digm of mammalogical anatomy —an inter- 
est in the incidental or abnormal. The idea 
of noting small pieces of information, such 
as unusual skulls (Thorpe, 1930), was passed 
downward through the main academic lin- 
eages along with the formulae for describing 
skulls, jaws, teeth, and coat colors. The orig- 
inal notations, published in early volumes 
of the Journal of Mammalogy, were very 
informal and seemingly were regarded as 
“news” to be shared with colleagues. Only 
very slowly did such incidental notes evolve 
into a more formal presentation, reaching 
their zenith in the 1960s. Some articles fo- 
cused on unusual features of anatomy that 
might have adaptive relevance (e.g., Breed, 
1981), but many focused on ‘‘abnormal”’ 
anatomical characteristics. The most strik- 
ing series of notes on anatomical abnor- 
malities appeared in a 10-year period that 
began in 1963. In that period, at least 17 
papers on dental abnormalities were pub- 
lished in the Journal of Mammalogy. A sur- 
prising number of these dealt with game 
species, especially cervids, and appear to 
have been incidental observations made in 
the course of other types of investigation. 
Since 1973, only two additional reports of 
dental abnormalities were published, sug- 
gesting either a change in interest or, per- 
haps, a new editorial policy. Indeed, the last 
decade apparently will mark the demise of 
the “note” as a means of recording inciden- 
tal observations of anatomical oddities. 


The Future of Mammalogical 
Anatomy 


In the previous sections I have described 
paradigms of academic anatomy practiced 
in North America and explained the origin 
of a special paradigm that I term mam- 


ZO2 PHILLIPS 


malogical anatomy. As we have seen, mam- 
malogical anatomy arose independently; the 
intellectual milieu, the format of presenta- 
tion, the selection of topics, the expectations 
of the practitioners—in fact, the para- 
digm—is a conglomerate of collection, tax- 
onomy, museum technique, and natural 
history. However, after 75 years, anatomy 
still has not been well-integrated into faunal 
mammalogy. That is to say, anatomy has a 
somewhat superficial relationship to main- 
line mammalogy. The general absence of 
anatomy in faunal mammalogy can be at- 
tributed to two factors: 1) the origins of 
mammalogical anatomy; and 2) the cultural 
and scientific gaps among mainline mam- 
malogy and medical anatomy and zoolog- 
ical morphology. The first factor is indeed 
ironic because many of the scientific prod- 
ucts of mammalogical anatomy are sub- 
merged in or identified with taxonomy and 
natural history. Mammalogical anatomists 
have unwittingly buried at least some of their 
work by publishing it in a taxonomic con- 
text or a taxonomic format. In a taxonomic 
context, descriptive anatomical data are rel- 
egated to the category of “‘characters”’ and, 
as a consequence, the data, or any discus- 
sion of functions or roles of anatomical fea- 
tures, are also lost in the body of the text. 
Mammalogical anatomy has been largely 
lost to medical anatomists and zoological 
morphologists for the same reasons that it 
has been overlooked by faunal mammalo- 
gists. Namely, articles in mammalogical 
anatomy often are misidentified as being of 
a purely taxonomic nature and thus are not 
consulted as sources of useful anatomical 
information. 

Having a retrospective on the past 75 years 
might offer some hints as to the future of 
mammalogical anatomy. An understanding 
of the relationships among mammalogy, its 
anatomical offspring, and medical anatomy 
and zoological morphology, might be useful 
to the next generation of mammalogical 
anatomists. It seems clear that the next step 
is integration. By this, I mean more than 
just integrating mammalogical anatomy into 


faunal mammalogy (although that alone is 
a worthy challenge). More importantly, 
mammalogical anatomy should begin to 
profit from its own intellectual and academ- 
ic heritage. The taxonomic perspective, for 
instance, should be employed to underscore 
the value of understanding relationships 
when designing experiments. Rather than 
being the instrument by which data are bur- 
ied and forgotten, systematics should be the 
reason why data are understandable in an 
evolutionary context. 


Acknowledgments 


I greatly appreciate the opportunity provided 
by my colleagues, E. C. Birney and J. R. Choate, 
to recount my understanding of the origins, his- 
tory, and paradigm of anatomy in mammalogy. 
The views expressed in my essay are strictly my 
own and should not be misconstrued to represent 
those of the ASM, editors, or various colleagues 
who were helpful to me while I was writing the 
manuscript. I appreciated the editorial advice of 
G. Lawrence Forman of Rockford College. My 
own interest in microanatomy was largely stim- 
ulated by Larry, who very kindly shared his ex- 
pertise in histology and histological technique 
when we both were graduate students at the Mu- 
seum of Natural History at The University of 
Kansas. Bernard Tandler, formerly of the De- 
partment of Oral Biology, School of Dentistry, 
Case Western Reserve University, helped me gain 
insight to medical anatomy and zoological mor- 
phology by sharing his own experiences as both 
a student and scientist. Christopher Horvath of 
the Philosophy Department and Department of 
Biological Sciences at Illinois State University 
provided very valuable insight to philosophical 
and ethical issues. Some of my ideas and insight 
came from discussions with scientists who now 
are deceased, in particular E. R. Hall and J. K. 
Jones, Jr. lam grateful for the scholarly tradition 
that they so willingly shared. I also wish to thank 
N. Doss, Illinois State University, for her con- 
siderable assistance with the preparation of the 
manuscript. Finally, I wish to acknowledge the 
assistance of K. M. Van de Graaff of Brigham 
Young University in reviewing the anatomical 
literature in the Journal of Mammalogy. 


ANATOMY 250 


Literature Cited 


AspotTtT, M. J., ET AL. 1984. Effect of photoperiod on 
growth and fat accretion in white-tailed doe fawns. 
The Journal of Wildlife Management, 48:776-787. 

ALLEN, H. 1880. The ethmoid bone in bats. Bulletin 
of the Museum of Comparative ZoGlogy, at Harvard 
College, 6:121-122. 

. 1885. On the embryos of bats. Contributions 
from the Zoological Laboratory of the University of 
Pennsylvania, 1:3-42. 

ALTENBACH, J. S. 1979. Locomotor morphology of 
the vampire bat, Desmodus rotundus. Special Pub- 
lication, American Society of Mammalogists, 6:1- 
SHE 

ANDERSON, S. 1968. New craniometer and sugges- 
tions for craniometry. Journal of Mammalogy, 49: 
21-228. 

AppEL, T. A. 1988. Organizing biology: The Ameri- 
can Society of Naturalists and its ‘“‘affiliated socie- 
ties,” 1883-1923. Pp. 87-120, in The American de- 
velopment of biology (R. Rainger, K. R. Benson, 
and J. Maienschein, eds.). University of Pennsyl- 
vania Press, Philadelphia, 380 pp. 

ATKESON, T. D., AND R. L. MARCHINTON. 1982. Fore- 
head glands of the white-tailed deer. Journal of 
Mammalogy, 63:613-617. 

Baber, R. S. 1965. A partition of variance in dental 
traits of the house mouse. Journal of Mammalogy, 
46:384-388. 

BENSON, K. R. 1988. From museum research to lab- 
oratory research: the transformation of natural his- 
tory into academic biology. Pp. 49-83, in The Amer- 
ican development of biology (R. Rainger, K. R. 
Benson, and J. Maienschein, eds.). University of 
Pennsylvania Press, Philadelphia, 380 pp. 

BirRNEY, E. C., AND R. M. Timm. 1975. Dental on- 
togeny and adaptation in Diphylla ecaudata. Journal 
of Mammalogy, 56:204-206. 

Biair, W. F. 1942. Systematic relationships of Pero- 
myscus and several related genera as shown by the 
baculum. Journal of Mammalogy, 23:196-204. 

Bonner, J. T. (ED). 1966. On growth and form by 
D’arcy Wentworth Thompson. Cambridge Univer- 
sity Press, London, 346 pp. 

BREED, W. G. 1981. Unusual anatomy of the male 
reproductive tract in Notomys alexis (Muridae). 
Journal of Mammalogy, 63:373-375. 

BRIAN, M., C. HickEy, AND M. B. FENTON. 1987. 
Scent-dispersing hairs (osmetricha) in some Pterop- 
idae and Molossidae (Chiroptera). Journal of Mam- 
malogy, 68:38 1-384. 

BropiE, F. W. 1974. Thomas Jefferson: an intimate 
history. W. W. Norton and Company, New York, 
594 pp. 

Bry skl, P. 1993. The evolutionary morphology of 
heteromyids. Pp. 357-385, in Biology of the Het- 
eromyidae (H. H. Genoways and J. H. Brown, eds.). 
Special Publication, American Society of Mammal- 
ogists, 10:1-719. 

Burt, W.H. 1960. Bacula of North American mam- 
mals. University of Michigan Museum of Zoology, 
Miscellaneous Publications, 113:1-76. 

Burt, W. H., AND F. S. BARKALOw, Jr. 


1942. A 


comparative study of the bacula of woodrats. Journal 
of Mammalogy, 23:287-297. 

CALFORD, M. B., GRAYDON, M. L., Huerta, M. F., 
KAAs, J. H., AND J. D. PettiGRew. 1985. A variant 
of the mammalian somatotrophic map in a bat. Na- 
ture, 313:477-479. 

CaRLETON, M. D. 1985. Macroanatomy. Pp. 116- 
175, in Biology of the New World Microtus (R. H. 
Tamarin, ed.). Special Publication, American Soci- 
ety of Mammalogists, 8:1-893. 

CARTMILL, M. 1985. Climbing. Pp. 73-88, in Func- 
tional vertebrate morphology (M. Hildebrand, D. M. 
Bramble, K. F. Liem, and D. B. Wake, eds.). The 
Belknap Press of Harvard University Press, Cam- 
bridge, 430 pp. 

CuHIAsson, R. B. 1957. The dentition of the Alaskan 
fur seal. Journal of Mammalogy, 38:310-319. 

Dapson, R. W., StupiER, E. H., BUCKINGHAM, M. J., 
AND A. L. Stupier. 1977. Histochemistry of odor- 
iferous secretions from integumentary glands in three 
species of bats. Journal of Mammalogy, 58:531-535. 

Diamonp, J. M. 1992. The red flag of optimality. 
Nature, 355:204—206. 

Doran, G. A., AND D. B. ALLBROOK. 1973. The tongue 
and associated structures in two species of African 
pangolins. Journal of Mammalogy, 54:909-923. 

DrybDen, G. L., AND C. H. Conaway. 1967. The 
origin and hormonal control of scent production in 
Suncus murinus. Journal of Mammalogy, 48:420- 
428. 

Duntap, T. R. 1988. Saving America’s wildlife. 
Princeton University Press, Princeton, New Jersey, 
214 pp. 

Eapige, W. R. 1938. The dermal glands of shrews. 
Journal of Mammalogy, 19:171-174. 

EISENBERG, J. F. 1981. The mammalian radiations: 
an analysis of trends in evolution, adaptation, and 
behavior. University of Chicago Press, Chicago, 610 
pp. 

Eriksson, M. 1981. The bulbourethral gland of the 
wood mouse, Apodemus sylvaticus. Journal of Mam- 
malogy, 62:375-378. 

Estes, R. D., CumMmiINnG, D. H. M., AND G. W. HERN. 
1982. New facial glands in domestic pig and wart- 
hog. Journal of Mammalogy, 63:618-624. 

FELDMAN, J. L., AND C. J. PHituips. 1984. Compar- 
ative retinal pigment epithelium and photoreceptor 
ultrastructure in nocturnal and fossorial rodents: the 
eastern woodrat and plains pocket gopher. Journal 
of Mammalogy, 65:231-245. 

Fis, F. E. 1982. Function of the compressed tail of 
surface swimming muskrats (Ondatra zibethicus). 
Journal of Mammalogy, 63:591-597. 

FLYNN, L. J., NEvo, E., AND G. HeTH. 1987. Incisor 
enamel microstructure in blind mole rats: adaptive 
and phylogenetic significance. Journal of Mammal- 
ogy, 68:500-507. 

FORMAN, G. L. 1972. Comparative morphological 
and histochemical studies of stomachs of selected 
American bats. University of Kansas Science Bul- 
letin, 59:591-729. 

FoRMAN, G.L., ANDC. J. PHILLIPS. 1988. Preparation 
and fixation of tissues for histological, histochemical, 
immunohistochemical, and electron microscopical 
studies. Pp. 405-424, in Ecological and behavioral 


254 PHILLIPS 


methods for the study of bats (T. H. Kunz, ed.). 
Smithsonian Institution Press, Washington, D.C., 533 


1993. The proximal colon of heteromyid ro- 
dents: possible morphological correlates to enhanced 
water conservation. Pp. 491-508, in Biology of the 
Heteromyidae (H. H. Genoways and J. H. Brown, 
eds.). Special Publication, American Society of 
Mammalogists, 10:1-719. 

FREEMAN, P. W. 1981. A multivariate study of the 
Family Molossidae (Mammalia, Chiroptera): mor- 
phology, ecology, and evolution. Fieldiana: Zoology, 
new series, 7:1-173. 

FREEMAN, P. W., AND C. A. LEMEN. 1991. Morpho- 
metrics of the Family Emballonuridae. Pp. 54-61, 
in Contributions to mammalogy in honor of Karl F. 
Koopman (T. A. Griffiths and D. Klingener, eds.). 
Bulletin of the American Museum of Natural His- 
tory, 432 pp. 

Genoways, H. H., AND J. H. BRown (eps.). 1993. 
Biology of the Heteromyidae. Special Publication, 
American Society of Mammalogists, 10:1-719. 

GiLL, A. E., AND K. BoLies. 1982. A heritable tooth 
trait varying in two subspecies of Microtus califor- 
nicus (Rodentia: Cricetidae). Journal of Mammal- 
ogy, 63:96-103. 

GINGERICH, P. D., AND K. D. Rose. 1979. Anterior 
dentition of the Eocene condylarth: convergence with 
the tooth comb of lemurs. Journal of Mammalogy, 
60:16-22. 

GINGERICH, P. D., AND D. A. WINKLER. 1979. Pat- 
terns of variation and correlation in the dentition of 
the red fox. Journal of Mammalogy, 60:691-704. 

GoLpsTEIN, B. 1972. Allometric analysis of relative 
humerus width and olecranon length in some un- 
specialized burrowing mammals. Journal of Mam- 
malogy, 53:148-156. 

GoLLey, F. B. 1960. Anatomy of the digestive tract 
of Microtus. Journal of Mammalogy, 41:89-99. 

GoopMaAN, B. 1991. Holy phylogeny! Did bats evolve 
twice? Science, 253:36. 

GREENBAUM, I. F., AND C. J. PHiLiips. 1974. Com- 
parative anatomy and general histology of the tongues 
of long-nosed bats with reference to infestation of 
oral mites. Journal of Mammalogy, 55:489-504. 

GREENBERG, G., BRINGAS, P., JR., AND H. C. SLAVKIN. 
1984. The epithelial genotype controls the pattern 
of extracellular enamel prism formation. Differen- 
tiation, 26:32-43. 

GRIFFITHS, T. A., AND A.L.SmitH. 1991. Systematics 
of emballonuroid bats (Chiroptera: Emballonuridae 
and Rhinopomatidae), based on hyoid morphology. 
Pp. 62-83, in Contributions to mammalogy in honor 
of Karl F. Koopman (T. A. Griffiths and D. Klin- 
gener, eds.). Bulletin of the American Museum of 
Natural History, 432 pp. 

HA.Lt, E. R. 1935. Certain osteological features of 
Euderma maculatum. Journal of Mammalogy, 15: 
68-70. 

. 1955. Handbook of mammals of Kansas. 
Miscellaneous Publications, Museum of Natural 
History, University of Kansas, 7:1-303. 

Harvey, E. B., AND L. E. ROSENBERG. 1960. An apo- 
crine gland complex of the pike. Journal of Mam- 
malogy, 41:213-219. 


HERRING, S. W. 1985. Morphological correlates of 
masticatory patterns in peccaries and pigs. Journal 
of Mammalogy, 66:603-617. 

HILDEBRAND, M. 1985a. Walking and running. Pp. 
38-57, in Functional vertebrate morphology (M. 
Hildebrand, D. M. Bramble, K. F. Liem, and D. B. 
Wake, eds.). The Belknap Press of Harvard Univer- 
sity Press, Cambridge, 430 pp. 

19855. Digging of quadrupeds. Pp. 89-109, 
in Functional vertebrate morphology (M. Hilde- 
brand, D. M. Bramble, K. F. Liem, and D. B. Wake, 
eds.). The Belknap Press of Harvard University Press, 
Cambridge, 430 pp. 

HOFFMAN, R. A., AND P. F. ROBINSON. 1966. Changes 
in the endocrine glands of white-tailed deer as af- 
fected by season, sex, and age. Journal of Mam- 
malogy, 47:266-280. 

Hotter, J. B., AND H. H. Hayes. 1977. Growth in 
white-tailed deer fawns fed varying energy and con- 
stant protein. The Journal of Wildlife Management, 
41:506-510. 

Homan, J. A., AND H. H. Genoways. 1978. An anal- 
ysis of hair structure and its phylogenetic implica- 
tions among heteromyid rodents. Journal of Mam- 
malogy, 59:740-760. 

Hoop, C.S., AND J. D. SMitH. 1982. Cladistical anal- 
ysis of female reproductive histomorphology in 
phyllostomatoid bats. Systematic Zoology, 31:241- 
251, 


. 1983. Histomorphology of the female repro- 
ductive tract in phyllostomoid bats. Occasional Pa- 
pers, The Museum, Texas Tech University, 86:1- 
38. 


1984. Histology of a sexually dimorphic in- 
tegumentary gland in Macroglossus lagochilus. Jour- 
nal of Mammalogy, 65:1-9. 

Horner, B.E. 1944. Anewcraniometer for carnivore 
and other mammalian crania. Journal of Mammal- 
ogy, 25:71-77. 

Horner, B. E., J. M. TAYLOR, AND H. H. PADYKULA. 
1964. Food habits and gastric morphology of the 
grasshopper mouse. Journal of Mammalogy, 45:513- 
535: 

Howe .t, A. B. 1929. Contributions to the compar- 
ative anatomy of the eared and earless seals (genera 
Zalophus and Phoca). United States National Mu- 
seum, Proceedings, 73:1-142. 

JANNETT, F. J., Jk. 1975. “Hip glands” of Microtus 
pennsylvanicus and M. longicaudus (Rodentia: Mu- 
ridae), voles “without” hip glands. Systematic Zo- 
ology, 24:171-175. 

Jones, J. K., Jk. 1991. Genealogy of Twentieth-cen- 
tury systematic mammalogists in North America: 
the descendants of Joseph Grinnell. Pp. 48-56, in 
Latin American mammalogy: history, biodiversity, 
and conservation (M. A. Mares and D. J. Schmidley, 
eds.). University of Oklahoma Press, Norman, 468 


pp. 

Jones, J. K., Jk., AND H. H. GeNoways. 1970. Chi- 
ropteran systematics. Pp. 3-21, in About bats: a chi- 
ropteran biology symposium (B. H. Slaughter and 
D. W. Walton, eds.). Southern Methodist University 
Press, Dallas, 339 pp. 

Jones, T. R., AND R. K. PLAKKE. 1981. The histology 
and histochemistry of the perianal scent gland of the 


ANATOMY 255 


reproductively quiescent black-tailed prairie dog 
(Cynomys ludovicianus). Journal of Mammalogy, 62: 
362-372. 

Kittie, R. A. 1981. The function of interlocking ca- 
nines in rain forest peccaries (Tayassuidae). Journal 
of Mammalogy, 62:459-469. 

KirKPATRICK, R. D., AND L. K. Sowts. 1962. Age 
determination of the collared peccary by tooth re- 
placement pattern. The Journal of Wildlife Manage- 
ment, 26:214-217. 

KEIN, D. R., MELDGAARD, M., ANDS. G. FANcy. 1987. 
Factors determining leg length in Rangifer tarandus. 
Journal of Mammalogy, 68:642-655. 

KLEVEZAL, G. A., AND A.C. Myrick, JR. 1984. Marks 
in tooth dentine of female dolphins as indicators of 
parturition. Journal of Mammalogy, 65:103-110. 

KLINGENER, D. 1963. Dental evolution of Zapus. 
Journal of Mammalogy, 44:248-260. 

KOHLSTEDT, S.G. 1988. Museums on campus: a tra- 
dition of inquiry and teaching. Pp. 15-47, in The 
American development of biology (R. Rainger, K. 
R. Benson, and J. Maienshein, eds.). University of 
Pennsylvania Press, Philadelphia, 380 pp. 

KOHLSTEDT, S. G. (ED.). 1991. The origins of natural 
science in America: the essays of George Brown 
Goode. Smithsonian Institution Press, Washington, 
411 pp. 

Krause, D. W., AND S. J. CARLSON. 1987. Prismatic 
enamel in multituberculate mammals: tests of ho- 
mology and polarity. Journal of Mammalogy, 68: 
755-765. 

Krutzscu, P. H. 1959. Variation in the os penis of 
tropical fruit bats. Journal of Mammalogy, 40:387- 
392. 


1962. Additional data on the os penis of 
Megachiroptera. Journal of Mammalogy, 43:34—-42. 

KusoTa, K., AND H. Horrucui. 1963. Anatomical 
studies of the salivary glands of the fur seal. Journal 
of Mammalogy, 44:349-364. 

Kuun, T. 1962. Structure of the scientific revolution. 
University of Chicago Press, Chicago, 173 pp. 

Lay, D. M. 1993. Anatomy of the heteromyid ear. 
Pp. 270-290, in Biology of the Heteromyidae (H. H. 
Genoways and J. H. Brown, eds.). Special Publica- 
tion, American Society of Mammalogists, 10:1-719. 

Layne, J.N. 1952. Theos genitalia of the red squirrel, 
Tamiasciurus. Journal of Mammalogy, 33:457-459. 

LipickER, W. Z. 1968. The phylogeny of New Guinea 
rodent genera based on phallic morphology. Journal 
of Mammalogy, 49:609-643. 

LOCHMILLER, R. L., HELLGREN, E. C., AND W. E. GRANT. 
1987. Physical characteristics of neonate, juvenile, 
and adult collared peccaries (Tayassu tajacu angu- 
latus) from south Texas. Journal of Mammalogy, 68: 
188-194. 

MAIENSCHEIN, J. 1988. Whitman at Chicago: estab- 
lishing a Chicago style of biology? Pp. 151-182, in 
The American development of biology (R. Rainger, 
K. R. Benson, and J. Maienschein, eds.). University 
of Pennsylvania Press, Philadelphia, 380 pp. 

Mares, M. A. 1991. How scientists can impede the 
development of their discipline: egocentrism, small 
pool size, and the evolution of “‘sapismo.” Pp. 57- 
75, in Latin American mammalogy: history, biodi- 
versity, and conservation (M. A. Mares and D. J. 


Schmidley, eds.). University of Oklahoma Press, 
Norman, 468 pp. 

Marks, S. A., AND A. W. ERICKSON. 1966. Age de- 
termination in black bears. The Journal of Wildlife 
Management, 30:389-410. 

MENNONE, A., C. J. PHILLIPS, AND D. E. Pumo. 1986. 
Evolutionary significance of interspecific differences 
in gastrin-like immunoreactivity in the pylorus of 
phyllostomid bats. Journal of Mammalogy, 67:373- 
384. 

Miter, G.S. 1895. On the introitus vaginae of cer- 
tain Muridae. Contributions from the Zodlogical 
Laboratory of the Museum of Comparative Zodlogy 
at Harvard College, 45:459-468. 

MossInG, T., AND L. KALLQuist. 1981. Variation in 
cutaneous glandular structures in reindeer. Journal 
of Mammalogy, 62:606-612. 

MULLER-SCHWARZE, D. 1983. Scent glands in mam- 
mals and their functions. Pp. 150-197, in Advances 
in the study of mammalian behavior (J. F. Eisenberg 
and D. G. Kleiman, eds.). Special Publication, 
American Society of Mammalogists, 7:1-753. 

NaGato, T., B. TANDLER, AND C. J. PHILLIPS. 1984. 
Unusual smooth endoplasmic reticulum in subman- 
dibular acinar cells in the male round-eared bat, To- 
natia sylvicola. Journal of Ultrastructure Research, 
87:275-284. 

NELSON, T. W., AND K. A. SHump, JR. 1978. Cranial 
variation and size allometry in agouti paca from Ec- 
uador. Journal of Mammalogy, 59:387-394. 

Novacek, M. J. 1991. Aspects of the morphology of 
the cochlea in microchiropteran bats: an investiga- 
tion of character transformation. Pp. 84-100, in 
Contributions to mammalogy in honor of Karl F. 
Koopman (T. A. Griffiths and D. Klingener, eds.). 
Bulletin of the American Museum of Natural His- 
tory, 432 pp. 

OzoGa, J. J., AND L. J. VERME. 1978. The thymus 
gland as a nutritional status indicator in deer. The 
Journal of Wildlife Management, 42:791-798. 

PATTERSON, B. D., AND C. S. THAELER, JR. 1982. The 
mammalian baculum: hypotheses on the nature of 
bacular variation. Journal of Mammalogy, 63:1-15. 

PuHILuips, C. J. 1971. The dentition of glossophagine 
bats: development, morphological characteristics, 
variation, pathology, and evolution. Miscellaneous 
Publications, Museum of Natural History, Univer- 
sity of Kansas, 54:1-138. 

. 1985a. Field fixation and storage of museum 

tissue collections suitable for electron microscopy. 

Acta Zoologica Fennica, 170:87-90. 

1985b. Microanatomy. Pp. 176-253, in Bi- 
ology of the New World Microtus (R. H. Tamarin, 
ed.). Special Publication, American Society of Mam- 
malogists, 8:1-893. 

PHILLIPs, C. J., AND B. A. OXBERRY. 1972. Compar- 
ative histology of molar dentitions of Microtus and 
Clethrionomys, with comments on dental evolution. 
Journal of Mammalogy, 53:1-21. 

PuHILuips, C. J., AND B. TANDLER. 1987. Mammalian 
evolution at the cellular level. Pp. 1-66, in Current 
mammalogy (H. H. Genoways, ed.). Plenum Press, 
New York, 1:1-519. 

PHILLIPS, C. J., J. K. JoNEs, JR., AND F. J. RADOVSKY. 
1969. Macronyssid mites in the oral mucosa of long- 


250 PHILLIPS 


nosed bats: notes on occurrence and associated pa- 
thology. Science, 165:1368-1369. 

Puiiuips, C. J., T. NAGATO, AND B. TANDLER. 1987a. 
Comparative ultrastructure and evolutionary pat- 
terns of acinar secretory product of parotid salivary 
glands in Neotropical bats. Pp. 213-229, in Studies 
in Neotropical mammalogy: essays 1n honor of Phil- 
ip Hershkovitz (B. D. Patterson and R. M. Timm, 
eds.). Fieldiana: Zoology, new series no. 39, 506 pp. 

PuiLuips, C. J., B. TANDLER, AND C. A. PINKSTAFF. 
1987b. Unique salivary glands in two genera of 
tropical microchiropteran bats: an example of evo- 
lutionary convergence in histology and histochem- 
istry. Journal of Mammalogy, 68:235-242. 

PuiLuips, C. J., B. STEINBERG, AND T. H. Kunz. 1982. 
Dentin, cementum, and age determination in bats: 
a critical evaluation. Journal of Mammalogy, 63: 
197-207. 

PHILuips, C. J., K. M. STUDHOLME, AND G. L. FORMAN. 
1984. Results of the Alcoa Foundation Suriname 
expeditions. VIII. Comparative ultrastructure of gas- 
tric mucosae in four genera of bats (Mammalia: Chi- 
roptera), with comments on gastric evolution. An- 
nals of Carnegie Museum, 53:71-117. 

PHILuirs, C. J., B. TANDLER, AND T. NAGATo. 1993. 
Evolutionary divergence of salivary gland acinar cells: 
a format for understanding molecular evolution. Pp. 
39-80, in Biology of the salivary glands (K. Dob- 
rosielski- Verona, ed.). CRC Press, Boca Raton, 461 
pp. 

Puitups, J. C. 1919. Skull measurements in the 
northern Virginia deer. Journal of Mammalogy, 
1:130-133. 

PIATIGORSKY, J., AND G. J. Wistow. 1989. Enzyme/ 
crystallins: gene sharing as an evolutionary strategy. 
Cell, 57:197-199. 

Powe LL, J. W. 1925. First through the Grand Can- 
yon. The MacMillian Company, New York, 820 pp. 

Purves, D., AND J. W. LICHTMAN. 1985. Geometrical 
differences among homologous neurons in mam- 
mals. Science, 228:298-302. 

Quay, W. B. 1962. Apocrine sweat glands in the 
angulus oris of microtine rodents. Journal of Mam- 
malogy, 43:303-310. 

1965. Comparative survey of the sebaceous 

and sudoriferous glands of the oral lips and angle in 

rodents. Journal of Mammalogy, 46:23-37. 

1968. The specialized posterolateral seba- 
ceous glandular regions in microtine rodents. Journal 
of Mammalogy, 49:427-445. 

Quay, W. B., AND D. MULLER-SCHWARZE. 1970. 
Functional histology of integumentary glandular 
regions in the black-tailed deer. Journal of Mam- 
malogy, 51:675-694. 

Quay, W. B., AND P. Q. TomicH. 1963. A specialized 
midventral sebaceous glandular area in Rattus ex- 
ulans. Journal of Mammalogy, 44:537-542. 

RAINGER, R., K. R. BENSON, AND J. MAIENSCHEIN (EDS. ). 
1988. The American development of biology. Uni- 
versity of Pennsylvania Press, Philadelphia, 380 pp. 

Ransom, A. G. 1965. Kidney and marrow fat as in- 
dicators of white-tailed deer condition. The Journal 
of Wildlife Management, 29:397-398. 

REDUKER, D. W. 1983. Functional analysis of the 


masticatory apparatus in two species of Myotis. Jour- 
nal of Mammalogy, 64:277-286. 

Ritey, M. A. 1985. An analysis of masticatory form 
and function in three mustelids. Journal of Mam- 
malogy, 66:519-528. 

RODRIGUEZ-COLUNGA, M. J., C. FERNANDEZ, C. 
RODRIGUEZ, D. TOLIVIA, AND A. MENENDEZ-PELAEZ. 
1992. Female Syrian hamster Harderian gland: de- 
velopment and effects of high environmental tem- 
perature and melatonin injections of histology and 
porphyrin deposits. The Anatomical Record, 232: 
293-300. 

Ryan, J. M. 1991. Comparative morphology of the 
glans penis in Molossus, Promops, and Eumops (Chi- 
roptera: Molossidae). Pp. 122-137, in Contributions 
to mammalogy in honor of Karl F. Koopman (T. A. 
Griffiths and D. Klingener, eds.). Bulletin of the 
American Museum of Natural History, 432 pp. 

SANDs, M. W., R. P. CoppiINGER, AND C. J. PHILLIPS. 
1977. Comparisons of thermal sweating and his- 
tology of sweat glands of selected canids. Journal of 
Mammalogy, 58:74—-78. 

SETON, E. T. 1919. For a methodic study of life- 
histories. Journal of Mammalogy, 1:67-69. 

SHADLE, A. R. 1936. The attrition and extrusive 
growth of the four major incisor teeth of domestic 
rabbits. Journal of Mammalogy, 17:15-21. 

SHorT, H. L. 1978. Analysis of cuticular scales on 
hairs using the scanning electron microscope. Jour- 
nal of Mammalogy, 59:261-268. 

SHortT, H. L., D. E. Mepin, AND A. E. ANDERSON. 
1965. Ruminoreticular characteristics of mule deer. 
Journal of Mammalogy, 46:196-199. 

SLAUGHTER, B. H., R. M. PINE, AND N. E. Pine. 1974. 
Eruption of cheek teeth in Insectivora and Carniv- 
ora. Journal of Mammalogy, 55:115-125. 

SmitH, J. D. 1972. Systematics of the chiropteran 
Family Mormoopidae. Miscellaneous Publications, 
Museum of Natural History, University of Kansas, 
56:1-132. 

SmitH, J. D., AND G. MApxkour. 1980. Penial mor- 
phology and the question of chiropteran phylogeny. 
Pp. 347-365, in Proceedings of the Fifth Interna- 
tional Bat Research Conference (D. E. Wilson and 
A. L. Gardner, eds.). Texas Tech University Press. 


STERLING, K. B. 1991. Two pioneering American 
mammalogists in Mexico: the field investigations of 
Edward William Nelson and Edward Alphonso 
Goldman, 1892-1906. Pp. 33-47, in Latin American 
mammalogy: history, biodiversity, and conservation 
(M. A. Mares and D. J. Schmidley, eds.). University 
of Oklahoma Press, Norman, 468 pp. 

STRANDSKOV, H. M. 1942. Skeletal variations in 
guinea pigs and their inheritance. Journal of Mam- 
malogy, 23:65-75. 

STUDHOLME, K. M., S. YAZULLA, AND C. J. PHILLIPS. 
1987. Interspecific differences in immunohisto- 
chemical localization of retinal neurotransmitters in 
four species of bats. Brain, Behavior and Evolution, 
30:160-173. 

Stupier, E. H., AND K. H. LAvorg. 1984. Microbial 
involvement in scent production in noctilionid bats. 
Journal of Mammalogy, 65:711-714. 

TAMARIN, R. H. (ED). 1985. Biology of the New World 


ANATOMY aos 


Microtus. Special Publication, American Society of 
Mammalogists, 8:1-893. 

TANDLER, B., AND C. J. PHiLiips. 1993a. Structure 
of serous cells in salivary glands. Microscopy Re- 
search and Techniques, 26:32-48. 

1993b. Giant mitochondria in the seromu- 
cous secretory cells of the accessory submandibular 
gland of the long-haired fruit bat, Stenonycteris lano- 
sus. The Anatomical Record, 237:157-162. 

TANDLER, B., T. NAGATO, AND C. J. PHILLIPS. 1986. 
Systematic implications of comparative ultrastruc- 
ture of secretory acini in the submandibular salivary 
gland in Artibeus (Chiroptera: Phyllostomidae). 
Journal of Mammalogy, 67:8 1-90. 

TANDLER, B., C. J. PHILLIPS, AND C. A. PINKSTAFF. In 
press a. Mucous droplets with multiple membranes 
in the accessory submandibular glands of long-winged 
bats. European Journal of Cell Biology. 

TANDLER, B., K. TOYOSHIMA, AND C. J. PHrLuips. In 
press b. Symbiotic bacteria in the accessory sub- 
mandibular salivary gland of the club-footed bat, 
Tylonycteris pachypus. Journal of Dental Research. 

TANDLER, B., C. J. PHityips, T. NAGATO, AND K. 
ToyYosHIMA. 1990. Ultrastructure diversity in chi- 
ropteran salivary glands. Pp. 31-52, in Ultrastruc- 
ture of the extraparietal glands of the digestive tract 
(A. Riva, P. M. Motta, and F. Testa Riva, eds.). 
Kluwer Academic Publishers, Boston, 278 pp. 

THorPE, R. 1930. A remarkable woodchuck skull. 
Journal of Mammalogy, 11:67. 

VALDIVvIESO, D., AND J. R. TAmsitt. 1964. The his- 


tology of the chest gland of the pale spearnosed bat. 
Journal of Mammalogy, 45:536-539. 

VAN DE GRAAFF, K. M., J. HARPER, AND G. E. GosLow, 
Jr. 1982. Analysis of posture and gait selection 
during locomotion in the striped skunk (Mephitis 
mephitis). Journal of Mammalogy, 63:582—590. 

VAUGHAN, T. A. 1959. Functional morphology of 
three bats: Eumops, Myotis, Macrotus. University of 
Kansas Publications, Museum of Natural History, 
12:1-153. 

. 1978. Mammalogy. W. B. Saunders Com- 
pany, Philadelphia, 522 pp. 

VERME, L. J. 1979. Influence of nutrition on fetal 
organ development in deer. The Journal of Wildlife 
Management, 43:79 1-796. 

WADE, O., AND P. T. GILBERT. 1940. The baculum 
of some Sciuridae and its significance in determining 
relationships. Journal of Mammalogy, 21:52-62. 

WERNER, H. J., AND D. M. Lay. 1963. Morphological 
aspects of the chest gland of the bat, Mo/ossus ater. 
Journal of Mammalogy, 44:552-555. 

WILkins, K. T., AND C. A. Woops. 1983. Modes of 
mastication in pocket gophers. Journal of Mam- 
malogy, 64:636-641. 

WILSON, D. E., AND J. F. EISENBERG. 1990. Origin 
and applications of mammalogy in North America. 
Pp. 1-35, in Current mammalogy (H. H. Genoways, 
ed.). Plenum Press, New York, 2:1-577. 

ZIEGLER, A.C. 1971. Dental homologies and possible 
relationships of Recent Talpidae. Journal of Mam- 
malogy, 52:50-68. 


PHYSIOLOGY 


Bruce A. WUNDER AND GREGORY L. FLORANT 


Introduction 


hysiology is at the same time a very old 
yet relatively new field of inquiry. 
Webster’s New World Dictionary defines 
physiology as “The branch of biology deal- 
ing with the functions and vital processes 
of living organisms or their parts and or- 
gans.”’ Karl Rothschuh (1973) has written 
a thorough history of physiological thought 
and the development of modern experi- 
mental methods in pursuing the subject. In 
it he notes that Aristotle (384-322 BC) used 
the term in its broadest sense to mean the 
study of knowledge about nature (from the 
Greek, physis, meaning knowledge of na- 
ture). Later, Greeks came to focus the mean- 
ing on the healing power of nature. Even as 
late as the 19th Century the term was still 
used in a broad sense. In 1848, Robley Dun- 
glison defined “‘biology” as simply “‘physi- 
ology” and in the minds of most the term 
meant “‘hygiene”’ (Appel, 1987a). It was not 
until the 1850s or 1860s that experimen- 
tation became a tradition, and physiology 
came to take on the meaning given in Web- 
ster as defined above. 

It is this topic, the study of organism func- 
tion, which we will review here in the time 
frame of the founding and development of 
the ASM. The literature on the topic is huge 


258 


SO we will try to focus primarily on physi- 
ology as it relates to wild mammals, but will 
not make a clean distinction as we frequent- 
ly need to reference what was being covered 
in the general physiological literature to put 
the wild mammal, or comparative studies, 
into perspective. Further, much of the ear- 
liest comparative literature comes from 
studies on invertebrates, as an early group 
of physiologists started work at the Woods 
Hole Marine Biological Labs and found in- 
vertebrates convenient forms for study. 


Physiology Background 


Biologists, natural historians, and medi- 
cal scientists dissected and reported on the 
anatomy of animals hundreds of years be- 
fore they started to experiment on and un- 
derstand the function of animals. Anatomy 
texts, with inferences to function, appeared 
in the time of the Greeks and the Middle 
Ages. However, the first texts in physiology, 
per se, were not written until the early 1800s 
in Europe. In North America, texts on phys- 
iology varied greatly from descriptive books 
on the human body intended primarily for 
medical students (e.g., Dunglison’s Human 


PHYSIOLOGY 


Physiology) to popular tracts on nutrition 
and hygiene. The first physiology text in 
North America was not published until 
1896, 9 years after formation of the Amer- 
ican Physiological Society (Appel, 19875). 
The concept of “‘experiment’”’ to under- 
stand function was first pursued in a modern 
sense in Germany by Ludwig and others, 
and early U.S. physiologists typically studied 
there (Adolph, 1987; Rothschuh, 1973). 
These early studies (around the turn of the 
century) were necessarily descriptive and 
primarily of medical focus. Only later did 
investigators broaden their scope to include 
organisms other than humans or lab ani- 
mals as models for humans. By 1887 there 
were several laboratories in North America 
where important research was being done 
and students could get good, thorough train- 
ing in experimental methods. Thus, in 1887, 
17 individuals decided to form the Amer- 
ican Physiological Society (APS). Interest- 
ingly, one of the societies they used as their 
model was the American Society of Natu- 
ralists (ASN, formed in 1883), primarily be- 
cause it was the first society that had re- 
stricted membership, a condition they felt 
was important. Further, at that time there 
was a closer tie between the disciplines of 
physiology and natural history. For exam- 
ple, the condition of membership in the ASN 
was ““Membership in this society shall be 
limited to Instructors in Natural History, 
Officers of Museums and other Scientific 
Institutions, Physicians and other persons 
professionally engaged in some branch of 
Natural History” (italics ours). Indeed, half 
the original members of the APS were also 
members of the ASN and three of the or- 
ganizers had been President of ASN! 
Physiological journals of North America 
are relatively young, although a little older 
than the Journal of Mammalogy (begun in 
1919-1920). The American Journal of 
Physiology was first published in 1898. The 
emphasis in stress physiology developed out 
of World War II brought the Journal of Ap- 
plied Physiology in 1948. The Journal of 
Comparative Physiology was first published 


259 


as Zeitschrift fiir Vergleichende Physiologie 
in 1924 and changed titles in 1972. Physi- 
ological Zoology started in 1928 and the 
early years focused primarily upon inver- 
tebrates; more vertebrate papers began to 
appear in the 1960s-1970s. As will be dis- 
cussed below, comparative physiology be- 
gan in earnest in the 1940s and truly flour- 
ished throughout the 1950s. Thus, many new 
specialty journals appeared after that (e.g., 
Comparative Biochemistry and Physiology, 
1960; Journal of Thermal Biology, 1975). 
The later application of physiology to eco- 
logical questions was covered in ecological 
journals (e.g., Ecology, Oikos, Oecologica, 
and Functional Ecology). 


Review Methods 


In order to evaluate how thought and sub- 
jects of investigation in physiology have 
changed over the 75 years since formation 
of the ASM, we reviewed several sources. 
There is a very informative history of the 
American Physiological Society that covers 
some aspects of this history (Brobeck et al., 
1987). In addition, we reviewed articles in 
the Journal of Mammalogy and picked two 
review journals (Physiological Reviews, be- 
gun in 1921, and Annual Review of Physi- 
ology, begun in 1939) to give us some idea 
of what the leaders in the field during par- 
ticular times thought were important topics 
to be reviewed and how authors approached 
and reviewed those topics. To focus on 
comparative physiology, we picked several 
journals in addition to the Journal of Mam- 
malogy to evaluate for topical coverage— 
Journal of Cellular and Comparative Phys- 
iology, Physiological Zoology, and Journal 
of Comparative Physiology. We divided 
physiological topics into a number of cat- 
egories (Table 1) following evaluation of the 
table of contents for chapter headings in six 
current physiology texts and placed papers 
into one of these categories. We realize this 
is not perfect but we found the list too large 
to be manageable otherwise. This necessar- 


260 


ily led to some groupings; for example, the 
heading water balance includes some papers 
on ion balance, but if the paper was decid- 
edly focused on the role of the kidney, then 
we placed it in kidney. Energetics includes 
papers on metabolism. Many papers cov- 
ered temperature regulation and energetics 
(and may have touched on evaporation as 
a thermoregulatory mechanism). Here we 
tried to decide what the major focus of the 
paper was and categorize it accordingly. We 
started out with digestion as a single topic 
and found that we needed to group nutrition 
with it since they were frequently related. 
Early papers on the nervous system simply 
described brain electrical activity or brain 
waves and later papers emphasized cellular 
mechanisms related to neurotransmitters 
and ion channel function. Here, again, we 
needed to decide whether the focus of the 
paper was neuronal or some cellular func- 
tion using neural tissue and that decided the 
category for us. 


1920-1940 


During this period physiology bloomed 
as a discipline in North America. Several 
early physiologists who had been building 
their own apparatus for experimentation 
developed companies to produce that ap- 
paratus (e.g., W. T. Porter—the Harvard 
Apparatus Company; Ellen Robinson, who 
married Albert Grass, and founded the Grass 
Instrument Company) making expansion of 
certain fields possible in many labs simul- 
taneously (Appel, 1987c). Much of the re- 
search during this period emphasized san- 
itation, temperature regulation, and 
nutrition as they related to human health. 
Physiological Reviews was initiated in 1921 
and many early articles during the 1920s 
related to topics such as levels of blood com- 
ponents (e.g., sugar), vitamin research, di- 
gestion, and absorption of food. Much of 
this work had its genesis and direction fo- 
cused from needs of World War I. The san- 


WUNDER AND FLORANT 


TABLE |1.— Categories employed for grouping 
of physiological topics. 


Temperature Regulation 

Energetics 

Hibernation 

Lipids 

Water Balance (including ion regulation) 
Urine 

Kidney 

Evaporation 

Endocrinology 

Reproduction (including the endocrinology thereof) 
Digestion/Nutrition 
Blood/Heart/Circulation 
Respiration/Lung Function 

CNS (neurophysiology in general) 
Muscle 

Cell (including molecular) 

Other 


itation and nutritional requirements of ar- 
mies directed many lines of research. In 
1924, Boothby and Sandiford wrote a com- 
prehensive paper in Physiological Reviews 
on basal metabolism in mammals empha- 
sizing humans, but citing the vast literature 
(> 100 articles) by Benedict on the subject. 
In 1922, at meetings of the American Phys- 
iological Society, F. G. Banting and C. H. 
Best presented their Nobel-winning re- 
search on the role of insulin in regulation 
of blood sugar. Work on the definition, role, 
and function of endocrine glands flourished. 
Most of the studies on such systems con- 
sisted of removing the gland to observe re- 
sults and infer function from the resulting 
change in function. 

In the physiological literature there was 
little emphasis on comparative physiology 
or using forms other than humans or ani- 
mals as models for humans. Because there 
was so little emphasis on comparative as- 
pects of physiology, very little work was re- 
ported on wild vertebrates in journals such 
as Journal of Mammalogy. In the Journal 
there are only about eight papers in the 1920s 
that could be defined generally as physio- 
logical, and five of those were on reproduc- 
tion. Most of the work on reproduction em- 
phasized description of reproductive cycles 


PAY STOLOGY 261 


and timing of reproduction, which might be 
considered population biology today. Things 
expanded somewhat in the 1930s with ca. 
35-40 papers appearing in the Journal. Most 
(16) of these concerned food habits with a 
little work on the actual physiology (digest- 
ibility) of digestion, but most simply listed 
food habits. Reproduction was still a strong 
field with 10 papers. Many articles de- 
scribed reproduction cycles, but a few pur- 
sued questions related to development. Be- 
nedict’s study on the physiology of the 
elephant appeared in 1938, and there was 
early work on the role of the pineal in pho- 
toperiodic mechanisms. F. G. Hall’s early 
work on adaptations to altitude was pub- 
lished in 1937, and there were ca. five pa- 
pers on respiration in porpoises, as inves- 
tigators pushed them as a novel system to 
understand respiration. Early work describ- 
ing electrical activity and effects of shock 
on brains of beaver and kangaroo rats ap- 
peared. Studies of temperature regulation 
patterns and metabolism in humans were 
expanding due to interests in nutrition fol- 
lowing World War I, and four papers on 
temperature regulation patterns and novel 
mechanisms (e.g., torpor) were published in 
the Journal of Mammalogy. Two of the pa- 
pers were descriptions of low body temper- 
atures in sloths by R. K. Enders from 
Swarthmore and one paper was on hiber- 
nation. As early as 1935, A. Brazier Howell 
and I. Gersh wrote a short paper indicating 
that kangaroo rats (given as Dipodomys mo- 
havensis) could exist without free drinking 
water and speculated on the role of succu- 
lent vegetation and metabolic water in their 
adaptations. They even performed histo- 
logical studies on the kidneys and speculat- 
ed about reabsorption. 


1940s 


The subject of comparative physiology 
expanded greatly in the 1940s. The early 
part of the decade saw a focus of physiology 
on national needs associated with the war 


efforts of World War II. This single event 
greatly expanded opportunity, support, and 
questions about physiology more than any 
other factor up to this time. With the de- 
velopment of aviation, new questions about 
adjustment to altitude arose. With more 
emphasis on submarine warfare, the Navy 
(through the Office of Naval Research) be- 
came interested in torpor (hence hiberna- 
tion) as a possible way of maintaining crews 
underwater for long periods of time. The 
“off-duty” crew would use less oxygen in a 
reduced metabolic state, allowing subma- 
rines to remain under the sea for longer 
times. Thus, money became available for 
studies of metabolism, temperature regu- 
lation, and water balance (how could troops 
better adjust to desert or jungle condi- 
tions?). The standard topics of nutrition, di- 
gestion, circulation (especially hemody- 
namic shock), endocrine function, and 
neural control flourished with new and bet- 
ter equipment and support. These latter ar- 
eas took on a more mechanistic flavor fol- 
lowing the earlier descriptive work. 

Two groups greatly spurred the work in 
comparative physiology. Laurence Irving 
was trying to build a program at Swarth- 
more and brought to the U.S. several stu- 
dents of the renowned Danish physiologist, 
August Krogh. In 1939, he helped Per Scho- 
lander obtain a Fulbright Fellowship and 
persuaded him to come to Swarthmore 
(Scholander, 1978). In 1946 Irving brought 
Bodil (Krogh’s daughter) and Knut Schmidt- 
Nielsen to Swarthmore. Here, Scholander, 
always clever at designing and building 
equipment, developed the “‘Scholander sy- 
ringe,”’ which allowed gas analysis in very 
small samples of fluid (a “micro” Van Slyke 
apparatus). In 1947 Irving suggested that 
the group go to southern Arizona to study 
water metabolism in kangaroo rats. From 
that experience Bodil Schmidt-Nielsen went 
on to a distinguished career studying the 
physiology of the kidney and its role (along 
with other excretory organs) in regulating 
osmolality and volume of extra- and intra- 
cellular compartments. She served as Pres- 


262 WUNDER AND FLORANT 


ident of the American Physiology Society 
from 1975 to 1976 (Brobeck, 1987). Knut 
Schmidt-Nielsen continued his outstanding 
career working on adaptations of a variety 
of animals to aridity focusing on tempera- 
ture regulation and water balance. Irving 
himself founded and was the first Director 
of the Institute of Arctic Biology at the Uni- 
versity of Alaska following a career focused 
on study of adaptations of mammals and 
birds to Arctic conditions. He became in- 
terested in the Arctic at the urging of Scho- 
lander, who previously had worked in 
Greenland doing botanical studies as a stu- 
dent. Scholander later became interested in 
questions related to diving physiology, 
which is what led him to Irving’s lab (Scho- 
lander, 1978). 

These collaborations resulted in the early 
classical papers describing how Arctic birds 
and mammals are better insulated than 
tropical forms, yet their patterns of metab- 
olism and temperature regulation are sim- 
ilar (Scholander et al., 1950a, 1950 5, 1950c). 
At this time such studies consisted primar- 
ily of exposing animals to different ambient 
temperatures and measuring their body 
temperatures. However, Scholander also 
developed an experimental way to measure 
insulation via heat flow through skins using 
a hot plate as heat source. Metabolism was 
tediously measured using a spirometer and 
taking gas samples periodically for analysis 
with the Scholander syringe or Haldane ap- 
paratus. The primary focus was the use of 
comparative material to ask questions about 
how animals might be best adapted to par- 
ticular, stressful environmental conditions. 

About this same time (1945-1948), a 
group at Harvard consisting of O. P. Pear- 
son, George Bartholomew, Peter Morrison, 
and G. E. Folk, Jr. was finishing their Ph.D.s 
and developed similar interests in asking 
questions about how animals were adapted 
to specific environmental stressors. Togeth- 
er with the Swarthmore group (Morrison 
went to Swarthmore before moving to the 
University of Wisconsin) they and their stu- 
dents were a dominant force in comparative 


physiology (especially mammals) for the 
next 30-40 years. 

Morrison began studying temperature 
regulation of Central American mammals 
in the 1940s and went to Swarthmore where 
he interacted some with R. K. Enders. He 
quickly moved to the University of Wis- 
consin and later went to Alaska where he 
became Director of the Institute of Arctic 
Biology following Larry Irving in the late 
1960s. In the 1940s, the emphasis was still 
upon measurement of temperature response 
patterns to varying environmental temper- 
atures. Some labs began to look at mecha- 
nisms by which heat was conserved or lost 
(see Scholander et al., 1950a, 1950, 1950c), 
but the methods were difficult and tedious. 
Pearson published some of the earliest pa- 
pers on metabolism and temperature reg- 
ulation of shrews, as did Morrison. Both 
were intrigued with the observation that the 
animals were reported to eat prodigious 
quantities and, hence, should have high me- 
tabolism. With small size they should have 
a large surface area for heat loss relative to 
their mass for active metabolism. Moving 
to southern California (UCLA), Bartholo- 
mew began work on desert forms using both 
birds and mammals, but emphasizing birds 
for his early work. However, in the 1950s 
he made trips to Alaska to study marine 
mammals. After some early reports on tem- 
perature and respiratory rates in these forms 
he emphasized study of behavior, a then 
emerging field. His mammal work, together 
with that of his students, focused on water 
balance and temperature regulation with at- 
tention to torpor as an energy-saving ad- 
aptation. 

The field of temperature regulation was 
changing in the physiological arena also. In 
the early 1940s, the Annual Review of Phys- 
iology had a section entitled ““Temperature 
Regulation,” but starting in 1943 it was 
changed to “Heat and Cold” and in 1948 
an entire paper on factors influencing sweat- 
ing was published. Most of the studies then 
looked at factors that influenced body tem- 
perature, such as ambient temperature, ra- 


PHYSIOLOGY 263 


% OF TOTAL 


ev oe ye epee nn er-. 
se Sate eve scaessetrasss 
Spemcs tems eles Cle em ca Ss Mol lacy a 
Sear seat rocvt fens 3 (o) 
setts 42SESSF=SZECE 
toe = 5s 
2c 3 br S oc 2) o.8 0 
me Ws 2 > Oe lt stgz 
eo oI WM Ueese ac 

a 4 ego & a N 

ae = 
E a oo 26 
Ay My eo OU 
oo mm 

i=) 


Fic. 1.—Percentage of papers published on 
various topics in the Annual Review of Physiol- 
ogy comparing the decades of the 1940s, 1950s, 
and 1960s. 


diant heat loads, and effects of different 
clothing and activity levels on body tem- 
perature. These, of course, were in response 
to needs associated with World War II. The 
mid-1940s also produced papers on applied 
aviation medicine and anoxia in aviation. 


1950s 


The impetus in comparative physiology 
begun in the 1940s continued into the 1950s. 
Much comparative work was done on water 
balance and thermoregulation of wild forms 
emphasizing Arctic and desert forms, with 
some studies on adaptation to high altitude. 
The general paradigm was to study an an- 
imal best adapted to deal with a particular 
environmental stressor. This work was a 
spin-off from the interest in stress physiol- 
ogy brought on by World War II and the 
available funding associated with that. In 
the “‘mechanistic”’ physiological literature 
there was a subtle shift in areas of emphasis. 
Figure | shows the relative percentage of 
papers in different subject areas published 
in the Annual Review of Physiology for the 
period 1940-1960 by decade. Temperature 


regulation made up ca. 5% of the literature 
in the 1940s and 1950s, then declined, while 
water balance (including ion balance) in- 
creased from 2% to 5% of the literature. The 
big increases were in work on endocrine sys- 
tems and shifts to cellular approaches to 
mechanism. The dramatic change in en- 
docrine coverage was a shift to specific gland 
function, their hormones, and mode of ac- 
tion versus “the endocrine system” dis- 
cussed in earlier reviews. Early papers fo- 
cused on the pituitary and its role in 
reproduction, along with thyroid and effects 
on growth and metabolism. These presaged 
the work on cellular mode of action that 
later were emphasized in the 1960s. 

The comparative approach shows dra- 
matically in the number and sorts of reviews 
written from the mid-1950s. Before this 
time, most studies emphasized the rat or 
humans, but in 1953 many papers with a 
comparative theme appeared in Annual Re- 
view. Starting in 1953, there was an article 
on comparative physiology of invertebrate 
muscle. From then to 1960 each issue had 
at least one paper with a definite compar- 
ative approach (e.g., sense organs, respira- 
tion in invertebrates, nutrition and feeding 
in vertebrates) with a comprehensive re- 
view by F. E. J. Fry on temperature com- 
pensation mechanisms for metabolism in 
poikilotherms in 1958. This followed a re- 
view on energetics in 1956 by Max Kleiber. 
In 1957 Kayser, who had worked on the 
subject since the 1930s, presented a defin- 
itive review on hibernation. Most of the 
work on hibernation to that point was de- 
scriptive regarding torpor patterns and body 
temperature shifts. Some early workers (e.g., 
Benedict and Lee, 1938; Lyman, 1948) had 
looked at metabolism in hibernators, but it 
was not until the 1950s and the develop- 
ment of the paramagnetic oxygen analyzer 
that such studies increased greatly in num- 
ber. Charles P. Lyman and others reported 
on studies of nerve conduction, electrical 
activity of the cerebral cortex, circulation, 
and function of endocrine glands of hiber- 
nators. 


264 


The early work on water balance was ex- 
panded to include wild forms and total wa- 
ter budgets during the 1950s. Early work 
had focused on movement of water through 
skin, sweating mechanisms, and amounts of 
water needed by animals (Adolph and Dill, 
1938: Dill et al., 1933; Tennent, 1946; Vor- 
hies, 1945), and on structure and concen- 
trating capacity of the kidney (Sperber, 
1944). Although Howell and Gersh (1935 
and see above) early pointed out that kan- 
garoo rats needed little or no water, and the 
Schmidt-Nielsens expanded upon that in the 
late 1940s, 1t was not until the 1950s that 
studies focused on compartmentalizing wa- 
ter balance. Bodil and Knut Schmidt-Niel- 
sen (see review 1n 1952) presented a ““com- 
plete’’ account of water balance for “‘the” 
kangaroo rat and reported a value for pul- 
monary water loss (actually evaporative wa- 
ter loss). Later in the decade, Robert M. 
Chew expanded the work and included many 
other desert rodents, as did Bartholomew 
and his students (Dawson, Hudson, 
MacMillen). 


1960s 


The comparative trend begun in the late 
1940s expanded even more in the 1960s. 
Throughout the decade each volume of the 
Annual Review of Physiology had at least 
one article with a comparative approach, 
starting with Clyde Manwell’s paper on 
blood pigments in 1960. An article by Don 
Farner on photoperiodic mechanisms in 
birds appeared in 1961 along with Florey’s 
paper on comparative transmitter sub- 
stances in neurophysiology (always a large 
topic for review). Vernberg reviewed what 
was known about adjustment to different 
geographic regions with a 1962 paper on 
latitudinal effects on physiological proper- 
ties of populations (most of his work was 
on marine invertebrates, but it stimulated 
interest in vertebrates, including mammals) 
and it introduced a new technique—trans- 
plantation, which was used later in the de- 


WUNDER AND FLORANT 


% OF TOTAL 


co Ce 
sv Es QeevesB®soostsecugys 2 
-— SS =< = = 2 So 2 ie Soir 
ao = 2925 5 = So Soa = SRE is) 
sos 3S ok se a 
> } --} “ao lkse ec = = 
wm 2 5 ares ‘Sone 
ee . avnZz Ss S25 2 
= & i 2 > 2 a~ S =a 2 
& r. ~ 

P = as <3) fe = = = EG 
= = es) os a2 
E = eo 
= =D = 

= 

a 


Fic. 2.—Percentage of papers published on 
various topics in the Annual Review of Physiol- 
ogy comparing the decades of the 1960s, 1970s, 
and 1980s. 


cade by Ray Hock and others to study ad- 
aptation to altitude in deer mice. Articles 
on navigation by animals, comparative 
physiology of nutrition of vertebrates and 
invertebrates, and hormones in fish (by 
Hoar) added to J. Aschoff’s classic paper on 
diurnal rhythms. Water and ion balance be- 
came more important topics than in the past. 
Bodil Schmidt-Nielsen reviewed mecha- 
nisms by which invertebrates dilute urine 
in the Annual Review of Physiology in 1963, 
a comparative paper on invertebrate excre- 
tory organs appeared in 1967, and G. Parry 
reviewed osmotic and ionic regulation (sys- 
tem level studies) in 1968. In 1961 Max 
Kleiber again reviewed energetics, empha- 
sizing cellular energy transfer and metabolic 
control mechanisms over organismal met- 
abolic rates, size, and ties to temperature 
regulation and food as in his 1956 review. 
Studies of thermoregulatory patterns and 
mechanisms of thermoregulation under 
stress conditions for wild animals became 
more common. H. T. Hammel reviewed this 
topic in a 1968 paper in Annual Review of 
Physiology. In 1964 the first, and only, 
Handbook of Physiology: Adaptation to the 
Environment was published by the Ameri- 


PHYSIOLOGY 265 


% OF TOTAL 


Hy 

i 

Hi 

i 

i 

i 

t 

' 

z 

' 

ig 

3 

Fy 

i 

g 

i 
= 
7) 
= 
=) 


Energetic: ae 
Blood/Heart = 


Water Balance —xaaeessmen 


Temp. Regulation -==essssessssnmnsenss 


A i fe i 
o.2 eon GH Ge ww won Ya 
ou ee eS oe cwo wo 
a=, =| = Sacto Saou? 
sa oc sls 
eSg5Sfeetsessi 
= 
be “Ms Eas ete 
o Qerosag oes 
2 ove Zs cate 
= = CO 1a, 3s 4 
{ao} MuUYE ES a 
=& oo _ N 
a ial a7 
+ v0 
oD i= 4 
i=) 


Fic. 3.—Percentage of papers published on 
various topics in Physiological Zoology compar- 
ing the decades of the 1960s, 1970s, and 1980s. 


can Physiological Society giving a state-of- 
the-art coverage for comparative physiol- 
ogy and human adjustment to stress brought 
on by environmental variables. In 1962 the 
first International Symposium of Hiberna- 
tion and Cold Physiology was held in As- 
pen, Colorado (the 9th meeting was held in 
1993 at Crested Butte, Colorado). Fig. 1 
suggests that this topic of thermoregulation 
declined in coverage from the 1940s to the 
1960s. Generally that is true with a reduc- 
tion in human “stress” work after World 
War II, but Fig. 2 shows an increase in the 
1970s and an increase in studies of ener- 
getics. The increase in 1970 reflects more 
papers on comparative topics (more wild 
species) and the increase in papers on en- 
ergetics reflects an increase in papers inves- 
tigating the mechanisms of thermoregula- 
tion and the role of metabolism in those. 
As noted, frequently it was difficult to sep- 
arate a paper into one or the other category 
of thermoregulation or energetics. In the 
1940s and early 1950s that was not so dif- 
ficult because energetics papers usually re- 
lated to total energy turnover, or need, and 
related more to nutrition and body size ef- 
fects than to mechanisms of thermoregu- 


% OF PHYSIOLOGY 


Energetics Ssh 
Lipids Ss 
Blood/Heart BSSSsss 
Other SSS 


4% A ee) ie 4 | 
e ec o > & > na oD 
° } = o © &® © = i] 
= = Se ae oo Oa a) 3 
a a = SZ ses 6 ¥ Go =] 
-_ GI be 3 jee} ~ 
S E aM 6 £ 5 ec 
a cr) > fo & } a S 
2 - ec a yy pe = = 
mo — Se ot > ° a s = 
pa a -. wm Dv ve 
: = =) oc @ a a= 
a w a 
i= 7) 
oO v 
a i-4 


Digestion/Nutrition PSS Sieve 


Fic. 4.—Percentage of papers published on 
various topics in physiology among the total pa- 
pers on physiology published in the Journal of 
Mammalogy comparing the decades of the 1960s, 
1970s, and 1980s. 


lation. Also many new journals appeared 
during this decade allowing more places for 
authors to submit work on these topics (e.g., 
Journal of Comparative Biochemistry and 
Physiology, Journal of Thermal Biology) and 
coverage of these topics in journals such as 
Physiological Zoology (Fig. 3) increased. 
The increased coverage was reflected in 
the Journal of Mammalogy (Fig. 4). Over 
20% of the papers in physiology concerned 
thermoregulation. Papers on energetics in- 
creased from 5% of all physiological cov- 
erage in the 1960s to 12% in the 1970s and 
20% in the 1980s. During the 1960s inves- 
tigators revived the paradigm of Justus von 
Leibig (from the 1800s) that physiology sets 
limits to distribution patterns. Thus, com- 
parative physiology evolved into the fields 
of comparative (mechanistic) physiology, 
which selected organisms because they might 
best show the mechanisms at work in an 
organ system under stress, and physiologi- 
cal ecology, a new field developed to inves- 
tigate how animal function and distribution 
might be restricted through physiological 
limitation to the environment. Thus, in 1963 
Brian McNab’s (a student of Peter Morri- 


266 WUNDER AND FLORANT 


son) paper on the relation between home 
range and energy needs of mammals ap- 
peared. In the Journal of Mammalogy many 
papers published during this decade took on 
the emphasis of the interaction between 
physiology and “limitations.” We find E. 
W. Jameson, Jr., writing about body mass 
effects and hibernation (how fat must in- 
dividuals be before they can enter torpor?) 
and L. Getz investigating salt tolerance and 
aridity tolerance in voles and their relation 
to competition and habitat use and selec- 
tion. Negus and Pinter published one of their 
first papers of a 15—20-year search for plant 
compounds that affect reproduction in voles 
(Negus was later joined by Berger in this 
work, which culminated in a 1981 paper in 
Science [Berger et al.] identifying the com- 
pound and a 1987 review of the topic). Fur- 
ther, Christian and Davis wrote about ad- 
renal function, reproduction, stress, and vole 
cycles in Microtus pennsylvanicus. No mat- 
ter what the physiological system (water 
balance-kidney; stress-adrenal; reproduc- 
tion-endocrine/gonads), the paradigm for 
questions in this decade became limitation 
on some ecological parameter. 


1970s 


As was the case in earlier decades, the 
strong topics for coverage in the physiolog- 
ical literature during the 1970s were endo- 
crinology, circulation and respiration, and 
topics in neurobiology. The urinary system 
and kidney received less coverage than be- 
fore. A look at Fig. 2 suggests that cell phys- 
iology received less attention in the 1970s 
than in the 1960s. However, that is mis- 
leading because much of the approach in 
endocrinology and neurobiology was mo- 
lecular and cellular, with work on hormone 
receptors and mode of receptor function re- 
ceiving much attention. In neurobiology, our 
understanding of impulse transmission and 
the cellular basis of nerve synapses and 
nerve/muscle interaction was being eluci- 
dated. 


In comparative physiology there was a 
continuation of the ecological emphasis be- 
gun in the 1960s. Thermoregulation (in- 
cluding hibernation), energetics, water bal- 
ance and kidney function, and reproduction 
continued strong or increased in coverage. 
At this time more papers on vertebrates in 
general, and mammals in particular, ap- 
peared in Physiological Zoology. Prior to 
1960 much of the coverage in this journal 
was on invertebrates. As can be seen in Fig. 
3, the topics listed above increased during 
the 1970s, just as they did among physiol- 
ogy papers published in the Journal of Mam- 
malogy (Fig. 4). Thermoregulation, ener- 
getics, and water balance were all topics that 
expanded in coverage during this decade 
(Fig. 4). However, there were changes in 
approach and paradigms within which these 
physiological data were interpreted. 

Within the field of thermoregulation and 
energetics, papers took on a new level of 
sophistication. Instead of just documenting 
more species for patterns of ability to ther- 
moregulate in extreme environments (e.g., 
hot or cold, dry or wet), or evaluating the 
effects of body mass, there was a shift to- 
ward studying effects of, and cost for, var- 
ious activities such as locomotion or repro- 
duction, and an incorporation of broader 
factors influencing thermoregulation. The 
field of biophysics came of age following 
publication of David Gates’ Energy Ex- 
change in the Biosphere a decade earlier 
(Gates, 1962). Aaron Moen applied these 
techniques to deer and Heller and Gates 
used them to describe thermal physiology 
as a factor influencing chipmunk distribu- 
tion along an altitudinal gradient on the 
eastern slope of the Sierra Nevada moun- 
tains in California. Here was use of physi- 
ology coupled with behavior to describe 
mechanisms of competitive exclusion for 
these distribution patterns. There was also 
increased emphasis on mechanisms of ther- 
moregulation. Work on brown adipose tis- 
sue as a means of warming small mammals 
(first described as a heat generating tissue 
by Smith and Horwitz in 1969) took on 


PHYSIOLOGY 267 


more importance and was investigated by 
Heldmaier in Germany and Lynch and 
Wunder in the U.S. The role of various 
structures and mechanisms to modulate en- 
ergy exchange with the environment (using 
biophysics and heat transfer concepts and 
equations) became more in vogue for study 
(e.g., Cena and Clark, 1973; Heller, 1972). 

Energetics studies were applied more at a 
population level (following the lead of 
McNab, 1963) in an attempt to explain a 
variety of processes (e.g., home range sizes, 
reproductive costs, population growth). En- 
ergy became a currency to be used for de- 
scribing behavior and to try to predict the 
consequences of population processes. The 
Polish school, led by Ladd Grodzinski, was 
quite active during this time writing on en- 
ergetics, and reproduction and population 
growth in a variety of mammals varying in 
size from voles to roe deer. Grodzinski and 
Wunder (1975) reviewed the topic of en- 
ergetics in small mammals. McNab contin- 
ued in the vein of his 1960s work using 
energetics to discuss the distribution of 
vampire bats and other mammals. He then 
went on to develop ideas about how life 
history traits, such as food habits and body 
mass, might influence energetics in mam- 
mals. 

There was also a shift during this decade 
to study energetics of animals in the field. 
While mechanistic studies still used meta- 
bolic rates measured as steady states during 
rest or some specific activity with oxygen 
analyzers in the lab, there was a new isotopic 
technique introduced to study integrated 
metabolism in the field. Much of the con- 
ceptual development and early validation 
work on these techniques to study metab- 
olism and water turnover was done by Lif- 
son in the 1950s and 1960s. However, the 
technique was expensive and required spe- 
cial equipment. Thus, few studies were un- 
dertaken until Ken Nagy, at UCLA, ac- 
quired access to the expensive isotopes and 
equipment to measure them. Much collab- 
orative work was done, culminating in his 
review paper a decade later (Nagy, 1987) 


summarizing and scaling the allometric re- 
lationships of field metabolism in birds and 
mammals. 

During this decade it was also realized 
that energy, per se, may not be the only 
limitation or major currency for evaluating 
performance of mammals in the field, and 
nutritional ecology took on a new impor- 
tance. George Batzli became a dominant fig- 
ure studying small herbivores. Realizing that 
energy is important to the lives of animals, 
these studies suggest that certain secondary 
chemicals in food may influence energy 
availability to mammals and some energy 
sources may have limited availability. For 
that reason most of the studies focused on 
herbivores, because they eat a high energy 
density food (plants) in which the energy is 
not readily available to mammals because 
it is tied up as cellulose and hemicellulose 
and vertebrates lack the enzymes necessary 
to break these down. Thus, digestion and 
digestive processes become critical to make 
these foods available for herbivores. There 
was a tremendous literature available from 
animal science where such processes had 
been studied for decades to enhance food 
production (e.g., Kleiber, Baldwin, Van 
Soest), but, with few exceptions, most stud- 
ies of wild forms did not reference this lit- 
erature to any great extent. 

Studies of water balance still focused pri- 
marily upon animals living in arid regions 
(work by MacMillen and Hinds). Like the 
studies on energetics, however, there was a 
new push to learn how animals were truly 
challenged in the wild and, hence, radioiso- 
topes were introduced to study water turn- 
over in the field (see papers in the Journal 
of Mammalogy by Nagy and by Bradford). 
In the latter part of the decade and into the 
early 1980s, Christian (1979, 1980) inves- 
tigated the role of water in reproduction, 
demographics, and habitat use by small des- 
ert rodents. Previously there was specula- 
tion that moisture may be important in these 
processes, but no one had sorted out mois- 
ture from energetics despite the fact that 
most desert forms obtain their moisture 


268 WUNDER AND FLORANT 


from their food. Christian simply intro- 
duced small watering stations in the field 
and found that the reproductive season was 
prolonged for some species, some actually 
showed numerical population increases, and 
there were habitat shifts to use of drier, more 
open habitat if moisture was present. Inter- 
estingly, little has been done with this tech- 
nique in application to other species or hab- 
itats. 

Overall during this decade there was a 
strong emphasis on environment factors and 
attempts to see how animals actually per- 
formed in the field. Tied to this was an in- 
terest in how performance of mammals 
shifts seasonally, regardless of whether one 
was studying temperature regulation, en- 
ergetics, water balance, or reproduction. 


1980s 


In the general physiological literature this 
was a time when many new, specialty jour- 
nals were started or expanded having been 
initiated during the 1970s. Thus, papers in 
many fields were being shifted to these spe- 
cialty journals and we found analysis of 
trends in a discipline harder to document 
using the standard review journals that we 
had used up to this decade. Endocrinology 
continued as a strong field with much more 
emphasis on molecular mechanism and ties 
to genetic control than had been the case 
earlier. Cardiac and circulatory function re- 
mained a strong area of research, with ca. 
25% of the papers in Physiological Review 
being published in this area (Fig. 2). Most 
topics had molecular and cellular orienta- 
tion and the general topic itself increased in 
coverage in Physiological Review from 
<10% to >15% of total papers. 

Physiological Zoology changed editors in 
the 1970s from T. Park, who had been in- 
volved with the journal since its early days, 
to C. L. Prosser and J. E. Heath. Thus, many 
more papers on vertebrates, and mammals 
in particular, were published in the 1980s. 


The area of emphasis was energetics and, 
secondarily, thermoregulation (Fig. 3). 
Within thermoregulation there was a resur- 
gence of interest in hibernation and torpor. 
This was a very topical subject in the 1950s 
and early 1960s, but seemed to lack focus 
in the late 1960s to late 1970s, except for 
papers on cellular mechanisms and tissue 
tolerance to cold. However, in the late 1970s 
there was renewed interest at the organismal 
level stimulated by work showing that the 
sorts of fuels burned during torpor may in- 
fluence lengths of torpor bouts and that dif- 
ferent kinds of fats (saturated versus poly- 
unsaturated) might be used differentially 
during torpor periods. Thus, mammals may 
need to seek certain nutrients prior to hi- 
bernation. Kenagy and Geiser, Florant, and 
later Frank added to this area. French de- 
veloped insight into the effects of body size 
on torpor bout lengths and optimal tem- 
peratures for torpidity. Many of the ener- 
getics papers of the decade relate to other 
aspects of a species’ biology, such as pop- 
ulation processes and costs for various be- 
haviors or reproduction, adding to the in- 
formation started in the 1970s. This was 
also a time when the field shifted to examine 
how body size (mass) influenced energetics 
and many other functions of organisms. 
Three major books on allometry were pub- 
lished emphasizing how body mass con- 
strains and allows organisms (mammals in 
particular) to function (Calder, 1984; Pe- 
ters, 1983; Schmidt-Nielsen, 1984). 

Water balance of mammals became a top- 
ic of less emphasis, but some fine work on 
total water budgets and their significance for 
distribution limits or function was pub- 
lished by MacMillen, and MacMillen and 
Hinds. This work grew from the early stud- 
ies of Bartholomew and Chew first pub- 
lished in the 1950s and 1960s. In these 
papers the relationship of water to ther- 
moregulation was stressed as much as the 
use of water for general life processes and 
as a means of effecting ion balance. 

Within the Journal of Mammalogy, pa- 
pers on thermoregulation remained strong 


PHYSIOLOGY 269 


at >15% of the total papers in physiology 
published (Fig. 4). Energetics received re- 
newed interest for investigation, increasing 
from around 10% to over 20% of the total 
physiology papers published. Reproduction 
remained steady at ca. 12%. The topic that 
truly took on a new interest was digestive 
biology and nutrition (Fig. 4). Many inves- 
tigators began to study how the process of 
digestion might limit energy acquisition by 
mammals, especially small herbivores, and 
how nutrition might influence herbivore- 
plant interactions and animal performance. 

Those studying thermoregulation contin- 
ued the trends of the 1970s, applying this 
theme to limits on distribution and perfor- 
mance in mammals. Many studies linked 
thermoregulation to energetics so the shift 
to more energetics papers was, in part, a 
slightly different emphasis on thermoregu- 
lation. Many studies had, as part of their 
focus, adjustment to different seasons (work 
by Wunder and Merritt and colleagues), and 
mechanisms for those shifts (work by Hill, 
Kenagy, MacMillen, Wunder, French, Har- 
low, Dertin, McNab, and Cranford). 

As mentioned above, along with these 
studies of energetics and the theme of limits 
to distribution and performance, many in- 
vestigators began to study how energy 
sources and allocation pathways were fueled 
by animals. That is, what were their foods 
and how were nutrients obtained? George 
Batzli and his students had studied such 
questions for about two decades and, in the 
1980s, began to look more closely at the role 
of secondary chemicals in food, in addition 
to rate processes and digestive efficiencies. 
Wunder and students showed that small 
herbivores (e.g., voles) could change gut size 
to better or more quickly process food, and 
many related papers followed. Two major 
texts on the topics of nutrition appeared, in 
addition to many symposia volumes, es- 
pecially on ruminant herbivores. Peter Van 
Soest’s book, Nutritional Ecology of the Ru- 
minant (1982), set the stage for Charlie 
Robbins’ book, Wildlife Feeding and Nu- 
trition (1983). Both are used as a basis for 


posing questions about how various mam- 
mals adjust to novel foods or environments 
compared to more studied forms. Recent 
work is beginning to focus on limits to en- 
ergy processing and the trade-off of the roles 
of digestion and assimilation with tissue uti- 
lization of substrates, or behavior of feed- 
ing. Over the next decade, we hope there 
will be a more complete understanding of 
how mammals, especially herbivores, uti- 
lize the myriad of plants available to them, 
and how plant—animal interactions are in- 
fluenced by physical and biological factors 
such as thermoregulation, energy needs, and 
nutrient needs for reproduction. 


Epilogue 


We have attempted to give a brief over- 
view of how physiology in general, but es- 
pecially comparative physiology of wild 
mammals, has shifted, waxed, and waned 
over the past 75 years as the ASM has grown. 
We suspect that the recent fervor for cell 
and molecular approaches will stimulate an 
understanding of not only mechanisms of 
process, but also how those processes relate 
to the fundamental biology (life histories) 
of the wild mammals possessing them. Such 
knowledge will be useful not only for un- 
derstanding ourselves and our functions, but 
also how mammals function in ecosystems, 
and how they might adjust to changes in 
those ecosystems as we witness climatic and 
other environmental changes. 


Literature Cited 


Apo pH, E. F. 1987. Physiology flourishes in Amer- 
ica. Pp. 1-10, in History of the American Physio- 
logical Society (J. R. Brobeck, O. E. Reynolds, and 
T. A. Appel, eds.). Waverly Press, Baltimore, Mary- 
land, 533 pp. 

Apo pH, E. F., AND D. B. Ditt. 1938. Observations 
on water metabolism in the desert. American Journal 
of Physiology, 123:369-372. 

AppeL, T. A. 1987a. Founding. Pp. 11-29, in History 
of the American Physiological Society (J. R. Bro- 
beck, O. E. Reynolds, and T. A. Appel, eds.). Waver- 
ly Press, Baltimore, Maryland, 533 pp. 


270 WUNDER AND FLORANT 


. 19876. First quarter century, 1887-1912. Pp. 
31-62, in History of the American Physiological So- 
ciety (J. R. Brobeck, O. E. Reynolds, and T. A. Ap- 
pel, eds.). Waverly Press, Baltimore, Maryland, 533 


. 1987c. Second quarter century, 1913-1937. 
Pp. 63-96, in History of the American Physiological 
Society (J. R. Brobeck, O. E. Reynolds, and T. A. 
Appel, eds.). Waverly Press, Baltimore, Maryland, 
533 pp. 

BENEDICT, F. G., AND R. C. LEE. 1938. Hibernation 
and marmot physiology. Carnegie Institution of 
Washington Publication, 497:1-239. 

BERGER, P. J., N. C. NEGus, E. H. SANDERS, AND P. D. 
GARDNER. 1981. Chemical triggering of reproduc- 
tion in Microtus montanus. Science, 214:69-70. 

Broseck, J.R. 1987. Presidents, 1963-1987. Pp. 177- 
280, in History of the American Physiological So- 
ciety (J. R. Brobeck, O. E. Reynolds, and T. A. Ap- 
pel, eds.). Waverly Press, Baltimore, Maryland, 533 


BROBECK, J. R., O E. REYNoLps, AND T. A. APPEL 
(EDs.). 1987. History of the American Physiological 
Society. Waverly Press, Baltimore, Maryland, 533 


pp. 

CALDER, W. A., III. 1984. Size, function, and life 
history. Harvard University Press, Cambridge, Mas- 
sachusetts, 431 pp. 

Cena, K., AND J. A. CLARK. 1973. Thermographic 
measurements of surface temperatures of animals. 
Journal of Mammalogy, 54:1003-1007. 

CHRISTIAN, D. P. 1979. Comparative demography of 
three namib desert rodents: responses to the provi- 
sion of supplementary water. Journal of Mammal- 
ogy, 60:679-690. 

1980. Vegetative cover, water resources, and 
microdistributional patterns in a desert rodent com- 
munity. Journal of Animal Ecology, 49:807-8 16. 

Dit, D. B., A. V. Bock, AND H. T. Epwarps. 1933. 
Mechanisms for dissipating heat in man and dog. 
American Journal of Physiology, 104:36-43. 

Gates, D. M. 1962. Energy exchange in the bio- 
sphere. Harper and Row Publishers, New York, 151 


GRODZINSKI, W., AND B. A. WUNDER. 1975. Ecolog- 
ical energetics of small mammals. Pp. 173-204, in 
Small mammals: their productivity and population 
dynamics (F. B. Golley, K. Petrusewicz, and L. Rysz- 
kowski, eds.). Cambridge University Press, Great 
Britain, 451 pp. 

HeLier, H. C. 1972. Measurements of convective 
and radiative heat transfer in small mammals. Jour- 
nal of Mammalogy, 53:289-295. 

Howe Lt, A. B., AND I. GersH. 1935. Conservation 
of water by Dipodomys. Journal of Mammalogy, 16: 
1-9. 

Lyman, C. P. 1948. Oxygen consumption and tem- 


perature regulation of hibernating hamsters. Journal 
of Experimental Zoology, 109:55-78. 

McNas, B. K. 1963. Bioenergetics and the deter- 
mination of home range size. The American Natu- 
ralist, 97:133-140. 

Nay, K. A. 1987. Field metabolic rate and food 
requirement scaling in mammals and birds. Ecolog- 
ical Monographs, 57:111-128. 

Necus, N. C., AND P. J. BERGER. 1987. Mammalian 
reproductive physiology: adaptive responses to 
changing environments. Pp. 149-173, in Current 
mammalogy (H. H. Genoways, ed.). Plenum Pub- 
lishing, New York, 1:1-519. 

Peters, R. H. 1983. The ecological implications of 
body size. Cambridge University Press, Cambridge, 
Great Britain, 328 pp. 

Rossins, C. T. 1983. Wildlife feeding and nutrition. 
Academic Press, New York, 343 pp. 

ROTHSCHUH, K. 1973. History of physiology (Trans- 
lated and edited by G. B. Risse). Robert E. Kriegen 
Publishing Co., Huntington, New York, 379 pp. 

SCHMIDT-NIELSEN, K. 1984. Scaling: why is animal 
size sO important? Cambridge University Press, 
Cambridge, Great Britain, 241 pp. 

SCHMIDT-NIELSEN, K., AND B. SCHMIDT-NIELSEN. 1952. 
Water metabolism of desert mammals. Physiological 
Reviews, 32:135-166. 

SCHOLANDER, P. F. 1978. Rhapsody in science. An- 
nual Review of Physiology, 40:1-17. 

SCHOLANDER, P. F., V. WALTERS, R. Hock, AND L. 
IRvING. 1950. Body insulation of some arctic and 
tropical mammals and birds. Biological Bulletin, 99: 
225-236. 

SCHOLANDER, P. F., R. Hock, V. WALTERS, F. JOHNSON, 
AND L. IRvING. 1950. Heat regulation in some arc- 
tic and tropical mammals and birds. Biological Bul- 
letin, 99:237-258. 

SCHOLANDER, P. F., R. Hock, V. WALTERS, AND L. 
IrvinG. 1950. Adaptation to cold in arctic and 
tropical mammals and birds in relation to body tem- 
perature, insulation, and basal metabolic rate. Bio- 
logical Bulletin: 99:259-271. 

Smitu, R. E., AND B. A. Horwitz. 1969. Brown fat 
and thermogenesis. Physiological Review, 49:330- 
425. 

SPERBER, I. 1944. Studies on the mammalian kidney. 
Zool. Bidrag Fran Uppsala, 22:249-431. 

TENNENT, D. M. 1946. A study of the water losses 
through the skin in the rat. American Journal of 
Physiology, 145:436-440. 

Van Sosst, P. J. 1982. Nutritional ecology of the 
ruminant. Comstock Publishing Associates, Cornell 
University Press, Ithaca, New York, 373 pp. 

Voruies, C. T. 1945. Water requirements of desert 
animals in the southwest. University of Arizona Ag- 
ricultural Experiment Station Technical Bulletin, 107: 
487-525. 


REPRODUCTION 


OLIVER P. PEARSON AND G. J. KENAGY 


Introduction 


W: find ourselves in the early 1990s 
enriched by fascinating accounts of 
the reproductive biology of hundreds of 
kinds of mammals. We believe that we have 
a sophisticated understanding of the ways 
in which various reproductive mechanisms 
and strategies represent fitness. Almost all 
of this knowledge has been gained in the last 
100 years. 

When the ASM was founded in 1919, de- 
tailed information about reproductive pat- 
terns and mechanisms was available for only 
a handful of domesticated and wild animals, 
as well as for humans. Further understand- 
ing was severely handicapped by the prim- 
itive state of the science of endocrinology 
and the absence of yet-to-be-discovered in- 
sights in cell and molecular biology. In the 
years immediately following the founding 
of ASM, anatomists, physiologists, geneti- 
cists, psychologists, medical researchers, and 
biochemists all began contributing ideas and 
data to the emerging discipline of mam- 
malian reproductive biology (Fig. 1). 

To illustrate the relative collective effort 
that mammalogists have directed into stud- 
ies of reproduction, we tallied all the articles 
published in every fifth volume of the Jour- 
nal of Mammalogy from 1920 through 1990. 


Pa 


{ Le ay ip = 
~ J 
1 ak 
/ 
/ ! 
Sa { { 
segame cs w= (Bim, \ 
Vas SSS et 
0° SarKtocs (| 
Wy \) “&@e PomeSqure \ 
[ WY ene \ 
Nf ok vont? \ \ 
KY /\y 
mh | 
GS | y 
| 
A\ iM 
(de \ (A Wy Ae 


Overall, 8% of the 1,846 papers dealt with 
reproduction. The percentage increased, 
however, during this 70-year interval. For 
the first 25 years only ca. 4% of the articles 
dealt with reproduction, but this increased 
to ca. 11% in the last 30 years (Fig. 2). 
The development of our knowledge of the 
reproductive biology of mammals resulted 
from the vision of a few pioneers whose 
discoveries and teachings spread and mul- 
tiplied while passing through a variety of 
institutions. The favorable climate for this 
flowering was found within institutions that 
drew their support from medical interests, 
agricultural and livestock interests, labora- 
tory-animal needs, entrepreneurs who saw 
commercial opportunities, and surely, from 
the traditional “‘pure”’ scientists— those who 
could not rest until they found out whether 
some exotic species was an induced ovu- 
lator or why some other species had such a 
long gestation. We shall focus on only a few 
of the institutions and people who played 
important roles during the early decades of 
the 20th-Century flowering of reproductive 
biology. By mid-century, when the number 
of participating scientists became so nu- 
merous, we shall call attention to new ideas 
and approaches that have been shaped by 


No 
~I 
i) 


120 


100 


80 


60 


40 


Number of Papers 


20 


O D 


1880 1890 1900 


Marshall, 1910 


1910 


PEARSON AND KENAGY 


NN 
oO 

oO ie 

m ® 

ge re) < 

335 

o— £ 

9 bo 

Ais 

E= 

1920 1930 1940 


Year of Citation 


Fic. 1.—The growth of research on mammalian reproduction. Dates of appearance of 1,362 papers 
cited by Asdell (1946). Five significant publications or events are noted, as discussed in the text. 


more recent generations of innovative peo- 
ple. In the more recent time period we have, 
for simplicity, been highly selective and 
general, indicating ideas of importance rath- 
er than describing them in detail, and we 
have not generally presented these modern 
trends in terms of specific people. More time 
will tell which recent approaches will have 
the most enduring value in the history of 
mammalian reproductive biology. 


EARLY 20TH CENTURY 
The Cambridge Legacy 


The flowering of interest in mammalian 
reproduction reflected in Figure 1 can be 
traced to Great Britain, where a founding 
figure was Walter Heape. After a late start 
in science, Heape found himself teaching 
anatomy at Cambridge, where he co-au- 
thored a textbook of embryology. He soon 
obtained grant support and thereafter de- 


voted himself full time to research. Working 
at a time when even the sex-determination 
mechanism of mammals was unknown, he 
successfully transplanted fertilized ova from 
one rabbit to another in the early 1890s, 
developed artificial insemination in 1897, 
and in 1901 published an impressive sum- 
mary and synthesis (Heape, 1901). He fitted 
humans and dozens of species of wild and 
domesticated mammals into a common 
framework of reproductive categories using 
now-familiar terms (British spellings) such 
as oestrus, pro-oestrum, metoestrum, 
dioestrum, polyoestrous, and monoestrous. 
Had he remained longer in teaching, he no 
doubt would have become the leader of a 
“‘school’’; but, his impact seems to have been 
mostly through his publications. 

Heape’s work inspired F. H. A. Marshall, 
a lecturer at the School of Agriculture in 
Cambridge, to publish his influential book 
on physiology of reproduction (1910). In the 
introduction, Marshall acknowledged, “I 
take this opportunity of recording my in- 


REPRODUCTION 273 


n=1847 


O N 


a 


Frequency (percent) 
Oo 


NO 


1920 1930 1940 


Oo 


1950 


1960 1970 1980 1990 


Year of Publication 


Fic. 2.—Increase in relative frequency of articles on reproduction appearing in the Journal of 
Mammalogy. Frequency distribution is shown for articles containing important information on re- 
production (7 = 1,847), sampled every fifth year from 1920 through 1990. 


debtedness to Mr. Walter Heape, through 
whose influence I was first led to realise the 
importance of generative physiology both 
in its purely scientific and in its practical 
aspects.” 

Marshall’s book was a wide-ranging ac- 
cumulation of information on breeding sea- 
sons, estrous cycles, uterine cycles, ovarian 
changes, spermatogenesis, the testes and 
ovaries as endocrine organs, the placenta, 
and other topics, all with reference to hu- 
mans, laboratory animals, livestock, and 
wild animals. The 1910 and 1922 editions 
of this book became the “bible” to a gen- 
eration of biologists who were to outline the 
diversity of reproductive strategies in mam- 
mals. 

A second book by Marshall on reproduc- 
tive physiology appeared in 1925. It ap- 
peared in time to influence the people, 
mostly British, who published reproductive 
studies in the 1930s and 1940s. The closing 


paragraphs of this second book call atten- 
tion to global population concerns, Mal- 
thus, contraception, and eugenics. Reading 
those paragraphs two generations later brings 
one face to face with the fact that during the 
intervening 70 years we have not resolved 
those old yet vital ethical concerns. Fur- 
thermore, still newer discoveries have cre- 
ated yet more concerns undreamed of by 
Marshall. 

Even before Marshall’s books, British bi- 
ologists were otherwise prepared for a flow- 
ering of reproductive studies. W. H. Cald- 
well, a Cambridge scholar on an expedition 
in 1884 to one of the colonies (Australia), 
sent the famous cable ““Monotremes ovip- 
arous, Ovum meroblastic” not to Great Brit- 
ain or “the Continent,” but to another out- 
post of the United Kingdom, Canada and 
the city of Montreal, where the British As- 
sociation for the Advancement of Science 
was holding its annual meeting (Burrell, 


274 


1927). In this geographically widespread and 
receptive climate, Marshall’s books provid- 
ed a foundation on which subsequent gen- 
erations could build. The pages of the Pro- 
ceedings of the Zoological Society of London, 
the Philosophical Transactions of the Royal 
Society of London, and other prestigious 
publications are forever enriched by im- 
portant reproductive studies by other lu- 
minaries such as E. C. Amoroso, J. R. Ba- 
ker, F. W. R. Brambell, R. Deanesly, J. 
Hammond, J. P. Hill, L. H. Matthews, A. 
S. Parkes, I. W. Rowlands, and Sir Solly 
Zuckerman. Within little more than a de- 
cade in the 1930s and early 1940s they de- 
scribed the intricacies and novelties in the 
reproductive cycles of shrews and bats, 
hedgehogs and hyaenas, kangaroos and fer- 
rets, wildcats and moles, and gibbons and 
rabbits. In the coming decades these re- 
searchers were followed by P. H. Leslie, J. 
L. Davies, B. Weir, and many others. A 
third edition of Marshall’s book, delayed by 
World War II and published as three vol- 
umes with many chapter authors, appeared 
between 1952 and 1966 under the editor- 
ship of A. S. Parkes. Marshall had died in 
1949, but he had contributed to many of 
the chapters. A fourth edition appeared in 
1984, edited by G. E. Lamming. 

The Zoological Society of London under 
the presidency of Sir Solly Zuckerman be- 
came one of the institutions that had a great 
impact on the development of studies of 
reproduction. In 1963, with support from 
industry (the Wellcome Trust), a research 
center was established, with an emphasis on 
studies of mammalian reproduction—I. W. 
Rowlands was its first director. 

Perhaps the ultimate fruition of the Cam- 
bridge rootstock came in 1960, when the 
Society for the Study of Fertility, with sup- 
port from the Wellcome Trust, founded the 
Journal of Reproduction and Fertility. C. R. 
Austin was editor and A. S. Parkes Chair 
of the Editorial Board. In the first issue, 
Parkes pointed out that the “‘output of lit- 
erature on reproduction and fertility is 
mounting rapidly owing to the increasing 
number of scientifically based clinical stud- 


PEARSON AND KENAGY 


ies, the greater importance attached to pro- 
ductivity in farm animals, the extension of 
field and laboratory studies to additional 
species, and the growing realization of the 
urgent need for finding means of controlling 
fertility in man.” For three decades this 
journal has advanced in a distinguished 
manner the research interests of reproduc- 
tive biologists. The first issue of volume 1 
contains Hilda Bruce’s description of a 
pheromonal influence on reproduction that 
came to be known as the Bruce Effect (Bruce, 
1960). A recent number (1988, no. 1), ed- 
ited in Cambridge by Barbara Weir, with E. 
J. C. Polge as Chair of the Executive Com- 
mittee, contains articles on the reproduc- 
tion of no less than 16 genera of mammals. 


The Johns Hopkins Legacy 


Returning to 19th-Century North Amer- 
ica, the Johns Hopkins University was es- 
tablished in Baltimore, Maryland, in 1876. 
The goal of the biology program was to pro- 
vide students with hands-on, laboratory- 
oriented research training rather than the 
traditional lecture-til-full system; this suc- 
cessful model was eventually adopted by 
many North American universities (Ben- 
son, 1987). Thomas Huxley had been con- 
sulted extensively during the planning of the 
curriculum, and he recommended one of his 
proteges from Cambridge, H. Newell Mar- 
tin, a physiologist, to be the first professor 
in the new Biology Department. The ap- 
pointment of W. K. Brooks, a morphologist, 
followed immediately. The department 
subsequently produced an impressive array 
of scholars including E. B. Wilson, T. H. 
Morgan, E. G. Conklin, and R. G. Harrison, 
who all made a great impact on biology. The 
great influence of biology at Johns Hop- 
kins on the discipline of mammalian repro- 
duction was accomplished through a variety 
of the university’s satellite programs, 1n- 
cluding the Medical School, the School of 
Hygiene and Public Health, the Institute for 
Biological Research, and, beyond the uni- 
versity itself, the Department of Embryol- 


REPRODUCTION 21D 


ogy of the Carnegie Institution of Washing- 
ton. 

The Medical School opened in 1893. It 
was headed by Franklin Mall, who had been 
head of the Anatomy Department at the 
University of Chicago, an institution that 
had been modelled after Johns Hopkins. 
Many of the movers and shakers in the dis- 
cipline of reproductive biology, such as Os- 
car Riddle, B. Bartelmez, Carl Moore, W. 
C. Young, Karl Lashley, and Frank Beach, 
were eventually trained at Johns Hopkins. 
Mall served as Professor of Anatomy at the 
Hopkins Medical School and encouraged 
development of at least 20 future professors 
ofanatomy; three of them are especially per- 
tinent to this review: George Wislocki, Her- 
bert Evans, and George Corner. 

Of these three, anatomist-histologist 
George Wislocki went from Johns Hopkins 
to the Medical School at Harvard Univer- 
sity. He and his colleagues and students, 
such as Roy Greep, E. B. Astwood, E. W. 
Dempsey, Don Fawcett, Helen Deane, and 
William Wimsatt, spread the base of species 
studied to even more remote corners of the 
Class Mammalia. 

Herbert Evans moved from Johns Hop- 
kins to the Medical School of the University 
of California in Berkeley, where he founded 
the Institute of Experimental Biology. Dur- 
ing nearly four decades he and members of 
the Institute accomplished a remarkable 
amount of important research. One of the 
first achievements was the 1922 monograph 
on the estrous cycle in the rat, coauthored 
by zoology professor Joseph Long (Long and 
Evans, 1922). They had created the Long- 
Evans strain of laboratory rat and, using the 
newly discovered vaginal smear technique, 
revealed the formerly unknown details of 
the estrous cycle of this laboratory animal. 
As pointed out by A. S. Parkes (1969), one 
has only to review the 10 abstracts by Long 
and Evans in the 1920 Anatomical Record, 
followed by 13 abstracts in the 1921 Ana- 
tomical Record by Evans and his associates, 
to be awed by the sweep of Evans’ early 
contributions to reproductive anatomy and 
physiology. This was only a beginning, and 


was followed in 1931 by a monograph on 
reproduction in the dog (with H. H. Cole of 
the Department of Animal Sciences of the 
University of California at Davis), dem- 
onstrations of the pituitary gland as an en- 
docrine organ, description of the growth 
hormone, and discovery of vitamin E and 
its role in reproduction (Parkes, 1969). 

The third of this trio, George Corner, went 
to the Medical School at the University of 
Rochester, where he became widely known 
for his studies of the menstrual cycle of 
monkeys, the role of the corpus luteum as 
an endocrine organ and, with Willard Allen, 
the purification of the hormone progester- 
one. In 1940, Corner returned to Johns 
Hopkins and became Director of the De- 
partment of Embryology of the Carnegie In- 
stitution of Washington. His former pro- 
fessor, Mall, had been the first Director 
(1914), and two of his Hopkins teachers, 
Florence Sabin and Warren Lewis, also had 
distinguished careers at Carnegie. 

The greatest impact of Johns Hopkins on 
the discipline of mammalian reproduction 
was through the Department of Embryology 
of the Carnegie Institution. It became the 
most important center of reproductive stud- 
ies in the United States. At one time or 
another it included important anatomists 
and physiologists such as Warren Lewis, 
George Streeter, Oscar Riddle, Chester 
Heuser, Arthur Hertig, John Rock, Carl 
Hartman, George Bartelmez, Sam Reyn- 
olds, George Corner, Robert Enders, and 
Harland Mossman (Fig. 3). 

Two other administrative units that add- 
ed strength to the Johns Hopkins University 
were the School of Hygiene and Public 
Health, created in 1918, and its offshoot, 
the Institute of Biological Research. The lat- 
ter was headed by Raymond Pearl and then 
was absorbed by the School of Hygiene and 
Public Health after Pearl’s death in 1940. 
Pearl was a biometrician. He applied sta- 
tistics to the birth, life, and death rates of 
populations, especially humans. He had 
wide-ranging influence through the two 
journals that he founded: Human Biology 
and Quarterly Review of Biology. 


276 PEARSON AND KENAGY 


a 
ro 
c oxy 
ot TT 
e i . al 
ig 
ee o- ‘ 
= * as 
>. gag é 
4 oe a, 
& . J a5 4 


Fic. 3.—Photograph at the Carnegie Institution of Washington, Department of Embryology, Bal- 
timore, 1931. Left to right: George Streeter, Robert Enders, Chester Heuser, Josephine Ball, Carl 
Hartman, P. Mihalic, Warren Lewis, Sam Reynolds. 


The Carnegie group became known for 
exquisite studies of the embryology of hu- 
mans, rhesus monkeys, and other mam- 
mals, published in the Contributions to Em- 
bryology of the Carnegie Institution. Many 
of the studies were beautifully illustrated by 
the noted medical illustrator James Di- 
dusch. Indeed, the first paper in volume 1 
is by Mall himself (Mall, 1915). The Car- 
negie group moved inevitably into endo- 
crine studies at a time when the exciting 
interplay of hormones produced by gonads, 
pituitary, and placenta was just being dem- 
onstrated. 


Other Legacies 


The significant role played by anatomists 
at medical schools during the development 
of our understanding of mammalian repro- 
duction is illustrated also by research and 


teaching at many such institutions. Much 
of the research was directed not at human 
problems but at a truly comparative un- 
derstanding. While teaching at Cornell 
Medical College, Stockard and Papanico- 
laou (1917) discovered the utility of the 
vaginal smear in guinea pigs and in humans 
(the Pap smear). Harland Mossman, after a 
brief stay at Carnegie, had a long career of 
teaching and research at the Medical School 
at the University of Wisconsin, and pub- 
lished several influential books on human 
embryology (Hamilton et al., 1945, 1962); 
comparative morphology of the mamma- 
lian ovary (Mossman and Duke, 1973); and 
fetal membranes of vertebrates (Mossman, 
1987). The potential of academic institu- 
tions was demonstrated by this small nu- 
cleus at Wisconsin; when an international 
symposium on the comparative biology of 
reproduction in mammals was convened in 
1964 in London, eight of the 30 contribu- 


REPRODUCTION 217 


tors held advanced degrees from the Uni- 
versity of Wisconsin. Further aspects of the 
development of North American reproduc- 
tive physiology in the early 20th Century 
are presented by Clarke (1987). 

Another radiation directly traceable to the 
Carnegie group was into a government- 
sponsored program to understand the re- 
productive performance of commercially 
important fur-bearing mammals. This pro- 
gram was led by Frank Ashbrook in the 
Division of Fur Resources, U.S. Depart- 
ment of Agriculture (later Fish and Wildlife 
Service of the Department of Interior). 
Studies were conducted on the reproduction 
of fur seals, martens, minks, foxes, nutrias, 
and muskrats. Some of these studies were 
carried out at Swarthmore College near 
Philadelphia under the leadership of Robert 
Enders, who had spent a stimulating post- 
doctoral period at the Carnegie Institution. 
In addition to his own research on the mink 
and other fur-bearing animals, he used this 
major project, beginning in the 1940s, to 
introduce numerous students to research on 
mammalian reproduction. Some of them, 
chronologically, were David Bishop (sperm 
physiology), David Davis (rat populations, 
stress), Oliver Pearson (reproductive cy- 
cles), Bent Boving (implantation), Hewson 
Swift (Sertoli cells), Duncan Chiquoine 
(germ cells), Allen Enders (implantation), 
William Tietz (embryogenesis), Edward 
Wallach (ovarian physiology), Phil Myers 
(rodent and bat reproduction), and Anne 
Hirschfield (dynamics of ovarian follicles). 
Many of these students and more recently 
their own students continue searching for 
insights into reproductive biology. 


Further Notable Publications 


North American researchers were influ- 
enced by the excitement over reproductive 
biology at Cambridge and other European 
sources in two ways—by reading the Eu- 
ropean literature and by direct contact with 
researchers in North America who had been 


exposed earlier to the ideas and approaches 
in Europe. For example, workers such as 
Asdell, Bissonnette, Chang, and Pincus spent 
early parts of their careers at Cambridge. 
Meanwhile, North Americans published 
most of their own work in American jour- 
nals. Three journals of great importance to 
reproductive biology were the American 
Journal of Anatomy (founded 1901), the An- 
atomical Record (1908), and The Journal of 
Experimental Zoology (1904). All three were 
managed by the Wistar Institute in Phila- 
delphia. 

A book of undoubted importance in the 
development of reproductive biology ap- 
peared in 1932, with a second edition in 
1939. Professor Edgar Allen at the Univer- 
sity of Missouri, who had published on the 
early embryology of humans in the Carnegie 
Institution Contributions to Embryology, 
assembled a collection of coherent reviews 
by 21 distinguished collaborators that was 
published under the title of ““Sex and Inter- 
nal Secretions.” It was dedicated to A. D. 
Mead, one of the members of the staff in 
anatomy at the University of Chicago in its 
early days. This book enabled a new gen- 
eration of students to approach reproduc- 
tive studies with a more solid foundation 
in the new science of endocrinology than 
was available to the generation weaned on 
Marshall’s book. W. C. Young edited a third 
edition in 1961. 

Studies of reproduction in farm livestock 
were conducted largely by federal agencies 
and by universities with an agricultural em- 
phasis, both in Europe and the United States. 
A milestone of this radiation, which dem- 
onstrated the coming-of-age of comparative 
reproductive biology, was the appearance in 
1946 of “Patterns of Mammalian Repro- 
duction” by S. A. Asdell. Asdell came as a 
postdoc from England to Corner’s labora- 
tory at the University of Rochester Medical 
School, and later became a professor in the 
Department of Animal Husbandry at Cor- 
nell University. Asdell realized that “a 
beneficial purpose would be served if the 
available information on mammalian re- 


120 


100 


10.¢] 
Oo 


122) 
© 


a 
© 


Number of Papers 


Marshall, 1910 
Marshall, 1922 


NO 
O 


1880 


Marshall, 1925 


PEARSON AND KENAGY 


Asdell, 1946 


1980 


1940 1960 


Year of Citation 


Fic. 4.—A century of research on mammalian reproduction. Dates of appearance of 2,346 papers 
cited by Mossman (1987). Significant publications or events are noted, as in Figure 1. 


production were brought together, species 
by species... .”” After paying tribute to his 
great predecessor at Cambridge, F. H. A. 
Marshall, Asdell displayed the fruits of the 
labors of hundreds of authors who had stud- 
ied the reproduction of about 382 genera 
and 850 species of mammals. In those pages 
one could find information ranging from the 
unilateral functioning of platypus ovaries 
(page 37) to the number of spermatozoa in 
the ejaculate of the donkey (page 405). An 
expanded version of Asdell’s book has ap- 
peared recently (Hayssen et al., 1993). 

We have compiled a distribution of the 
dates of 1,362 literature citations in Asdell’s 
1946 book (Fig. 1), covering the late 19th 
and first half of the 20th centuries. Studies 
in mammalian reproduction clearly blos- 
somed beginning in the 1920s. The dates of 
Marshall’s books, of the founding of the 
ASM, and of Allen’s book on sex and in- 
ternal secretions are indicated as reference 
points. The decrease in number of citations 
in the mid-1940s is partially the result of 


World War II, in addition to the decline 
expected for bibliographic truncation at the 
approach of the publication date of Asdell’s 
book. Figure 4 illustrates further prolifera- 
tion of work in the 1960s and 1970s, fol- 
lowing three-quarters of a century over 
which the early historical background was 
built. 

Another serial, Biology of Reproduction, 
was Started in the United States in 1969, 
edited by H. H. Cole. It too continues to 
publish scores of papers each year in com- 
parative reproductive anatomy and physi- 
ology. 

With growing audiences of university stu- 
dents, in addition to the research specialists, 
a parade of text books in reproductive bi- 
ology arrived on the scene in the 1960s, 70s, 
and 80s. These books, mostly dealing with 
all vertebrates, rather than exclusively with 
mammals, are general and broad enough to 
be useful as texts for undergraduate zoology 
courses in reproduction, yet most of them 
also contain sufficient synthesis and over- 


REPRODUCTION 219 


view along with the observational detail to 
make them useful in a personal research 
library. Sadleir’s book (1969) on mammals 
emphasizes reproductive ecology, breeding 
patterns, and responses to environmental 
conditions; his later book (1973) on verte- 
brates is much broader in scope and surveys 
general reproductive patterns and compar- 
ative anatomy and physiology. Nalban- 
dov’s book (three editions: 1958, 1964, and 
1976) emphasizes the reproductive physi- 
ology and anatomy of mammals and birds. 
Van Tienhoven (1968, 1983) treats the 
physiology and anatomy of all vertebrates. 
Two of the most recent books on vertebrate 
reproduction (Bliim, 1986; Jameson, 1988) 
are organized thematically, rather than tax- 
onomically and, in addition to the usual 
information on reproductive patterns, anat- 
omy, and physiology, provide a greater 
comparative and evolutionary perspective 
and greater integration of behavioral themes 
with all these areas. 

Finally, and with exclusive attention to 
mammals, several well-produced text 
sources are available. Austin and Short 
(1972-1986, eight books, two editions) have 
produced a series of booklets that cover 
pretty much the full range of topics in mam- 
malian reproductive biology, with well il- 
lustrated examples of research results along 
with the conceptual developments. Bron- 
son’s (1989) single-volume book offers an 
extensive and well laid out analysis of the 
regulatory processes that comprise mam- 
malian reproductive physiology set in an 
environmental context. Flowerdew (1987) 
provides a useful blend of fundamentals of 
reproductive physiology with the biology of 
free-living mammalian populations. Clear- 
ly, and without our being able to mention 
all such existing books, a great variety of 
general reading has become available on 
mammalian reproductive biology. Proba- 
bly one of the strongest recent areas of in- 
tegrative bridging in the field of reproduc- 
tive biology has been between physiology 
and behavior (see Eisenberg and Wolff, 
1994), 


Perhaps the most impressive evidence of 
the growth and vigor of reproductive biol- 
ogy as a general discipline is the appearance 
in 1963 and subsequent growth of the Bib- 
liography of Reproduction. It is published 
monthly in Cambridge, England, by a con- 
sortium of reproductive societies in Great 
Britain, the United States, and Australia. 
The editors estimated that the annual pro- 
duction of papers (in 1990) on the repro- 
ductive biology and clinical sciences of ver- 
tebrates including humans may be on the 
order of 20,000. 

In view of such a torrent of research, the 
impact of a relatively few institutions and 
individuals (as we have selected), along with 
their academic offspring, on the early de- 
velopment of the discipline of mammalian 
reproduction becomes quickly lost in the 
distal branches of the family trees. While 
emphasizing, and even exaggerating, the 
roles of only a few individuals, we have 
omitted untold other early researchers and 
teachers, many of whom worked in other 
countries. Thus we admit that it would be 
impossible to trace a balanced and objective 
presentation of all the research schools and 
their modern ramifications in a short article 
such as this. The rest of our review will thus 
simply identify the appearance of a selected 
series of what we believe to be important 
research trends in reproduction that have 
developed during the final third of the 20th 
Century. 


LATE 20TH CENTURY 


We find ourselves at the end of the 20th 
Century in a stream of fast-moving devel- 
opments and continued new discoveries in 
reproductive biology as we mark the 75th 
anniversary of the founding of the ASM. 
The most comprehensive new treatise on 
mammalian reproduction (Knobil and Neill, 
1988) in the latter part of the century has 
appeared in two volumes, 2,413 pages, and 
60 contributed chapters, each containing 
from several hundred to a thousand refer- 


280 PEARSON AND KENAGY 


ences. This new work, inspired by Allen’s 
(1932) original “Sex and Internal Secre- 
tions,’ was edited and produced in the 
United States, with most of the authors from 
North America and many from elsewhere 
around the world. It provides a strong focus 
on cells, tissues, and neuroendocrine phys- 
iology, yet extends as far as reproductive 
behavior. 

Our breadth of understanding of repro- 
ductive physiology and behavior in an evo- 
lutionary context can only continue to im- 
prove and become more meaningful. For 
the entire first century after Darwin’s writ- 
ings, challenges in the form of important 
questions in evolutionary theory resulted in 
important refinements and a maturity in our 
current view. Therefore, the potential rel- 
evance of integrative and evolutionary 
thinking at present is greater than ever. The 
tendency of so many scientists to specialize 
SO narrowly offers a new challenge: to over- 
come narrow specialization by seeking 
breadth of understanding in the context of 
evolutionary biology. 

As an example, a seemingly simple ques- 
tion remains of interest: why is reproduc- 
tion typically sexual, rather than asexual, 
and why are there two, rather than some 
other number, of sexes (Short, 1994)? Ideas 
concerning this theoretical and evolution- 
ary question can lead us in our search for 
the still unresolved issues of the mecha- 
anisms of sex determination and sexual dif- 
ferentiation, which lie at the level of the 
molecular biology of gene function (Mc- 
Laren, 1991). 


Reproduction, Neuroendocrinology, 
and Molecular Biology 


Mammalian reproduction is comprised 
of a great array of processes: gamete pro- 
duction and release, mating behaviors, fer- 
tilization, implantation, development, pla- 
cental function, parturition, lactation, and 
parental care. Our understanding of each of 
these processes has developed strongly in 
conjunction with the identification of hor- 


mones that control them (Knobil and Neill, 
1988). Study of hormones has been a major 
paradigm of reproductive physiology since 
the middle of the century, with the advent 
of radioimmunoassays for measuring hor- 
mone concentrations and the perfection of 
biochemical techniques for characterizing 
hormone structure and function. Under- 
standing the integration of nervous system 
output, including secretion by neurons of 
small peptide hormones that stimulate fur- 
ther hormonal signals that enter the blood 
stream, has provided the challenge to elu- 
cidate the role of the brain, hypothalamus, 
and pituitary in neuroendocrine regulation 
(Everett, 1988). Despite the general appli- 
cability of the neuroendocrine paradigm, it 
should also be useful in elucidating excep- 
tional patterns and modes of reproduction. 
For example, the arrest and later reactiva- 
tion of embryonic development occurs in 
special cases (“delayed implantation” and 
““embryonic diapause’’) where the delay may 
be associated with either lactation for earlier 
young that precede the arrested embryo(s), 
or with environmental factors that allow 
birth to occur at an appropriate time (Ren- 
free and Calaby, 1981). 

Through the extensive series of neuroen- 
docrine regulatory schemes that have been 
unveiled by research on mammalian repro- 
duction, the general field has served as a 
model for study of neuroendocrinology. One 
of the newest directions for this research has 
been the molecular neuroendocrinology of 
gene expression, i.e., identifying and quan- 
tifying the first gene products associated with 
hormone production. This new research 
trend amounts to another dimension in in- 
tegrative reproductive biology, namely elu- 
cidating functional (physiological) aspects 
of molecular biology. 


Environmental Physiology and 
Regulatory Processes 


Use of environmental information and 
environmental stimulation or inhibition of 
reproductive function represents one of the 


REPRODUCTION 281 


most popular themes in reproductive re- 
search. We will mention only a few high- 
lights of our current understanding of this 
area, for which Bronson’s (1989) book pro- 
vides a useful view. 

The time course over which seasonally 
breeding mammals respond and the cues 
used differ between the sexes and according 
to different stages in the overall reproduc- 
tive program, beginning with activation of 
the gonads and extending through final as- 
pects of postnatal care and termination of 
breeding condition (Wingfield and Kenagy, 
1991). An enormous literature on the initial 
predictive effects of day length in stimulat- 
ing gonadal recrudescence and thus prepar- 
ing mammals for the onset of breeding 
(Bronson, 1989; Farner, 1985; Wingfield and 
Kenagy, 1991) has probably lead to an over- 
impression of the importance of ‘“‘photo- 
period”’ in breeding, at least in part because 
the effect is so consistent, easily obtainable, 
and the first to occur in a series of steps. 
Actually, not all mammals are “photope- 
riodic,” 1.e., capable of differential response 
to long versus short days. A small number 
of species (most notably ground squirrels 
and their relatives in the tribe Marmotini 
of the squirrel family Sciuridae) show per- 
sistent endogenous cycles of reproductive 
function in the experimental absence of sea- 
sonal changes in day length (Gwinner, 1986). 

The mammalian mechanism of photo- 
reception that drives the initial response of 
the reproductive system begins with the eyes 
and then a connection through the retino- 
hypothalamic tract, a neural circuit from the 
retina to the brain that is distinct from the 
visual pathway (Rusak and Morin, 1976; 
Stetson and Watson-Whitmyre, 1976). The 
information on day length is processed in 
the suprachiasmatic nuclei of the hypo- 
thalamus, and signals are then sent through 
the brainstem and a spinal ganglion and back 
to another site in the brain, the pineal gland. 
Finally the daily rhythmic secretion of mel- 
atonin by the pineal plays an important role 
in the regulation of reproduction in re- 
sponse to changes in day length (Binkley, 
1988; Hoffmann, 1981; Reiter, 1984). Some 


of the earliest pioneering work with the pi- 
neal was that of Wilbur Quay (1956), who 
studied seasonal and sexual variation in the 
pineal of Peromyscus. 

Many aspects of environmental infor- 
mation besides day length, including the so- 
cial context of an animal in its population, 
provide supplementary stimuli that syn- 
chronize, integrate, and modify the repro- 
ductive responses at all stages of the breed- 
ing cycle (Wingfield and Kenagy, 1991). 
Manipulations of simulated environmental 
conditions have been conducted to observe 
these responses, often including hormonal 
measurements, to environmental factors 
such as food supply (quantity and quality), 
water availability, temperature, and the so- 
cial setting and attendant cues (Bronson, 
1989; Wingfield and Kenagy, 1991). Some 
of the most useful research has involved 
species that can be studied both in the field, 
for correlative analysis, and in the labora- 
tory, where simulation and manipulation 
can be carried out. Comparative field in- 
vestigation of multiple species has illustrat- 
ed that diverse patterns of breeding occur 
even in the same environment, and that 
body size, phylogeny, and specific adapta- 
tions of species account for the differences 
in timing and intensity of reproduction 
(Kenagy and Bartholomew, 1985). Such field 
observations have indicated the potential 
for each species to utilize different cues and 
to respond with different sensitivities to the 
entire range of environmental factors. 

One of the most obvious and direct phys- 
iological responses that involves regulation 
of reproductive function is the availability 
of appropriate amounts and quality of nu- 
trients and energy. Nutritional plane and 
energy balance act directly on the animal’s 
metabolism and the assessment of body 
condition (I’Anson et al., 1991). Research 
in this area involves integration of data on 
general metabolism and metabolic hor- 
mones, as well as relevant organs such as 
the thyroid and adrenals, with the neuroen- 
docrinology of the hypothalamic-pituitary- 
gonadal axis. The mammal’s assessment of 
its nutritive plane and energy balance ap- 


282 


pears to play a direct day-by-day role in the 
onset and maintenance of reproductive 
function. 

A mechanism of reproductive stimula- 
tion in mammalian herbivores that has re- 
mained an attractive research subject is the 
possibility that fresh green food contains a 
gonadotropic chemical signal (Friedman and 
Friedman, 1939). A natural plant com- 
pound, 6-methoxy-2 benzooxazolinone (6- 
MBOA), has more recently been identified 
and shown experimentally to stimulate re- 
productive function (Berger et al., 1981). 
Since the initial demonstration of this effect, 
similar results have been obtained in several 
rodent species. However, much remains to 
be learned about this effect, the extent of its 
occurrence among rodents, and the strength 
of interactions between the effect of 
6-MBOA and other environmental factors 
that promote reproductive responses. It 
could be argued, for example, that because 
food is available already at the time it is 
being consumed, there would be no need 
for a predictive cue. The fact that a com- 
pound of interest has been identified has 
opened the door to new research possibili- 
ties. 


Reproductive Energy Expenditures 


The study of energy relations in repro- 
duction has continued to develop in pop- 
ularity (Loudon and Racey, 1987). Energy 
is a meaningful reflection of allocation to 
reproduction and the relative functional sig- 
nificance of both physiological and behav- 
ioral work; it is often considered to be a 
currency that might represent fitness. The 
reproductive energy allocations of small 
mammals are of particular interest because 
of the extreme maternal intake and expen- 
diture that must be required to support a 
litter whose requirements eventually far ex- 
ceed those of the mother herself (Pearson, 
1944). Considerable impetus to the analysis 
of energy use in animals came from the ef- 
forts of two researchers in American agri- 
cultural university settings at the middle of 


PEARSON AND KENAGY 


the 20th Century. Both S. Brody (1945) of 
the University of Missouri and Max Kleiber 
(1961) of the University of California at Da- 
vis produced important books that present 
the usefulness of energy analysis. 

Attention recently focused on measuring 
the energy allocated to reproduction and 
growth in the context of life histories of free- 
living animals. It is clear that for many spe- 
cies the peak of all energy expenditures is 
reached by females during lactation (Ofte- 
dal, 1984). Some of the earlier attention to 
“reproductive energetics’ that addressed 
only the basal, nonreproductive rates of en- 
ergy expenditure will not remain as useful 
as newer, more explicit approaches (Loudon 
and Racey, 1987). A more direct approach 
that seeks to quantify reproductive energy 
expenditure and intake as they approach 
peaks may allow us to understand energetic 
bottlenecks associated with reproduction 
and even thereby the impact of reproduc- 
tive expenditures on fitness costs (Daan et 
al., 1991; Kenagy et al., 1990). 


Olfaction and Regulation of 
Reproduction 


Mammals generally rely to a much greater 
extent on the use of air-borne chemical in- 
formation concerning their environment and 
their conspecifics than do most other ver- 
tebrates. Olfactory sensation and “‘phero- 
mones” are particularly important in repro- 
ductive behavior and physiology, which has 
made mammals the most important re- 
search model for the study of olfaction 
(Booth and Signoret, 1992; Marchlewska- 
Koj, 1984; Vandenbergh, 1988). Next to 
research on mammalian olfaction, that on 
insects is far greater than on all the other 
vertebrate classes. The function of air-borne 
chemicals (pheromones) to prime other in- 
dividuals by influencing their physiology and 
behavior probably extends across most 
mammalian orders; pheromones play a role 
not only in priming the initial (puberty or 
recrudescence) and mating stages of repro- 
duction, but extend through the time of lac- 


REPRODUCTION 283 


tation and mother-young relations, and be- 
yond that to the level of recognizing the 
identity of individuals within a population 
(Booth and Signoret, 1992). 

Substantial documentation is available for 
pheromonal influences such as the cancel- 
lation of pregnancy due to the odors of a 
strange male (the classical “‘Bruce Effect’; 
Bruce, 1960), the accelerated onset of pu- 
berty in females due to the odors of males, 
and the inhibition of onset of female repro- 
duction by the odors of other females or 
family (Vandenbergh, 1988). The impact of 
this field of research has been substantiated 
by study of these kinds of processes in the 
field, which represents an important con- 
tribution to population biology and behav- 
ior. 


Behavior and Neuroendocrinology 


During the last quarter of the 20th Cen- 
tury the contributions of studies of neu- 
roendocrine mechanisms to the under- 
standing of reproductive behavior have 
become extremely important. The popular- 
ity of such research derives from its ability 
to address ecological and evolutionary 
questions with the approaches of neuro- 
biology and molecular biology (Crews, 
1992). Such a potential for integrative ex- 
ploration with a focus at the organismal lev- 
el reflects back to a view that prevailed at 
the founding of the ASM in 1919. It is grat- 
ifying, in light of the enormity and diversity 
of the modern biological research enter- 
prise, that modern mammalogists have the 
opportunity to foster interest in the per- 
spective of mammals as organisms. 

Research on the diversity of reproductive 
patterns and their mechanisms of neuroen- 
docrine control has produced valuable evo- 
lutionary insights. For example, certain bats 
have temporally dissociated the time of 
mating from the time of gametogenesis by 
allowing hibernation to intervene; the gen- 
eration of neural and endocrine signals that 
direct this program modification illustrates 
the adaptive adjustments that can evolve 


within the constraints of the mammalian 
system (Crews, 1992). As another example, 
we have accumulated information on over 
50 species of primates alone concerning var- 
ious patterns of neuroendocrine regulation 
that sustain the diversity of sexual behavior 
strategies within this group (Dixson, 1983). 

An area of mammalian reproductive bi- 
ology that has relied on integration of phys- 
iological and neuroendocrine analyses going 
back to the middle of this century, and even 
earlier, is represented by the classic rodent 
population studies of Christian and Davis 
(1956). The potential interaction of the ad- 
renal glands (and glucocorticoid hormones) 
with somatic and reproductive condition 
became apparent with the advent of the 
“stress” concept by Selye (1936). Neuroen- 
docrine mechanisms of reproductive func- 
tion and the interaction of this with stress 
physiology have thus been a long-standing 
aspect of research on small-mammal pop- 
ulation regulation (Lee and McDonald, 
1985). The most recent research has dem- 
onstrated the action of glucocorticoids in 
establishing a behavioral basis for differ- 
ences among individuals within popula- 
tions (Sapolsky, 1992). 


Marsupials 


Mammalian diversity has provided a ba- 
sis for comparative functional studies as well 
as evolutionary analysis. In this regard the 
marsupials represent a most remarkable 
payload of fascinating subject matter. J. P. 
Hill, C. G. Hartman, and G. B. Sharman 
were the earlier pioneers of marsupial re- 
productive biology. Since their time, re- 
search has been conducted by many others, 
especially in Australia, both at the univer- 
sities and at other institutions, particularly 
the Commonwealth Scientific and Indus- 
trial Research Organization (CSIRO) and 
its Division of Wildlife and Ecology, known 
earlier by other names. ‘“‘Reproductive 
Physiology of Marsupials” (Tyndale-Biscoe 
and Renfree, 1987) is an excellent mono- 
graphic review of this research, answering 


284 PEARSON AND KENAGY 


many earlier questions concerning patterns 
and mechanisms, and raising new questions 
for future research. Many of the most re- 
markable contributions to marsupial repro- 
ductive endocrinology involve the process 
of embryonic diapause, originally identified 
by G. B. Sharman (1955). This process has 
since been shown in macropod marsupials 
to include simultaneous maintenance of two 
or three young of different ages by a mother 
and the production of milk of two different 
types out of different teats to support young 
of different ages (Tyndale-Biscoe and Ren- 
free, 1987). 

The evolutionary question as to why mar- 
supials quickly pass through uterine embry- 
onic life and then so greatly prolong lacta- 
tion, as the major avenue of matrotrophy 
for development, remains open. One idea, 
now dispelled by recent immunological 
research (Rodger et al., 1985), was that mar- 
supial mothers have a short gestation be- 
cause the trophoblast lacks the immuno- 
suppressive capability that would allow it 
to remain in the uterus without being re- 
jected by the mother as “foreign”? tissue. 
Fetal immunosuppression had already been 
recognized as a basis for eutherian maternal 
recognition of pregnancy and retention of 
young in the uterus, and was only demon- 
strated recently in marsupials (Rodger et al., 
1985). New arguments for the evolutionary 
predilection of marsupials for lactation over 
placentation must be developed and sup- 
ported. It is clear that the marsupial mode 
of reproduction is adaptive and should not 
be considered “‘primitive”’ or “inferior” — 
which was an inappropriate notion that dates 
back to the earliest discoveries of the pouch 
mode of nurturing extremely immature 
newborn. 


Reproductive Technologies 


Experimental reproductive biology has 
both agricultural and medical applications. 
Manipulations of hormones, cells, and tis- 
sues were underway by the mid 20th Cen- 


tury, whereas genetic (transgenic) manipu- 
lation did not arise in the applied context 
until the 1980s. 

Many aspects of reproduction have been 
manipulated to increase production by farm 
animals (Betteridge, 1986). These include 
artificial insemination, induction of estrus, 
synchronization of estrus or ovulation in 
groups, embryo transfer and manipulation, 
and in vitro fertilization; development of 
diagnostic tests has improved the usefulness 
of all these techniques. Genetic engineering, 
the insertion of specialized hormone-pro- 
ducing genes in transgenic animals, is being 
tested actively for applications such as en- 
hancement of milk and meat production by 
growth hormone (Pursel et al., 1989). 

Other manipulations of mammalian re- 
production are being developed in wildlife 
conservation or management and in pest 
control. Captive breeding programs, which 
often include artificial insemination or em- 
bryo transfer, have been the only apparent 
alternative for maintaining some rare spe- 
cies, either in zoos or wildlife sanctuaries. 
On the other hand, explorations are being 
made of means to curb female reproduction, 
for example, in elephant populations that 
are overcrowded due to habitat destruction; 
in this case the antigestagenic steroid RU486 
has been proposed as an abortion agent 
(Short, 1992). Finally, artificial steroid hor- 
mones that produce infertility or disturb 
normal function have recently been pro- 
posed to control pest populations of wild 
rodents (Gao and Short, 1993). 

Human reproductive technology ad- 
dressed birth control as a first priority and 
achieved this in the 1950s; control of the 
human population had been established as 
a goal of public planning (Austin and Short, 
1986). Recently, immunological techniques 
have been applied to fertility control in the 
form of the “pregnancy vaccine” (Wang and 
Heap, 1992). Enhancement of fertility rep- 
resents a growing enterprise of the 1980s 
and 1990s, with in vitro fertilization and 
manipulations of embryos becoming more 
important bases of attempted therapies. Fi- 


REPRODUCTION 285 


nally, and with even greater ethical reser- 
vations, we are moving in the 1990s in the 
direction of genetic manipulations, gender 
manipulations, and transgenic innovation. 
Clearly the creativity of scientists and the 
demands of at least some members of so- 
ciety will drive us further. In this realm our 
ethical and legal systems have much catch- 
ing up to do, as we struggle to deal with 
‘‘what science has wrought.” 


Natural History and the Future 


Certainly scientific cleverness and crea- 
tivity will spur us on to new vistas in re- 
productive biology. Approaching the end of 
the century, we are well equipped with tech- 
nological potential to make new discover- 
ies. It is reassuring to know that natural his- 
tory and biodiversity remain part of the stuff 
from which we can extract discoveries. For 
example, this 75th anniversary year of the 
ASM we will learn of something that seems 
to violate a simple generality of mammalian 
parental care standards, and it was discov- 
ered serendipitously by unsuspecting inves- 
tigators in Malaysia, who had set up mist 
nets for birds (Francis et al., 1994). The 
discovery was a population of fruit bats (Dy- 
acopterus spadiceus) with males that had 
actively lactating mammary glands, yet also, 
later discovered, actively spermatogenic 
testes. Nature will certainly continue to sur- 
prise us and teach us, even as we enter the 
21st Century. 

We hope that the present historical syn- 
opsis of some of the highlights of mam- 
malian reproductive biology over the past 
75 years will offer some insights to mam- 
malogists both young and old. From the 
standpoint of the ASM, some aspects of the 
early beginnings were provincially North 
American in scientific character. Another 
important trend in the history of science, 
along with the modernization of travel and 
communication, has been the internation- 
alization of science. As modern scientists 
we have much available in the way of sci- 


entific resources to enhance our future pur- 
suits and a whole world in which to do so, 
yet as mammalogists we also have our an- 
imals. Being oriented to the biology of the 
Class Mammalia, we can distinguish our- 
selves by continuing to seek the insights that 
will come from continued attention to these 
animals and their natural history and di- 
versity. 


Acknowledgments 


We thank K. Benson and M. Wake for histor- 
ical references and A. Enders for providing Fig- 
ure 3: 


Literature Cited 


ALLEN, E. 1932. Sex and internal secretions. First ed. 
Williams and Wilkins, Baltimore, 951 pp. 

. 1939. Sex and internal secretions. Second ed. 
Williams and Wilkins, Baltimore, 1,346 pp. 

ASDELL, S. A. 1946. Patterns of mammalian repro- 
duction. Comstock Publishing Company, Ithaca, New 
York, 437 pp. 

Austin, C. R., AND R. V. SHoRT. 1972-1986. Re- 
production in mammals. First and second eds. Cam- 
bridge University Press, New York, Books 1-8. 

1986. Reproduction in mammals. Manipu- 
lating reproduction. Cambridge University Press, 
New York, Book 5, 235 pp. 

Benson, K.R. 1987. H. Newell Martin, W. K. Brooks, 
and the reformation of American biology. American 
Zoologist, 27:759-771. 

Bercer, P. J., N. C. NeGus, E. H. SANDERS, AND P. D. 
GARDNER. 1981. Chemical triggering of reproduc- 
tion in Microtus montanus. Science, 214:69-70. 

BeTTERIDGE, K. J. 1986. Increasing productivity in 
farm animals. Pp. 1-47, in Reproduction in mam- 
mals. Manipulating reproduction (C. R. Austin and 
R. V. Short, eds.). Cambridge University Press, New 
York, Book 5, 235 pp. 

BINKLEY, S. 1988. The pineal: endocrine and nonen- 
docrine functions. Prentice-Hall, Englewood Cliffs, 
New Jersey, 304 pp. 

BLim, V. 1986. Vertebrate reproduction. Springer- 
Verlag, Berlin, 405 pp. 

Bropy,S. 1945. Bioenergetics and growth. Reinhold 
Publishing Corporation, New York, 1,023 pp. 

Bronson, F. H. 1989. Mammalian reproductive bi- 
ology. University of Chicago Press, Chicago, 325 pp. 

Bruce, H. M. 1960. A block to pregnancy in the 
mouse caused by the proximity of strange males. 
Journal of Reproduction and Fertility, 1:96-103. 

BurRELL, H. 1927. The platypus. Angus and Rob- 
ertson, Sydney, 227 pp. 

CHRISTIAN, J. J., AND D. E. Davis. 1956. The rela- 


286 PEARSON AND KENAGY 


tionship between adrenal weights and population 
status in urban Norway rats. Journal of Mammalogy, 
37:475—-486. 

CLARKE, A. E. 1987. Research materials and repro- 
ductive science in the United States, 1910-1940. Pp. 
323-350, in Physiology in the American context, 
1850-1940 (G. L. Geison, ed.). American Physio- 
logical Society, Bethesda, Maryland, 403 pp. 

Crews, D. 1992. Behavioral endocrinology and re- 
production: an evolutionary perspective. Oxford Re- 
views in Reproductive Biology, 16:303-370. 

Daan, S., D. MASMAN, A. M. STRUKSTRA, AND G. J. 
KENAGY. 1991. Daily energy turnover during re- 
production in birds and mammals: its relationship 
to basal metabolic rate. Acta XX Congressus Inter- 
nationalis Ornithologici: 1976-1987. 

Drxson, A. F. 1983. The hormonal control of sexual 
behavior in primates. Oxford Reviews of Repro- 
ductive Biology, 5:131-219. 

EISENBERG, J. F., AND J. O. WoLFr. 1994. Behavior. 
Pp. 398-420, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication. The American Society of Mam- 
malogists, 1 1:1-433. 

Evans, H. M., AND H. H. Cote. 1931. An introduc- 
tion to the study of the oestrous cycle of the dog. 
Memoirs of the University of California, 9:66-118. 

EveERETT, J. W. 1988. Pituitary and hypothalamus: 
perspectives and overview. Pp. 1143-1233, in The 
physiology of reproduction (E. Knobil and J. D. Neill, 
eds.). Raven Press, New York (2 vols.), 2,413 pp. 

FARNER, D. S. 1985. Annual rhythms. Annual Re- 
view of Physiology, 47:65-82. 

Francis, C. M., E. L. P. ANTHONY, J. A. BRUNTON, 
AND T.H. Kunz. 1994. Lactation in male fruit bats 
(Pteropodidae). Nature. 

FRIEDMAN, M. H., AND G. S. FRIEDMAN. 1939. Go- 
nadotrophic extracts from green leaves. American 
Journal of Physiology, 125:486—-490. 

Gao, Y., AND R. V. SHorT. 1993. Use of an oestro- 
gen, androgen or gestagen as a potential chemoster- 
ilant for control of rat and mouse populations. Jour- 
nal of Reproduction and Fertility, 97:39-49. 

Gwinner, E. 1986. Circannual rhythms. Springer- 
Verlag, Berlin, 154 pp. 

HamILtTon, W. J., J. D. Boyp, AND H. W. MossMan. 
1945. Human embryology. Williams and Wilkins, 
Baltimore, 366 pp. 

HAyssEeN, V., ARI VAN TIENHOVEN, AND ANS VAN 
TIENHOVEN. 1993. Asdell’s patterns of mammalian 
reproduction. A compendium of species-specific data. 
Comstock Publishing Associates, Ithaca, New York, 
1,023 pp. 

HEAPE, W. 1901. The “sexual season” of mammals 
and the relation of “pro-estrum”’ to menstruation. 
Quarterly Journal of Microscopic Science, 44:1-70. 

HOFFMANN, K. 1981. Photoperiodism in vertebrates. 
Pp. 449-473, in Handbook of behavioral neuro- 
biology. Biological rhythms (J. Aschoff, ed.). Plenum 
Press, New York, volume 4, 563 pp. 

PAnson, H., D. L. Foster, G. R. Foxcrorr, AND P. 
J. Bootu. 1991. Nutrition and reproduction. Ox- 
ford Reviews of Reproductive Biology, 13:239-311. 

JAMESON, E. W. 1988. Vertebrate reproduction. Wi- 
ley, New York, 526 pp. 


KENAGY, G. J., AND G. A. BARTHOLOMEW. 1985. Sea- 
sonal reproductive patterns in five coexisting Cali- 
fornia desert rodent species. Ecological Monographs, 
55:371-397. 

Kenacy, G. J., D. MASMAN, S. M. SHARBAUGH, AND 
K. A. NAGy. 1990. Energy expenditure during lac- 
tation in relation to litter size in free-living golden- 
mantled ground squirrels. Journal of Animal Ecol- 
ogy, 59:73-88. 

KerBer, M. 1961. The fire of life. John Wiley & Sons, 
Inc., New York, 454 pp. 

KNoBIL, E., AND J. D. NEILL (EDs.). 1988. The phys- 
1ology of reproduction. Raven Press, New York, 1:1- 
1390 + 90, 2:1393-2413 + 90. 

LAMMING, G. E. 1984. Marshall’s physiology of re- 
production. Volume |, Reproductive cycles of ver- 
tebrates. Fourth ed. Churchill Livingstone, London, 
842 pp. 

Leg, A. K., AND I. R. MCDONALD. 1985. Stress and 
population regulation in small mammals. Oxford 
Reviews in Reproductive Biology, 7:261-304. 

Lona, J. A., AND H. M. Evans. 1922. The oestrous 
cycle in the rat and its associated phenomena. Mem- 
oirs of the University of California, 6:1-148. 

Loupon, A. S. I., AND P. A. RAcey. 1987. Repro- 
ductive energetics in mammals. Symposia of the 
Zoological Society (London), 57:1-371. 

McLaren, A. 1991. Sex determination in mammals. 
Oxford Reviews of Reproductive Biology, 13:1-33. 

MALL, F. P. 1915. On the fate of the human embryo 
in tubal pregnancy. Contributions to Embryology, 
1:3-104. 

MarsHALL, F.H. A. 1910. The physiology of repro- 
duction. First ed. Longmans, Green and Company, 
London, 706 pp. 

1922. The physiology of reproduction. Sec- 

ond ed. Longmans, Green and Company, London, 

770 pp. 

1925. An introduction to sexual physiology 
for biological, medical and agricultural students. 
Longmans, Green and Company, London, 167 pp. 

MossMANn, H. W. 1987. Vertebrate fetal membranes. 
Rutgers University Press, New Brunswick, New Jer- 
sey, 383 pp. 

MossMan, H. W., AND K. DuKE. 1973. Comparative 
morphology of the mammalian ovary. University of 
Wisconsin Press, Madison, 461 pp. 

NALBANDOv, A. V. 1958. Reproductive physiology: 
comparative reproductive physiology of domestic 
animals, laboratory animals and man. Freeman, San 
Francisco, 271 pp. 

OFTEDAL, O. T. 1984. Milk composition, milk yield 
and energy output at peak lactation: a comparative 
review. Symposia of the Zoological Society (Lon- 
don), 51:33-85. 

Parkes, A. S. 1969. Foreword: Dr. Herbert McLean 
Evans: an interview; selected publications; abstracts 
from Anatomical Record. Journal of Reproduction 
and Fertility, 19:1-49. 

PEARSON, O. P. 1944. Reproduction in the shrew 
(Blarina brevicauda Say). American Journal of Anat- 
omy, 75:39-93. 

PurseEL, V. G., ET AL. 1989. Genetic engineering of 
livestock. Science, 244:1281-1288. 

Quay, W. B. 1956. Volumetric and cytologic vari- 


REPRODUCTION 207 


ation in the pineal body of Peromyscus leucopus (Ro- 
dentia) with respect to sex, captivity and day length. 
Journal of Morphology, 98:471-495. 

REITER, R. J. (ED.). 1984. The pineal gland. Raven 
Press, New York, 382 pp. 

RENFREE, M. B., AND J. H. CALABy. 1981. Back- 
ground to delayed implantation and embryonic dia- 
pause. Journal of Reproduction and Fertility (Sup- 
plement), 29:1-9. 

RopGer, J. C., T. P. FLETCHER, AND C. H. 
TYNDALE-BiscozE. 1985. Active anti-paternal im- 
munization does not affect the success of marsupial 
pregnancy. Journal of Reproductive Immunology, 
8:249-256. 

Rusak, B., AND L. P. Morin. 1976. Testicular re- 
sponses to photoperiod are blocked by lesions of the 
suprachiasmatic nuclei in golden hamsters. Biology 
of Reproduction, 15:366-374. 

SADLEIR, R. M. F. S. 1969. The ecology of repro- 
duction in wild and domestic mammals. Methuen, 
London, 321 pp. 

1973. The reproduction of vertebrates. Ac- 
ademic Press, New York, 227 pp. 

SAPOLSKY, R. M. 1992. Stress, the aging brain, and 
the mechanisms of neuron death. MIT Press, Cam- 
bridge, Massachusetts, 429 pp. 

SeLye, H. 1936. A syndrome produced by diverse 
nocuous agents. Nature, 138:32. 

SHARMAN, G. B. 1955. Studies on marsupial repro- 
duction. II]. Normal and delayed pregnancy in Se- 
tonix brachyurus. Australian Journal of Zoology, 
3:56-70. 

SHORT, R. V. 1992. Elephants and birth control. New 
Scientist, 1 August, 21-23. 


1994. Why sex? Pp. 3-22, in The differences 
between the sexes (R. V. Short and E. Balaban, eds.). 
Cambridge University Press, Cambridge, 465 pp. 

STETSON, M. H., AND M. WATSON-WHITMYRE. 1976. 
Nucleus suprachiasmaticus: the biological clock in 
the hamster? Science, 191:197-199. 

STOCKARD, C. R., AND G. N. PAPANICOLAOU. 1917. 
The existence ofa typical oestrous cycle in the guinea 
pig—with a study ofits histological and physiological 
changes. American Journal of Anatomy, 22:225-265. 

TYNDALE-BiscogE, H., AND M. RENFREE. 1987. Re- 
productive physiology of marsupials. Cambridge 
University Press, Cambridge, 476 pp. 

VANDENBERGH, J. G. 1988. Pheromones and mam- 
malian reproduction. Pp. 1679-1696, in The phys- 
iology of reproduction (E. Knobil and J. D. Neill, 
eds.). Raven Press, New York, 1:1-1390 + 90, 2: 
1393-2413 + 90. 

VAN TIENHOVEN, A. 1968. Reproductive physiology 
of vertebrates. Saunders, Philadelphia, 498 pp. 

1983. Reproductive physiology of verte- 
brates. Second ed. Cornell University Press, Ithaca, 
New York, 491 pp. 

WANG, M.-W., AND R. B. Heap. 1992. Vaccination 
against pregnancy. Oxford Reviews of Reproductive 
Biology, 14:303-370. 

WINGFIELD, J. C., AND G. J. KENAGy. 1991. Natural 
regulation of reproductive cycles. Vertebrate Endo- 
crinology: Fundamentals and Biomedical Implica- 
tions, 4(B):181-241. 

Youna, W. C., (ED.) 1961. Sex and internal secre- 
tions. Third ed. Williams and Wilkins, Baltimore, 2 
volumes, 1,609 pp. 


MOLECULAR SYSTEMATICS 


RopDNEY L. HONEYCUTT AND TERRY L. YATES 


Introduction 


Be the founding of the American So- 
ciety of Mammalogists in 1919, Nut- 
tall (1904) wrote a paper on the use of blood 
immunology in comparative studies of an- 
imals. This paper was the prelude to later 
comparative serological papers (see Boy- 
den, 1942). By 1953 the molecular structure 
of DNA had been discovered (Watson and 
Crick, 1953), yet it was not until much later 
that systematists and evolutionary biolo- 
gists capitalized on this discovery and the 
earlier serological findings. Molecular sys- 
tematics is a young field that the founders 
of the ASM probably never imagined. Nev- 
ertheless, from the beginning, research on 
mammals played an important role in the 
development of the field of molecular sys- 
tematics, and in many cases mammalian 
taxa were used to investigate the patterns 
and processes of molecular evolution. It was 
not until the middle to late 1970s, however, 
that mammalogists began to use cladistic 
methodology and molecular characters in 
phylogeny reconstruction and the study of 
evolutionary processes. Once the applica- 
tion of these techniques began, the field of 
molecular mammalian systematics explod- 
ed and has been rapidly growing as a result 
of increased access to molecular techniques 
and computer technologies. 


The purpose of this chapter is to provide 
an historical account of mammalian mo- 
lecular systematics. We present this infor- 
mation in three parts. First, we describe sev- 
eral molecular techniques and discuss how 
these have been used in mammalian sys- 
tematics. Second, we discuss how mammals 
have been used to study the molecular evo- 
lutionary process, especially as it relates to 
the derivation of a molecular clock. Finally, 
we provide an overview of emerging issues 
and future directions in mammalian mo- 
lecular systematics. 


Molecular Techniques in 
Mammalian Systematics 


Protein Electrophoresis 


For the past 30 years, protein electropho- 
resis has been the most extensively used 
method by those interested in patterns of 
genetic variation within and between pop- 
ulations and species. The method allows for 
the recognition and quantification of allo- 
zyme differences for both enzymatic and 
nonenzymatic proteins. These differences 
are observed as changes in migration of pro- 
teins across an electric field, primarily as a 


MOLECULAR SYSTEMATICS 289 


consequence of changes in net charge (size 
and shape are minor factors as well) of the 
protein. These changes are genetically based 
and reflect underlying changes in amino acid 
sequence between products of alleles at the 
same locus. As a result, genetic variation at 
multiple loci can be examined and used as 
characters in comparative studies. 

The application of protein electrophoresis 
in evolutionary studies has been enhanced 
by a continual refinement of electrophoretic 
techniques (Harris and Hopkinson, 1976; 
Hunter and Markert, 1957; Murphy et al., 
1990; Selander et al., 1971; Shaw and Pra- 
sad, 1970). Two early technique papers, 
Harris and Hopkinson (1976) on humans 
and Selander et al. (1971) on Peromyscus 
polionotus, have continued to be important 
contributions to mammalogy because they 
provided the detailed conditions (e.g., stains 
and buffers) for examination of electropho- 
retic variation at many loci in mammalian 
species. Michael H. Smith was a coauthor 
on the Selander et al. (1971) paper, and he 
has continued to promote protein electro- 
phoresis by training and collaborating with 
a large number of mammalian systematists 
and evolutionary biologists. 

Most electrophoretic studies during the 
1960s and 1970s consisted of an examina- 
tion of genetic variation within populations 
and species, with an emphasis on popula- 
tion genetics and the selectionist versus neu- 
tralist controversy (Harris, 1966; Hubby and 
Lewontin, 1966; Hubby and Throckmor- 
ton, 1965). As indicated by Selander and 
Whittam (1983), the neutral theory provid- 
ed a null hypothesis for those interested in 
levels of diversity in structural genes. As a 
result, many of the earlier studies of allo- 
zyme variation attempted to interpret the 
observed levels of genetic heterozygosity and 
polymorphism found in species in light of 
neutral models as well as differences in se- 
lection pressures and life history strategies 
(Allendorf and Leary, 1986; Hedrick et al., 
1976; Lewontin, 1974: Nei and Graur, 1984; 
Nevo, 1978; Selander, 1977; Selander and 
Kaufman, 1973). Research on genetic vari- 


ation in mammals, using electrophoretic 
techniques, began in the middle 1960s and 
paralleled similar studies on other organ- 
isms. This research can be subdivided into: 
1) microevolutionary studies and studies of 
geographic variation; and 2) macroevolu- 
tionary studies. 

Microevolutionary studies. —The primary 
emphasis of early microevolutionary stud- 
ies ON mammals was on levels of genetic 
diversity within and between populations 
and the microevolutionary processes re- 
sponsible for the variation observed (e.g., 
random genetic drift, migration, population 
bottlenecks, selection). One of the more in- 
teresting debates pertaining to mammals re- 
lated to an interpretation of genetic varia- 
tion within and between populations and 
species, especially in fossorial mammals 
(Baccus et al., 1983; Kilpatrick and Crowell, 
1985; Nevo, 1985; Nevo and Shaw, 1972; 
Patton and Yang, 1977; Penny and Zim- 
merman, 1976; Sage et al., 1986; Schnell 
and Selander, 1981; Selander et al., 1974; 
Straney et al., 1976, 1979; Yates, 1983). Ev- 
itar Nevo and colleagues (Nevo, 1978, 1985; 
Nevo and Shaw, 1972; Nevo et al., 1974) 
found a correlation between biotic factors 
associated with the environment and allo- 
zyme polymorphism and heterozygosity in 
mammalian species, suggesting “‘adaptive 
relationships between genetic variability and 
spatial environmental heterogeneity.’ The 
low levels of genetic variation seen in fos- 
sorial mammals was interpreted as selection 
for homozygosity in a narrow subterranean 
niche. Other electrophoretic studies on pri- 
marily fossorial mammals disagreed with 
Nevo’s interpretations (Patton and Yang, 
1977; Penny and Zimmerman, 1976; Sage 
et al., 1986; Schnell and Selander, 1981; 
Selander et al., 1974). These studies re- 
vealed no positive relationship between 
“niche-width” and genetic variability in 
fossorial and non-fossorial mammals, sup- 
porting a more important role for historical 
factors related to fluctuating population size, 
founder events, and random drift. The data 


ZOU 


on this topic are still equivocal and little 
consensus has been achieved. 

Some of the more interesting studies of 
microgeographic variation in mammals us- 
ing electrophoresis have utilized genetic 
markers to examine both interactions be- 
tween hybridizing species or chromosome 
races (Baker et al., 1989a; Cothran and 
Zimmerman, 1985; Gentz and Yates, 1986: 
Greenbaum, 1981; Greenbaum and Baker, 
1976; Hafner et al., 1983; Heaney and 
Timm, 1985; Herd and Fenton, 1983; Nel- 
son et al., 1987; Patton et al., 1972, 1979a, 
19795; Smith and Patton, 1984; Sullivan et 
al., 1986) and the structure of mammalian 
populations as a result of social organization 
and dispersal patterns (Chesser, 1983; 
Gaines and Krebs, 1971; McCracken and 
Bradbury, 1977, 1981; Scribner et al., 1983; 
Smith et al., 1983; Wilkinson, 1985). The 
most extensive research on mammalian hy- 
brid zones has been conducted on hybrid- 
izing chromosomal races of Peromyscus leu- 
copus (Adkins et al., 1991; Baker et al., 1983: 
Nelson et al., 1987; Stangl, 1986), Uroder- 
ma bilobatum (Baker, 1981: Greenbaum, 
1981), and Geomys bursarius (Baker et al., 
1989a; Bradley et al., 1991a, 1991). These 
studies have characterized gene flow across 
hybrid zones using a combination of chro- 
mosomal and gene markers (both nuclear 
and mitochondrial) and have demonstrated 
that the dynamics of mammalian hybrid 
zones differ with respect to the origin of 
variation, the distribution of that variation, 
and the survival of hybrid individuals. 

Finally, patterns of both microgeographic 
and macrogeographic variation have been 
used in studies of threatened and endan- 
gered species of mammals (Bonnell and Se- 
lander, 1974; Chesser et al., 1980; Dragoo 
et al., 1990; Forman et al., 1986; Hafner 
and Yates, 1983; Hamilton et al., 1987; Kil- 
patrick et al., 1986; Newman et al., 1985; 
Sullivan and Yates, in press; Wayne et al., 
1986, 1991; Wayne and Jenks, 1991). Some 
of this research has focused on the overall 
level of genetic variation within species of 
mammals as a consequence of past popu- 


HONEYCUTT AND YATES 


lation bottlenecks and other demographic 
features, and other studies have attempted 
to discuss conservation issues (e.g., identi- 
fication of unique genetic stocks and deter- 
mination of population status) in light of 
observed levels of genetic variation. The 
classic studies by Stephen J. O’Brien and 
colleagues on genetic variation in the chee- 
tah (Newman et al., 1985; O’Brien et al., 
1983; 1985, 19874) and other felid species 
(O’Brien et al., 1986, 1987a, 1990: Packer 
et al., 1991; Roelke et al., 1993) have con- 
tributed greatly to our understanding of 
population bottlenecks and how genetics can 
be used in the conservation of mammalian 
species. These studies helped pave the way 
for a more routine use of genetic techniques 
and theory in conservation and manage- 
ment. 

Geographic variation.—Protein electro- 
phoresis also has been used to examine pat- 
terns of geographic variation in mammals, 
with the majority of the studies pertaining 
to patterns of variation in rodents. These 
geographic studies have focused on issues 
pertaining to the biogeographic history of 
relict populations (Hafner and Geluso, 1983; 
Smith et al., 1973), species such as pocket 
gophers that demonstrate fragmented pop- 
ulations and reduced gene flow (Hafner and 
Geluso, 1983; Hafner et al., 1987; Patton 
and Yang, 1977; Patton et al., 19795; Smith 
et al., 1983), species demonstrating a mon- 
tane distribution (Sullivan, 1985), the bio- 
geography of species that have a more ex- 
tended distribution (Nadler et al., 1973; 
Svoboda et al., 1985), and an examination 
of speciation patterns within a genus (Nevo 
et al., 1974; Patton, 1985). In more recent 
years, electrophoretic data have been com- 
bined with other genetic, morphologic, and 
ecologic data in an effort to identify recent 
or historical factors responsible for ob- 
served patterns of geographic variation (Av- 
ise et al., 1979c; Nelson et al., 1987; Nevo 
et al., 1993: Smith and Patton, 1988). 

Allozyme variation has been used to 
compare differences in the overall level of 
genetic variation between island and main- 


MOLECULAR SYSTEMATICS Pie 


land populations of the same species as well 
as taxa endemic to islands (Aquadro and 
Kilpatrick, 1981; Avise et al., 19744; Berry, 
1964; Kilpatrick, 1981; Patton, 1984). 
Again, most of these studies have involved 
rodent populations and, as indicted by Kil- 
patrick (1981), the overall pattern of vari- 
ation is one whereby insular populations are 
more monomorphic than mainland popu- 
lations. These results suggest that the level 
of variation on islands is related to the re- 
cency of colonization, the number of colo- 
nizations, the immigration rate between the 
island and mainland, and the effects of 
founder events and genetic drift. These con- 
clusions may also hold true for insular pop- 
ulations on continental land masses as well. 
Macroevolutionary studies.—As indicat- 
ed by Avise (1974) and Buth (1984), protein 
electrophoresis is a valuable tool for ad- 
dressing taxonomic issues in mammals and 
determining the relationships among taxa. 
A large number of electrophoretic studies 
have been used to identify species bound- 
aries, identify cryptic species, compare sib- 
ling species, and determine the taxonomic 
status of particular species (some of which 
are threatened or endangered; Dragoo et al., 
1990). For instance, Peter Baverstock and 
colleagues (Adams et al., 1982, 1987; Bav- 
erstock et al., 1977, 1983, 1984) have used 
protein electrophoresis to identify cryptic 
species of bats, rodents, and marsupials in 
Australia. Similar studies have been con- 
ducted on Nearctic mammal genera includ- 
ing Lasiurus (Baker et al., 1988), Geomys 
(Burns et al., 1985), Spermophilus (Cothran 
etal., 1977; Hafner and Yates, 1983; Nadler 
etal., 1982), Blarina (Tolliver and Robbins, 
1987), and insectivores in general (Tolliver 
et al., 1985). Some studies, such as those on 
Peromyscus comanche (Johnson and Pack- 
ard, 1974), Peromyscus hooperi (Schmidly 
etal., 1985), Peromyscus maniculatus/Pero- 
myscus melanotis (Bowers et al., 1973), and 
arid-land foxes (Dragoo et al., 1990) were 
taxonomically focused with the primary role 
being the determination of the taxonomic 
status of a particular population or race. 


Some of the earliest systematic studies 
employing protein electrophoresis per- 
tained to the derivation of phylogenetic re- 
lationships among mammalian taxa. The 
rodent genus Peromyscus has received con- 
siderable attention over the years (Avise et 
al., 1974a, 1974b, 1979c; Bowers et al., 1973; 
Kilpatrick and Zimmerman, 1975; Patton 
et al., 1981; Rennert and Kilpatrick, 1986; 
Robbins et al., 1985; Schmidly et al., 1985; 
Zimmerman et al., 1975, 1978), and elec- 
trophoresis has helped resolve many taxo- 
nomic problems within this diverse genus. 
Robert Baker and colleagues (Arnold et al., 
1982, 1983a; Baker et al., 1981; Koop and 
Baker, 1983) have conducted a large num- 
ber of electrophoretic studies on phyllos- 
tomoid bats, both within and among genera. 
These studies are significant because they 
incorporated a cladistic approach (outgroup 
approach of Baverstock et al., 1979; Patton 
and Avise, 1983; Patton et al., 1981) to the 
analysis of allozyme data. In addition, these 
studies examined phylogenetic hypotheses 
using multiple data sets and discussed issues 
of taxonomic congruence (see Mickevich 
and Johnson, 1976). These studies, in com- 
bination with immunological, chromosom- 
al, and morphological data, resulted in a 
revised phylogenetic classification for the 
bat family Phyllostomidae (Baker et al., 
19895). 

Phylogenetic studies also have been con- 
ducted on a large number of other mam- 
malian taxa, including rodents (Arnold et 
al., 1983; Best et al., 1986; Cook and Yates, 
in press; Hafner, 1982; Hafner et al., 1981; 
Honeycutt and Williams, 1982; Janecek et 
al., 1992; Johnson and Selander, 1971; Nel- 
son et al., 1984; Woods, 1982; Zimmerman 
and Nejtek, 1977), insectivores (George, 
1986; Yates and Greenbaum, 1982; Yates 
and Moore, 1990), and carnivores (Wayne 
and O’Brien, 1987). Although these studies 
vary in the analytical approach chosen, the 
resultant phylogenies have been used to ad- 
dress hypotheses related to the biogeogra- 
phy and speciation. In this regard, studies 
designed to examine coevolution among 


292 HONEYCUTT AND YATES 


mammalian hosts and their parasites are 
some of the more innovative in terms of 
using molecular phylogenies to examine 
evolutionary processes (Gardner, 1991; 
Hafner and Nadler, 1988, 1990: Reduker et 
al., 1987). 

Concluding remarks concerning electro- 
phoresis. — Protein electrophoresis is still the 
most cost-effective and rapid approach for 
assessing patterns of genetic variation, and 
it is very important in areas where little is 
known about the taxonomy of specific 
groups. In short, if one is interested in vari- 
ation within a genus, electrophoresis has 
been and will continue to be the best starting 
point for the assessment of genetic variation 
and species-level differences. Having said 
this, we must add that the analysis of allo- 
zyme data has changed significantly over 
the past 20 years. Phenetic analyses utilizing 
distance estimates (Nei, 1972; Rogers, 1972) 
and clustering approaches that assume rate 
constancy have been shown to be inappro- 
priate (Buth, 1984; Farris, 1972, 1985; Mi- 
yamoto and Cracraft, 1991; Swofford and 
Berlocher, 1987; Swofford and Olsen, 1990). 
Today, allozyme data can be analyzed more 
objectively using either distance approaches 
that do not assume rate constancy (Farris, 
1972; Felsenstein, 1982, 1990; Fitch and 
Margoliash, 1967) or cladistic approaches 
(Farris, 1988; Patton et al., 1981; Swofford, 
1990; Swofford and Berlocher, 1987; Swof- 
ford and Olsen, 1990) that treat loci (or al- 
leles) as character states. If one peruses the 
papers that have been published on mam- 
mals over the past 20 years, the trend to- 
ward a cladistic approach in phylogeny re- 
construction is apparent. 

Finally, one of the major contributions 
that an interest in allozyme variation con- 
tributed to mammalogy is the formation of 
frozen tissue collections at several major 
museums including: 1) Texas Cooperative 
Wildlife Collection, Texas A&M Univer- 
sity; 2) Museum of Vertebrate Zoology, 
University of California at Berkeley; 3) Mu- 
seum of Southwestern Biology, University 
of New Mexico; 4) The Museum, Texas Tech 


University; 5) San Diego Zoo; 6) Section of 
Mammals, Carnegie Museum of Natural 
History; and 7) Natural History Museum, 
Louisiana State University. In addition, 
there are a large number of laboratories that 
have considerable frozen tissue holdings. 
One can affirm that frozen tissue collections 
are today an important resource to the 
mammalogical community (Dessauer et al., 
1990) and more curatorial research, such as 
that conducted by Moore and Yates (1983), 
is needed. In addition, all those involved in 
studies that include collection of specimens, 
such as surveys and inventories, should be 
encouraged to collect tissue samples. Not 
only are resources limited and these speci- 
mens are valuable to continued molecular 
systematic efforts, but those values that ap- 
ply to long term storage and maintenance 
of more traditional museum specimens ap- 
ply to these specimens as well. 


Immunology 


One of the oldest molecular techniques 
for evaluating relationships among mam- 
malian taxa is comparative immunology 
(Boyden, 1942; Gerber and Leone, 1971; 
Goodman, 1963; Leone and Wiens, 1956; 
Levine and Moody, 1939; Nuttall, 1904), 
and this technique was perfected by Allan 
Wilson and Vincent Sarich at the University 
of California at Berkeley. Wilson, Sarich, 
and colleagues published a considerable 
number of papers on the rates of protein 
evolution and the relationships among 
mammals and other vertebrates (Carlson et 
al., 1978; Cronin and Sarich, 1975; Hafner, 
1982: Honeycutt and Sarich, 1987a, 1987b; 
Honeycutt et al., 1981; Maxson et al., 1975; 
Pierson et al., 1986; Sarich, 1969a, 19695, 
1973, 1977, 1985; Sarich and Cronin, 1976; 
Sarich and Wilson, 1967a, 1967b; Wilson 
and Sarich, 1969). Most of these studies dealt 
with intraordinal relationships among 
mammalian genera and families and em- 
ployed primarily the immunological tech- 
niques of precipitin and microcomplement 


MOLECULAR SYSTEMATICS 


fixation (MC’F). The two major molecules 
examined in these studies were albumin and 
transferrin, and an immunological distance, 
depicting the amount of amino acid differ- 
ence between molecules from different taxa, 
was determined. This quantitative estimate 
of immunological distance was determined 
by the degree of reactivity between anti- 
bodies and antigens from different species 
based on comparisons of homologous and 
heterologous reactions (Maxson and Max- 
son, 1990). 

In many cases both albumin and trans- 
ferrin were shown to evolve in a clocklike 
manner within mammalian orders, and the 
early studies on primates employed this 
clock in estimating divergence times for 
specific taxa such as the hominoid primates 
(Sarich and Wilson, 1967a, 1967b; Wilson 
and Sarich, 1969). One exception to the al- 
bumin clock was found by Arnold et al. 
(1982) and Honeycutt and Sarich (1987a) 
for phyllostomoid bats, with considerable 
rate heterogeneity observed among lineages. 
Although immunological distance data have 
been criticized (Farris, 1985), the overall 
usefulness of these data to mammalian sys- 
tematics has been verified, with phylogenies 
from albumin and transferrin being congru- 
ent, in most cases, with other molecular and 
non-molecular data (Arnold et al., 1982; 
Baker etal., 1989a; Dene et al., 1978; Prager 
and Wilson, in press; Sarich, 1985, in press; 
Sarich and Cronin, 1976). In at least two 
cases (Baker et al., 19895; Kirsch, 1977), 
the phylogenetic trees shown by immuno- 
logical data were used in combination with 
other data to revise the classification of 
mammalian groups. 


Amino Acid Sequences 


The most thorough molecular studies of 
interordinal relationships in mammals have 
been conducted by Morris Goodman, Jaap 
Beintema, Wilfried De Jong, and colleagues 
using amino acid sequence data from ap- 
proximately 10 polypeptides (Beintema et 


209 


al., 1973, 1991; Beintema and Lenstra, 1982: 
Czelusniak et al., 1990; De Jong, 1982; De 
Jong et al., 1977, 1981; Dene et al., 1982; 
Goodman, 1976a, 1976b; Goodman et al., 
1982, 1985, 1987; Miyamoto and Good- 
man, 1986; Romero-Herrera et al., 1978). 
One of the major strengths of amino acid 
sequences is that the data can be analyzed 
cladistically. A maximum parsimony pro- 
cedure was introduced by Moore et al. (1973) 
to find ancestral codons which minimize the 
number of mutations over a given network 
of species. This approach operates on the 
principle that the genetic code is redundant 
and, therefore, the number of possible co- 
dons at a particular node in a network will 
be minimized. The procedure works back- 
wards from a derived network and deter- 
mines ancestral codons for particular nodes. 
The overall objective of this procedure is to 
obtain a network or phylogeny of sequences 
that minimizes the total number of nucle- 
otide replacements (NR score). Goodman 
and colleagues have used this procedure for 
years to examine the relationships of eu- 
therian mammals and primate taxa. 

There have been criticisms of the maxi- 
mum parsimony approach used by Good- 
man (Kimura, 1981), as well as the resultant 
trees derived from this approach or amino 
acid sequence data in general (Wyss et al., 
1987). Goodman (1981) addressed some of 
Kimura’s original criticisms. Issues raised 
by Wyss et al. (1987), concerning incongru- 
ence among phylogenies derived from dif- 
ferent polypeptide sequences and between 
sequence phylogenies and those derived 
from morphological characters, are some- 
what harder to address. As indicated by 
Honeycutt and Adkins (1993), one critical 
problem with amino acid sequence data 1s 
that the numbers and kinds of taxa repre- 
sented by different genes vary. In addition, 
some genes are more conservative than oth- 
ers in terms of the overall amount of amino 
acid sequence differences between taxa, an 
observation related to functional con- 
straints on the molecule. Both of these fac- 


294 HONEYCUTT AND YATES 


tors may contribute to a certain amount of 
incongruence. 

More recent studies (Graur et al., 1991; 
Lietal., 1990, 1992) of relationships among 
eutherian orders still rely on amino acid se- 
quence data. In one case, the issue of rodent 
monophyly has been challenged (Graur et 
al., 1991; Li et al., 1992). Honeycutt and 
Adkins (1993) discussed these data at length 
and suggested that in all of these recent stud- 
ies the results are equivocal. 


Nucleotide Sequences 


Most recent studies on the molecular sys- 
tematics of mammals have focused on pat- 
terns of nucleotide sequence divergence in 
both the nuclear and mitochondrial ge- 
nomes, and advances in molecular tech- 
nology have made these studies consider- 
ably easier. These comparisons can be 
divided into two major categories, those us- 
ing indirect estimates of nucleotide se- 
quence divergence and those employing a 
direct sequencing method. 

DNA/DNA hybridization provides a 
quantitative estimate of sequence differ- 
ences between single copy nuclear DNAs 
from two or more taxa. This indirect esti- 
mate of sequence divergence is based on 
differences between the melting tempera- 
tures of a hybrid duplex DNA (heterodu- 
plex) and DNA from a single species (homo- 
duplex). The methodology used is based on 
earlier studies of reassociation kinetics 
(Britten and Kohne, 1968; Kohne et al., 
1972), and in recent years this method has 
been employed extensively in studies of bird 
phylogenies (Sibley and Ahlquist, 1981). In 
fact, Sibley and Ahlquist have published nu- 
merous papers on avian systematics and 
have even provided a classification of birds 
based upon their findings (Sibley et al., 
1988). 

The results and interpretations of DNA/ 
DNA hybridization studies have been chal- 
lenged by several individuals (Cracraft, 
1987; Sarich et al., 1989). Some of these 


criticisms arose in direct response to the 
findings of Sibley and Ahlquist (1984) on 
hominoid primate relationships. These crit- 
icisms pertained to the appropriateness of 
estimates of divergence based on T;)H, a 
measure of melting differences that includes 
the non-hybridizing portion of the melting 
profile. Although many of the issues raised 
by these criticisms have not been complete- 
ly answered, DNA/DNA hybridization 
studies have been conducted on mammals 
(Arnason and Widegren, 1986; Brownell, 
1983; Catzeflis et al., 1987; Kirsch et al., 
1990a, 19906, 1991, 1993; Springer and 
Kirsch, 1989, 1991; Springer and Krajews- 
ki, 1989). By far the most extensive research 
on mammals has been conducted by John 
Kirsch and colleagues at the University of 
Wisconsin on marsupials, and these studies 
have provided an excellent assessment of 
earlier criticisms and potential problems 
with the technique. 

Another indirect method of estimating 
nucleotide sequence divergence involves an 
examination of restriction site variation in 
mitochondrial genomes and nuclear genes 
(for details, see Melnick et al., 1992). In this 
technique, DNA is digested with restriction 
endonucleases that specify combinations of 
primarily four and six base pair sequences. 
These restriction endonucleases cleave at 
specific sites and, when digested, the DNA 
is separated by gel electrophoresis and ei- 
ther labelled directly in the case of mito- 
chondrial DNA (mtDNA) or probed with 
specific cloned DNA fragments. These re- 
sultant fragment patterns can be used di- 
rectly to estimate sequence divergence or 
converted to restriction site maps, making 
the estimate of sequence divergence more 
straightforward (for more details see Li and 
Graur, 1991; Melnick et al., 1992). 

The analysis of restriction fragment or site 
variation among mtDNAs has been the most 
popular approach in most studies involving 
mammals, and it is impossible to do justice 
in this review to the many studies that have 
been done. As indicated by several research- 
ers (Avise et al., 1984; Brown, 1983, 1985; 


MOLECULAR SYSTEMATICS 295 


Brown et al., 1979, 1982), mammalian 
mtDNA is maternally inherited and evolves, 
on average, much faster than nuclear genes. 
These features have made this molecule ex- 
ceedingly useful in studies of geographic 
variation and the biogeography of mam- 
mals (Avise et al., 1979a, 1979b, 1987; Cann 
etal., 1987; Patton and Smith, 1992; Riddle 
et al., 1993; Riddle and Honeycutt, 1990; 
Wayne et al., 1992), patterns of speciation 
(Nevo et al., 1993), interactions among hy- 
bridizing taxa (Baker et al., 1989a; Carr et 
al., 1986; Nelson et al., 1984), and phylo- 
genetic studies (Ferris et al., 1981, 1983; 
George and Ryder, 1986; Honeycutt et al., 
1987). Although Allan Wilson, Wesley 
Brown, Robert Lansman, and John Avise 
introduced the technique of restriction en- 
zyme analysis of mtDNA to evolutionary 
biologists, today there are laboratories all 
over the world involved in this type of re- 
search. 

Restriction site analysis of mammalian 
nuclear DNA has not been as extensive, with 
most studies focusing on the ribosomal DNA 
(rDNA) repeat (see Hillis and Dixon, 1991, 
for a review). In terms of mammalian stud- 
ies, two recent studies involving the higher 
level systematics of bats (Baker et al., 1991) 
and relationships among rodent taxa (AI- 
lard and Honeycutt, 1991) have been con- 
ducted. In both these studies, restriction site 
variation at the rDNA repeat provided little 
resolution, with most variation restricted to 
the nontranscribed spacer region. 

Direct sequencing of mammalian mito- 
chondrial and nuclear genes is fast becom- 
ing the method of choice for those interested 
in relationships at higher taxonomic levels 
(see review by Honeycutt and Adkins, 1993). 
By far, the bulk of data is from the mito- 
chondrial cytochrome c oxidase subunit II 
gene (Adkins and Honeycutt, 1991, in press; 
Disotell et al., 1992; Ruvolo et al., 1991), 
the cytochrome b gene (Irwin et al., 1991; 
Sudman and Hafner, 1992), the ND4 and 
NDS genes (Brown et al., 1982; Hayasaka 
etal., 1988), and the 12S and 16S ribosomal 
RNA genes (Allard and Honeycutt, 1992; 


Allard et al., 1991b, 1992: Hixson and 
Brown, 1986; Kraus and Miyamoto, 1991; 
Mindell et al., 1991; Miyamoto and Boyle, 
1989; Miyamoto et al., 1989, 1990). These 
data have been used to address questions 
pertaining to relationships among taxa 
within primarily the orders Primates, Ar- 
tiodactyla, and Rodentia, and in several 
cases issues pertaining to interordinal rela- 
tionships were addressed. Two of the more 
interesting debates concerning ordinal level 
relationships involved the question of chi- 
ropteran monophyly and relationships 
among orders in the superorder Archonta, 
and in these studies both nuclear and mi- 
tochondrial gene sequences were used to test 
conflicting hypotheses (Adkins and Honey- 
cutt, 1991; Ammerman and Hillis, 1992; 
Bailey et al., 1992; Honeycutt and Adkins, 
1993; Mindell et al., 1991; Stanhope et al., 
1992). 

Research in molecular systematics on 
mammals using nuclear gene sequences has 
lagged behind studies of mitochondrial gene 
sequences. The most extensive data exist for 
rDNA genes, and these data have consid- 
erable potential for higher level questions 
(Hillis and Dixon, 1991; Mindell and Hon- 
eycutt, 1990). One exception to the more 
extensive rDNA studies has been the con- 
sistent research efforts of Morris Goodman 
and colleagues with respect to determining 
the relationships among eutherian mam- 
malian orders using single copy genes or 
pseudogenes (Bailey et al., 1992; Koop and 
Goodman, 1988; Koop et al., 1986; Stan- 
hope et al., 1992). As indicated by Honey- 
cutt and Adkins (1993), morphology has not 
been able to resolve the relationships among 
eutherian orders (Novacek, 1992; Shoshani, 
1986; Simpson, 1945) and, if nucleotide se- 
quence data are to contribute to this issue, 
considerably more information is needed. 


Molecular Clock Concept 


The analysis of morphological change in 
mammals has revealed irregularity in the 


296 HONEYCUPY AND YATES 


evolutionary process, with different lineages 
demonstrating mosaic evolution in terms of 
the overall rate of morphological evolution. 
This mosaic evolution reflects the overall 
adaptive radiation observed for mammals, 
especially in terms of the diversity in form 
and function seen for higher categories. In 
contrast to phenotypic evolution, molecules 
(both proteins and nucleic acids) of mam- 
mals and other organisms presumably 
evolve in a neutral fashion, demonstrating 
a rather constant rate of change through 
evolutionary time and across diverse tax- 
onomic groups (Brown et al., 1982; Easteal, 
1985, 1990; Kimura, 1983; Sarich and Wil- 
son, 1967a, 1967b; Wilson et al., 1977; Zu- 
kerkandl and Pauling, 1965). Some of the 
principles of the neutral theory were derived 
to distinguish between evolution at the mor- 
phological and molecular level. These prin- 
ciples relate to both the elimination of del- 
eterious mutations and fixation of variation 
through selective neutrality as opposed to 
positive Darwinian selection and the over- 
all rate of evolution observed for particular 
molecules as a consequence of the level of 
structural and functional constraints placed 
on these molecules. 

An outgrowth of the neutral theory is the 
idea of a molecular clock, which sees the 
evolutionary process at the molecular level 
as arandom process with a constant average 
rate of change (Fitch and Langley, 1976; 
Kimura, 1983; Li and Graur, 1991; Wilson 
et al., 1977; Zukerkandl and Pauling, 1965). 
In fact, one might say that the observation 
of a molecular clock has provided support 
for the neutral theory. By necessity, the mo- 
lecular clock is a statistical clock, and it as- 
sumes a linear relationship between time 
since evolutionary divergence and molec- 
ular divergence. Obviously, the best test for 
a clock is one that evaluates the regularity 
of overall rates of divergence through time, 
and this test is best applied in a phylogenetic 
context (Fitch and Langley, 1976). 

When evaluating rates of molecular evo- 
lution, several analytical approaches can be 
applied. One approach, the relative rate test, 
first introduced by Sarich and Wilson 


(1967a, 1967b) and expanded upon by oth- 
ers (Li and Graur, 1991; Li et al., 1987; 
Mindell and Honeycutt, 1990; Wu and Li, 
1985), is a test for rate uniformity. It re- 
quires no knowledge of divergence times be- 
tween species but does presuppose branch- 
ing order in that an outside reference species 
or outgroup is required for the examination 
of lineages sharing a common point of di- 
vergence. The test is actually a comparison 
of the magnitude of change along two lin- 
eages subsequent to divergence from a com- 
mon ancestor. It has been suggested that 
more than one outside reference species be 
used to minimize the effects of back mu- 
tations and convergent substitutions (Bev- 
erley and Wilson, 1984). The effects of such 
homoplasy increase over evolutionary time, 
thus the need for several calibration points 
(Gingerich, 1986). 

Another method, the star phylogeny ap- 
proach (Kimura, 1983), is a test that con- 
siders a case where all species diverge at the 
same point in time from a common ancestor 
and compares the observed and expected 
variances in rate under the Poisson process. 
This approach might be valid for mam- 
malian orders but the estimates are proba- 
bly minimal as a result of dichotomous 
branching (Nei, 1987). Gillespie (1986) has 
modified this approach to take into account 
branching. 

Langley and Fitch (1974) introduced a 
third procedure that requires knowing the 
branching order. In this procedure expected 
branch lengths are calculated using maxi- 
mum likelihood, and then a test for rate 
heterogeneity is employed using chi-square 
analysis. 

Finally, the absolute rate can be estimated 
by calculating substitutions along each 
branch length in a phylogeny and calibrating 
the evolutionary rate based on dates from 
either the fossil record or biogeography 
(Beverly and Wilson, 1984; Maxson et al., 
1975; Sarich and Wilson, 1967a, 1967b). 

What is the evidence for a molecular 
clock? First, the evolutionary rate of diver- 
gence in amino acid sequence has been 
shown to be linear with time. This has been 


MOLECULAR SYSTEMATICS 207 


demonstrated for many proteins in mam- 
mals, including globins (Kimura, 1983; Li 
et al., 1985; Zukerkandl and Pauling, 1965). 
Although the overall rates between proteins 
may differ, this difference can be explained 
in terms of functional constraints and is 
consistent with the neutral theory (Kimura, 
1983). Second, a large body of data on al- 
bumin immunology in mammals has re- 
vealed an overall relationship between rate 
of divergence and time (Carlson et al., 1978; 
Sarich, 1977), and this albumin/transferrin 
clock has been used extensively in compar- 
isons of times of mammalian divergence. 
Finally, at the level of nucleotide sequence 
in both mitochondrial and nuclear genes, 
certain types of substitutions demonstrate 
clock-like behavior in terms of their diver- 
gence over time (Brown et al., 1982; Bulmer 
et al., 1991; Easteal, 1985, 1990; Hasegawa 
et al., 1985; Kimura, 1983; Mindell and 
Honeycutt, 1990; Miyamoto and Boyle, 
1989: Vawter and Brown, 1986). In mam- 
mals there also is evidence of clock-like be- 
havior of estimates of divergence derived 
from DNA/DNA hybridization (Catzeflis et 
al., 1987; Sibley and Ahlquist, 1984). 
Although there is some confirmation of 
rates of amino acid and nucleotide substi- 
tutions being linear with time, there are 
many exceptions that challenge the gener- 
ality of a molecular clock. First, differential 
rates of evolution have been observed for 
both nuclear and mitochondrial genes (Ad- 
kins and Honeycutt, 1991; Bajaj et al., 1984; 
Britten, 1986; Gillespie, 1991; Goodman et 
al., 1975; Holmes, 1991; Liand Graur, 1991; 
Li et al., 1985, 1987; Romero-Herrera et 
al., 1978; Wu and Li, 1985). Second, both 
distance estimates from DNA/DNA hy- 
bridization and synonymous substitution 
rates in genes suggest a generation time ef- 
fect for mammals and other animals in terms 
of overall rates of divergence at the level of 
nucleotide substitutions (Britten, 1986; Li 
and Graur, 1991; Li et al., 1985; Wu and 
Li, 1985). Recently, a relationship between 
substitution rate differences, body size, and 
metabolic rates in mammals and other or- 
ganisms has been found (Martin and Pal- 


umbi, 1993). Finally, in the case of an elec- 
trophoretic clock (Nei, 1971; Sarich, 1977; 
Smith and Coss, 1984), rates calculated from 
the same overall genetic distances from dif- 
ferent mammals and other organisms vary 
as much as 20-fold (Avise and Aquadro, 
1982). Therefore, the idea of using an al- 
bumin clock to set the electrophoretic clock 
is clearly suspect (Sarich, 1977). 

As Hills and Moritz (19905) pointed out, 
molecular divergence and time are corre- 
lated to an extent. The question, however, 
pertains to the amount of error associated 
with any time estimate derived from the 
magnitude of divergence separating taxa and 
the various means of clock calibration. In 
terms of the latter, paleontological and bio- 
geographical estimates of time since diver- 
gence have associated errors and, in addi- 
tion, using a calibrated rate from one set of 
taxa (e.g., between the rodent taxa Mus and 
Rattus) to determine time since divergence 
in an unrelated set of taxa (e.g., another or- 
der of mammals) can clearly create error if 
the overall rate or pattern of divergence dif- 
fers for the same gene between the two un- 
related groups. Although the error associ- 
ated with an estimate of absolute time can 
be great, assessments of relative rates of mo- 
lecular divergence are very useful to those 
interested in the processes of molecular evo- 
lution and the use of molecular characters 
in phylogeny reconstruction. Clearly, mam- 
mals provide an excellent model for study- 
ing either of these two aspects of evolution. 


Emerging Issues and Future 
Directions 


Several major developments over the past 
three decades have had a profound impact 
on systematic and evolutionary biology. 
First, cladistic analysis has become the pri- 
mary methodological approach used in phy- 
logeny reconstruction, and it has provided 
an objective framework for deriving clas- 
sifications, studying biogeography, and in- 
vestigating speciation, cospeciation, and 


298 HONEYCUTT AND YATES 


other evolutionary processes (Baker et al., 
1989a; Brooks and McLennan, 1991; El- 
dredge and Cracraft, 1980; Hafner and Nad- 
ler, 1988, 1990; McKenna, 1975; Riddle and 
Honeycutt, 1990). Second, the ability to test 
hypotheses pertaining to the patterns and 
processes of evolution have been enhanced 
by the development of more sophisticated 
analytical procedures and more accessible 
computer software and hardware (Farris, 
1988; Felsenstein, 1990; Miyamoto and 
Cracraft, 1991; Swofford, 1990; Swofford 
and Olsen, 1990). Third, genetics and mo- 
lecular biology have provided information 
that has broadened our view as to the role 
of selection and neutrality in the evolution- 
ary process (Gillespie, 1991; Kimura, 1983; 
Li and Graur, 1991; Nei, 1987). Finally, 
variation at the level of genes, gene prod- 
ucts, and nucleotide sequences has provided 
a suite of literally thousands of indepen- 
dently evolving characters that can be used 
to examine diversity within populations, 
species, and higher taxa (Hillis and Moritz, 
1990a; Honeycutt and Adkins, 1993). All 
of the above events have contributed di- 
rectly to the ever increasing use of molecular 
characters in systematic and evolutionary 
studies, and today molecular systematics and 
molecular evolution are two of the fastest 
growing areas of research in systematic and 
evolutionary biology. 

Recent advances in molecular biology 
have provided an easy-to-use set of tools 
for mammalogists interested in the origin 
and diversification of mammalian taxa. The 
polymerase chain reaction (Allard et al., 
1991a; Higuchi and Ochman, 1989; Kocher 
etal., 1989; Saiki et al., 1988) and improved 
methods for obtaining nucleotide sequence 
information (Maxam and Gilbert, 1980; 
Sanger et al., 1977) are revolutionizing the 
fields of molecular evolutionary biology and 
systematics. Literally thousands of molec- 
ular characters can be used to address ques- 
tions of higher level relationships among 
mammalian families and orders and, in 
combination with morphological data, one 
can begin to unravel the secret of the mam- 


malian radiations. One of the most exciting 
areas of research is the use of ancient DNA, 
extracted from museum specimens and fos- 
sils, to provide a historical perspective on 
the genetics of populations and the rela- 
tionships among extinct and extant forms 
of mammals (Higuchi et al., 1984; Paabo, 
1989; Paabo et al., 1988, 1989; Shoshani et 
al., 1985; Thomas et al., 1990). As these 
techniques become more refined, we may 
one day be able to address questions per- 
taining to the early origin of mammals. 

A major challenge to all mammalogists 
interested in molecular systematics pertains 
to data analysis, as can be seen by recent 
publications on the subject (Felsenstein, 
1981, 1984, 1988; Miyamoto and Cracraft, 
1991; Swofford and Olsen, 1990). This issue 
will become even more important as the 
amount of sequence data increases, and sev- 
eral questions pertaining to molecular data 
and the analysis of those data must be ad- 
dressed. Some of these questions are (for a 
more detailed discussion on mammalian 
molecular systematics see Honeycutt and 
Adkins, 1993): 1) What criteria should be 
used in selecting the correct molecule and 
experimental approach? 2) Should one use 
equal or unequal weighting schemes in an 
analysis of molecular data? 3) How impor- 
tant is the selection of an outgroup, and 
what criteria should be used in selecting out- 
groups? 4) Which methodological approach 
to estimating evolutionary trees should be 
used, and are there situations when one par- 
ticular method might be superior to the more 
accepted method? 5) How does one evaluate 
the reliability of trees derived from molec- 
ular sequences, and what factors can influ- 
ence the accuracy of a cladogram? and 6) 
How does one consider total evidence when 
evaluating phylogenetic hypotheses, and 
what are some explanations for incongru- 
ence among trees derived from different 
molecular and non-molecular characters? 

Finally, questions pertaining to the evo- 
lutionary process are being addressed using 
a phylogenetic framework (Brooks and 
McLennan, 1991). For instance, the orga- 


MOLECULAR SYSTEMATICS 


nization and evolution of communities are 
being examined using a combination of bio- 
geography, phylogenetics, and molecular 
characters (Avise et al., 1987; Riddle and 
Honeycutt, 1990; Riddle et al., 1993). As 
indicated earlier, the process of cospeciation 
is being studied by comparing the phylog- 
enies of both the mammalian hosts and their 
parasites (Hafner and Nadler, 1988, 1990; 
Reduker et al., 1987). Phylogenies also offer 
a means of evaluating the evolution of com- 
plex behavior in mammals (Honeycutt, 
1992). Aside from questions pertaining to 
organismal evolution, gene trees derived 
from mammals offer a means of examining 
convergent evolution at the molecular level 
(Stewart and Wilson, 1987) and the mech- 
anisms responsible for producing variation 
(Bradley et al., 1993). Interest in all these 
areas will increase in the future, and as our 
knowledge of the molecular genetics of the 
developmental process increases, we may 
begin to examine the origin of morpholog- 
ical form and function of mammals by 
studying the underlying patterns of devel- 
opment at the level of genes and gene prod- 
ucts. 


Acknowledgments 


We thank R. D. Bradley, J. Salazar-Bravo, and 
an anonymous reviewer for helpful comments 
on this manuscript. 


Literature Cited 


Apams, M., P. R. BAverstTock, C. H. S. WATTS, AND 
T. REARDON. 1987. Electrophoretic resolution of 
species boundaries in Australian Microchiroptera. 
II. The Pipistrellus group (Chiroptera: Vespertilion- 
idae). Australian Journal of Biological Science, 40: 
163-170. 

Apams, M., P. R. BAvERSTOCK, C. R. TIDEMANN, AND 
D. P. Woopsipe. 1982. Large genetic differences 
between sibling species of bats, Eptesicus, from Aus- 
tralia. Heredity, 48:435-438. 

Apkins, R. M., AND R. L. Honeycutr. 1991. Mo- 
lecular phylogeny of the superorder Archonta. Pro- 


209 


ceedings of the National Academy of Sciences (USA), 

88:10317-10321. 

In press. Evolution of the primate cyto- 
chrome c oxidase subunit II gene. Journal of Mo- 
lecular Evolution, 38. 

ADKINS, R. M., K. McBEgE, C. A. PorTER, AND R. J. 
BAKER. 1991. Hybrid breakdown in Peromyscus 
leucopus and examination of the recombinational— 
breakdown model. Journal of Mammalogy, 72:535- 
541. 

ALLARD, M. W., AND R. L. Honeycutt. 1991. Ri- 
bosomal DNA variation within and between species 
of rodents, with emphasis on the genus Onychomys. 
Molecular Biology and Evolution, 8:71-84. 

ALLARD, M. W., AND R. L. Honeycutt. 1992. Nu- 
cleotide sequence variation in the mitochondrial 12S 
tRNA gene and the phylogeny of African mole-rats 
(Rodentia: Bathyergidae). Molecular Biology and 
Evolution, 9:27-40. 

ALLARD, M. W., D. L. ELLSworTH, AND R. L. 
Honeycutr. 199la. The production of single- 
stranded DNA suitable for sequencing using the 
polymerase chain reaction. BioTechniques, 10:24- 
26. 

ALLARD, M. W., M. M. Miyamoto, AND R. L. 
Honeycutt. 19916. Tests for rodent polyphyly. 
Nature, 353:610-611. 

ALLARD, M. W., M. M. MryAmoto, L. JARECKI, F. 
Kraus, AND M. R. TENNANT. 1992. DNA system- 
atics and evolution of the artiodactyl family Bovidae. 
Proceedings of the National Academy of Sciences 
(USA), 89:3972-3976. 

ALLENDORF, F. W., AND R. F. LEARY. 1986. Hetero- 
zygosity and fitness in natural populations of ani- 
mals. Pp. 57-76, in Conservation biology: the sci- 
ence of scarcity and diversity (M. E. Soule, ed.). 
Sinauer Associates, Sunderland, 584 pp. 

AMMERMAN, L. K., AND D. M. Huis. 1992. A mo- 
lecular test of bat relationships: monophyly or di- 
phyly? Systematic Biology, 41:222-—232. 

AQUADRO, D. F., AND C. W. KILPATRICK. 1981. Mor- 
phological and biochemical variation and differen- 
tiation in insular and mainland deer mice (Pero- 
myscus maniculatus). Pp. 214-230, in Mammalian 
population genetics (M. H. Smith and J. Joule, eds.). 
University of Georgia Press, Athens, 380 pp. 

ARNASON, U., AND B. WIDEGREN. 1986. Pinniped 
phylogeny enlightened by molecular hybridization 
using highly repetitive DNA. Molecular Biology and 
Evolution, 3:356-365. 

ARNOLD, M. L., R. J. BAKER, AND R. L. HONEYCUTT. 
1983a. Genic differentiation and phylogenetic re- 
lationships within two New World bat genera. Bio- 
chemical Systematics and Ecology, 1 1:295-303. 

ARNOLD, M. L., L. W. Rossrns, R. K. CHESSER, AND 
J. C. Patron. 1983. Phylogenetic relationships 
among six species of Reithrodontomys. Journal of 
Mammalogy, 64:1 28-132. 

ARNOLD, M. L., R. L. HONEycuTT, R. J. BAKER, V. M. 
SARICH, AND J. K. Jones, JR. 1982. Resolving a 
phylogeny with multiple data sets: a systematic study 
of phyllostomoid bats. Occasional Papers, The Mu- 
seum, Texas Tech University, 77:1-15. 

AvIsE, J.C. 1974. Systematic value of electrophoretic 
data. Systematic Zoology, 23:137-148. 


300 HONEYCUTT AND YATES 


AvisE, J. C., AND C. F. AQUADRO. 1982. A compar- 
ative summary of genetic distances in the verte- 
brates. Patterns and correlations. Evolutionary Bi- 
ology, 15:151-185. 

AvliseE, J.C., R. A. LANSMAN, ANDR. O. SHADE. 19790. 
The use of restriction endonucleases to measure mi- 
tochondrial DNA sequence relatedness in natural 
populations. I. Population structure and evolution 
in the genus Peromyscus. Genetics, 92:279-295. 

AvIsE, J. C., J. E. NEIGEL, AND J. ARNOLD. 1984. De- 
mographic influences on mitochondrial DNA lin- 
eage survivorship in animal populations. Journal of 
Molecular Evolution, 20:99-105. 

AvIsE, J. C., M. H. SMITH, AND R. K. SELANDER. 1974a. 
Biochemical polymorphism and systematics in the 
genus Peromyscus. VI. The boylii species group. 
Journal of Mammalogy, 55:751-763. 

AvIsE, J. C., M. H. SMITH, AND R. K. SELANDER. 1979c. 
Biochemical polymorphism and systematics in the 
genus Peromyscus. VII. Geographic differentiation 
in members of the truei and maniculatus species 
groups. Journal of Mammalogy, 60:177-192. 

AvIsE, J. C., C. GIBLIN-DAvIDson, J. LAERM, J. C. 
PATTON, AND R. A. LANSMAN. 1979a. Mitochon- 
drial DNA clones and matriarchal phylogeny within 
and among geographic populations of the pocket go- 
pher, Geomys pinetis. Proceedings of the National 
Academy of Sciences (USA), 76:6694-6698. 

AvisE, J. C., J. F. SHAPIRA, S. W. DANIEL, C. F. AQUAD- 
RO, ANDR. A. LANSMAN. 1983. Mitochondrial DNA 
differentiation during the speciation process in Pero- 
myscus. Molecular Biology and Evolution, 1:38-56. 

AvisE, J. C., M. H. Smitu, R. K. SELANDER, T. E. 
LAWLOR, AND P. R. RAMsEy. 19746. Biochemical 
polymorphism and systematics in the genus Pero- 
myscus. V. Insular and mainland species of the sub- 
genus Haplomylomys. Systematic Zoology, 23:226— 
238. 

AvisE, J. C., ET AL. 1987. Intraspecific phylogeogra- 
phy: the mitochondrial bridge between population 
genetics and systematics. Annual Review of Ecology 
and Systematics, 18:489-522. 

Baccus, R., N. RYMAN, M. H. SmitH, C. REUTERWALL, 
AND D. CAMERON. 1983. Genetic variability and 
differentiation of large grazing mammals. Journal of 
Mammalogy, 64:109-120. 

BaILey, W. J., J. L. SuuGHTOM, AND M. GOODMAN. 
1992. Rejection of the “flying primate” hypothesis 
by phylogenetic evidence from the e-globin gene. 
Science, 256:86-89. 

Basas, M., T. BLUNDELL, AND S. Woop. 1984. Evo- 
lution in the insulin family: molecular clocks that 
tell the wrong time. Biochemical Society Sympo- 
sium, 49:45—54. 

BAKER, R. J. 1981. Chromosome flow between chro- 
mosomally characterized taxa of a volant mammal, 
Uroderma bilobatum (Chiroptera: Phyllostomati- 
dae). Evolution, 35:296—-305. 

BAKER, R. J., R. L. HONEYcUTT, AND R. A. VAN DEN 
BusscHE. 1991. Examination of monophyly of bats: 
restriction map of the ribosomal DNA cistron. Bul- 
letin of the American Museum of Natural History, 
206:42-53. 

BAKER, R. J., C. S. Hoop, AND R. L. HONEycUuUTT. 
1989b. Phylogenetic relationships and classification 


of the higher categories of the New World bat family 
Phyllostomidae. Systematic Zoology, 38:228—238. 

Baker, R. J., J. C. PATron, H. H. GENoways, AND J. 
W. BICKHAM. 1988. Genic studies of Lasiurus (Chi- 
roptera: Vespertilionidae). Occasional Papers, The 
Museum, Texas Tech University, 117:1-15. 

BAKER, R. J., L. W. Rossins, F. B. STANGL, JR., AND 
E. C. Birney. 1983. Chromosomal evidence for a 
major subdivision in Peromyscus leucopus. Journal 
of Mammalogy, 64:356-359. 

BAKER, R. J.,S. K. Davis, R. D. BRADLEY, M. J. HAM- 
ILTON, AND R. A. VAN DEN BusSCHE. 1989a. Ri- 
bosomal-DNA, mitochondrial-DNA, chromosom- 
al, and allozymic studies on a contact zone in the 
pocket gopher, Geomys. Evolution, 43:63-75. 

BAKER, R. J., R. L. Honeycutt, M. L. ARNOLD, V. M. 
SARICH, AND H. H. GeNoways. 1981. Electropho- 
retic and immunological studies of the relationship 
of the Brachyphyllinae and the Glossophaginae. 
Journal of Mammalogy, 62:665-672. 

BAVERSTOCK, P. R., M. ADAMS, AND M. ARCHER. 1984. 
Electrophoretic resolution of species boundaries in 
the Smithopsis murina complex (Dasyuridae). Aus- 
tralian Journal of Zoology, 32:823-832. 

BAVERSTOCK, P. R., C. H. S. Watts, AND S. R. COLE. 
1977. Electrophoretic comparisons between the al- 
lopatric populations of five Australian pseudomyine 
rodents (Muridae). Australian Journal of Biological 
Science, 30:471-485. 

BAVERSTOCK, P. R., S. R. Coe, B. J. RICHARDSON, AND 
C.H. Watts. 1979. Electrophoresis and cladistics. 
Systematic Zoology, 28:214-219. 

BAVERSTOCK, P. R., M. ApAmMs, M. ARcHER, N. L. 
McKENZIE, AND R. How. 1983. An electrophoretic 
and chromosomal study of the dasyurid marsupial 
genus Ningaui archer. Australian Journal of Zoology, 
31:381-392. 

BEINTEMA, J. J., AND J. A. LENsTRA. 1982. Evolution 
of mammalian pancreatic ribonucleases. Pp. 43-73, 
in Macromolecular sequences in systematics and 
evolutionary biology (M. Goodman, ed.). Plenum 
Press, New York, 418 pp. 

BEINTEMA, J. J., K. RODEWALD, G. BRAUNITZER, J. CZE- 
LUSNIAK, AND M. GOODMAN. 1991. Studies on the 
phylogenetic position of the Ctenodactylidae (Ro- 
dentia). Molecular Biology and Evolution, 8:151- 
154. 

BEINTEMA, J. J., A. J. SCHEFFER, H. VAN Duk, G. W. 
WELLING, AND H. Zwiers. 1973. Pancreatic ribo- 
nuclease: distribution and comparisons in mam- 
mals. Nature New Biology, 241:76-78. 

Berry, R. J. 1964. The evolution of an island pop- 
ulation of the house mouse. Evolution, 18:468-483. 

Best, T. L., R. M. SULLIVAN, J. A. Cook, AND T. L. 
Yates. 1986. Chromosomal, genic, and morpho- 
logic variation in the agile kangaroo rat, Dipodomys 
agilis (Rodentia: Heteromyidae). Systematic Zool- 
ogy, 35:311-324. 

BEVERLEY, S. M., ANDA.C. Witson. 1984. Molecular 
evolution of Drosophila and the higher Diptera. II. 
A time scale for fly evolution. Journal of Molecular 
Evolution, 21:1-13. 

BONNELL, M. L., AND R. K. SELANDER. 1974. Ele- 
phant seals: genetic variation and near extinction. 
Science, 184:908-909. 


MOLECULAR SYSTEMATICS 301 


Bowers, J. H., R. J. BAKER, AND M. H. Situ. 1973. 
Chromosomal, electrophoretic, and breeding studies 
of selected populations of deer mice (Peromyscus 
maniculatus) and black-eared mice (P. melanotis). 
Journal of Mammalogy, 27:378-386. 

Boypen, A. 1942. Systematic serology: a critical ap- 
preciation. Part I. Systematic serology and its rela- 
tion to general biology. Physiological Zoology, 25: 
109-145. 

BRADLEY, R. D., J. J. Butt, A. D. JOHNSON, AND D. 
M. Hirus. 1993. Origin ofa novel allele in a mam- 
malian hybrid zone. Proceedings of the National 
Academy of Sciences (USA), 90:8939-8941. 

BRADLEY, R. D.,S. K. DAvis, AND R. J. BAKER. 1991 a. 
Genetic control of premating-isolating behavior: Ka- 
neshiro’s hypothesis and asymmetrical sexual selec- 
tion in pocket gophers. Journal of Heredity, 82:192- 
196. 

BRADLEY, R. D., S. K. Davis, S. F. Lockwoop, J. W. 
BICKHAM, AND R. J. BAKER. 19916. Hybrid break- 
down and cellular-DNA content in a contact zone 
between two species of pocket gophers (Geomys). 
Journal of Mammalogy, 72:697-705. 

Britten, R. J. 1986. Rates of DNA sequence evo- 
lution differ between taxonomic groups. Science, 231: 
1393-1398. 

Britten, R. J., AND D. E. KoHNE. 1968. Repeated 
sequences in DNA. Science, 161:529-540. 

Brooks, D. R., AND D. A. MCLENNAN. 1991. Phy- 
logeny, ecology, and behavior. University of Chicago 
Press, Chicago, 434 pp. 

Brown, W. M. 1983. Evolution of animal mito- 
chondrial DNA. Pp. 62-88, in Evolution of genes 
and proteins (M. Nei and R. K. Koehn, eds.). Sinauer 
Associates, Sunderland, Massachusetts, 331 pp. 

. 1985. The mitochondrial genome of animals. 
Pp. 95-130, in Molecular evolutionary genetics (R. 
MacIntyre, ed.). Plenum Press, New York, 610 pp. 

Brown, W. M., M. GeorGE, JR., AND A. C. WILSON. 
1979. Rapid evolution of animal mitochondrial 
DNA. Proceedings of the National Academy of Sci- 
ences (USA), 76:1967-1971. 

Brown, W. M., E. M. PRAGER, A. WANG, AND A. C. 
Witson. 1982. Mitochondrial DNA sequences of 
primates: tempo and mode of evolution. Journal of 
Molecular Evolution, 18:225-239. 

BROWNELL, E. 1983. DNA/DNA hybridization stud- 
ies of muroid rodents: symmetry and rates of mo- 
lecular evolution. Evolution, 38:1034-1051. 

Butmer, M., K. H. WoLFE, AND P. M. SHARP. 1991. 
Synonymous nucleotide substitution rates in mam- 
malian genes: implications for the molecular clock 
and the relationships of mammalian orders. Pro- 
ceedings of the National Academy of Sciences (USA), 
88:5974-5978. 

Burns, J. C., J. R. CHOATE, AND E. G. ZIMMERMAN. 
1985. Systematic relationships of pocket gophers 
(genus Geomys) on the central Great Plains. Journal 
of Mammalogy, 66:102-118. 

Butnu, D.G. 1984. The application of electrophoretic 
data in systematic studies. Annual Review of Ecol- 
ogy and Systematics, 15:501-522. 

CANN, R. L., M. STONEKING, AND A. C. WILSON. 1987. 
Mitochondrial DNA and human evolution. Nature, 
325:31-36. 


CaRLSON, S. S., A. C. WILSON, AND R. D. MAxson. 
1978. Do albumin clocks run on time? Science, 200: 
1183-1185. 

Carr, S. M., S. W. BALLINGER, J. N. Derr, L. H. 
BLANKENSHIP, AND J. W. BICKHAM. 1986. Mito- 
chondrial DNA analysis of hybridization between 
sympatric white-tailed deer and mule deer in West 
Texas. Proceedings of the National Academy of Sci- 
ences (USA), 83:9576-9580. 

CATZEFLIS, F. M., F. H. SHELDON, J. E. ALQUIST, AND 
C. S. SiBLEy. 1987. DNA-DNA hybridization ev- 
idence of the rapid rate of rodent DNA evolution. 
Molecular Biology and Evolution, 4:242-253. 

CuEsseR, R. K. 1983. Genetic variability within and 
among populations of the black-tailed prairie dog. 
Evolution, 37:320-331. 

CHESSER, R. K., M. H. SMITH, AND I. L. BRIsBIN, JR. 
1980. Management and maintenance of genetic 
variability in endangered species. International Zoo 
Yearbook, 20:146-154. 

Cook, J. A., AND T. L. YATEs. In press. Systematic 
relationships of the Bolivian Tuco-tucos, genus 
Ctenomys (Rodentia: Ctenomyidae). Journal of 
Mammalogy. 

COTHRAN, E. G., AND E. G. ZIMMERMAN. 1985. Elec- 
trophoretic analysis of the contact zone between 
Geomys breviceps and Geomys bursarius. Journal of 
Mammalogy, 66:489-497. 

CoTHRAN, E. G., E. G. ZIMMERMAN, AND C. F. NADLER. 
1977. Genic differentiation and evolution in ground 
squirrel subgenus /ctidomys (genus Spermophilus). 
Journal of Mammalogy, 58:610-622. 

CRACRAFT, J. 1987. DNA hybridization and avian 
phylogenetics. Evolutionary Biology, 21:47-96. 

Cronin, J. E., AND V. M. SaricH. 1975. Molecular 
systematics of the New World monkeys. Journal of 
Human Evolution, 4:357-375. 

CZELUSNIAK, J., ET AL. 1990. Perspectives from ami- 
no acid and nucleotide sequences on cladistic rela- 
tionships among higher taxa of Eutheria. Current 
Mammalogy, 2:541-567. 

De Jonc, W. W. 1982. Eye lens proteins and verte- 
brate phylogeny. Pp. 75-114, in Macromolecular se- 
quences in systematics and evolutionary biology (M. 
Goodman, ed.). Plenum Press, New York, 418 pp. 

DE Jonc, W. W., J. T. GLEAVES, AND D. BOULTER. 
1977. Evolutionary changes of a-crystallin and the 
phylogeny of mammalian orders. Journal of Molec- 
ular Evolution, 10:123-135. 

De Jona, W. W., A. ZWEERS, AND M. GoopMAN. 1981. 
Relationship of aardvark to elephants, hyraxes and 
sea cows from a-crystallin sequences. Nature, 292: 
117-119. 

Dene, H., M. GoopMan, M. C. MCKENNA, AND A. E. 
ROMERO-HERRERA. 1982. Ochotona princeps (pika) 
myoglobin: an appraisal of lagomorph phylogeny. 
Proceedings of the National Academy of Sciences 
(USA), 79:1917-1920. 

Dene, H., M. GOODMAN, AND W. S. PRYCHODKO. 1978. 
An immunological examination of the systematics 
of the Tupaioidea. Journal of Mammalogy, 59:697-— 
706. 

Dessauer, H. C., C. J. COLE, AND M. A. HAFNER. 1990. 
Collection and storage of tissues. Pp. 25-41, in Mo- 
lecular systematics (D. M. Hillis and C. Moritz, eds.). 


302 HONEYCUTT AND YATES 


Sinauer Associates, Sunderland, Massachusetts, 588 


DISOTELL, T. R., R. L. HONEYcuTT, AND M. RuVOLo. 
1992. Mitochondrial DNA phylogeny of the Old 
World monkey tribe Papionini. Molecular Biology 
and Evolution, 9:1-13. 

Dracoo, J. W., J. R. CHOATE, AND T. L. YATEs. 1990. 
Evolutionary and taxonomic relationships among 
North American arid-land foxes. Journal of Mam- 
malogy, 71:318-332. 

EASTEAL, S. 1985. Generation time and the rate of 
molecular evolution. Molecular Biology and Evo- 
lution, 2:450-453. 

1990. The pattern of mammalian evolution 
and the relative rate of molecular evolution. Genet- 
ics, 124:165-173. 

ELDREDGE, N., AND J. CRACRAFT. 1980. Phylogenetic 
patterns and the evolutionary process: method and 
theory in comparative biology. Columbia University 
Press, New York. 

Farris, J.S. 1972. Estimating phylogenetic trees from 
distance matrices. American Naturalist, 106:645- 
668. 


zy L985: 
2:144-157. 
1988. HENNIG86: program and documen- 
tation. State University of New York, New York. 
FELSENSTEIN, J. 1981. Evolutionary trees from DNA 
sequences: a maximum likelihood approach. Journal 
of Molecular Evolution, 17:368-376. 
1982. Numerical methods for inferring evo- 
lutionary trees. Quarterly Review of Biology, 57:379- 
404. 


Distance data revisited. Cladistics, 


1984. Distance methods for inferring phy- 

logenies: a justification. Evolution, 38:16-24. 

1988. Phylogenies for molecular sequences: 

inference and reliability. Annual Review of Genetics, 

22:521-565. 

. 1990. PHYLIP: phylogenetic inference pack- 
age, version 3.3. Program and documentation. De- 
partment of Genetics, University of Washington, Se- 
attle, Washington. 

FERRIS, S. D., A. C. WILSON, AND W. M. Brown. 1981. 
Evolutionary trees for apes and humans based on 
cleavage maps of mitochondrial DNA. Proceedings 
of the National Academy of Sciences (USA), 78: 
2432-2436. 

Ferris, S. D., R. D. SAGE, E. M. PRAGER, U. RITTE, 
AND A. C. WiLson. 1983. Mitochondrial DNA 
evolution in mice. Genetics, 105:681-721. 

Fircu, W. M., AND C. H. LANGLEY. 1976. Evolu- 
tionary rates in proteins: neutral mutations and the 
molecular clock. Pp. 197-219, in Molecular anthro- 
pology (M. Goodman, R. E. Tashian, and J. H. Tash- 
ian, eds.). Plenum Press, New York, 466 pp. 

Fitcu, W. M., AND E. MARGOLIASH. 1967. Construc- 
tion of phylogenetic trees. Science, 155:279-284. 

FORMAN, L., ET AL.. 1986. Genetic variation within 
and among lion tamarins. American Journal of Phys- 
ical Anthropology, 71:1-11. 

Gaines, M.S., AND C.J. Kress. 1971. Genetic changes 
in fluctuating vole populations. Evolution, 25:702- 
123: 

GARDNER, S. L. 1991. Phyletic coevolution between 
subterranean rodents of the genus Ctenomys (Ro- 


dentia: Hystricognathi) and nematodes of the genus 
Paraspidodera (Heterakoidea: Aspidoderidae) in the 
Neotropics: temporal and evolutionary implica- 
tions. Zoological Journal of the Linnean Society, 102: 
169-201. 

GENTZ, E. J., AND T. L. YATES. 1986. Genetic iden- 
tification of hybrid camelids. Zoo Biology, 5:349- 
354. 

GeEorGE, M., JR., AND O. A. RyperR. 1986. Mito- 
chondriai DNA evolution in the genus Equus. Mo- 
lecular Biology and Evolution, 3:535-546. 

GeorGE, S. B. 1986. Evolution and historical bio- 
geography of soricine shrews. Systematic Zoology, 
35:153-162. 

GERBER, J. D., AND C. A. LEONE. 1971. Immunologic 
comparisons of the sera of certain phyllostomatid 
bats. Systematic Zoology, 20:160-166. 

GILtespiE, J. H. 1986. Variability of evolutionary 
rates of DNA. Genetics, 113:1077-1091. 

. 1991. The causes of molecular evolution. Ox- 
ford University Press, New York, 336 pp. 

GINGERICH, P. D. 1986. Temporal scaling of molec- 
ular evolution in primates and other mammals. Mo- 
lecular Biology and Evolution, 3:205-221. 

GoopMAN, M. 1963. Serological analysis of the sys- 
tematics of recent hominoids. Human Biology, 35: 
377-424. 

1976a. Protein sequences in phylogeny. Pp. 

141-159, in Molecular evolution (F. J. Ayala, ed.). 

Sinauer Associates, Sunderland, Massachusetts, 277 

pp. 


1976b. Toward a genealogical description of 

the primates. Pp. 321-353, in Molecular anthropol- 

ogy (M. Goodman, R. E. Tashian, and J. H. Tashian, 
eds.). Plenum Press, New York, 466 pp. 

1981. Globin evolution was apparently very 
rapid in early vertebrates: a reasonable case against 
the rate-constancy hypothesis. Journal of Molecular 
Evolution, 17:114—120. 

GcGODMAN, M., J. CZELUSNIAK, AND J. E. BEEBER. 
1985. Phylogeny of Primates and other eutherian 
orders: a cladistic analysis using amino acid and nu- 
cleotide sequence data. Cladistics, 1:171-185. 

GoopMan, M., G. W. Moore, ANDG. Matsupba. 1975. 
Darwinian evolution in the genealogy of hemoglo- 
bin. Nature, 253:603-608. 

GoopMaN, M., J. CZELUSNIAK, B. F. Koop, D. A. TA- 
GLE, AND J. L. SLIGHTOM. 1987. Globins: a case 
study in molecular phylogeny. Cold Spring Harbor 
Symposium on Quantitative Biology, 52:875-890. 

Goopman, M., A. E. ROMERO-HERRERA, H. DENE, J. 
CZELUSNIAK, AND R. E. TASHIAN. 1982. Amino acid 
sequence evidence on the phylogeny of primates and 
other eutherians. Pp. 115-191, in Macromolecular 
sequences in systematic and evolutionary biology 
(M. Goodman, ed.). Plenum Press, New York, 418 


pp. 
Graur, D., W. A. Hipe, AND W.-H. Li. 1991. Is the 
guinea-pig a rodent? Nature, 351:649-652. 
GREENBAUM, I. F. 1981. Genetic interactions be- 
tween hybridizing cytotypes of the tent-making bat 
(Uroderma bilobatum). Evolution, 35:306-321. 
GREENBAUM, I. F., AND R. J. BAKER. 1976. Evolu- 
tionary relationships in Macrotus (Mammalia: Chi- 


MOLECULAR SYSTEMATICS 505 


roptera): biochemical variation and karyology. Sys- 
tematic Zoology, 25:15-25. 

Harner, D. J., AND K. N. GeL_uso. 1983. Systematic 
relationships and historical zoogeography of the des- 
ert pocket gopher, Geomys arenarius. Journal of 
Mammalogy, 64:405-413. 

HAFNeER, D. J., K. E. PETERSEN, AND T. L. YATEs. 1981. 
Evolutionary relationships of jumping mice (genus 
Zapus) of the Southwestern United States. Journal 
of Mammalogy, 62:501-512. 

HAFNER, D. J., AND T. L. YATES. 1983. Systematic 
status of the Mojave ground squirrel, Spermophilus 
mohavensis (subgenus Xenospermophilus). Journal 
of Mammalogy, 64:397-404. 

Harner, J. C., D. J. HAFNER, J. L. PATTON, AND M. 
F. SmirH. 1983. Contact zones and the genetics of 
differentiation in the pocket gopher Thomomys bot- 
tae (Rodentia: Geomyidae). Systematic Zoology, 32: 
1-20. 

Harner, M.S. 1982. A biochemical investigation of 
geomyoid systematics (Mammalia: Rodentia). Son- 
derdruck aus Zoologie fur zoologischen Systematik 
und Evolutionforschung, 20:1 18-130. 

Harner, M. S., J. C. HAFNER, J. L. PATTON, AND M. 
F. SmitH. 1987. Macrogeographic patterns of ge- 
netic differentiation in the pocket gopher Thomomys 
umbrinus. Systematic Zoology, 36:18-34. 

Harner, M. S., AND S. A. NADLER. 1988. Phyloge- 
netic trees support the coevolution of parasites and 
their hosts. Nature, 332:258-259. 

1990. Cospeciation in host-parasite assem- 
blages: comparative analysis of rates of evolution 
and timing of cospeciation events. Systematic Zo- 
ology, 39:192-204. 

HamILton, M. J., R. K. CHESSER, AND T. L. BEST. 
1987. Genetic variation in the Texas kangaroo rat, 
Dipodomys elator Merriam. Journal of Mammalogy, 
68:775-781. 

Harris, H. 1966. Enzyme polymorphism in man. 
Proceedings of the Royal Society London Series B, 
164:298-310. 

Harris, H., AND D. A. HopKINsSoN. 1976. Handbook 
of enzyme electrophoresis. North-Holland Publish- 
ing Company, Amsterdam, Netherlands, looseleaf, 
unpaged. 

HaseGcawa, M., H. KIsHINO, AND T.-A. YANO. 1985. 
Dating the human-ape splitting by a molecular clock 
of mitochondrial DNA. Journal of Molecular Evo- 
lution, 22:160—-174. 

HayASAKA, K., T. GoyoBoriI, AND S. Horar. 1988. 
Molecular phylogeny and evolution of primate mi- 
tochondrial DNA. Molecular Biology and Evolution, 
5:626-644. 

HEANEY, L. R., AND R. M. Timm. 1985. Morphology, 
genetics, and ecology of pocket gophers (genus Geo- 
mys) in a narrow hybrid zone. Biological Journal of 
the Linnean Society, 25:301-317. 

Heprick, P. W., M. E. GINEVAN, AND E. P. Ewina. 
1976. Genetic polymorphism in heterogeneous en- 
vironments. Annual Review of Ecology and System- 
atics, 7:1-32. 

HERD, R. M., AND M. B. FENTON. 1983. An electro- 
phoretic, morphological, and ecological investiga- 
tion of a putative hybrid zone between Myotis lu- 
cifugus and Myotis yumanensis (Chiroptera: 


Vespertilionidae). Canadian Journal of Zoology, 61: 
2029-2050. 

HiGucul, R., B. BowMAN, M. FREIBERGER, O. A. RYDER, 
AND A. C. WiLson. 1984. DNA sequences from 
the quagga, an extinct member of the horse family. 
Nature, 312:282-284. 

HiGucui, R. G., AND H. OCHMAN. 1989. Production 
of single-stranded DNA templates by exonuclease 
digestion following the polymerase chain reaction. 
Nucleic Acids Research, 17:5865. 

Hiiuis, D. M., AND M. T. Drxon. 1991. Ribosomal 
DNA: molecular evolution and phylogenetic infer- 
ence. The Quarterly Review of Biology, 66:41 1-453. 

Hiius, D. M., AND C. Moritz (EpDs.). 1990a. Molec- 
ular systematics. Sinauer Associates, Sunderland, 
Massachusetts, 588 pp. 

Hiius, D. M., AND C. Moritz. 19906. An overview 
of applications of molecular systematics. Pp. 502- 
515, in Molecular systematics (D. M. Hillis and C. 
Moritz, eds.). Sinauer Associates, Sunderland, Mas- 
sachusetts, 588 pp. 

Hixson, J. E., AND W. M. Brown. 1986. A compar- 
ison of the small mbosomal RNA genes from the 
mitochondrial DNA of the great apes and humans: 
sequence, structure, evolution, and phylogenetic im- 
plications. Molecular Biology and Evolution, 3:1- 
18. 

Homes, E. C. 1991. Different rates of substitution 
may produce different phylogenies of the eutherian 
mammals. Journal of Molecular Evolution, 33:209- 
215. 

Honeycutt, R. L. 1992. Naked mole-rats. American 
Scientist, 80:43-53. 

Honeycutt, R. L., AND R. M. Apxkins. 1993. Higher 
level systematics of eutherian mammals: an assess- 
ment of molecular characters and phylogenetic hy- 
potheses. Annual Review of Ecology and System- 
atics, 24:279-305. 

Honeycutt, R. L., AND V. M. SARiIcH. 1987a. Al- 
bumin evolution and subfamilial relationships among 
New World leaf-nosed bats (family Phyllostomidae). 
Journal of Mammalogy, 68:508-517. 

1987b. Monophyly and molecular evolution 
within three phyllostomid bat genera. Journal of 
Mammalogy, 68:518-525. 

Honeycutt, R. L., AND S. L. WILLIAMS. 1982. Genic 
differentiation in pocket gophers of the genus Pap- 
pogeomys, with comments on intergeneric relation- 
ships in the subfamily Geomyinae. Journal of Mam- 
malogy, 63:208-217. 

Honeycutt, R. L., S. Y. E>DwArRDs, K. NELSON, AND 
E. Nevo. 1987. Mitochondrial DNA variation and 
the phylogeny of African mole rats (Rodentia: Bathy- 
ergidae). Systematic Zoology, 36:280-293. 

Honeycutt, R. L., I. F. GREENBAUM, R. J. BAKER, AND 
V.M. SaricH. 1981. Molecular evolution of vam- 
pire bats. Journal of Mammalogy, 62:805-811. 

Hussy, J. L., AND R. C. Lewontin. 1966. A molec- 
ular approach to the study of genic heterozygosity 
in natural populations. I. The number of alleles at a 
different loci in Drosophila pseudoobscura. Genetics, 
54:577-594. 

Hussy, J. L., AND L. H. THROCKMORTON. 1965. Pro- 
tein differences in Drosophila. 11. Comparative spe- 


304 HONEYCULT AND YATES 


cies genetics and evolutionary problems. Genetics, 
52:203-215. 

Hunter, R. L., AND C. L. MARKERT. 1957. Histo- 
chemical demonstration of enzymes separated by 
zone electrophoresis in starch gels. Science, 125:1294— 
1295. 

Irwin, D. M., T. D. KocHER, AND A.C. WiLson. 1991. 
Evolution of the cytochrome b gene in mammals. 
Journal of Molecular Evolution, 32:128-144. 

JANECEK, L. L., R. L. Honeycutt, I. L. RAUTENBACH, 
B. H. Erasmus, S. REIG, AND D. A. SCHLITTER. 1992. 
Allozyme variation and systematics of African mole- 
rats (Rodentia:Bathyergidae). Biochemical System- 
atics and Ecology, 20:401-416. 

JOHNSON, G. L., AND R. L. PACKARD. 1974. Electro- 
phoretic analysis of Peromyscus comanche Blair, with 
comments on its systematic status. Occasional Pa- 
pers, The Museum, Texas Tech University, 24:1- 
16. 

Jounson, W. E., AND R. K. SELANDER. 1971. Protein 
variation and systematics in kangaroo rats (genus 
Dipodomys). Systematic Zoology, 20:377-405. 

KILpaTRick, C. W. 1981. Genetic structure of insular 
populations. Pp. 28-60, in Mammalian population 
genetics (M. H. Smith and J. Joule, eds.). University 
of Georgia Press, Athens, 380 pp. 

KILPATRICK, C. W., AND K. L. CROWELL. 1985. Genic 
variation of the rock vole, Microtus chrotorrhinus. 
Journal of Mammalogy, 66:94-101. 

KILPATRICK, C. W., AND E. G. ZIMMERMAN. 1975. 
Genetic variation and systematics of four species of 
mice of the Peromyscus boylii species group. System- 
atic Zoology, 24:143-162. 

KILPATRICK, C. W., S. C. FORREST, AND T. W. CLARK. 
1986. Estimating genetic variation in the black- 
footed ferret—a first attempt. Great Basin Naturalist, 
8:145-149. 

Kimura, M. 1981. Was globin evolution very rapid 
in its early stages? A dubious case against the rate- 
constancy hypothesis. Journal of Molecular Evolu- 
tion, 17:110-113. 

1983. The neutral theory of molecular evo- 
lution. Cambridge University Press, Cambridge, En- 
gland, 367 pp. 

Kirscu, J. A.W. 1977. The comparative serology of 
Marsupialia, and a classification of marsupials. Aus- 
tralian Journal of Zoology, Supplementary Series, 
52:1-152. 

Kirscn, J. A. W., R. E. BLEISwEIss, AND A. W. DIck- 
ERMAN. 1993. DNA/DNA hybridization studies of 
carnivorous marsupials. III. Relationships among 
species of Didelphis (Didelphidae). Journal of Mam- 
malian Evolution, 1:75-97. 

Kurscu, J. A. W., A. W. DICKERMAN, O. A. REIG, AND 
M. S. Sprincer. 1991. DNA hybridization evi- 
dence for the Australian affinity of the American 
marsupial Dromiciops australis. Proceedings of the 
National Academy of Sciences (USA), 88:10465- 
10469. 

Kirscu, J. A. W., C. KRAsEwSKI, M. S. SPRINGER, AND 
M. ArcHER. 1990a. DNA-DNA hybridization 
studies of carnivorous marsupials. II. Relationships 
among dasyurids (Marsupialia: Dasyuridae). Aus- 
tralian Journal of Zoology, 38:673-696. 

Kirscu, J. A. W., M. S. SPRINGER, C. KRAJEWSKI, M. 


ARCHER, K. APLIN, AND A. W. DICKERMAN. 1990b. 
DNA/DNA hybridization studies of carnivorous 
marsupials. I: The intergeneric relationships of ban- 
dicoots (Marsupialia: Perameloidea). Journal of Mo- 
lecular Evolution, 30:434-448. 

Kocuer, T. D., ET AL. 1989. Dynamics of mito- 
chondrial DNA evolution in animals: amplification 
and sequencing with conserved primers. Proceedings 
of the National Academy of Sciences (USA), 86: 
6196-6200. 

KOHNE, D. E., J. A. CHISCON, AND B. H. Hover. 1972. 
Evolution of primate DNA sequences. Journal of 
Human Evolution, 1:627-644. 

Koop, B. F., AND R. J. BAKER. 1983. Electrophoretic 
studies of relationships of six species of Artibeus 
(Chiroptera: Phyllostomidae). Occasional Papers, 
The Museum, Texas Tech University, 83:1-12. 

Koop, B. F., AND M. GOODMAN. 1988. Evolutionary 
and developmental aspects of two hemoglobin 
B-chain genes (e™“ and B™) of opossum. Proceedings 
of the National Academy of Sciences (USA), 85: 
3893-3897. 

Koop, B., M. GOODMAN, P. Xu, K. CHAN, AND J. L. 
SLIGHTOM. 1986. Primate n-globin DNA sequenc- 
es and man’s place among the great apes. Nature, 
319:234-238. 

Kraus, F., AND M. M. MryAmoto. 1991. Rapid 
cladogenesis among the pecoran ruminants: evi- 
dence from mitochondrial DNA sequences. System- 
atic Zoology, 40:117-130. 

LANGLEY, C. H., AND W. M. Fitcu. 1974. An ex- 
amination of the constancy of the rate of molecular 
evolution. Journal of Molecular Evolution, 3:161- 
aie 

LEONE, C. A., AND A. L Wiens. 1956. Comparative 
serology of carnivores. Journal of Mammalogy, 37: 
11-23. 

Levine, H. P., AND P. A. Moopy. 1939. Serological 
investigation of rodent relationships. Physiological 
Zoology, 12:400-411. 

Lewontin, R. C. 1974. The genetic basis of evolu- 
tionary change. Columbia University Press, New 
York, 346 pp. 

Li, W.-H., AND D. Graur. 1991. Fundamentals of 
molecular evolution. Sinauer Associates, Sunder- 
land, Massachusetts, 284 pp. 

Li, W.-H., C.-C. Luo, AND C.-I. Wu. 1985. Evolution 
of DNA sequences. Pp. 1-84, in Molecular evolu- 
tionary genetics (R. J. MacIntyre, ed.). Plenum Press, 
New York, 610 pp. 

Li, W.-H., M. TANIMURA, AND P. M. SHARP. 1987. 
An evaluation of the molecular clock hypothesis us- 
ing mammalian DNA sequences. Journal of Molec- 
ular Evolution, 25:330-342. 

Li, W.-H., M. Gouy, P. M. SHARP, C. O’HUIGIN, AND 
Y.-W. YANG. 1990. Molecular phylogeny of Ro- 
dentia, Lagomorpha, Primates, Artiodactyla, and 
Carnivora and molecular clocks. Proceedings of the 
National Academy of Sciences (USA), 87:6703-6707. 

Li, W.-H., W. A. Hipe, A. ZHARKIKH, D.-P. MA, AND 
D. Graur. 1992. The molecular taxonomy and 
evolution of the guinea pig. Journal of Heredity, 83: 
174-181. 

Martin, A. P., AND S. R. PALUMBI. 1993. Body size, 
metabolic rate, generation time, and the molecular 


MOLECULAR SYSTEMATICS 305 


clock. Proceedings of the National Academy of Sci- 
ences (USA), 90:4087-409 1. 

Maxa, A. M., AND W. GILBERT. 1980. Sequencing 
end-labeled DNA with base-specific chemical cleav- 
ages. Methods of Enzymology, 65:499-559. 

Maxson, L. R., AND R. D. MAxson. 1990. Proteins 
II: immunological techniques. Pp. 127-155, in Mo- 
lecular systematics (D. M. Hillis and C. Moritz, eds.). 
Sinauer Associates, Sunderland, Massachusetts, 588 


pp. 

Maxson, L. R., V. M. SARICH, AND A. C. WILSON. 
1975. Continental drift and the use of albumin as 
an evolutionary clock. Nature, 255:397-399. 

McCRACKEN, G. F., AND J. W. BRADBURY. 1977. Pa- 
ternity and genetic heterozygosity in the polygynous 
bat, Phyllostomus hastatus. Science, 198:303-306. 

1981. Social organization and kinship in the 
polygynous bat Phyllostomus hastatus. Behavioral 
Ecology and Sociobiology, 8:1 1-34. 

McKenna, M.C. 1975. Toward a phylogenetic clas- 
sification of Mammalia. Pp. 21-46, in Phylogeny of 
the Primates (W. P. Luckett and F. S. Szalay, eds.). 
Plenum, New York, 483 pp. 

MELNIck, D. J.,G. A. HOELZER, AND R. L. HONEYCUTT. 
1992. Mitochondrial DNA: its uses in anthropo- 
logical research. Pp. 179-233, in Molecular appli- 
cations in biological anthropology (E. J. Devor, ed.). 
Cambridge University Press, Cambridge, England, 
258 pp. 

MickeEvicu, M. F., AND M. S. JOHNSON. 1976. Con- 
gruence between morphological and allozyme data 
in evolutionary inference and character evolution. 
Systematic Zoology, 25:260—270. 

MINDELL, D. P., AND R. L. Honeycutr. 1990. Ri- 
bosomal RNA in vertebrates: evolution and phy- 
logenetic applications. Annual Review of Ecology 
and Systematics, 21:541-566. 

MINDELL, D. P., C. W. Dick, AND R. J. BAKER. 1991. 
Phylogenetic relationships among megabats, micro- 
bats, and primates. Proceedings of the National 
Academy of Sciences (USA), 88:10322-10326. 

Mryamoto, M. M., AND S. M. Boye. 1989. The 
potential importance of mitochondrial DNA se- 
quence data to eutherian mammal phylogeny. Pp. 
437-450, in The hierarchy of life (B. Fernholm, K. 
Bremer, and H. Jornvall, eds.). Elsevier, New York, 
500 pp. 

Miyamoto, M. M., AND J. CRACRAFT (EDS.). 1991. 
Phylogenetic analysis of DNA sequences. Oxford 
Press, New York, 358 pp. 

Mryamorto, M. M., AND M. GoopMAN. 1986. Biomo- 
lecular systematics of eutherian mammals: phylo- 
genetic patterns and classification. Systematic Zo- 
ology, 35:230-240. 

Mryamorto, M. M., F. KRAUS, ANDO. Z. RYDER. 1990. 
Phylogeny and evolution of antlered deer deter- 
mined from mitochondrial DNA sequences. Pro- 
ceedings of the National Academy of Sciences (USA), 
87:6127-6131. 

Mryamoto, M. M., S. M. TANHAUSER, AND P. J. LAIPIs. 
1989. Systematic relationships in the artiodacty] tribe 
Bovini (family Bovidae), as determined from mi- 
tochondrial DNA sequences. Systematic Zoology, 38: 
342-349. 

Moore, D. W., AND T. L. YATES. 1983. Rate of pro- 


tein inactivation in selected mammals following 
death. Journal of Wildlife Management, 47:1166- 
1169. 

Moore, G. W., J. BARNABAS, AND M. GOODMAN. 1973. 
A method for constructing maximum parsimony an- 
cestral amino acid sequences on a given network. 
Journal of Theoretical Biology, 38:459-485. 

Murpny, Rosert W., J. W. Sites, D. G. BUTH, AND 
C. H. HAuFLER. 1990. Proteins I: isozyme electro- 
phoresis. Pp. 45-126, in Molecular systematics (D. 
M. Hillis and C. Moritz, eds.). Sinauer Associates, 
Sunderland, Massachusetts, 588 pp. 

NADLER, C. F., N. N. Vorontsov, R. S. HOFFMANN, 
I. I. FORMICHOVA, AND C. F. NADLER, JR. 1973. 
Zoogeography of transferrins in Arctic and long-tailed 
ground squirrel populations. Comparative Biochem- 
istry and Physiology, B. Comparative Biochemistry, 
44:33-40. 

NADLER, C. F., R. S. HOFFMAN, N. N. Vorontsov, J. 
W. Kowpp., L. DEUTSCHAND, AND I. SUKERKNIK. 
1982. Evolution in ground squirrels. II. Biochem- 
ical comparisons on Holartic populations of Sper- 
mophilus. Zeitschrift Saugetierkund, 47:198-215. 

Nel, M. 1971. Interspecific gene differences and evo- 
lutionary time estimated from electrophoretic data 
on protein identity. American Naturalist, 105:385- 
398. 


. 1972. Genetic distance between populations. 

The American Naturalist, 106:283-292. 

1987. Molecular evolutionary genetics. Co- 
lumbia University Press, New York, 512 pp. 

Ne, M., AND D. GraAur. 1984. Extent of protein 
polymorphism and the neutral mutation theory. 
Evolutionary Biology, 17:73-118. 

NELson, K., R. J. BAKER, AND R. L. Honeycutt. 1987. 
Mitochondrial DNA and protein differentiation be- 
tween hybridizing cytotypes of the white-footed 
mouse, Peromyscus leucopus. Evolution, 41:864—872. 

NELson, K., R. J. BAKER, AND H. S. SHELLHAMMER. 
1984. Test of alternative hypotheses concerning the 
origin of Reithrodontomys raviventris: genetic anal- 
ysis. Journal of Mammalogy, 65:668-673. 

Nevo, E. 1978. Genetic variation in natural popu- 
lations: patterns and theory. Theoretical Population 
Biology, 13:121-177. 

1985. Ecological and populational correlates 
of allozyme polymorphisms in mammals. Acta 
Zoologica Fennica, 170:25-29. 

Nevo, E., AND C. R. SHAW. 1972. Genetic variation 
in a subterranean mammal, Spalax ehrenbergi. Bio- 
chemical Genetics, 7:235-241. 

Nevo, E., Y. J. Kim, C. R. SHAW, AND C. S. THAELER, 
Jr. 1974. Genetic variation, selection, and speci- 
ation in Thomomys talpoides pocket gophers, Evo- 
lution, 28:1—23. 

Nevo, E., R. L. HoNEycutt, H. YONEKAWA, K. NELSON, 
AND N. Hanzawa. 1993. Mitochondrial DNA 
polymorphism in subterranean mole-rats of the Spa- 
lax ehrenbergi superspecies in Israel, and its periph- 
eral isolates. Molecular Biology and Evolution, 10: 
590-604. 

NEwMan, A., ET AL. 1985. Biochemical genetic vari- 
ation in eight endangered or threatened felid species. 
Journal of Mammalogy, 66:256-267. 


306 HONEYCUTT AND YATES 


Novacek, M. J. 1992. Mammalian phylogeny: shak- 
ing the tree. Nature, 356:121-125. 

NutTALL, G. H. F. 1904. Blood immunity and blood 
relationships. Cambridge University Press, Cam- 
bridge, England. 

O’Brien, S. J., D. E. Witpt, AND M. BusH. 1986. 
The cheetah in genetic peril. Scientific American, 
May, 1986:84-92. 

O’Brien, S. J., D. E. Witpt, D. GOLDMAN, C. R. 
MERRIL, AND M. BusH. 1983. The cheetah is de- 
pauperate in genetic variation. Science, 221:459-462. 

O’BrIEN, S. J., ET AL. 19875. East African cheetahs: 
evidence for two population bottlenecks? Proceed- 
ings of the National Academy of Sciences (USA), 84: 
508-511. 

O’BRIEN, S. J., ET AL. 1987a. Biochemical genetic 
variation in geographic isolates of African and Asi- 
atic lions. National Geographic Research, 3:1 14-124. 

O’BrieN, S. J., ET AL. 1985. Genetic basis for species 
vulnerability in the cheetah. Science, 227:1428-1434. 

O’Brien, S. J., ET AL. 1990. Genetic introgression 
within the Florida panther Felis concolor coryi. Na- 
tional Geographic Research, 6:485-494. 

PaaBo, S. 1989. Ancient DNA: extraction, charac- 
terization, molecular cloning, and enzymatic ampli- 
fication. Proceedings of the National Academy of 
Sciences (USA), 86:1939-1943. 

PAABo, S., J. A. GIFFORD, AND A. C. WILSON. 1988. 
Mitochondrial DNA sequences from a 7000-year- 
old brain. Nucleic Acid Research, 16:9775-9787. 

Passo, S., R. HiGucHI, AND A. C. WILSON. 1989. 
Ancient DNA and the polymerase chain reaction. 
Journal of Biological Chemistry, 264:9709-971 2. 

Packer, C., A. E. Pusey, H. Rowtey, D. A. GILBERT, 
J. MARTENSON, AND S. J. O'BRIEN. 1991. Case study 
of a population bottleneck: lions of the Ngorongoro 
Crater. Conservation Biology, 5:219-230. 

Patton, J. L. 1984. Genetical processes in the Ga- 
lapagos. Biological Journal of the Linnean Society, 
21:97-111. 

1985. Population structure and the genetics 
of speciation in pocket gophers, genus Thomomys. 
Acta Zoologica Fennica, 170:109-114. 

Patton, J. C., AND J. C. Avise. 1983. An empirical 
evaluation of qualitative Hennigian analyses of pro- 
tein electrophoretic data. Journal of Molecular Evo- 
lution, 19:244-254. 

Patton, J. L., AND M. F. SmitH. 1992. MtDNA phy- 
logeny of Andean mice: a test of diversification across 
ecological gradients. Evolution, 46:174-183. 

Patton, J. L., AND S. Y. YANG. 1977. Genetic vari- 
ation in Thomomys bottae pocket gophers: macro- 
geographic patterns. Evolution, 31:697—720. 

Patton, J. C., R. J. BAKER, AND J. C. AviseE. 1981. 
Phenetic and cladistic analyses of biochemical evo- 
lution in peromyscine rodents. Pp. 288-308, in 
Mammalian population genetics (M. H. Smith and 
J. Joule, eds.). University of Georgia Press, Athens, 
Georgia, 380 pp. 

Patton, J. L., R. K. SELANDER, AND M. H. SMITH. 
1972. Genic variation in hybridizing populations 
of gophers (genus Thomomys). Systematic Zoology, 
21:263-270. 

Patton, J. L., J.C. HAFNER, AND M.S. HAFNER. 1979a. 
Hybrid zones in Thomomys bottae pocket gophers: 


genetic, phenetic, and ecologic concordance patterns. 
Evolution, 33:860-876. 

Patton, J. L., M. S. HAFNER, J. C. HAFNER, AND M. 
F. SmitH. 1979b. Hybrid zones in Thomomys bot- 
tae pocket gophers: genetic, phenetic, and ecologic 
concordance patterns. Evolution, 33:860-876. 

PENNY, D. F., AND E. G. ZIMMERMAN. 1976. Genic 
divergence and local population differentiation by 
random drift in the pocket gopher genus Geomys. 
Evolution, 30:473-483. 

Pierson, E. D., V. M. SARIcH, J. M. LOWENSTEIN, M. 
J. DANIEL, AND W. E. Ratney. 1986. A molecular 
link between the bats of New Zealand and South 
America. Nature, 323:60-63. 

PRAGER, E. M., AND A. C. WiLson. In press. Infor- 
mation content of immunological distances. Meth- 
ods In Enzymology. 

REDUKER, D. W., D. W. DuSZYNSKI, AND T. L. YATES. 
1987. Evolutionary relationships among Eimeria 
spp. (Apicomplexa) infecting cricetid rodents. Ca- 
nadian Journal of Zoology, 65:722-735. 

RENNERT, P. D., AND C. W. KILPATRICK. 1986. Bio- 
chemical systematics of populations of Peromyscus 
boylii. 1. Populations from east-central Mexico with 
low fundamental numbers. Journal of Mammalogy, 
67:48 1-488. 

RIDDLE, B. R., AND R. L. Honeycutr. 1990. His- 
torical biogeography in North American arid regions: 
an approach using mitochondrial-DNA phylogeny 
in grasshopper mice (genus Onychomys). Evolution, 
44:1-15. 

Ripp_e, B. R., R. L. HONEYcUTT, AND P. L. LEE. 1993. 
Mitochondrial DNA phylogeography in northern 
grasshopper mice (Onychomys leucogaster): the in- 
fluence of Quaternary climatic oscillations on pop- 
ulation dispersion and divergence. Molecular Ecol- 
ogy, 2:183-193. 

Rossins, L. W., M. H. Smitu, M. C. WooTeEN, AND R. 
K. SELANDER. 1985. Biochemical polymorphism 
and its relationship to chromosomal and morpho- 
logical variation in Peromyscus leucopus and Pero- 
myscus gossypinus. Journal of Mammalogy, 66:498- 
510. 

RoeLkE, M. E., J. S. MARTENSON, AND S. J. O'BRIEN. 
1993. The consequences of demographic reduction 
and genetic depletion in the endangered Florida pan- 
ther. Current Biology, 3:340-350. 

Rocers, J. S. 1972. Measures of genetic similarity 
and genetic distance. Studies in genetics VII. Uni- 
versity of Texas Publications, 7213:145-153. 

ROMERO-HERRERA, A. E., H. LEHMANN, K. A. JOYSEY, 
AND A. E. Fripay. 1978. On the evolution of myo- 
globin. Philosophical Transactions of the Royal So- 
ciety of London, 283:61-163. 

Ruvo o, M., T. R. DisoTeELL, M. W. ALLARD, W. M. 
BRowNn, AND R. L. Honeycutt. 1991. Resolution 
of the African hominoid trichotomy by use of a mi- 
tochondrial gene sequence. Proceedings of the Na- 
tional Academy of Sciences (USA), 88:1570-1574. 

SAGE, R. D., J. R. CONTRERAS, V. G. RoIG, AND J. L. 
PATTON. 1986. Genetic variation in the South 
American burrowing rodents of the genus Ctenomys 
(Rodentia: Ctenomyidae). Zeitschrift Saugetier- 
kunde, 51:158-172. 

Saiki, R. K., ETAL. 1988. Primer-directed enzymatic 


MOLECULAR SYSTEMATICS 307 


amplification of DNA with a thermostable DNA 
polymerase. Science, 239:487-491. 

SANGER, F., S. NICKLEN, AND A. R. COULSON. 1977. 
DNA sequencing with chain terminating inhibitors. 
Proceedings of the National Academy of Sciences 
(USA), 74:5463-5467. 

SaricH, V. M. 1969a. Pinniped origins and the rate 
of evolution of carnivore albumins. Systematic Zo- 
ology, 18:286-295. 

. 1969b. Pinniped phylogeny. Systematic Zo- 

ology, 18:416-422. 

. 1973. The giant panda isa bear. Nature, 245: 

218-220. 

1977. Rates, sample sizes, and the neutrality 

hypothesis for electrophoresis in evolutionary stud- 

ies. Nature, 256:24-28. 

. 1985. Rodent macromolecular systematics. 

Pp. 423-452, in Evolutionary relationships among 

rodents: a multidisciplinary analysis (W. P. Luckett 

and J.-L. Hartenberger, eds.). Plenum, New York, 

721 pp. 

In press. Some results of 25 years with the 
blood of mammals. /n Mammal phylogeny: placen- 
tals (F. S. Szalay, M. J. Novacek, and M. C. Mc- 
Kenna, eds.). Springer-Verlag, New York. 

SARICH, V. M., AND J. E. CRONIN. 1976. Molecular 
systematics of the primates. Pp. 141-170, in Molec- 
ular anthropology (M. Goodman, R. E. Tashian, and 
J. H. Tashian, eds.). Plenum Press, New York, 466 
pp. 

SaricH, V. M., AND A. C. WILSON. 1967a. Rates of 
albumin evolution in primates. Proceedings of the 
National Academy of Sciences (USA), 58:142-148. 

1967b. Immunological time scale for hom- 
inid evolution. Science, 158:1200-1203. 

SARICH, V. M., C. W. SCHMID, AND J. MARKS. 1989. 
DNA hybridization as a guide and phylogenies: a 
critical analysis. Cladistics, 5:3-32. 

SCHMIDLY, D. J., M. R. Lee, W. S. Mont, AND E. G. 
ZIMMERMAN. 1985. Systematics and notes on the 
biology of Peromyscus hooperi. Occasional Papers, 
The Museum, Texas Tech University, 97:1-40. 

SCHNELL, G. D., AND R. K. SELANDER. 1981. Envi- 
ronmental and morphological correlates of genetic 
variation in mammals. Pp. 60-99, in Mammalian 
population genetics (M. H. Smith and J. Joule, eds.). 
University of Georgia Press, Athens, 380 pp. 

SCRIBNER, K. T., R. K. CHESSER, AND R. J. WARREN. 
1983. Spatial and temporal genetic variability of 
the eastern cottontail on west Texas playa basins. 
Journal of Mammalogy, 64:287-294. 

SELANDER, R. K. 1977. Genic variation in natural 
populations. Pp. 21-45, in Molecular evolution (F. 
J. Ayala, ed.). Sinauer Associates, Sunderland, Mas- 
sachusetts, 277 pp. 

SELANDER, R. K., AND D. W. KAUFMAN. 1973. Genic 
variability and strategies of adaptation in animals. 
Proceedings of the National Academy of Sciences 
(USA), 70:1875-1877. 

SELANDER, R. K., AND T. S. WHITTAM. 1983. Protein 
polymorphism and the genetic structure of popula- 
tions. Pp. 89-114, in Evolution of genes and proteins 
(M. Nei and R. K. Koehn, eds.). Sinauer Associates, 
Sunderland, Massachusetts, 331 pp. 

SELANDER, R. K., D. W. KAUFMAN, R. J. BAKER, AND 


S. L. WitiiAmMs. 1974. Genic and chromosomal 
differentiation in pocket gophers of the Geomys bur- 
Sarius group. Evolution, 28:557-564. 

SELANDER, R. K., M. H. Smitu, S. Y. YANG, W. E. 
JOHNSON, AND J. R. GENTRY. 1971. Biochemical 
polymorphism and systematics in the genus Pero- 
myscus. I. Variation in the old-field mouse (Pero- 
myscus polionotus). Studies in genetics VI. Univer- 
sity of Texas Publications, 7103:49-90. 

SHAW, C. R., AND R. PRASAD. 1970. Starch gel elec- 
trophoresis of enzymes: a compilation of recipes. 
Biochemical Genetics, 4:297-320. 

SHOSHANTI, J. 1986. Mammalian phylogeny: compar- 
ison of morphological and molecular results. Mo- 
lecular Biology and Evolution, 3:222-242. 

SHOSHANI, J., J. M. LOWENSTEIN, D. A. WALz, AND M. 
GoopMAN. 1985. Proboscidean origins of masto- 
don and woolly mammoth demonstrated immuno- 
logically. Paleobiology, 1 1:429-437. 

S1BLEY, C. G., AND J. E. AHLQuistT. 1981. The phy- 
logeny and relationships of the ratite birds as indi- 
cated by DNA-DNA hybridization. Pp. 301-335, in 
Evolution today (G. G. E. Scudder and J. L. Reveal, 
eds.). Carnegie-—Mellon University, Pittsburgh, 
Pennsylvania. 

S1BLEY, C. G., AND J. E. AHLQuistT. 1984. The phy- 
logeny of the hominoid primates, as indicated by 
DNA-DNA hybridization. Journal of Molecular 
Evolution, 20:2-15. 

SIBLEY, C. G., J. E. AHLQUIST, AND B. L. MONROE, JR. 
1988. Aclassification of the living birds of the world, 
based on DNA/DNA hybridization studies. Auk, 105: 
409-423. 

Stmpson, G. G. 1945. The principles of classification 
and a classification of mammals. Bulletin of the 
American Museum of Natural History, 85:1-350. 

Smitu, D. G., AND R. G. Coss. 1984. Calibrating the 
molecular clock: estimates of ground squirrel diver- 
gence made using fossil and geological time markers. 
Molecular Biology and Evolution, 1:249-259. 

SmiTH, M. F., AND J. L. PATron. 1984. Dynamics of 
morphological differentiation: temporal impact of 
gene flow in pocket gopher populations. Evolution, 
38:1079-1087. 

1988. Subspecies of pocket gophers: causal 
bases for geographic differentiation in Thomomys 
bottae. Systematic Zoology, 37:163-178. 

Smitu, M. H., R. K. SELANDER, AND W. E. JOHNSON. 
1973. Biochemical polymorphism and systematics 
in the genus Peromyscus. III. Variation in the Florida 
deer mouse (Peromyscus floridanus), a Pleistocene 
relict. Journal of Mammalogy, 54:1-13. 

SmiTu, M. F., J. L. Patron, J. C. HAFNER, AND D. J. 
HaAFNeER. 1983. Thomomys bottae pocket gophers 
of the central Rio Grande Valley, New Mexico: local 
differentiation, gene flow, and historical biogeogra- 
phy. Occasional Papers, Museum Southwestern Bi- 
ology, 2:1-16. 

SPRINGER, M. S., AND J. A. W. Kirscu. 1989. Rates 
of single-copy DNA evolution in phalangeriform 
marsupials. Molecular Biology and Evolution, 6:33 1- 
341. 

1991. DNA hybridization, the compression 

effect, and the radiation of diprotodontian marsu- 

pials. Systematic Zoology, 40:131-151. 


308 HONEYCUTT AND YATES 


SPRINGER, M. S., AND C. KRAJEWSKI. 1989. DNA 
hybridization in animal taxonomy: a critique from 
first principles. Quarterly Review of Biology, 64:29 1- 
318. 

STANGL, F. G., JR. 1986. Aspects of a contact zone 
between two chromosomal races of Peromyscus leu- 
copus (Rodentia: Cricetidae). Journal of Mammal- 
ogy, 67:465-473. 

STANHOPE, M. J., J. CZELUSNIAK, J.-S. Si, J. NICKERSON, 
AND M. GoopMAN. 1992. A molecular perspective 
on mammalian evolution from the gene encoding 
interphotoreceptor retinoid binding protein, with 
convincing evidence for bat monophyly. Molecular 
Phylogenetics and Evolution, 1:148-160. 

STEWART, C.-B., AND A. C. WILSON. 1987. Sequence 
convergence and functional adaptation of stomach 
lysozymes from foregut fermenters. Cold Spring 
Harbor Symposium Quantitative Biology, 52:891- 
899. 

STRANEY, D. O., M. H. Smitu, R. J. BAKER, AND I. F. 
GREENBAUM. 1976. Biochemical variation and genic 
similarity of Myotis velifer and Macrotus californi- 
cus. Comparative Biochemistry and Physiology, A. 
Comparative Biochemistry, 54:243-248. 

STRANEY, D. O., M. H. Smitu, I. F. GREENBAUM, AND 
R. J. BAKER. 1979. Biochemical genetics. Pp. 157- 
176, in Biology of bats of the New World family 
Phyllostomatidae, Part III (R. J. Baker, J. K. Jones, 
Jr., and D. C. Carter, eds.). Special Publications, The 
Museum, Texas Tech University, 16:1-441. 

SupDMAN, P. D., AND M. S. HAFNER. 1992. Phyloge- 
netic relationships among Middle American pocket 
gophers (genus Orthogeomys) based on mitochon- 
drial DNA sequences. Molecular Phylogenetics and 
Evolution, 1:17-25. 

SULLIVAN, R. M. 1985. Phyletic, biogeographic, and 
ecologic relationships among montane populations 
of least chipmunks (Eutamias minimus) in the 
southwest. Systematic Zoology, 34:419-448. 

SULLIVAN, R. M., D. J. HAFNER, AND T. L. YATES. 
1986. Genetics of a contact zone between two chro- 
mosomal forms of the grasshopper mouse (genus 
Onychomys): a reassessment. Journal of Mammal- 
ogy, 67:640-659. 

SULLIVAN, R. M., AND T. L. YATEs. In press. Popu- 
lation genetics and conservation of relict populations 
of the red squirrel (genus 7amiasciurus). Special 
Publications, Smithsonian Institution. 

SvopopA, P. L., J. R. CHOATE, AND R. K. CHESSER. 
1985. Genetic relationships among southwestern 
populations of the Brasilian free-tailed bat. Journal 
of Mammalogy, 66:444-450. 

SwoFForD, D. L. 1990. PAUP: phylogenetic analysis 
using parsimony, version 3.0. Illinois Natural His- 
tory Survey, Champaign, Illinois. 

SWOFFORD, D. L., AND S. H. BERLOCHER. 1987. In- 
ferring evolutionary trees from gene frequency data 
under the principle of maximum parsimony. Sys- 
tematic Zoology, 36:293-325. 

SWOFFORD, D. L., AND G. J. OLSEN. 1990. Phylogeny 
reconstruction. Pp. 411-501, in Molecular system- 
atics (D. M. Hillis and C. Moritz, eds.). Sinauer As- 
sociates, Sunderland, Massachusetts, 588 pp. 

THomas, W. K., S. PAABO, F. X. VWILLABLANCA, AND 
A. C. Witson. 1990. Spatial and temporal conti- 


nuity of kangaroo rat populations shown by sequenc- 
ing mitochondrial DNA from museum specimens. 
Journal of Molecular Evolution, 31:101-112. 

TOLLIVER, D. K., AND L. W. Rossins. 1987. Genetic 
variability within Blarina carolinensis, and among 
three sympatric species of shrews (Insectivora: Soric- 
idae). Journal of Mammalogy, 68:387-390. 

TOLLIVER, D. K., M. H. SMITH, AND R. H. LEFrwicu. 
1985. Genetic variability in Insectivora. Journal of 
Mammalogy, 66:405-410. 

VAWTER, L., AND W. M. Brown. 1986. Nuclear and 
mitochondrial DNA comparisons reveal extreme rate 
variation in the molecular clock. Science, 234:194— 
196. 

Watson, J. D., AND F. H. C. Crick. 1953. Molecular 
structure of nucleic acids. Nature, 25:737. 

WAYNE, R. K., ANDS. M. JENKS. 1991. Mitochondrial 
DNA analysis implying extensive hybridization of 
the endangered red wolf Canis rufus. Nature, 351: 
365-368. 

WAYNE, R. K., AND S. J. O'BRIEN. 1987. Allozyme 
divergence within the Canidae. Systematic Zoology, 
36:339-355. 

WAYNE, R. K., N. LEHMAN, M. W. ALLARD, AND R. 
L. Honeycutt. 1992. Mitochondrial DNA vari- 
ability of the gray wolf: genetic consequences of pop- 
ulation decline and habitat fragmentation. Conser- 
vation Biology, 6:558-569. 

WAYNE, R. K., L. FORMAN, A. K. Newman, J. M. 
SIMONSON, AND S. J. O'BRIEN. 1986. Genetic mon- 
itors of zoo populations: morphological and electro- 
phoretic assays. Zoo Biology, 5:215-232. 

Wayne, R. K., ET AL. 1991. Conservation genetics 
of the endangered Isle Royale gray wolf. Conserva- 
tion Biology, 5:41-S1. 

Witey, E. O. 1981. Phylogenetics: the theory and 
practice of phylogenetic systematics. John Wiley & 
Sons, New York, 439 pp. 
divergence within the Canidae. Systematic Zoology, 
36:339-355. 

Witey, E. O. 1981. Phylogenetics: the theory and 
practice of phylogenetic systematics. John Wiley & 
Sons, New York, 439 pp. 

WILKINSON, G. S. 1985. The social organization of 
the common vampire bat. II. Mating system, genetic 
structure, and relatedness. Behavioral Ecology and 
Sociobiology, 17:123-134. 

Wiison, A. C., S. S. CARLSON, AND T. J. WHITE. 1977. 
Biochemical evolution. Annual Review of Biochem- 
istry, 46:573-639. 

Witison, A. C., AND V. M. SaricH. 1969. A molecular 
time scale for hominid evolution. Proceedings of the 
National Academy of Sciences (USA), 63:1088-1093. 

Woops, C. A. 1982. The history and classification of 
South American hystricognath rodents: reflections 
on the far away and long ago. Pp. 377-392, in Mam- 
malian biology in South America (M. A. Mares and 
H. H. Genoways, eds.). University of Pittsburgh Press, 
Pittsburgh, Pennsylvania, 539 pp. 

Wu, C.-I., AND W.-H. Li. 1985. Evidence for higher 
rates of nucleotide substitution in rodents than in 
man. Proceedings of the National Academy of Sci- 
ences (USA), 82:1741-1745. 

Wyss, A. R., M. J. NovACEK, AND M. C. MCKENNA. 
1987. Amino acid sequence versus morphological 


MOLECULAR SYSTEMATICS 309 


data and the interordinal relationships of mammals. of the genus Peromyscus (Rodentia). Comparative 
Molecular Biology and Evolution, 4:99-116. Biochemistry and Physiology, B. Comparative Bio- 
YATES, T. L. 1983. The mole that keeps its nose clean. chemistry, 52:541-54S. 
Natural History, 92:54-61. ZIMMERMAN, E. G., C. W. KILPATRICK, AND B. J. HART. 
YATES, T. L., ANDI. F. GREENBAUM. 1982. Biochem- 1978. The genetics of speciation in the rodent genus 
ical systematics of North American moles (Insectiv- Peromyscus. Evolution, 32:565-579. 
ora: Talpidae). Journal of Mammalogy, 63:368-374. ZIMMERMAN, E. G., AND M. E. NEJTEK. 1977. Ge- 
YATES, T. L., AND D. W. Moore. 1990. Speciation netics and speciation of three semispecies of Neot- 
and evolution in the family Talpidae (Mammalia: oma. Journal of Mammalogy, 587:39 1-402. 
Insectivora). Pp. 1-22, in Evolution of subterranean ZUKERKANDL, E., AND L. PAULING. 1965. Evolution- 
mammals at the organismal and molecular levels (E. ary divergence and convergence in proteins. Pp. 97- 
Nevo and O. Reig, eds.). Liss/Wiley, New York, 422 166, in Evolving genes and proteins (V. Bryson and 
pp. H. J. Vogel, eds.). Academic Press, New York, 629 
ZIMMERMAN, E. G., B. J. HART, AND C. W. KILPATRICK. pp. 


1975. Biochemical genetics of truei and boylei groups 


CYTOGENETICS 


ROBERT J. BAKER AND MARK S. HAFNER 


Introduction 


hen we were invited to prepare a re- 
view of the field of cytogenetics for 
the 75th anniversary of the ASM we had 
several discussions on the breadth and na- 
ture of the subject. This sent us scurrying 
to A Dictionary of Genetics (King and Stans- 
field, 1990:98) to determine the exact def- 
inition of the word cytogenetics: cytogenet- 
ics—the science that combines the methods 
and findings of cytology and genetics. This 
definition failed to provide us with the res- 
olution that we desired. Pertaining to the 
field of mammalogy, the word cytogenetics 
is a synonym for karyology, chromosomal 
evolution, or chromosome biology. 
Chromosomes, or collectively, the karyo- 
type, are subcellular morphological entities, 
and this chapter on cytogenetics is the only 
such chapter devoted to a single cellular or- 
ganelle. Why then is the karyotype accorded 
such an important position in mammalogy? 
Several books have been written on this 
subject, of which two of the best are M. J. 
D. White’s Animal Cytology and Evolution 
(1973) and Modes of Speciation (1978b). 
This organelle (the chromosome) has been 
implicated in many biological phenome- 
na including speciation (Baker and Bick- 
ham, 1986; Bush et al., 1977; White, 1968, 
1978a), rapid morphological change (Wil- 


310 


son et al., 1974), gene duplication (White, 
19785), and sex determination (Bull, 1983; 
Ohno, 1967). However, there may even be 
more basic reasons that the karyotype has 
been important to mammalogy. Before the 
advent of molecular biology, there were few 
easily quantified characters that provided 
systematic resolution among closely related 
species. The karyotype represents such a 
character. In addition, most karyological 
techniques are adaptable to field conditions 
and require minimal expense; therefore, it 
is not surprising that a number of mam- 
malogists have chosen to specialize in this 
area. As is the case with many other sub- 
disciplines, the field of cytogenetics extends 
far beyond the classical limits of mammal- 
ogy. For example, cytogenetics has impor- 
tant implications in the fields of carcino- 
genesis, mutagenesis, and medicine. Herein, 
however, we restrict our report to cytoge- 
netics as related to the science of mam- 
malogy. 


Conceptual Development of 
the Field 


The field of cytogenetics was essentially 
nonexistent prior to 1919. Until the 1950s, 


CYTOGENETICS ell 


no reliable methods were available to de- 
termine diploid number or morphology of 
chromosomes. Although the theory that he- 
redity was chromosomally based was de- 
veloped in the 1890s, this discovery had 
little immediate impact on the field of mam- 
malogy. A brief review of the history of our 
understanding of the karyotype of the hu- 
man provides insight into the state of the 
methods available during this time. In the 
early 1920s, the diploid number for Homo 
sapiens was commonly described as 24. In 
1923, T. S. Painter reported the diploid 
number was 48 with an XX/XY sex-deter- 
mining system (Painter, 1923). Not until 
1956 (Tjio and Levan, 1956) was the correct 
diploid number (46) determined. The sig- 
nificance of the difficulty in documenting 
the human karyotype is that early methods 
were tedious, subjective, and labor inten- 
sive, and they could not be adapted easily 
to the type of survey work that mammal- 
ogists usually conduct. Nevertheless, by 
1951, two significant lists of chromosomal 
data had been generated that together de- 
scribed the diploid or haploid numbers of 
approximately 175 species of mammals 
(Makino, 1951; Matthey, 1950). As verified 
by more recent studies, the majority of these 
descriptions were reasonably accurate. 
Even though technical aspects of the field 
of cytogenetics were rather primitive until 
the mid-1950s, some theoretical and con- 
ceptual aspects of the field were remarkably 
current as early as the 1920s. The following 
quote is from Painter (1925:407—408): 


“In the present paper a good deal of atten- 
tion has been given to chromosome num- 
bers, yet at the same time it has been fully 
realized that numbers per se are of second- 
ary importance. The significant point is that 
as far as we can gauge it, the total amount 
of chromatin in the different mammalian 
groups 1s about the same, and there has been 
a remarkabie stability in the chromosome 
associations. Inferentially, we may surmise 
that the total number of genes is about the 


same in all groups. In their chromosome 
constitution, the mammals have shown 
themselves, so far at least, comparable to 
an order of insects. 

If my general conclusion is a valid one, 
then we may expect that the plotting of 
chromosome maps in the eutheria will go 
forward with comparative rapidity, because 
linkage values established in one group or 
species can be applied to other forms... . 
Transverse fragmentation or end to end fu- 
sion will occasionally upset these relations, 
but on the whole they should prove the same 
in different forms, and enable us eventually 
to plot the chromosome maps of the euth- 
Cilawe 


Painter’s (1925) insights into chromo- 
somal evolution and the future of mam- 
malian cytogenetics were remarkably pre- 
scient, especially considering the dearth of 
actual data that existed in the field of cy- 
togenetics in the mid-1920s. We encourage 
the student of cytogenetics to review Pain- 
ter’s article in its entirety. 

In the 1960s, there was a burst of activity 
in the field of cytogenetics that produced 
accurate diploid numbers and descriptions 
of karyotypes for a wide variety of mam- 
malian taxa. Interpretation of these new data 
was influenced strongly by prevailing views 
of chromosome evolution in the 1950s and 
1960s. For example, it was widely held that 
most or all chromosome rearrangements re- 
duced fertility (i.e., fitness); hence, karyo- 
typic differences were generally viewed as 
indicators of species distinctiveness. For this 
reason, the first examples of chromosomal 
polymorphism discovered within taxa that 
behaved otherwise as biological species 
(Ford et al., 1957) received considerable at- 
tention. Of course there are several exam- 
ples where numerous chromosomal poly- 
morphisms exist in naturally occurring 
populations and these demonstrate rather 
conclusively that fitness reduction in het- 
erozygotes can be minimal if not nonexis- 
tent (Koop et al., 1983; Nachman, 1992a, 


piZ BAKER AND HAFNER 


1992b; Nachman and Myers, 1989; Stangl, 
1986). 

Most mammalian cytogeneticists of the 
1960s also assumed that karyotypes iden- 
tical in gross morphology were also identical 
at the level of gene order. Of course G-band- 
ing has shown that similar nonbanded kar- 
yotypes may underestimate amounts of 
chromosomal evolution by several orders 
of magnitude (Baker and Bickham, 1980; 
Haiduk et al., 1981). Breakage points in 
chromosomes were assumed to be stochas- 
tic, such that the independent occurrence of 
the same rearrangement in separate lineages 
was considered highly improbable and con- 
vergent evolution would not be a problem 
in cytogenetics. The significance of this con- 
clusion is that when two taxa shared a chro- 
mosomal rearrangement identified by 
G-bands, its usefulness as a synapomorphy 
was almost beyond question. This too has 
been shown to be inaccurate by several ex- 
amples, including chromosome 6 in 30 spe- 
cies of Peromyscus, which may have been 
rearranged as many as seven times (Stangl 
and Baker, 1984). The strongest evidence 
that the same chromosomal rearrangement 
can occur repeatedly comes from studies of 
human families that have unusual rear- 
rangements (such as the 11q;22q; Fraccaro 
et al., 1980) that have arisen independently 
in many families from widely separated geo- 
graphic origins. Chromosomal evolution 
was thought to be a highly ordered and time- 
dependent process (John and Lewis, 1966; 
for review see Baker et al., 1987). Therefore, 
taxa distinguished by a large number of 
chromosomal differences were thought to be 
distantly related. Examples such as the fol- 
lowing two document that little time or ge- 
netic distance is required in some cases 
where extensive chromosomal evolution has 
occurred. 1) Despite the karyotypic differ- 
ences between the species of Muntiacus (one 
with 2n = 6, 7 and the other with 2n = 46), 
viable offspring are produced by interspe- 
cific crosses of the two (Wurster and Be- 
nirschke, 1970). 2) Reithrodontomys mega- 
lotisand R. zacatacae have widely divergent 
karyotypes distinguished by over 30 rear- 


rangements, but the two are not distin- 
guished by any differences in allozymes at 
30 loci (Hood et al., 1984; Nelson et al., 
1984). 

In the 1960s, chromosomes were be- 
lieved to be stable structures and exchanges 
between nonhomologous chromosomes 
were thought to be rare. Barbara Mc- 
Clintock’s Nobel Prize-winning work (1978) 
provided the first insights into an excep- 
tionally dynamic process of exchange among 
nonhomologous chromosomes. Although 
the syntenic groups shared by various or- 
ders of mammals (O’Brien et al., 1985) in- 
dicate a measure of stability in the karyo- 
type, nonetheless it is widely documented 
that the exchange of transposable elements, 
heterochromatin, and other pieces of DNA, 
such as ribosomal genes, between nonho- 
mologous chromosomes is a common pro- 
cess (Arnheim et al., 1980; Dover, 1982; 
Hamilton et al., 1990, 1992; Wichman et 
al., 1991, 1992). Concepts about chromo- 
somal evolution and the forces that result 
in chromosomal conservatism in some lin- 
eages and rapid change in others are being 
revised continually (Baker et al., 1987; 
Bradley and Wichman, in press; Grapho- 
datsky, 1989; Patton and Sherwood, 1983; 
Wichman et al., 1991, 1992). The primary 
focus at this time reflects recent technolog- 
ical advances associated with molecular bi- 
ology, which has permitted more sophisti- 
cated experiments and testing of the 
molecular based hypotheses associated with 
cytogenetics. 


Technological Advances 


Although the microscope was invented in 
1590 by Hans and Zacharias Janssen in 
Holland (King and Stansfield, 1990), in- 
struments powerful enough to observe chro- 
mosomes were not designed until the 1800s. 
It was not until 1888 that the term chro- 
mosome was introduced by Wilhelm Wal- 
deyer. The X chromosome was documented 
in 1891 by Henking, who also described its 
meiotic behavior. Henking (1891) used the 


CYTOGENETICS eB: 


term ““X”’ because the function of the chro- 
mosome was unknown. The concept of the 
X chromosome’s involvement in sex deter- 
mination was developed by McClung (1901, 
1902). The Y chromosome was first de- 
scribed by Wilson (1909). In 1901, Mont- 
gomery associated maternal and paternal 
chromosomes into pairs (homologous chro- 
mosomes) and related this to Mendel’s ge- 
netic laws. By 1903 the role of the chro- 
mosome in heredity was demonstrated 
conclusively by Sutton (1902, 1903). 

One technical difficulty in examining 
chromosome morphology and number 
stems from the fact that the cellular space 
is small and the methods used to examine 
chromosomes before 1960 involved 
squashing cells between a microscope slide 
and a coverslip (Hsu, 1979). The end result 
of this procedure was poorly spread masses 
of chromosomes whose depth extended be- 
yond the normal field of focus for light mi- 
croscopes. Therefore, chromosomal counts 
were made by following within the mass of 
chromosomes an individual chromosome 
through several focal lengths. Needless to 
say, this process was exceedingly tedious and 
often inaccurate. 

A technical breakthrough that was of par- 
amount importance in determining chro- 
mosomal morphology was the hypotonic 
pretreatment of cells to enlarge the cells and 
aid in the ability to see each chromosome 
of the karyotype as an independent unit in 
a single field of focus. Hsu (1979) calls this 
the hypotonic miracle in his well-written 
documentation of this discovery. Although 
the effects of hypotonic treatment of cells 
were described by Slifer in 1934, the sig- 
nificance of her discovery to the field of cy- 
togenetics went unnoticed for almost two 
decades. In 1952, three papers (Hsu, 1952; 
Hughes, 1952; Makino and Nishimura, 
1952) were published describing the hypo- 
tonic pretreatment phenomenon. Ultimate- 
ly, hypotonic pretreatment was combined 
with another methodological breakthrough, 
the blaze-dry method (Scherz, 1962), to 
spread the chromosomes effectively from a 
single cell into a broader field for easier 


viewing of chromosomal detail. Students of 
cytogenetics who are interested in the his- 
tory and development of this field should 
read Hsu’s (1979) account. 

Another major methodological break- 
through in the field of cytogenetics was 
Krishnan’s (1968) discovery that mitotic in- 
hibitors such as Colchicine and vinblastine 
sulfate (Velban) arrest cell division at the 
metaphase plate. Mitotic inhibitors had been 
used commonly in plant genetics long be- 
fore they were applied to mammalian cy- 
togenetics. For example, Blakeslee and 
Avery demonstrated as early as 1937 that 
Colchicine induced polyploidy in plants. 

Techniques for preferential staining of 
particular regions of chromosomes (col- 
lectively called ‘“‘banding’’ techniques) 
stemmed from work by Caspersson et al. 
(1968, 1970) and Pardue and Gall (1970). 
Those generally acknowledged as producing 
the first Q-bands are Caspersson et al. (1968, 
1970), and production of the first C-bands 
is credited to Pardue and Gall (1970) and 
Arrighi and Hsu (1971). G-bands were first 
documented by Seabright (1971) and Sum- 
ner et al. (1971), R-bands were developed 
by Dutrillaux and Lejeune (1971), and stains 
specific for nucleolar organizing regions 
(NORs) are credited to Matsui and Sasaki 
(1973). Modern techniques for in situ hy- 
bridization stemmed from work by Gall and 
Pardue (1969) and John et al. (1969). In situ 
hybridization techniques advanced even 
further with the introduction of nonradioac- 
tive antibody probes visualized with en- 
zymes or fluorescent dyes (Frommer et al., 
1988; Langer et al., 1981; Manuelidis et al., 
1982; Pinkel et al., 1986). A modern review 
of chromosome banding and other cytoge- 
netic methods was provided by Sumner 
(1990). 


Cytogenetic Studies: Insights 
from the Journal of 
Mammalogy 


There are more than 9,000 scientific jour- 
nals that deal with the biological sciences. 


314 BAKER AND HAFNER 


In 1992 alone, nearly 7,000 articles in the 
field of cytogenetics were published in no 
fewer than 627 different journals (Macgre- 
gor, 1993). Because of the revolution in mo- 
lecular biology, the scope of cytogenetics is 
ever expanding. We feel that valuable in- 
sights into the nature of the science of mam- 
malogy can be gained by examination of 
publications in the Journal of Mammalogy 
that appeared during this period of expan- 
sion of the science of cytogenetics. Approx- 
imately 130 studies emphasizing chro- 
mosomes or using cytogenetic data or tech- 
niques have been published in the Journal 
of Mammalogy since its inception. Included 
in these studies are the first descriptions of 
karyotypes of roughly 284 species of mam- 
mals, including the first karyotypes reported 
for many mammalian genera and several 
families. As the following account will doc- 
ument, the Journal of Mammalogy played 
only a minor role in the early history of the 
field of cytogenetics. However, in 1967 it 
was thrust into the mainstream of mammal 
cytogenetic research, largely due to the im- 
provement of karyotyping techniques such 
as use of mitotic inhibitors and blaze-dry 
methods that improved the spreading of 
chromosomes. Since 1966 (Nadler, 1966; 
Nadler and Hughes, 1966; Singh and Mc- 
Millan, 1966), the Journal of Mammalogy 
has played an important role in the field of 
mammal cytogenetics, especially in the sub- 
fields of cytotaxonomy and cytosystematics. 

Readers of the Journal of Mammalogy 
were introduced to the nascent field of cy- 
togenetics in L. C. Dunn’s (1921) study of 
coat-color inheritance in rodents. This study, 
which also introduced many mammalogists 
to Mendelian genetics, reported that diploid 
numbers were known at that time for only 
four species of rodents: the mouse (Mus); 
the rat (Rattus); the guinea pig (Cavia); and 
the Old World rabbit (Oryctolagus; rabbits 
were then classified as rodents). Based on 
this fragmentary evidence, Dunn (1921:139) 
made a remarkably insightful speculation, 
““... there is some slight evidence that in 
the evolution of rodents a fractionation of 


chromosomes may have occurred, for the 
mice and rats have 19 (haploid) while the 
guinea-pigs have 28.’ This comment was 
all the more remarkable considering that the 
entire concept of organic evolution was open 
to question when Dunn published this work. 
With reference to the haploid-number sim- 
ilarity between Mus and Rattus, Dunn (1921: 
139) commented: “Whether this is due to 
a community of descent in the terms of cur- 
rent evolutionary theory or to relationship 
through some other cause is one of the ques- 
tions which genetics, aided by the chro- 
mosome notation, may be expected at some 
time to answer.” 

Seventy-two years later Science published 
a genome issue showing a genetic linkage 
map of Mus (Copeland et al., 1993) docu- 
menting exactly the kinds of results pre- 
dicted by Dunn (1921). Copeland et al. 
(1993) calculated that based on linkage 
maps, the mouse and the human have un- 
dergone approximately 150 chromosomal 
rearrangements since they last shared a 
common ancestor (Nadeau and Taylor, 
1984). 

The first figure of chromosomes pub- 
lished in the Journal of Mammalogy was a 
camera lucida drawing of meiotic prophase 
tetrads of the house mouse, Mus musculus 
(Hoy and Berkowitz, 1931). Although this 
article described a relatively simple method 
for fixation and preservation of chromo- 
somes in the field, it did not catalyze the 
intense interest in mammalian chromo- 
somes anticipated by its authors. To the 
contrary, this article was followed by a hi- 
atus of almost 30 years, during which time 
no cytogenetic paper was published in the 
Journal of Mammalogy. 

As noted above, two landmark books were 
published during this time in the rapidly 
expanding field of cytogenetics: Matthey’s 
(1950) Les Chromosomes des Vertebres, and 
Makino’s (1951) An Atlas of the Chromo- 
some Numbers in Animals. Although these 
books were primarily compendia of diploid 
and fundamental numbers known at that 
time, Matthey (1950) speculated on the po- 


CYTOGENETICS 315 


tential systematic value of chromosomes in 
the Mammalia. Johnson and Ostenson 
(1959) were the first to publish a paper in 
the Journal of Mammalogy that empha- 
sized the potential usefulness of chromo- 
somes as taxonomic characters. Their study 
was primarily a review of taxonomic meth- 
ods available in 1959, and they reported no 
new mammalian karyotypes. However, 
Johnson and Ostenson (1959:573) referred 
to Matthey’s (1952) pioneering studies of 
microtine chromosomes and stated: “Such 
a fundamental difference as in chromo- 
somes [between two voles, Microtus agrestis 
and M. pennsylvanicus| must be regarded 
as strong evidence of species difference.” 
This was the first of many such statements 
to appear in the Journal of Mammalogy sig- 
naling the taxonomic importance of cyto- 
genetic characters. 

The first figure of a mitotic-metaphase 
karyotype to be published in the Journal of 
Mammalogy appeared in volume 47 (Nad- 
ler and Hughes, 1966). This karyotype of a 
ground squirrel (Spermophilus spilosoma) 
was remarkably clear and showed in con- 
siderable detail individual chromosomal el- 
ements. The same year, Nadler (1966) pub- 
lished the first diagram to appear in the 
Journal showing hypothetical chromosom- 
al changes that occurred during the evolu- 
tionary history of a mammalian lineage (in 
this case, the ground squirrel subgenus Sper- 
mophilus). Nadler’s (1966) paper was among 
the first to bring cytogenetic evidence to bear 
on higher-order questions in the field of 
mammalian systematics, a field that, before 
that time, had been dominated by morpho- 
logical and paleontological studies. 

Before 1967, articles on mammalian cy- 
togenetics were published in a wide variety 
of outlets including The American Natu- 
ralist, Anatomical Record, Chromosoma, 
Experientia, Journal of Genetics, Journal of 
Morphology, Proceedings of the Society of 
Experimental Biology and Medicine, and a 
myriad of other books, journals, proceed- 
ings, and reports. In an effort to organize 
the rapidly expanding literature in this field, 


Hsu and Benirschke published in 1967 their 
important compendium titled, 4n Atlas of 
Mammalian Chromosomes. 

Methodological breakthroughs in the field 
of cytogenetics in 1967 triggered a major 
thrust in this research area worldwide. In- 
strumental in development of these new 
methods was James L. Patton, then a grad- 
uate student at the University of Arizona. 
The University of Arizona was a nucleus 
for this type of activity at this time with 
Patton and Robert J. Baker focusing on 
mammalian cytogenetics. Fortunately, Pat- 
ton and Baker chose to publish many of 
their earliest cytogenetic studies in the Jour- 
nal of Mammalogy (e.g., Baker and Patton, 
1967; Patton, 1967; Patton and Hsu, 1967), 
which in concert with others (Nadler, 1966; 
Nadler and Hughes, 1966; Singh and Mc- 
Millan, 1966) brought the Journal into the 
mainstream of cytogenetics research. Baker 
and Patton’s seminal contributions to the 
field of mammalian cytogenetics and, in 
particular, their development of convenient 
techniques for use in the field (e.g., Baker, 
1970; Patton, 1967), are still widely cited 
in the cytogenetics literature. 

An analysis of the rate of publication of 
cytogenetic studies in the Journal of Mam- 
malogy from the time of the journal’s in- 
ception (1920) to the present (Fig. 1) illus- 
trates the enormous surge in this field that 
began in the 1960s. For example, no cyto- 
genetic studies appeared in the Journal be- 
tween 1961 and 1965; in contrast, 22 such 
articles appeared for the time period of 1966 
to 1970. Similarly, no new karyotypes were 
described in the Journal during the first half 
of the 1960s, whereas the karyotypes of 85 
species of mammals were reported there for 
the first time between 1966 and 1970. 

Most cytogenetic studies published in the 
Journal of Mammalogy in the late 1960s 
and early 1970s were descriptive in nature, 
and most authors linked—explicitly or im- 
plicitly—chromosomal differentiation with 
taxonomic distinctness. For example, Shell- 
hammer (1967:549) stated (with respect to 
two species of harvest mice, Reithrodonto- 


316 BAKER AND HAFNER 


mys) that: “the karyotypes... are different 
enough to suggest that the two are in the 
terminal stages of speciation.’’ However, as 
the karyotypes of more and more species of 
mammals were reported in the Journal and 
elsewhere, it became apparent that chro- 
mosomal variation in mammals was not 
necessarily linked to the process of specia- 
tion and that chromosomal variation, in 
general, was much more complex than had 
been envisioned by earlier workers in the 
field. In a study that described the karyo- 
types of 32 species of vespertilionid bats, 
Baker and Patton (1967:283) stated: ““From 
the few studies of mammalian karyotypes 
that have thus far been made, it appears 
obvious that the degree of karyotypic vari- 
ation encountered at a given taxonomic lev- 
el. ..1is in itself highly variable from mam- 
malian group to group.” 

Thus began a period of intensive surveys 
of chromosomal variation in mammals, 
which was the subject of several articles 
published in the Journal beginning in 1968 
(e.g., Blanks and Shellhammer, 1968; Lee 
and Zimmerman, 1969; Nelson-Rees et al., 
1968). Although intraspecific chromosomal 
polymorphism had been known since Ford 
et al.’s (1957) classic study of shrews (Sorex 
araneus), the genetic consequences and evo- 
lutionary significance of chromosomal 
polymorphism were only poorly under- 
stood even a decade later. For example, 
Blanks and Shellhammer (1968:729), whose 
article in the Journal of Mammalogy was 
the first report of supernumerary chromo- 
somes in mammals, stated candidly: ““We 
do not understand the mode of inheritance 
of the small chromosomes... .”” Not sur- 
prisingly, this period of intensive karyolog- 
ical surveys (1966-1970) generated a large 
gap between data and theory in the field of 
mammalian cytogenetics. This, in turn, led 
to a certain amount of disillusionment on 
the part of workers attempting to solve tax- 
onomic problems using chromosomal evi- 
dence. For example, Sutton and Nadler 
(1969:534) stated: ““Chromosomes are of 
limited value for the solution of taxonomic 


35 


> 
N 


Number of Karyotypes Described ai s 


Number of Cytogenetics Publications—_{, oN 


Number of Publications 
Number of Karyotypes 


0 T T T —- 
1925 1930 1935 1940 1945 1950 1955 1960 1965 1970 1975 1980 1985 1990 


Year 


Fic. 1.—Number of cytogenetics publications 
and number of new karyotypes described in the 
Journal of Mammalogy from 1919 to 1990. The 
axes are scaled differently to show that the rate 
of publication of cytogenetics research increased 
throughout the 1970s, whereas the rate of pub- 
lication of new karyotypes has declined consis- 
tently since 1970. 


problems and they are of little help in es- 
tablishing relationships between species and 
subspecies of the genus Eutamias [chip- 
munks].”’ 

As the number of studies reporting intra- 
specific chromosomal variation in mam- 
mals increased, there was growing confu- 
sion in the literature with respect to the terms 
‘“‘seographic variation’ and “‘polymor- 
phism.”’ Fortunately, Lee and Zimmer- 
man’s (1969) chromosomal study of cotton 
rats (Sigmodon) stemmed the tide of grow- 
ing confusion by carefully distinguishing be- 
tween geographic variation (“‘. . . differences 
in karyotype between [presumably conspe- 
cific] organisms from different localities 
...’) and chromosomal polymorphism 
(““... variation within a geographically lo- 
calized, panmictic population.’’) (Lee and 
Zimmerman, 1969:335-—336). 

Patton and Dingman’s (1968) cytogenetic 
study of natural hybridization between the 


CYTOGENETICS od7 


pocket gophers Thomomys bottae and T. 
umbrinus was published in volume 49 of 
the Journal of Mammalogy. Although their 
taxonomic conclusion (that 7. bottae and T. 
umbrinus are distinct species) was contro- 
versial and was rejected by certain leading 
mammalogists of the time (e.g., Hall, 1981: 
469), they demonstrated for the first time 
the value of chromosomes in analyses of 
genetic introgression in mammals. Patton 
and Dingman’s (1968) taxonomic conclu- 
sion was bolstered 5 years later by a detailed 
analysis of meiosis in bottae x umbrinus 
hybrids (Patton, 1973). This work set the 
standard for cytogenetic studies of mammal 
hybrid zones for many years. 

From 1967 through 1972, most major 
publications in the field of mammalian cy- 
togenetics and reports of significant meth- 
odological and conceptual breakthroughs in 
the field were published in the journals 
Chromosoma, Cytogenetics, Experientia, 
Science, and Nature. During the same pe- 
riod, most studies describing the karyotypes 
of mammal species were published in Mam- 
malian Chromosomes Newsletter. Perhaps 
as a result, the number of karyotypes de- 
scribed in the Journal of Mammalogy began 
to decline in the early 1970s (from its peak 
in the late 1960s) and has continued to de- 
cline (Fig. 1). However, as more and more 
chromosomal data accumulated in the early 
1970s making large-scale syntheses possi- 
ble, noteworthy publications in the field of 
mammalian cytogenetics began to appear 
with increasing frequency in the journals 
Evolution, Hereditas, Systematic Zoology, 
and Journal of Mammalogy (Fig. 1). One 
particularly important contribution that ap- 
peared in the Journal of Mammalogy during 
this period was Nadler et al.’s (1971) study 
of prairie dog (Cynomys) evolution; this was 
the first of many studies published in the 
Journal that used combined chromosomal 
and biochemical evidence to address a sys- 
tematic problem. 

The early 1970s witnessed a renaissance 
in the field of cytogenetics that was cata- 
lyzed by the development of techniques for 


banding chromosomes that increased dra- 
matically the taxonomic usefulness of kar- 
yotypes. In his chromosomal study of kan- 
garoo rats (Dipodomys), Stock (1974) 
published the first figure of a metaphase 
karyotype stained for constitutive hetero- 
chromatin (““C-bands’’) and the first figure 
of a Geimsa-banded karyotype (““G-bands’’) 
to appear in the Journal. Stock’s contribu- 
tion was followed soon thereafter by a study 
that used banded karyotypes to document 
chromosomal conservatism in rodents 
(Mascarello et al., 1974a), and another that 
used banded karyotypes to confirm the role 
of Robertsonian mechanisms in the origin 
of chromosomal diversity in woodrats (Ne- 
otoma; Mascarello et al., 19745). Four years 
later, Mascarello (1978) introduced readers 
of the Journal of Mammalogy to yet another 
staining procedure (Ag-As silver staining), 
which was used to visualize nucleolus or- 
ganizing regions on individual chromo- 
somes. 

Development of these new staining pro- 
cedures in the mid-1970s triggered a burst 
of activity on the part of mammalian cy- 
tosystematists, and as a result the number 
of cytogenetic studies published in the Jour- 
nal of Mammalogy peaked between 1976 
and 1980 (Fig. 1). During this period, 
Greenbaum and Baker (1978) published the 
first article in the Journal that used C- and 
G-banded karyotypes to deduce the prim- 
itive karyotype for a group of mammals (in 
this case, white-footed mice of the genus 
Peromyscus). This was among the first stud- 
ies published anywhere in which a cytoge- 
netic analysis was viewed in the context of 
phylogenetic systematics. Bickham’s (1979) 
study of the chromosomal variation in ves- 
pertilionid bats used cladistic methods to 
produce a phylogeny of these taxa using 
G-banded karyotypes. 

The frequency of appearance of cytoge- 
netic publications in the Journal of Mam- 
malogy declined steadily during the 1980s 
and continues to decline today (Fig. 1). This 
trend probably reflects the general shift away 
from morphological and cytogenetic meth- 


318 BAKER AND HAFNER 


ods toward use of molecular methods by 
large numbers of mammalian biologists. 
This decline in frequency of cytogenetic 
studies during the 1980s has occurred de- 
spite the recent introduction of new and 
promising cytogenetic techniques. Notable 
among these new techniques are fluores- 
cent-banding procedures (Bickham, 1987) 
and flow cytometric studies of nuclear-DNA 
content (Burton and Bickham, 1989). The 
first color photo published in the Journal of 
Mammalogy appeared in an article by Ba- 
ker et al. (1992) that documented the num- 
ber of ribosomal gene sites in bats using 
fluorescent in situ hybridization. Although 
these new developments have failed, thus 
far, to reinvigorate the field of mammalian 
cytogenetics within the pages of the Journal 
of Mammalogy, there is little doubt that the 
next generation of mammalogists will re- 
discover the value of cytogenetic characters 
for genetic and systematic inquiry. 

Geographic and taxonomic coverage. — 
Published literature in the Journal of Mam- 
malogy shows a strong emphasis on North 
American species. This geographic bias is 
likewise reflected in the set of 130 studies 
categorized herein as cytogenetic research. 
For example, 102 of the 130 studies (78%) 
published between 1920 and 1990 in the 
field of cytogenetics have dealt exclusively 
with North American species. Of the re- 
maining 35 studies, 22 (17%) have involved 
Central or South American species, 6 (5%) 
have focused on African species, 5 (4%) on 
Asian species, and 2 (2%) on Australian or 
New Zealand species. 

All 79 karyotypes published in the Jour- 
nal of Mammalogy during its initial 50 years 
of existence (1920-1969) were from either 
rodents (14 studies/45 species) or bats (three 
studies/34 species). This trend was broken 
in 1970 when Holden and Eabry published 
the karyotypes of two species of rabbits (Sy/- 
vilagus). The first cetacean (Kulu et al., 1971) 
and artiodactyl (Nadler, 1971) karyotypes 
were published in volume 52, and the first 
carnivore karyotype appeared a year later 
(Wurster-Hill, 1973). Yates and Schmidly 


80 Taxonomic Representation for Major Orders of Mammals 


O Percentage of all mammal species 


604 ll Percentage of all karyotypes published in 


Journal of Mammalogy 


Percentage 


Fic. 2.—Taxonomic bias in the cytogenetics 
literature published in the Journal of Mammal- 
ogy. For each of the nine orders of mammals 
listed, the bar on the left represents the percent- 
age of all extant mammalian species that belong 
to that order and the bar on the right indicates 
the percentage of all karyotypes published in the 
Journal of Mammalogy relating to species of that 
order. Note that rodents are over-represented in 
the cytogenetics literature, whereas all other 
groups, except bats, are under-represented rela- 
tive to their species abundance. 


(1975) reported the first insectivore karyo- 
type, and the first marsupial karyotype ap- 
peared almost a decade later (Seluja et al., 
1984). Surprisingly, no other mammalian 
order is represented by karyotypes pub- 
lished in the Journal. 

Considering that bat species (Chiroptera) 
comprise approximately 22% of all living 
species of mammals (Anderson and Jones, 
1984), it seems appropriate that roughly 22% 
of all karyotypes that have appeared in the 
Journal of Mammalogy are from species of 
bats (Fig. 2). In contrast, rodents comprise 
approximately 42% of extant mammal spe- 
cies, yet nearly 70% of all karyotypes re- 
ported in the Journal are of rodents. This 
striking taxonomic bias in favor of rodents 


CYTOGENETICS 319 


is probably a consequence of the fact that 
most rodents are small and easily captured 
and karyotyped. 


Summary and Conclusions 


The field of cytogenetics was in its infancy 
when the ASM was founded in 1919. Per- 
haps in part because the Journal of Mam- 
malogy was not yet widely known in inter- 
national circles, early workers in the field of 
mammalian cytogenetics chose to publish 
results of their studies in journals with wider 
readership; hence the Journal played only a 
minor role in the early development of the 
field. In the 1960s, methodological advanc- 
es developed by several mammalogists, in- 
cluding Charles F. Nadler, James L. Patton, 
and Robert J. Baker, finally brought the 
Journal of Mammalogy into the main- 
stream of cytogenetics research. 

The future of cytogenetic studies is es- 
pecially promising. Recent advances in 
chromosome painting (Lengauer et al., 1990, 
1991), which can provide resolution to ho- 
mologous chromosomal regions among dis- 
tantly related taxa, should permit survey 
type work among various groups of mam- 
mals. Polymerase chain reaction amplifi- 
cation of chromosomal loci with conserved 
primers (Koch et al., 1991; Terkelsen et al., 
1993) should also be readily adaptable to 
the types of investigations that are valuable 
to the science of mammalogy. The use of 
multi-color in situ hybridizations (Reid et 
al., 1992; Scherthan et al., 1992) will permit 
examination of the order of genes on a chro- 
mosome during a single experiment. Chro- 
mosomal banding through computerized 
images using fluorescent dyes (K. L. Bowers, 
pers. comm.; Volpi and Baldini, 1993; Ward 
et al., 1991) will greatly facilitate identifi- 
cation of chromosomes without the nu- 
merous replications required by the old 
trypsin methods. The development of in situ 
probes from DNA libraries should provide 
countless loci to be mapped. These ad- 
vances indicate that we are only beginning 


to see the methodological improvements 
that will aid in cytogenetic analyses. Amid 
this technological growth, we note that there 
is a tremendous number of mammals for 
which karyotypic data are not available. 
Survey work in these areas is also desirable. 


Literature Cited 


ANDERSON, S., AND J. K. JONES, JR. (EDS.). 1984. Or- 
ders and families of Recent mammals of the world. 
John Wiley & Sons, New York, 686 pp. 

ARNHEIM, N., M. KrYSTAL, R. SCHMICKEL, G. WILSON, 
O. RyDER, AND E. Zimmer. 1980. Molecular evi- 
dence for genetic exchange among ribosomal genes 
on nonhomologous chromosomes in man and apes. 
Proceedings of the National Academy of Sciences, 
USA, 77:7323-7327. 

ARRIGHI, F. E., AND T. C. Hsu. 1971. Localization 
of heterochromatin in human chromosomes. Cyto- 
genetics, 10:81-86. 

BAKER, R. J. 1970. Karyotype trends in bats. Pp. 65- 
96, in Biology of bats (W. A. Wimsatt, ed.). Aca- 
demic Press, New York, 1:1—406. 

BAKER, R. J., AND J. W. BICKHAM. 1980. Karyotypic 
evolution in bats: evidence of extensive and conser- 
vative chromosomal evolution in closely related taxa. 
Systematic Zoology, 29:239-251. 

. 1986. Speciation by monobrachial centric fu- 
sions. Proceedings of the National Academy of Sci- 
ences, USA, 83:8245-8248. 

BAKER, R. J., AND J. L. PATTON. 1967. Karyotypes 
and karyotypic variation of North American ves- 
pertilionid bats. Journal of Mammalogy, 48:270- 
286. 

BAKER, R. J., M. B. QUMSIYEH, AND C.S. Hoop. 1987. 
Role of chromosomal banding patterns in under- 
standing mammalian evolution. Pp. 67-96, in Cur- 
rent mammalogy (H. H. Genoways, ed.). Plenum 
Press, New York, 1:1-519. 

BAKER, R. J., M. MALTBIE, J. G. Owen, M. J. HAm- 
ILTON, AND R. D. BRADLEY. 1992. Reduced num- 
ber of ribosomal sites in bats: evidence for a mech- 
anism to contain genome size. Journal of 
Mammalogy, 73:847-858. 

BICKHAM, J. W. 1979. Chromosomal variation and 
evolutionary relationships of vespertilionid bats. 
Journal of Mammalogy, 60:350-363. 

1987. Chromosomal variation among seven 
species of lasiurine bats (Chiroptera: Vespertilioni- 
dae). Journal of Mammalogy, 68:837-842. 

BLAKESLEE, A. F., AND A. G. Avery. 1937. Methods 
of inducing doubling in chromosomes in plants. 
Journal of Heredity, 28:392-411. 

BLANKS, G. A., ANDH.S.SHELLHAMMER. 1968. Chro- 
mosome polymorphism in California populations of 
harvest mice. Journal of Mammalogy, 49:726-731. 

BRADLEY, R. D., AND H. A. WICHMAN. In press. Rap- 
idly evolving repetitive DNAs in a conservative ge- 
nome: a test of factors that affect chromosomal evo- 
lution. Chromosome Research. 


320 BAKER AND HAFNER 


BuLt, J.J. 1983. Evolution of sex determining mech- 
anisms. Benjamin/Cummings, London, 316 pp. 
BurTON, D. W., AND J. W. BICKHAM. 1989. Hetero- 
chromatin variation and DNA content conservatism 
in Geomys attwateri and Geomys breviceps (Roden- 
tia: Geomyidae). Journal of Mammalogy, 70:580- 

591. 

Busu, G. L., S. M. Case, A. C. WILSON, AND J. L. 
PATTON. 1977. Rapid speciation and chromosomal 
evolution in mammals. Proceedings of the National 
Academy of Sciences, USA, 74:3942-3946. 

CASPERSSON, T., L. ZECH, AND C. JOHANSON. 1970. 
Differential banding of alkylating fluorochromes in 
human chromosomes. Experimental Cell Research, 
60:315-319. 

CASPERSSON, T., ET AL. 1968. Chemical differentia- 
tion along metaphase chromosomes. Experimental 
Cell Research, 49:219-222. 

COPELAND, N.G., ETAL. 1993. A genetic linkage map 
of the mouse: current applications and future pros- 
pects. Science, 262:57-66. 

Dover, G. A. 1982. Molecular drive: a cohesive mode 
of species evolution. Nature, 299:111-117. 

Dunn, L. C. 1921. Unit character variation in ro- 
dents. Journal of Mammalogy, 2:125-140. 

DuTRILLAUX, B., AND J. LEJEUNE. 1971. Sur une nou- 
velle technique d’analyse du caryotype humain. 
Comptes Rendus de |’Academie des Sciences (Paris) 
D, 272:2638-2640. 

Forp, C. E., J. L. HAMERTON, AND G. B. SHARMAN. 
1957. Chromosome polymorphism in the common 
shrew. Nature, 180:392-393. 

Fraccaro, M., ET AL. 1980. The 11q;22q translo- 
cation: a European collaborative analysis of 43 cases. 
Human Genetics, 56:21-51. 

FRomMER, M., C. PAUL, AND P. C. VINCENT. 1988. 
Localization of satellite DNA sequences on human 
metaphase chromosomes using bromodeoxyuri- 
dine-labelled probes. Chromosoma, 97:11-18. 

GALL, J. G., AND M. L. PARDuE. 1969. Formation 
and detection of RNA-DNA hybrid molecules in 
cytological preparations. Proceedings of the National 
Academy of Sciences, USA, 63:378-383. 

GRAPHODATSKY, A. S. 1989. Conserved and variable 
elements of mammalian chromosomes. Pp. 95-123, 
in Cytogenetics of animals (C. R. E. Halnan, ed.). 
CAB International Press, Wallingford, Oxon, United 
Kingdom, 519 pp. 

GREENBAUM, I. F., AND R. J. BAKER. 1978. Deter- 
mination of the primitive karyotype for Peromyscus. 
Journal of Mammalogy, 59:820-834. 

Harpuk, M. W., R. J. BAKER, L. W. Rossins, AND D. 
A. SCHLITTER. 1981. Chromosomal evolution in 
African Megachiroptera: G- and C-band assessment 
of the magnitude of change in similar standard kar- 
yotypes. Cytogenetics and Cell Genetics, 29:221-232. 

Hatt, E. R. 1981. The mammals of North America. 
Second ed. John Wiley & Sons, New York, 1:1-600 
+ 90. 

Hami.ton, M. J., R. L. HONEYCUTT, AND R. J. BAKER. 
1990. Intragenomic movement, sequence amplifi- 
cation and concerted evolution in satellite DNA in 
harvest mice, Reithrodontomys: evidence from in 
situ hybridization. Chromosoma, 99:321-329. 

Hamitton, M. J., G. HonG, AND H. A. WICHMAN. 


1992. Intragenomic movement and concerted evo- 
lution of satellite DNA in Peromyscus: evidence from 
in situ hybridization. Cytogenetics and Cell Genet- 
ics, 60:40-44. . 

HENKING, H. 1891. Uber Spermatogenese bei Pyr- 
rhocoris apterus. Z. wiss. Zoology, 51:685. 

HOLpDEN, E., AND H. S. EAsry. 1970. Chromosomes 
of Sylvilagus floridanus and Sylvilagus transitionalis. 
Journal of Mammalogy, 51:166-168. 

Hoop, C. S., L. W. Rossins, R. J. BAKER, AND H. S. 
SHELLHAMMER. 1984. Chromosomal studies and 
evolutionary relationships of an endangered species, 
Reithrodontomys raviventris. Journal of Mammalo- 
gy, 65:655-667. 

Hoy, W.E., Jr., AND K. BERLowiTz. 1931. A method 
for the fixation of mammalian chromosomes in the 
field. Journal of Mammalogy, 12:397-400. 

Hsu, T.C. 1952. Mammalian chromosomes in vitro. 
I. The karyotype of man. Journal of Heredity, 43: 
172 


. 1979. Human and mammalian cytogenetics: 
an historical perspective. Springer-Verlag, New York, 
186 pp. 

Hsu, T. C., AND K. BENIRSCHKE (EDS.). 1967. Anatlas 
of mammalian chromosomes. Volumes 1-4. Spring- 
er-Verlag, New York, 1967-1970. 

HuaGues, A. 1952. Some effects of abnormal tonicity 
on dividing cells in chick tissue cultures. Quarterly 
Journal of Microscopical Science, 93:207-220. 

Joun, B., AND K. R. Lewis. 1966. Chromosome vari- 
ability and geographic distribution in insects. Sci- 
ence, 152:711-721. 

Joun, H., M. L. BirNsTEIL, AND K. W. Jones. 1969. 
RNA-DNA hybrids at cytological levels. Nature, 223: 
582-587. 

Jounson, M. L., AND B. T. OsTENSON. 1959. Com- 
ments on the nomenclature of some mammals of the 
Pacific northwest. Journal of Mammalogy, 40:571- 
S17. 

Kina, R. C., AND W. D. STANSFIELD. 1990. A dictio- 
nary of genetics. Fourth ed. Oxford University Press, 
New York, 406 pp. 

Kocu, J., J. HinDKJAER, J. MORGENSEN, S. KOLVRAA, 
AND L. BoLunb. 1991. An improved method for 
chromosome specific labeling of alpha satellite DNA 
in situ using denatured double stranded DNA probes 
as primers in a PRimed IN Situ labeling (PRINS) 
procedure. Genetic Analysis and Technical Appli- 
cations, 8:171-178. 

Koop, B. F., R. J. BAKER, AND H. H. GeNoways. 1983. 
Numerous chromosomal polymorphisms in a nat- 
ural population of rice rats (Oryzomys: Cricetidae). 
Cytogenetics and Cell Genetics, 35:131-135. 

KRISHNAN, A. 1968. Time-lapse and ultrastructure 
studies on the reversal of mitotic arrest induced by 
vinblastine sulfate in Earle’s L cells. Journal of Na- 
tional Cancer Institute, 41:581-596. 

Kutvu, D. D., I. VEOMAT, AND R. S. Sparkes. 1971. 
Cytogenetic comparisons of four species of ceta- 
ceans. Journal of Mammalogy, 52:828-832. 

LANGER, P. R., A. A. WALDROP, AND D. C. WARD. 
1981. Enzymatic synthesis of biotin-labeled poly- 
nucleotides: novel nucleic acid affinity probes. Pro- 
ceedings of the National Academy of Sciences, USA, 
78:6633-6637. 


CYTOGENETICS 321 


Lee, M. R., AND E. G. ZIMMERMAN. 1969. Robert- 
sonian polymorphism in the cotton rat, Sigmodon 
fulviventer. Journal of Mammalogy, 50:333-339. 

LENGAUER, C., ET AL. 1991. Painting of defined chro- 
mosomal regions by in situ suppression hybridiza- 
tion of libraries from laser-microdissected chromo- 
somes. Cytogenetics and Cell Genetics, 56:27-30. 

LENGAUER, C., H. RIETHMAN, AND T. CREMER. 1990. 
Painting of human chromosomes with probes gen- 
erated from hybrid cell lines by PCR with Alu and 
L1 primers. Human Genetics, 86:1-6. 

Maccrecor, H.S. 1993. Chromosome research-look 
forward to 2001. Chromosome Research, 1:5-7. 

Makino, S. 1951. An atlas of the chromosome num- 
bers in animals. The Iowa State College Press, Ames, 
290 pp. 

MAKINO, S., AND I. NISHIMURA. 1952. Water pre- 
treatment squash technic: a new and simple practical 
method for the chromosome study of animals. Stain 
Technology, 27:1-7 

MANUELIDIS, L., P. R. LANGER-SAFER, AND D. C. WARD. 
1982. High-resolution mapping of satellite DNA 
using biotin-labeled DNA probes. Journal of Cell 
Biology, 95:619-625. 

MASCARELLO, J. T. 1978. Chromosomal, biochemi- 
cal, mensural, penile, and cranial variation in desert 
woodland woodrats (Neotoma lepida). Journal of 
Mammalogy, 59:477-495. 

MASCARELLO, J. T., A. D. Stock, AND S. PATHAK. 
1974a. Conservatism in the arrangement of genetic 
material in rodents. Journal of Mammalogy, 55:695- 
704. 

MASCARELLO, J. T., J. W. WARNER, AND R. J. BAKER. 
1974b. A chromosome banding analysis of the 
mechanisms involved in the karyological divergence 
of Neotoma phenax (Merriam) and Neotoma mi- 
crops Baird. Journal of Mammalogy, 55:831-834. 

Matsul, S., AND M. SASAKI. 1973. Differential stain- 
ing of nucleolus organizers in mammalian chro- 
mosomes. Nature, 246:148-150. 

MATTHEY, R. 1950. Les chromosomes des vertébrés. 
Université F. Rouge-Lausanne, 356 pp. 

. 1952. Chromosomes de Muridae (Microtinae 
et Cricetinae). Chromosoma, 5:1 13-138. 

McC iintock, B. 1978. Mechanisms that rapidly re- 
organize the genome. Stadler Symposium, 10:25-47. 

McCLuna, C. E. 1901. Notes on the accessory chro- 
mosome. Anatomischer Anzeiger, 20:220-226. 

1902. The accessory chromosome-sex deter- 
minant. Biological Bulletin, 3:43-84. 

Montcomery, T. H. 1901. A study of the chromo- 
somes of the germ cells of the Metazoa. Transactions 
of the American Philosophical Society, 20:154—236. 

NACHMAN, M. W. 1992a. Geographic patterns of 
chromosomal variation in South American marsh 
rats, Holochilus brasiliensis and H. vulpinus. Cyto- 
genetics and Cell Genetics, 61:10-16. 

. 19926. Meiotic studies of Robertsonian poly- 
morphisms in the South American marsh rat, Holi- 
chilus (sic) brasiliensis. Cytogenetics and Cell Ge- 
netics, 61:17—24. 

NACHMAN, M. W., AND P. Myers. 1989. Exceptional 
chromosomal mutations in a rodent population are 
not strongly underdominant. Proceedings of the Na- 
tional Academy of Sciences, USA, 86:6666-6668. 


NADEAU, J. H., AND B. A. TAYLor. 1984. Length of 
chromosomal segments conserved since divergence 
of man and mouse. Proceedings of the National 
Academy of Sciences, USA, 81:814-818. 

NADLER, C. F. 1966. Chromosomes and systematics 
of American ground squirrels of the subgenus Sper- 
mophilus. Journal of Mammalogy, 47:579-596. 

. 1971. Chromosomes of the Dall sheep, Ovis 
dalli dalli (Nelson). Journal of Mammalogy, 52:46 1- 
463. 

NADLER, C. F., AND C. E. HUGHEs. 1966. Chromo- 
somes and taxonomy of the ground squirrel subgenus 
Ictidomys. Journal of Mammalogy, 47:46-53. 

NADLER, C. F., R. S. HOFFMANN, AND J. J. P1zzZiIMENTI. 
1971. Chromosomes and serum proteins of prairie 
dogs and a model of Cynomys evolution. Journal of 
Mammalogy, 52:545-555. 

NELSoNn, K., R. J. BAKER, H. S. SHELLHAMMER, AND R. 
K. CHEssER. 1984. Test of alternative hypotheses 
concerning the origin of Reithrodontomys raviven- 
tris: genetic analysis. Journal of Mammalogy, 65: 
668-673. 

NELsSOn-REES, W. A., A. J. KNIAZEFF, R. J. BAKER, AND 
J. L. Patron. 1968. Intraspecific chromosome 
variation in the bat, Macrotus waterhousii Gray. 
Journal of Mammalogy, 49:706-7 12. 

O’Brien, S. J., H. N. SEUANEZ, AND J. E. WoMACK. 
1985. On the evolution of genome organization in 
mammals. Pp. 519-589, in Molecular evolution ge- 
netics (R. S. MacIntyre, ed.). Plenum Press, New 
York, 610 pp. 

Ouno, S. 1967. Sex chromosomes and sex-linked 
genes. Springer-Verlag, New York, 192 pp. 

PAINTER, T. S. 1923. Studies in mammalian sper- 
matogenesis. II. The spermatogenesis of man. Jour- 
nal of Experimental Zoology, 37:29 1-336. 

1925. A comparative study of chromosomes 
in mammals. The American Naturalist, 59:385—408. 

PARDUE, M.L., ANDJ.G. GALL. 1970. Chromosomal 
localization of mouse satellite DNA. Science, 168: 
1356-1358. 

PaTTon, J. L. 1967. Chromosome studies of certain 
pocket mice, genus Perognathus (Rodentia, Heter- 
omyidae). Journal of Mammalogy, 48:27-37. 

1973. An analysis of natural hybridization 
between the pocket gophers, Thomomys bottae and 
Thomomys umbrinus, in Arizona. Journal of Mam- 
malogy, 54:561-584. 

PaTTon, J. L., AND R. E. DINGMAN. 1968. Chro- 
mosome studies of pocket gophers, genus Thomo- 
mys. I. The specific status of Thomomys umbrinus 
(Richardson) in Arizona. Journal of Mammalogy, 
49:1-13. 

PaTTon, J. L., AND T. C. Hsu. 1967. Chromosomes 
of the golden mouse Peromyscus (Ochrotomys) nut- 
talli (Harlan). Journal of Mammalogy, 48:637-639. 

PaTTon, J. L., AND S. W. SHERWOOD. 1983. Chro- 
mosome evolution and speciation in rodents. An- 
nual Review of Ecology and Systematics, 14:139- 
158. 

PINKEL, D., T. STRAUME, AND J. W. GRAY. 1986. Cy- 
togenetic analysis using quantitative, high-sensitiv- 
ity, fluorescence hybridization. Proceedings of the 
National Academy of Sciences, USA, 83:2934-2938. 

Reip, T., A. BALDINI, T. C. RAND, AND D. C. WARD. 


S22 BAKER AND HAFNER 


1992. Simultaneous visualization of seven different 
DNA probes by in situ hybridization using combi- 
natorial fluorescence and digital imaging microsco- 
py. Proceedings of the National Academy of Sci- 
ences. USA, 89:1388-1392. 

SCHERTHAN, H., M. KOHLER. P. VoGt, K. VON MALSCH, 
AND D. ScHweizeR. 1992. Chromosomal in situ 
hybridization with double-labeled DNA: signal am- 
plification at the probe level. Cytogenetics and Cell 
Genetics, 60:4-7. 

SCHERZ, R. G. 1962. Blaze drying, by igniting the 
fixative, for improved spreads of chromosomes in 
leucocytes. Stain Technology, 37:386. 

SEABRIGHT, M. 1971. A rapid banding technique for 
human chromosomes. Lancet. 2:971-972. 

SELusA, G. A., M. V. Dt TOMAso, N. BRUM-ZORRILLA, 
AND H. Carposo. 1984. Low karyotypic variation 
in two didelphids (Marsupialia): karyogram and 
chromosome banding analysis. Journal of Mam- 
malogy. 65:702-707. 

SHELLHAMMER, H. S. 1967. Cytotaxonomic studies 
of the harvest mice of the San Francisco Bay region. 
Journal of Mammalogy, 48:549-556. 

StncuH, R. P., AND D. B. MCMILLAN. 1966. Karyo- 
types of three subspecies of Peromyscus. Journal of 
Mammalogy. 47:261-265. 

SuiFeR, E.H. 1934. Insect development: VI. The be- 
havior of grasshopper embryos in anisotonic, bal- 
anced salt solution. Journal of Experimental Zool- 
ogy, 67:137-157. 

STANGL, F. B., JR. 1986. Aspects of a contact zone 
between two chromosomal races of Peromyscus leu- 
copus (Rodentia: Cricetidae). Journal of Mammal- 
ogy. 67:465-473. 

STANGL, F. B., JR.. AND R. J. BAKER. 1984. Evolu- 
tionary relationships in Peromyscus: congruence in 
chromosomal, genic. and classical data sets. Journal 
of Mammalogy. 65:643-654. 

Stock, A. D. 1974. Chromosome evolution in the 
genus Dipodomys and its taxonomic and phyloge- 
netic implications. Journal of Mammalogy, 55:505- 
526. 

SuMNER, A. T. 1990. Chromosome banding. Unwin 
Hyman, London, 434 pp. 

SuMNER, A. T., E. H. EvANs, AND R. A. BUCKLAND. 
1971. New technique for distinguishing between hu- 
man chromosomes. Nature. New Biology, 232:31- 
32. 

Sutton, D. A., AND C. F. NADLER. 1969. Chromo- 
somes of the North American chipmunk genus FEu- 
tamias. Journal of Mammalogy, 50:524-535. 

Sutton, W.S. 1902. On the morphology of the chro- 
mosome group in Brachystola magna. Biological 
Bulletin, 4:24-39. 

. 1903. The chromosomes in heredity. Bio- 

logical Bulletin, 4:231-251. 


TERKELSEN, C., J. KocH, S. KOLVRAA, J. HINDKJAER, 
S. PEDERSEN, AND L. BoLUND. 1993. Repeated 
primed in situ labeling: formation and labeling of 
specific DNA sequences in chromosomes and nuclei. 
Cytogenetics and Cell Genetics, 63:235—237. 

Tno, J. H., AND A. LEVAN. 1956. The chromosome 
number of man. Hereditas, 42:1-6. 

Votp!, E. V..AND A. BALDINI. 1993. MULTIPRINS: 
a method for multicolor primed in situ labeling. 
Chromosome Research, 1:257-260. 

WALDEYER, W. 1888. Uber Karyokinese und ihre 
Beziehung zu den Befruchtungsvorgangen. Archiv 
fiir Mikroskopische Anatomie und Entwicklungs- 
mechanik, 32:1. 

Warp, D. C., P. LicHTER, A. BoyLe, A. BALDINI, J. 
MENNINGER, AND S.G. BALLARD. 1991. Gene map- 
ping by fluorescent in situ hybridization and digital 
imaging microscopy. Pp. 291-303 in Etiology of hu- 
man disease at the DNA level (J. Lindsten and U. 
Petersson, eds.). Raven Press, Limited, New York, 
316 pp. 

Waite, M. J.D. 1968. Models of speciation. Science. 
159:1065-1070. 

1973. Animal cytology and evolution. Wil- 

liam Clowes and Sons, London, 961 pp. 

. 1978a. Chain processes in chromosomal spe- 

ciation. Systematic Zoology, 27:285-298. 

1978b. Modes of speciation. Freeman, San 
Francisco. 455 pp. 

WICHMAN. H. A., R. A. VAN DEN BusscHE, M. J. HAM- 
ILTON, AND R. J. BAKER. 1992. Tranposable ele- 
ments and the evolution of genome organization in 
mammals. Genetica, 86:287-293. 

WICHMAN, H. A., C. T. PAYNE, O. A. RyDer, M. J. 
HAMILTON, M. MALTBIE, AND R. J. BAKER. 1991. 
Genomic distribution of heterochromatic sequences 
in equids: implications to rapid chromosomal evo- 
lution. Journal of Heredity, 82:369-377. 

Witson, A. C., V. M. SARICH, AND L. R. MAXSON. 
1974. The importance of gene rearrangement in 
evolution: evidence from studies on rates of chro- 
mosomal, protein, and anatomical evolution. Pro- 
ceedings of the National Academy of Sciences, USA, 
71:3028-3030. 

Witson. E. B. 1909. Recent researchers on the de- 
termination and heredity of sex. Science, 29:53. 
WuorstTer, D. H., AND K. BENIRSCHKE. 1970. Indian 
muntjac, .Wuntiacus muntijak: a deer with a low dip- 
loid chromosome number. Science, 168:1364-1366. 

WuourstTer-Hit, D. 1973. Chromosomes of eight spe- 
cies from five families of Carnivora. Journal of 
Mammalogy, 54:753-760. 

Yates, T. L.. AND D. J. ScHMIDLy. 1975. Karyotype 
of the eastern mole (Sca/opus aquaticus), with com- 
ments on the karyology of the family Talpidae. Jour- 
nal of Mammalogy, 56:902-905. 


POPULATION ECOLOGY 


WILLIAM Z. LIDICKER, JR. 


Introduction 


he term “‘population” traces its roots to 
“people” (Latin populus), which is a 
collection of human beings. Later it took on 
the meaning of collections of (usually sim- 
ilar) things. In biology it defines a group of 
individuals of the same species (kind). Of- 
ten such groups live in a prescribed place 
and can be distinguished operationally from 
other similar groups by partial or complete 
discontinuities 1n space or time or both. It 
is important, however, to recognize that such 
“natural” groupings are not essential to the 
“concept” of population; any arbitrarily 
designated group of individuals of the same 
species is sufficient. Once a population is 
designated, it is then possible to investigate 
whether it can also be defined by spatial or 
temporal discontinuities. Much confusion 
results from confounding these two objec- 
tives. 

Increasingly, biologists find it useful to 
view the living world (the biosphere) as be- 
ing organized on different levels of com- 
plexity that can be hierarchically arranged. 
Such a holistic perspective is by no means 
universally accepted as useful, and in fact 
this view has progressed rather slowly and 
fitfully over the past century. The history of 
population ecology as an intellectual disci- 


3235 


pline is inextricably connected to that de- 
velopment (Allen and Starr, 1982; MclIn- 
tosh, 1985; O’Neill et al., 1986). 

The concept population fits into the hi- 
erarchy of biotic complexity above the level 
of the individual organism and below that 
of the community. The concept community 
is biotically much more complex than pop- 
ulation because it concerns a universe (sys- 
tem) that includes more than one species 
(kind) of living organism. It is often difficult 
to distinguish studies at the population and 
community levels because populations al- 
most universally live with and interact with 
other kinds of living organisms. Neverthe- 
less, a distinction can generally be made on 
the basis of whether the study 1s focused on 
a single species or more than one. This is 
the same distinction made by the old terms 
“‘autecology”’ and “synecology.”’ Moreover, 
populations can be viewed conceptually in 
isolation even if this is rarely realistic, and 
one can certainly focus attention on one spe- 
cies at a time. A bacterial culture in a test 
tube is an example of the former and a study 
of the causes of mortality in a population 
of deer is an example of the latter. The con- 
cept population also fits into a hierarchy of 
evolutionary units (Brandon and Burian, 


324 LIDICKER 


1984; Eldridge, 1985; Lewontin, 1970; 
Salthe, 1985; Vrba and Eldridge, 1984). 

This chapter reviews how research on 
mammals over the last 75 years has influ- 
enced population ecology and considers how 
developments in ecology generally have im- 
pacted mammalogy. One of the central is- 
sues 1n population ecology is that of pop- 
ulation regulation, and this will therefore 
constitute a major thread through this chap- 
ter. A second theme will concern matura- 
tion of the concept “‘population”’ along with 
the recognition of population processes as 
being real biological phenomena above the 
level of the individual organism. I use the 
metaphor of a tree to organize this chapter. 
First I discuss the historical underpinnings 
(roots: pre-1930), followed by a review of 
the early research on population processes 
in mammals (trunk: 1930-1070). Next is an 
overview of modern foci in the field 
(branches: 1970 forward), and finally, I give 
brief comments on future perspectives 
(buds). Note that flowers and fruit are left 
for the future. In the context of this book, 
the emphasis has been on North American 
contributions, although I fully acknowledge 
the immense importance of others to this 
history. 


Roots: Initial Thoughts 


The question of what regulates the num- 
bers of organisms all began of course with 
a focus on a mammal, Homo sapiens. 
Thomas Malthus (1798) pointed out that 
populations have the capacity to increase 
exponentially but, except for brief episodes, 
do not do so. Therefore, negative forces 
(checks and balances) must operate on pop- 
ulations so as to counter the tendency to 
increase toward infinity. This insight was 
critical to the ontogeny of Charles Darwin’s 
thinking about evolution, and an essential 
ingredient in the development of our un- 
derstanding of evolution by natural selec- 
tion. In ecology, however, Malthus’ pio- 
neering contribution to the analysis of 
population processes languished until early 


in the 20th Century when ecology really 
started to blossom as a discipline (McIn- 
tosh, 1985). 

Like the roots of a majestic chestnut, the 
origins of mammalian population ecology 
are deep, intricate, numerous, and nourish- 
ing. Formal discussions of population birth, 
death, and growth rates were published in 
the first few years of this century (Lotka, 
1907, and references therein). One influ- 
ential paper that is often credited with the 
beginning of modern population theory (at 
least in North America) was published in 
1911 by two economic entomologists work- 
ing on gypsy moths (Howard and Fiske, 
1911). They clearly defined density equilib- 
rium and attributed its achievement to “‘fac- 
ultative agents” that increased proportion- 
ally in their suppressing effects as density 
increased. Thus, it was an interest in eco- 
nomically important insects and their con- 
trol that was the impetus for quantitative 
thinking about population growth. Ento- 
mologists were soon joined by mathemati- 
cal theorists in the development of quan- 
titative models for population processes 
(Lotka, 1925; Pearl, 1927; Volterra; 1926; 
1931), but these efforts were slow to influ- 
ence ecologists generally and mammalogists 
in particular. Early ecology texts hardly 
mentioned population regulation at all 
(Chapman, 1931; Shelford, 1913, 1929). 

In the early part of this century, mam- 
malogists were preoccupied with faunal sur- 
veys and documenting the occurrences and 


- distribution of species and subspecies of 


mammals (Hamilton, 1955; Miller, 1929). 
Population-level thinking was not much in 
evidence, and in fact most systematists har- 
bored a typological philosophy. A common 
view was that if a specimen were demon- 
strably different from “‘typical’’ individuals, 
it should be given a formal scientific name 
so that the fact of its uniqueness would not 
be lost to the scientific community. As in- 


_ formation accumulated on geographic vari- 


ation within species and within popula- 
tions, these views were gradually replaced 
by the realization that populations are not 
collections of identical individuals, and that 


POPULATIONS 325 


these assemblages of individuals also have 
features beyond those of the individuals that 
make them up. 

Mammalogists also gradually became 
more interested in ecological questions, es- 
pecially as information on life histories was 
acquired. In this they were encouraged by 
several leaders including Cabrera (1922), 
Seton (1929), Hamilton (1939), and Bour- 
liére (1951). Wildlife managers also played 
a critical role in this transition, because they 
were interested in questions of population 
regulation and control. Their approach, 
however, was normally to identify impor- 
tant mortality factors, and not to view pop- 
ulations in any quantitative way (Leopold, 
1933; Trippensee, 1948). They also popu- 
larized the notion of ‘‘optimal density,” not 
only as an ideal of management technology, 
but also as a natural state of some popula- 


tions (Bates, 1950; Dasmann, 1964; Elton, ~ 


1927; Howard, 1965; Leopold, 1933). The 
idea was that densities stabilized below a 
subsistence level so that body size, health, 
growth, and fecundity would be maximal. 
There was no recognition of the difficulties 
such idealism posed for natural selection at 
the individual level, although professional 
managers could strive for such a goal. 
Another root of critical importance to fu- 
ture population ecology was the gradual de- 
velopment of holistic philosophy. The name 
and formal description date from Smuts 
(1926), but the roots are deep and pervasive 
(Forbes, 1880; Semper, 1881), and include 
Forbes’ “‘microcosm” (1887) and the infa- 
mous “‘vitalism”’ of earlier times. Also ho- 
listic philosophy has been a dominant thread 
in many Eastern cultures for at least 2,500 
years (Barnett, 1982; Konishi and Ito, 1973; 
Lidicker, 1988). A few well-known early 
ecologists struggled with holistic notions 
(Clements and Shelford, 1939; Elton, 1930: 
30; Friederichs, 1927, 1930; Gause, 1934: 
2; Thienemann, 1939), but were largely un- 
successful because of a combination of the 
difficulty of the idea, lack of formal termi- 
nology for systems concepts, lack of a data 
base for population and community pro- 
cesses, and the spectacular successes of re- 


ductionist approaches to research (Lidicker, 
1978). One example will illustrate this sit- 
uation. When Clements (1905, 1916) and 
especially Clements and Shelford (1939) 
used the metaphor of “‘complex organism” 
to express the idea that communities rep- 
resented a higher order of biological orga- 
nization than that of individuals, the idea 
was received with hostility. Today we rec- 
ognize their supra-organism as an expedient 
metaphor for an idea almost all ecologists 
now accept, but only in the suitable format 
of modern jargon. E. P. Odum deserves con- 
siderable credit for encouraging holistic 
thinking, especially through his influential 
ecology texts beginning in 1953 (Odum, 
1953). 

The final major “root” to be mentioned 
is that of genetics and evolution. These two 
disciplines developed independently of 
ecology until recent decades. Of course, there 
were notable exceptions such as Charles El- 
ton, who was very much an evolutionary 
biologist as well as an ecologist (Crowcroft, 
1991: McIntosh, 1985). For the most part 
mammalogists thought about evolution in 
terms of phylogenies and adaptations, but 
not much about population-level processes. 
With the ““modern synthesis” in the 1940s, 
evolution and genetics (especially popula- 
tion genetics) were brought together and 
provided a more appropriate framework for 
synthesis with ecology (Brown and Wilson, 
1994). Still, the entrenched notion that eco- 
logical time frames are very much shorter 
than evolutionary time is still hampering us 
today. In 1969, I started to teach a lecture 
course in genetic ecology for graduate stu- 
dents, and remember well that for a number 
of years I spent the first lecture explaining 
and justifying such a radical interdisciplin- 
ary notion. 


The Trunk: Early Research on 
Population Processes 


Early research (1930s and 1940s) on 
mammalian population ecology emerged 


326 LIDICKER 


from research on life histories and on wild- 
life and forest management. Hamilton 
(1955), in his review of American mam- 
malogy, pointed out how important the in- 
vention and widespread use of the snap-trap 
was in encouraging life history studies and 
in making possible large collections of spec- 
imens. Still, populations were not viewed 
as entities with growth rates, birth rates, and 
the like. In Hamilton’s (1939) classic trea- 
tise on American mammals, only one brief 
chapter is devoted to populations. In this 
he debunked the “balance of nature’ as a 
fiction pointing to the ubiquitous variability 
in species numbers. Most of the chapter is 
devoted to cycles and mass outbreaks. 
Twelve years later, Gabrielson (1951) sim- 
ilarly allocated only one chapter to “pop- 
ulation controls” in his wildlife manage- 
ment text. He also attacked the balance of 
nature ideal, especially where human influ- 
ences are present, and briefly discussed in- 
terspecific competition, predation, damage 
to crops and habitat by wildlife, and the 
control of introduced plants. Trippensee’s 
text (1948) mainly discussed individual 
game species, followed by a section called 
‘‘Miscellaneous Wildlife Relationships,” 
with a chapter on “‘variations in numbers 
of wild animals” and one on “predator-prey 
relationships.” 

Toward the end of this period, main- 
stream ecologists at least were clear on the 
components of the population growth equa- 
tion (Allee et al., 1949; Cole, 1948; Park, 
1946). However, no coherent concept of 
populations being regulated by the quanti- 
tative interplay of births, deaths, and dis- 
persal rates was generally expressed. Trip- 
pensee (1948:386), for example, seems to 
have been unaware that an unrestrained 
positive biotic potential will produce ex- 
ponential growth toward infinity. Of course, 
any concept of community processes was 
even more vaguely perceived. While inter- 
specific competition, predation, and dis- 
eases were clearly thought important, no 
interacting network of interspecific inter- 
actions was envisioned. Trippensee (1948: 
398), nonetheless, did warn readers that 


“Predator relationships are complex and 
cannot be dealt with as simple phenomena,” 
and then illustrated the prevailing simpli- 
fied viewpoint with a table from Mendall 
(1944) classifying species of predators into 
four categories from ‘“‘distinctly beneficial” 
to “primarily detrimental.”’ 

It is interesting that “cycles” played such 
a prominent role in discussions of popula- 
tions even before Elton’s (1942) classic work 
on this subject. Hamilton’s (1939) analysis 
of multi-annual cycles is particularly thor- 
ough. He gives most space to sunspots as 
the causal agent, but in the end finds the 
evidence inadequate. Paraphrasing his views 
at that time, cyclic increases seemed to be 
the result of “abnormal” reproduction, and 
declines were caused by disease. In Trip- 
pensee’s (1948) chapter on variations in 
numbers, four out of 19 references cited have 
sunspots in the title, and he gives serious 
support to “cosmic factors’? as causative 
agents. Surprisingly, food was not consid- 
ered a critical factor at that time, except for 
lynx (Lynx canadensis) during crashes in 
snowshoe hares (Lepus americanus). Gen- 
erally the feeling was that population growth 
usually was checked far short of subsistence 
limitations (McAtee, 1936), a view that was 
consistent with the prevalent notion of ‘“‘op- 
timal densities.” Hamilton (1939:253) did, 
however, speculate that the “‘“abnormal” re- 
production that led to rodent outbreaks may 
have been abetted by a vitamin. 

The importance given to predation and 
disease as significant mortality agents went 
through an interesting transition at that time. 
Early wildlife biologists (e.g., Leopold, 1933) 
generally accepted predation and disease as 
major mortality agents. In this they were 
supported by the prevailing opinion among 
insect ecologists that parasites (including 
parasitoids) were the most important biotic 
mortality agents. A major shift in thinking 
can be attributed to the classical work of 
Errington (1946), whose primary research 
was on muskrats (Ondatra zibethicus). He 
professed that predators generally took only 
surplus prey, and therefore had no influence 
on density levels. This view of benign pre- 


POPULATIONS a2f 


dation gained rapid popularity, possibly fu- 
eled by a reaction to the vehement anti- 
predator stance of ranchers and government 
agencies. It reached an extreme form in the 
Cartwright Principle, which proclaimed that 
predators could save gallinaceous birds from 
extinction because when first nests were de- 
stroyed, birds re-nested at a more favorable 
time of the year and hence were more pro- 
ductive (Trippensee, 1948:392). This Er- 
ringtonian principle dominated thinking 
about predation among mammalian ecol- 
ogists almost to the present day, although, 
as I will point out, in recent decades im- 
portant modifications have been advanced. 

While mammalogical ecologists were thus 
occupied, insect ecologists were moving 
rapidly toward more rigorous and quanti- 
tative approaches to population regulation 
(Lidicker, 1978). Strongly influenced by the 
mathematical theorists active early in the 
century, they sought to fit environmental 
complexities into the relatively simple pop- 
ulation models that were being developed. 
They thus began to think clearly about how 
various factors can interact quantitatively 
to bring about changes in population num- 
bers. Some early and spectacular successes 
in biological control abetted this approach 
(Dunlap, 1981:31-35). The inherent risk in 
this path was that simple models led to sim- 
ple concepts of reality, and investigators 
were seduced into looking for single factor 
explanations of population changes. Tre- 
mendous advances in experimental biology 
made possible by reductionist approaches 
to research made the search for general and 
elegant explanations of biological phenom- 
ena especially tantalizing (Lidicker, 1988)). 
Mammalogists were, of course, not com- 
pletely isolated from this ferment. Hamil- 
ton (1939), for example, quotes the ento- 
mologist Uvarov (1931) at length regarding 
the balance of nature idea, and by the 1950s 
vertebrate ecologists generally had joined 
the fray. The Bureau of Population at Ox- 
ford under Charles Elton’s leadership was 
one of the centers of ferment and excitement 
that contributed to the developing synthesis 
(Crowcroft, 1991). 


As changes in numbers were seen increas- 
ingly clearly as the product of rate changes 
in the influences of various environmental 
“factors,” controversies quickly developed. 
It became widely appreciated in the 1930s 
that control of numbers required that neg- 
ative processes (environmental resistance) 
be positively related to population densi- 
ties. Some, however, were convinced that 
the relevant forces were abiotic factors and 
others were just as sure that they had to be 
biotic (Lidicker, 1978). On the one side were 
those most impressed with climate, weath- 
er, habitat, fire, and the like as determining 
numbers, with biotic factors being inciden- 
tal. Others were sure that biotic factors such 
as intra-specific competition, food, para- 
sites, and predators were all important, with 
the abiotic environment simply setting the 
stage for their actions. Advocates of the for- 
mer tended to view population densities as 
strongly variable, even stochastic, with local 
extinctions common. Champions of biotic 
control usually saw densities as carefully 
regulated about an equilibrium that, while 
not constant, was not random. 

Because of the association between abi- 
otic factors and failure to establish a fairly 
constant equilibrium density, and the cor- 
responding association between biotic fac- 
tors and density regulation, the term “‘den- 
sity independent factor’’ came to be applied 
to the abiotic and ‘“‘density dependent fac- 
tor’ to biotic influences. These terms were 
introduced by Smith (1935) and quickly be- 
came widely used. Unfortunately, they took 
on so many shades of meaning and innu- 
endo that semantic problems have plagued 
the subject ever since (Lidicker, 1978; Sol- 
omon, 1958). To summarize briefly, density 
dependence sometimes meant biotic fac- 
tors, sometimes density regulating, some- 
times simply that the factor’s effect changed 
with density, sometimes positively, some- 
times negatively, sometimes absolutely and 
sometimes proportionately, and sometimes 
it meant that the factor itself (not its effect) 
changed with density (responsiveness). 
Similarly, density independence meant 
whatever density dependence did not: abi- 


328 LIDICKER 


otic factors, non-regulating effects, effects 
that were unrelated to density, were con- 
stant numerically or proportionately, or were 
factors that were simply unresponsive 
themselves to density changes. Valiant ef- 
forts by leading ecologists failed to untangle 
this muddle (Schwertfeger, 1941; Solomon, 
1949; Thompson, 1939). 

Clarifying data were slow to accumulate. 
Because the questions were semantically 
mired, so were the answers. This was after 
all before the era of field experiments and 
hypothesis testing. Excellent laboratory 
studies were reported that clearly estab- 
lished that both biotic and abiotic factors 
could regulate numbers, but such infor- 
mation was easily dismissed by field ecol- 
ogists as irrelevant. Field researchers were 
generally searching for evidence to support 
their particular biases and almost always 
they succeeded. This situation led to a lot 
of argument and excitement, but little prog- 
ress toward clarifying the issues of the rel- 
ative importance of abiotic and biotic in- 
fluences, how they interacted, and how 
decimating effects changed quantitatively 
with density in field populations. 

A second circumstance that strongly in- 
fluenced the way that research on popula- 
tions was done in this era, and how ecolo- 
gists thought about the issues was the 
predominance of reductionist approaches. 
Not that very many ecologists thought ex- 
plicitly about what they were doing in these 
terms but, as already alluded to, holistic 
thinking was still embryonic and quite dif- 
ficult. Reductionism was achieving fantastic 
successes in cell and molecular biology, as 
well as physiology and medicine. All science 
students were taught that in good science 
one asks only “how” something works and 
not “why” it works the way it does. Natu- 
rally, ecologists wanted to be good scientists 
too. 

The emphasis on reductionism had sev- 
eral beneficial effects. It led to many good 
field and laboratory experiments and it en- 
couraged the practice of carefully studying 
the effects of various factors on a subject 


population one by one. This was, and re- 
mains, a powerful protocol. To suggest that 
it had its limitations remains controversial 
indeed (Gaines et al., 1991; Lidicker, 1991). 
In my view, however, the single-minded re- 
ductionist approach, without a complemen- 
tary systems (holistic) framework to guide 
it, ultimately limits understanding (Lidick- 
er, 1988a, 1988h; Macfadyen, 1975; Odum, 
1977). For the time and subject under dis- 
cussion, the important effect was to en- 
courage investigators to expect simple 
mechanisms for density regulation to be 
found. Not only were single key factors reg- 
ulating densities sought, but it was opti- 
mistically hoped that the answer once found 
could be extrapolated across time, across 
populations to the entire species, and then 
across species and even larger taxonomic 
groupings. After all, general properties of 
cells, biotic molecules, and genetic codes, 
were being reported regularly. In retrospect, 
we now know that this approach failed be- 
cause density regulation machinery turned 
out to be generally not simple, and single 
factor hypotheses are not amenable to this 
discovery (Hilborn and Stearns, 1982; Lid- 
icker, 1978:133; Smith, 1952). It is analo- 
gous to the futile search for the cause of 
cancer. 

With various investigators focusing on 
different aspects of density regulation, new 
controversies emerged. An important one 
that is only just now fading is whether ex- 
trinsic or intrinsic factors were most im- 
portant. That is, some argued that factors 
in the environment directly imposed regu- 
lation on the subject population, while oth- 
ers felt that changes in the organisms that 
constitute the population were the essential 
variables. It is surprising that ecologists 
could be so oblivious to the basic paradigm 
of their discipline, namely the organism- 
environment interaction system, and to the 
truism that both the properties of the or- 
ganisms and the environment change over 
space and time. Thus, while the intrinsic 
versus extrinsic argument was ultimately 
sterile, it did call attention to the impor- 


POPULATIONS 329 


tance of looking at the properties of both 
organism and environment in trying to un- 
derstand population processes (Lidicker, 
1978). 

Another development in the 1940s to 
which mammalogists made critical contri- 
butions was the acceptance of the life table 
concept in population ecology (Deevey, 
1947). It was, of course, introduced much 
earlier (Pearl, 1922), but failed to make much 
of an impact on vertebrate ecologists, prob- 
ably because the required data were too dif- 
ficult to acquire with existing technologies. 
Life tables served to focus attention on the 
attributes of various age and sex groups 
within populations, and eventually led to 
an appreciation for the age and sex structure 
of populations. The Leslie Matrix (Leslie, 
1945) for calculation of population growth 
is a familiar manifestation of this devel- 
opment. Thus intra-population demo- 
graphic variation was added to the increas- 
ing appreciation for genetic variation within 
populations to generate an increasingly re- 
alistic image of population phenomena. 
Modern population modelers continue to 
invoke structured populations in their mod- 
els (Boyce, 1977; Lomnicki, 1980; Schaffer, 
1974). One negative aspect of the enthusi- 
asm for life tables was the easy assumption 
that a particular life table characterized each 
species. In strict terms, a life table applies 
to a particular cohort of individuals born 


over a specified, and usually quite limited, . 


time and space. Confusion on this point 
continues. 

In the 1950s and 1960s, proponents of 
various classes of density-regulating factors 
tended to be viewed as “schools of thought.” 
The climatic school was not very popular 
among vertebrate ecologists (once sunspots 
were abandoned), but it was sometimes 
conceded that climatic factors could be crit- 
ical on the edges of species’ ranges. The 
availability of cover and nest sites were ad- 
mittedly part of what determined a species’ 
habitat, but were not often considered in 
determination of densities. Predation and 
parasitism had their champions, but mam- 


malian ecologists generally seemed to have 
lost interest in disease and the Erringtonian 
Principle diminished faith in the efficacy of 
predators (Errington, 1963; Howard, 1953). 

The extrinsic factor with the most wide- 
spread support was that of food. Lack (1954, 
1966) had eloquently argued for food lim- 
itation being the primary regulating factor. 
It was logical (all organisms required nutri- 
tion), and it fit into the emerging synthesis 
of evolutionary thinking in ecology (organ- 
isms should evolve so as to maximally use 
their food supplies). Detractors, however, 
pointed to contradictory evidence in spe- 
cific cases, to the potential (and frequently 
to evidence as well) for regulation by non- 
food factors, and to the necessity that con- 
sistent regulation by food requires optimal 
tracking by a population of its food re- 
sources. The food theory also became more 
sophisticated. While food quantity was 
stressed at first, nutrients later became rec- 
ognized as potentially limiting (Pitelka and 
Schultz, 1964). 

Other researchers turned their attention 
to intrinsic mechanisms. For some, self-reg- 
ulation made sense in that organisms would 
seem to be better off if they were not always 
at the point of exhausting their resources 
(e.g., Wynne-Edwards, 1962, 1965). Pru- 
dence demanded some measure of self con- 
trol. Others were disappointed that no ex- 
trinsic factor was found that fulfilled the 
hope of a general regulating factor. A tech- 
nique that became widely utilized at this 
time was to grow populations of small 
mammals in laboratory or outdoor enclo- 
sures. In this way a bridge between the lab- 
oratory and field was forged, and population 
processes could be studied in a circum- 
stance such that either intrinsic or extrinsic 
factors could be manipulated individually. 

One class of intrinsic factors that was 
studied extensively was that of physiologi- 
cal change associated with varying densi- 
ties. An early hypothesis of Chitty (1952, 
1955, 1958) that high densities led to phys- 
iological damage that increased mortality 
rates and moreover could be passed on to 


590 LIDICKER 


offspring during gestation or lactation was 
later abandoned by him (Chitty, 1960, 
1967). Christian (1950) introduced the 
intriguing idea that exhaustion of the ad- 
reno-pituitary system may be involved in 
population declines. High densities would 
feature a variety of stressors, he suggested, 
and hence the proximate causes of mortality 
would be non-specific. Later (Christian, 
1955a, 1955b, 1959, 1961; Christian and 
Davis, 1955) he expanded the model to sug- 
gest that high densities activated the stress 
resistance mechanisms of the body, even- 
tually resulting in their exhaustion. Re- 
duced reproductive competence and death 
soon followed. A related phenomenon was 
the “‘shock disease”’ widely associated with 
population crashes in snowshoe hares. As 
this was known to involve hypoglycemia 
and non-specific mortality agents, it could 
easily be fitted into the stress hypothesis. 
Trippensee (1948:392), however, thought 
shock disease was caused by a lack of min- 
erals in the diet. Many researchers pursued 
these ideas, and by the end of the 1960s the 
situation could be summarized as follows 
(Lidicker, 1978): the stress syndrome was 
real in laboratory situations, but was not 
found to be generally applicable to field pop- 
ulations. 

A second class of intrinsic factors to be 
proposed was that of behavioral changes 
with density. Territoriality, fighting, dis- 
persal, and cannibalism all could change 
with density and may be expected to have 
demographic consequences. Wynne-Ed- 
wards (1962, 1965, 1986) proposed that 
“‘epideictic displays’’ were a mechanism by 
which individuals communicated their den- 
sity circumstances to each other. As such, 
this notion was criticized for not making 
sense in the context of individual selection, 
but could be defended by involving group 
selection mechanisms (Wynne-Edwards, 
1986). The use of enclosed populations led 
to the discovery of behaviorally-mediated 
reproductive inhibition (Calhoun, 1949, 
1962: Crowcroft and Rowe, 1957; Davis, 


1949: Lidicker, 1965; Petrusewicz, 1957; 
Southwick, 1955). In fact, Petrusewicz 
(1957) startled ecologists with his evidence 
that in laboratory colonies of house mice 
(Mus musculus), a socially-inhibited group 
can be induced to resume reproduction sim- 
ply by moving it to a new cage, even a small- 
er one. Otherwise, phenotypic behavioral 
changes with density were mainly studied 
in more recent decades. 

Genotypic shifts in populations with den- 
sity changes were the third class of intrinsic 
factors contemplated seriously as regulating 
mechanisms. Led by Chitty (1960, 1967) 
and Krebs (1964, 1971), the stimulating idea 
was proposed that selective pressures vary- 
ing with density favored different genotypes 
at high versus low densities, and the cor- 
responding shifts in gene frequencies led to 
predictable demographic consequences. 
Such ideas had been suggested earlier for 
insect populations (Turner, 1960; Welling- 
ton, 1960; Wilbert, 1963), but Chitty and 
Krebs applied them specifically to density 
cycles of microtine rodents and suggested 
that aggressive versus docile behavior was 
the relevant behavior being selected. Later 
they hypothesized that, instead of aggres- 
sion, the behavior being selected was spac- 
ing behavior including dispersal (Krebs, 
1979a; Krebs et al., 1973). These ideas were 
so important that they strongly influenced 
the character and direction of research on 
small-mammal populations in subsequent 
decades. 

Over the roughly four decades covered in 
this section (1930s through 1960s), some 
general trends in the relative importance of 
mortality, natality, and movements in and 
out of populations (immigration and emi- 
gration, respectively; Lidicker, 1975) can be 
discerned. Of course, early in this period, 
mammalian researchers did not usually 
think of these processes as interacting vari- 
ables in a growth equation. Early emphasis 
was on mortality; reproduction was thought 
to be almost always “normal,” i.e., non- 
varying. In fact, Smith (1935), in his sem- 


POPULATIONS pol 


inal paper defining density dependence and 
independence, referred to density depen- 
dent factors as mortality agents only. Even 
Dasmann (1964) discussed density depen- 
dence only in terms of mortality. Gradually, 
the importance of reproduction gained ap- 
preciation, especially as data accumulated 
showing that it too could vary with density. 
At first, “abnormally” good reproduction 
was thought to produce population out- 
breaks (Hamilton, 1939:274), but then it 
became apparent that reproduction often 
declines with increasing density as well (see 
Howell, 1923, for a pioneering example). 
This new focus on reproduction reaches an 
extreme with demographers who tend to 
view human population growth rates as 
mainly influenced by birth rates and hardly 
at all by mortality, a tradition going back at 
least to Pearl (1925). 

Movements in and out of populations 
were not given much attention (but see 
Hamilton, 1953). Early on, dispersal was 
viewed as destabilizing because individuals 
were visualized as moving about in search 
of favorable circumstances, thus increasing 
the variability of local densities. Then, as 
growth equations entered the arena, growth 
rates were defined as birth rates minus death 
rates (r). This dogma swept through the text 
books and assured that immigration and 
emigration would not be considered seri- 
ously. When they were mentioned at all, 
they were dismissed as trivial or balanced 
between imports and exports and therefore 
ignorable. If significant emigration was ac- 
knowledged, it was lumped with mortality 
under the rubric “‘gross mortality.” Except 
for the paper by Howard (1960) postulating 
that both “genetic” and “environmental” 
dispersal may occur, and my own paper 
(Lidicker, 1962) suggesting that emigration 
should be examined for its possible effects 
in density regulation, the fervor of interest 
in dispersal came in later decades. 

I end this section with a caveat and men- 
tion of two exceptional individuals. For the 
four decades covered here, I have tried to 


portray major themes of intellectual devel- 
opment. As time progressed through the pe- 
riod, it becomes increasingly difficult to fol- 
low one thread. Our disciplinary “trunk” 
forms major branches and many more re- 
searchers are involved. Moreover, the av- 
erage intellect that one tries to describe is a 
statistical artifact drawn from a fairly small 
sample size. Each individual investigator is 
of course exceptional in at least some re- 
spects. An important exception to this av- 
erage intellect was Charles Elton, who some 
consider the father of mammalian popula- 
tion ecology (Berry, 1987). Not only was he 
an early architect of community concepts 
(e.g., Eltonian pyramids), but he was an ad- 
vocate of incorporating evolutionary think- 
ing in ecology long before this was routine. 
As early as 1930, he expressed the holistic 
view that a whole biological community 
could act as a unit of selection (Elton, 1930: 
30), and warned that “*. . . the modern ecol- 
ogist runs a risk of ... falling back upon a 
severely mechanistic view... based on the 
laws of physics and chemistry, solid in 
themselves, but unsatisfactory as a com- 
plete explanation of the life and mind of 
animals” (1930:9). Secondly, for the mam- 
malian ecologist, Elton’s treatise on voles, 
mice, and lemmings (1942) was where it all 
began. His Bureau of Population at Oxford 
was, moreover, the gestation site for nota- 
bles such as Dennis Chitty, Peter Crowcroft, 
Richard Miller, and Mick (H. M.) Southern, 
and also strongly influenced long-term vis- 
itors like Frank Pitelka (see also Crowcroft, 
1991). 

A second exceptional individual in this 
formative era was Kazimierz Petrusewicz 
(Lidicker, 1984). He established in 1952, in 
the rubble of World War II, a Department 
of Ecology within the Polish Academy of 
Sciences, which was elevated in 1971 to the 
status of an Institute. Petrusewicz was di- 
rector from 1956 to 1973, during which time 
Polish ecology became an internationally 
recognized center of excellence, with im- 
portant work on mammals. Mammalian re- 


932 LIDICKER 


search extended from the analysis of pop- 
ulation processes in laboratory settings to 
energetics, production, population regula- 
tion, dispersal, social behavior, and wildlife 
management. Petrusewicz himself was in- 
tensely interested in relating evolution to 
ecological processes, had a sophisticated ho- 
listic philosophy, and contemporaneously 
with Elton was writing papers on concepts 
of community structure. His influence on 
population ecology in Poland, eastern Eu- 
rope, and the world community was pro- 
found and long lasting (Lidicker, 1984). He 
was elected an Honorary Member of the 
American Society of Mammalogists in 1975 
(Taylor and Schlitter, 1994). 


The Modern Era: 
The Last Two Dozen Years 


Alluding to our botanical metaphor, we 
have now reached the stage in the devel- 
opment of our subject where we have 
branches, lots of branches, both major sup- 
ports, and idiosyncratic twigs. No longer can 
we imagine that there is but a single path 
or even a few major paths of intellectual 
ontogeny, and it becomes increasingly dif- 
ficult to review intellectual history by trac- 
ing the origin and transmission of key ideas 
and the influence of especially significant 
leaders in the process. Of course, there were 
these, but the abbreviated hindsight of his- 
tory and the huge dimensions and the es- 
tablishment make these leaders seem for 
now more like extenders of intellectual 
pseudopodia than creaters of new para- 
digms. 

Mammalian population ecology had in 
this period not only joined the mainstream 
(I should say maelstrom) of population ecol- 
ogy, but was providing a leading voice. It 
was and is vigorous, diverse, incredibly in- 
terdisciplinary, and has nurtured the ger- 
mination of new subdisciplines such as evo- 
lutionary ecology, behavioral ecology, 
community ecology, landscape ecology, and 
conservation biology. Still our enthusiasm 
cannot quite match that of R. J. Berry who 


wrote (1987:1) that “... the proper study 
of biology inevitably involves an investi- 
gation of the processes which affect popu- 
lations.” 

ASM programs.—The increasing atten- 
tion given to populations and community 
level phenomena, as well as the expanding 
diversity of subdisciplines in this field, are 
reflected in the programs of the annual 
meetings of the ASM. These programs allow 
us to monitor and assess the prevailing par- 
adigms over time among working mam- 
malogists, and thus to measure the net pro- 
gressions of the field (Also see Gill and 
Wozencraft, 1994). 

For this purpose, I classified all the papers 
in 16 programs covering 1926 to 1991. The 
classification was subjective and used 10 
major categories plus a number of subcat- 
egories. There were, or course, a few am- 
biguous or cryptic titles, and some papers 
could be placed into more than one cate- 
gory. Because of the scope of this chapter, 
I focused particularly on papers that seemed 
to reflect a population or community con- 
cept. Ecological papers judged to be at the 
organismal level were assigned to a “general 
life history” or “physiology and morphol- 
ogy’ category. The few titles with a land- 
scape perspective were lumped with com- 
munity ecology. A category of “behavioral 
ecology”’ was also recognized to include pa- 
pers that related behavior to ecological pro- 
cesses and that included group behavior such 
as mating systems or other social behavior. 
This scheme of categorization allows for the 
monitoring of research activities at the pop- 
ulation or higher levels, which is the subject 
of this chapter. Otherwise, the plethora of 
papers in general life history phenomena 
would obscure these patterns. 

The percentage of papers in ecology at the 
population or higher level is plotted over a 
66-year period (Fig. 1). There were no pa- 
pers in this category in 1926 and only two 
in 1938. These first in our sample were an 
address by Joseph Grinnell on “Effects of a 
wet year On mammalian populations,” and 
one by W. P. Taylor on “Significance of 
numbers in mammalian ecology.” There was 


POPULATIONS b Jee 


No. of papers 47 62 276 
44 39 33 38 39 58 76 104 112 203 227 247 270 


Percent 


Population 


Community 


Behavioral 


1920 1930 1940 1950 1960 1970 1980 1990, 


Year 


Fic. 1. Percentages of papers on ecological 
subjects presented at annual meetings of the ASM, 
based on 16 programs from 1926 to 1991. Eco- 
logical papers are allocated to behavioral, pop- 
ulation, and community categories based on their 
primary conceptual level. The dashed line for 
1947 indicates the percent of papers in popula- 
tion ecology when six papers in a symposium on 
populations are omitted. 


an increase to nine papers in 1947, but this 
was almost entirely the result of a sympo- 
sium on “Population, home range, and ter- 
ritories in mammals.” Interestingly, five out 
of the nine papers were on techniques and 
another (by Durward L. Allen) was titled 
“Purposes of population studies.” If these 
six are subtracted, the percentage of ecology 
papers drops from 30.3 to 12.1% (Fig. 1). 
This symposium and one at the society’s 
1950 meeting on the dynamics of mam- 
malian populations mark the beginning of 
a steady increase in the proportion of papers 
given on these topics, which reached a peak 
of 31.3% in 1981 and declined moderately 
after that. 

Papers recognizable as community-level 
started in the 1954 program and increased 
rapidly after 1969. One paper was assigned 
to behavioral ecology in 1947, but the next 
one was not until 1961, and the third was 
in 1974. After 1947, the proportion of pop- 
ulation-level papers varied hardly at all (6.5- 
18.3%), with changes in the ecological of- 
ferings being due to the addition of com- 
munity and behavioral ecology contribu- 
tions (Fig. 1). 


Importance of new techniques.—An im- 
portant contributor to the success of pop- 
ulation research in this period was the ar- 
rival of new and powerful techniques. 
Whereas the snaptrap and livetrap were the 
technical “work horses” of the previous era, 
they were soon supplemented by an im- 
pressive list of innovations. Following 
World War II, radioactive isotopes became 
readily available and were used to follow 
individuals, determine pedigrees, reveal 
movements, and measure various demo- 
graphic parameters (Stenseth and Lidicker, 
1992a). Because of health hazards to the 
investigators as well as to the research sub- 
jects and their environments, however, such 
isotopes are less commonly used now. 

A second technique was that of radio- 
tracking (Amlaner and MacDonald, 1980; 
McShea and Madison, 1992). At first this 
approach was restricted to large mammals, 
but with the increasing miniaturization of 
transmitters, radios with batteries have 
shrunk to where even mice can carry them 
successfully. Telemetry has provided a 
wonderful opportunity to follow the move- 
ments and activities of individual mam- 
mals, even through the guts of predators. 
When numerous individuals in the same 
population are being followed simulta- 
neously, it is also possible to reveal social 
interactions, and thereby to understand why 
certain movements are occurring in addi- 
tion to describing them. 

A more recent development is the use of 
fluorescent powders to track movements of 
nocturnal species (Kaufman, 1989). Under 
favorable circumstances these powders can 
reveal paths of movement by reflection of 
ultra-violet light. They have also been used 
to determine social bonds such as mother- 
juvenile and adult male-female relation- 
ships by detection of the transfer of small 
amounts of the powder between individuals 
(Ribble and Salvioni, 1990). 

Critically important has been the devel- 
opment of various biochemical techniques. 
Electrophoresis of blood and tissue proteins 
and enzymes has been used widely since the 
late 1960s, and has been an effective tool in 


334 LIDICKER 


assessing the genetic architecture of popu- 
lations and in measuring relatedness among 
groups. The analysis of mitochondrial DNA 
restriction enzyme fragments has also been 
useful for measuring relationships over a 
shorter time span than is usually possible 
with the allozymic variants coded by nu- 
clear DNA. This is because the mutation 
rate, and hence biochemical drift, is faster 
with certain sections of mitochondrial DNA 
than with nuclear. As of this writing, the 
most exciting new development is that of 
DNA-fingerprinting. Although a more dif- 
ficult and laborious technique, it has the 
potential for unequivocal individual iden- 
tification as well as for parental exclusion 
analysis. Thus it has tremendous promise 
in investigations requiring individual rec- 
ognition and knowledge of pedigrees. An- 
other new development with great promise 
is the polymerization chain reaction (PCR), 
which allows for amplification (multipli- 
cation) of small sections of DNA so that 
such fragments can be sequenced, com- 
pared, and relatedness judged. It has also 
opened up the possibility of using small 
amounts of DNA surviving in museum 
specimens and near-fossils to assess rela- 
tionships among taxa, and perhaps more 
relevant to the ecologist, is the possibility 
of charting genetic change in populations 
over relatively short periods of time. PCR 
techniques utilizing dinucleotide repeats 
called ‘“‘microsatellites” that are widely dis- 
tributed throughout the mammalian ge- 
nome may be rich sources of polymor- 
phisms and hence information on re- 
latedness among individuals because of their 
extensive and presumably neutral variabil- 
ity. New developments useful to the pop- 
ulation biologist can be predicted confi- 
dently. 

Finally, it is appropriate to call attention 
to the vast improvement in quantitative 
techniques available to the population bi- 
ologists. These include powerful computer 
software packages for organizing and ana- 
lyzing data, using multivariate statistics, 
clustering techniques, and the like. Even field 


methodologies for gathering demographic 
and other data are greatly improved (Ham- 
mond, 1987; Hiby and Jeffery, 1987; Mont- 
gomery, 1987; Smith et al., 1975; Ward et 
al., 1987). Mathematical modeling, both 
analytical and computer simulation, has 
benefited our understanding of population 
processes (Conley and Nichols, 1978; Dek- 
ker, 1975; Hestbeck, 1988; Stenseth, 1981, 
1983, 1986; Stenseth and Lidicker, 19925), 
and undoubtedly will play a large role in the 
future. It helps us to think clearly, to test 
the quantitative consequences of our ideas, 
and allows us to synthesize quantities of facts 
and relationships that would otherwise push 
beyond the limits of our mental capacities. 
Modeling only threatens progress when we 
view mathematical expressions as tem- 
plates of reality, or as substitutes for data, 
or confuse mathematical proof with careful 
testing of hypotheses. 

Intellectual foci.—1 have divided the in- 
tellectual history of our subject in the mod- 
ern era into six interconnected and over- 
lapping foci or themes. A single branch of 
inquiry is no longer realistic, and, more- 
over, the order in which I discuss them is 
completely arbitrary. These vignettes are in 
no way attempts to review these topics, each 
of which is a vast subject in itself. The most 
I can do here is attempt to connect each 
theme with the previous historical period, 
and to suggest major intellectual trends. As 
I have been a participant in this process, the 
risks of personal biases creeping into the 
analysis are greater than for the earlier pe- 
riods. My intention, nevertheless, is to be 
as objective as possible. One major area 
omitted here is that of life history evolution 
(Boyce, 1988). This is because I think of 
this field as more at the organismal than 
population level of analysis. Clearly, how- 
ever, the study of life history extends into 
the population level especially where gender 
differences in life history strategy or other 
polymorphisms occur. 

1) Spatial structuring of populations. As 
mentioned, population ecologists were gen- 
erally aware of the importance of age and 
sex structure within populations, continued 


POPULATIONS 6 o)5) 


to gather data on this, constructed life ta- 
bles, and increasingly emphasized cohort 
analysis rather than extrapolation over time 
or to species as a whole. Appreciation of 
spatial structure, however, was slower in 
coming. 

Contrary to common sense, populations 
of organisms were, at the beginning of this 
modern period, conceptualized as infinite 
in size and generally panmictic. Such ap- 
proximations were consistent with the the- 
ory of population genetics and evolution 
then prevailing and with the ubiquitous 
maps of species’ ranges. Although mam- 
malian ecologists generally realized that 
these simplifications were unrealistic, they 
did not, I think, appreciate that it mattered 
very much. In the summer of 1967, P. K. 
Anderson traveled extensively in the Soviet 
Union, and learned first hand about the 
views of several leading Soviet ecologists 
(particularly B. K. Fenyuk, T. V. Koshkina, 
N. P. Naumov, P. A. Panteleyev, I. Ya Pol- 
yakov, and S. S. Shvarts) concerning the 
spatial structuring of mammalian popula- 
tions and the ecological and genetic impor- 
tance attributed to this structuring. Inspired 
by these insights (as well as recent research 
on Mus musculus), Anderson (1970) wrote 
an important review on ecological structure 
and gene flow in small mammals in which 
he proposed that genetic and social frag- 
mentation was indeed the rule for species 
of small mammals and that this implied a 
dramatic change in the way we should view 
population biology. Shortly thereafter, 
Shvarts’ book (1969) on the evolutionary 
ecology of animals was translated into En- 
glish by A. E. Gill (Shvarts, 1977), and 
Hansson (1977) wrote his influential paper 
on the importance of heterogeneous land- 
scapes in the ecology of small mammals. It 
is important that these contributions ap- 
peared in an intellectual environment in 
which notions of environmental grain (Lev- 
ins, 1968) were being widely discussed, at 
least by evolutionary theorists. 

In 1978, a symposium on mammalian 
population genetics was held in conjunction 
with the annual meetings of the ASM. In 


reviewing the published volume from this 
symposium (Smith and Joule, 1981), it is 
apparent that even at this time, most atten- 
tion was given to temporal variation in ge- 
netic constitution of populations (e.g., 
Gaines, 1981) and the causes and signifi- 
cance of genetic variation within popula- 
tions (e.g., Schnell and Selander, 1981). Only 
one paper gave significant attention to 
spatial variation on the scale of habitat 
patches (Massey and Joule, 1981). 

Subsequently, the importance of spatial 
structuring became increasingly recognized 
as a critical demographic and genetic influ- 
ence. Currently, it is an extremely fashion- 
able topic of investigation (Hansson and 
Stenseth, 1988). Even models of density cy- 
cles of microtines now are incorporating 
habitat heterogeneity as a relevant variable 
(Bondrup-Nielsen and Ims, 1988; Gaines et 
al., 1991; Lidicker, 1985a, 1988a, 1991; 
Ostfeld et al., 1985). 

The culmination of this trend is the emer- 
gence of the subdiscipline of landscape ecol- 
ogy (Forman and Godron, 1986; Lidicker, 
19885), and its application to mammalian 
ecology (Bauchau and LeBoulengé, 1991; 
Lidicker et al., 1991; Merriam, 1990, 1991: 
Szacki and Liro, 1991; Wegner and Merri- 
am, 1990; Wolff, 1980). At this level of bi- 
ological complexity, systems composed of 
two or more habitat patches (community- 
types) are the subject of inquiry. Thus, the 
role of patch size, edge-to-area ratios, con- 
nectedness, and inter-patch fluxes are ex- 
plicitly investigated. Many new demo- 
graphic and evolutionary insights can be 
anticipated as a result of this advance. 

2) Dispersal. As pointed out, interest in 
dispersal was almost non-existent at the 
beginning of this modern era. Currently, it 
is one of the most vigorous areas of inquiry 
in mammalian ecology, marking a devel- 
opment that is clearly one of the most dra- 
matic of this period. Apart from some early 
signals (Andrzejewski et al., 1963; Howard, 
1960; Kalela, 1961; Lidicker, 1962), a bur- 
geoning interest developed in the late 1960s 
and 1970s (see Fenton and Thomas, 1985; 
Lidicker, 1975, 1985b; McCullough, 1985 


336 LIDICKER 


for early reviews). In January 1992, the BIo- 
sis electronic data base listed 7,240 refer- 
ences (Zoological Record, Online, 1978 to 
1991) indexed by the descriptor “‘dispers- 
ale 

Basically, what happened were two crit- 
ical intellectual breakthroughs: 1) the real- 
ization that movements into and out of pop- 
ulations (immigration and emigration, 
respectively) are critical components, along 
with births and deaths, of population dy- 
namics; and 2) the realization that if pop- 
ulations were not always panmictic and in- 
finite (see previous section), subpopulations 
must be connected genetically, demograph- 
ically, and behaviorally by dispersal. Thus, 
the study of dispersal became a critical in- 
gredient in questions ranging over physi- 
ology, behavior, evolution, epidemiology, 
and conservation biology as well as all levels 
of complexity in ecology (Stenseth and Lid- 
icker, 1992c). 

One important factor that helped start this 
avalanche of research on dispersal was the 
extensive use of confined populations (en- 
closures, islands) giving meaning to the 
fence-effect concept. Thus it was that the 
study of populations in which dispersal was 
absent helped us realize how important it 
was in unconfined situations (Lidicker, 
1979a). These studies, as well as a growing 
number on unenclosed populations, led to 
the explicit recognition that dispersal often 
occurred before conditions in the home 
habitat became economically desperate 
(“pre-saturation dispersal,’ Lidicker, 1975) 
and hence at least some dispersal was fa- 
vored by natural selection (“‘adaptive,” 
Stenseth, 1983); see Lidicker and Stenseth 
(1992) for summary of the factors motivat- 
ing dispersal. 

A second important element was the in- 
corporation of dispersal in models of mi- 
crotine rodent multi-annual cycles. Early 
papers (Krebs et al., 1973; Lidicker, 1973; 
Stenseth, 1978; Tamarin, 1978) led to 
widespread attention to dispersal by micro- 
tine ecologists and inspired numerous in- 


vestigations, empirical and theoretical, as to 
the role of dispersal in these cycles. 

3) Coactions. I use the term “‘coaction”’ 
as a brief equivalent to “interspecific inter- 
action” (Clements, 1916; Clements and 
Shelford, 1939; Haskell, 1949; Leary, 1985; 
Lidicker, 19796). Such community-level 
processes are appropriately reviewed in an- 
other chapter (Mares and Cameron, 1994), 
but it is important to comment here, albeit 
briefly, on several paradigm shifts occurring 
in recent decades. 

In the last section, I pointed out how the 
Erringtonian or benign predation view had 
become the prevailing one. This trend 
reached an extreme form in Howard’s (1965) 
extension of the Cartright Principle to 
mammalian predators. He advocated the 
view that in management of rodent pests, 
predators were a hindrance rather than a 
help because they stimulated rodent popu- 
lations to increase reproductive effort. 

Two other shifts in the way predation was 
viewed were more generally accepted. The 
first was that in spite of usually lower re- 
productive rates (than their prey), predators 
could reduce prey densities through func- 
tional rather than numerical responses to 
prey numbers (Keith and Windberg, 1978; 
Weaver, 1979). The second change was the 
realization that predators sometimes made 
their greatest impact, not on increasing prey 
populations, but on declining ones. Thus, 
they have an increasing effect as density falls 
(anti-regulating, de-stabilizing) and can drive 
prey densities to extremely low levels (Lid- 
icker, 1975, 1988a; MacLean et al., 1974; 
Maher, 1967: Newsome and Corbett, 1975; 
Pearson, 1966, 1971, 1985; Wagner and 
Stoddart, 1972). In the case of ungulates, 
well-documented examples of predator reg- 
ulation became available (Caughley, 1970; 
McCullough, 1979; Peterson and Page, 
1983). All of these developments reestab- 
lished predation as a potentially important 
influence in population regulation. 

The importance of parasitism in the pop- 
ulation biology of mammals went, as ex- 
plained, from the early assumption that it 


POPULATIONS 337 


was important to almost complete neglect. 
In recent decades a renewed interest is 
emerging. Partly this was fueled by theo- 
reticians (Anderson and May, 1979; Dietz 
and Schenzle, 1985; May and Anderson, 
1979; Mollison, 1977, 1987), who drew at- 
tention to the potential for demographic im- 
pact that parasites and disease can have. A 
second factor was the slowly increasing em- 
pirical evidence that parasites can regulate 
mammalian populations (Anderson, 1982; 
Anderson et al., 1981; Fenner, 1976; Greg- 
ory, 1991; Plowright, 1982; Ross, 1982; 
Scott, 1988). In my view, this is one area 
ripe for exploitation by interdisciplinary 
teams of investigators. 

The extent to which species of mammals 
enter into competitive coactions with each 
other and with non-mammals began to be 
explored vigorously by the beginning of this 
modern period. Early leaders were Rosen- 
zweig (Rosenzweig, 1966, 1973; Schroder 
and Rosenzweig, 1975), Grant (Grant, 1969, 
1972, 1978: Morris and Grant, 1972) and 
Brown (Brown, 1971; Brown and Davidson, 
1977; Brown et al., 1979; Davidson and 
Brown, 1980; Munger and Brown, 1981). 
The potentially exciting arena of coopera- 
tive coactions (mutualisms) remains to be 
explored in the future. 

4) Social behavior. Although a topic that 
is discussed more fully in another chapter 
(Eisenberg and Wolff, 1994), it is important 
to mention here that studies of social be- 
havior are an increasingly important part of 
mammalian population biology. Behavior 
has always been of interest to mammalo- 
gists, but until recently it was viewed simply 
as one element in the description of a spe- 
cies’ life history. In recent years social be- 
havior has been studied as a group process 
impacting in important ways and in turn 
being influenced by various aspects of evo- 
lutionary and ecological dynamics (Armi- 
tage, 1988; Berger, 1986, 1988; Cockburn, 
1988; Krebs and Davies, 1984; Mech, 1987: 
Sherman et al., 1991; Slobodchikoff, 1988; 
Smith and Ivens, 1984; Tamarin et al., 
1990). It is this view of behavior that I have 


included in “behavioral ecology.” It began 
as a serious trend in mammalian ecology 
about 1970 (Fig. 1). Examples of a few early 
contributors include King (1955), Eisenberg 
(1967), Hamilton (1971), Trivers (1971), 
Kleiman and Eisenberg (1973), Alexander 
(1974), and Barash (1974). Wilson’s (1975) 
influential opus on sociobiology stands as a 
monument to this critically important de- 
velopment. 

Important current themes in behavioral 
ecology include: 1) social signaling with spe- 
cial emphasis on the olfactory mode; 2) 
mating systems; 3) kin recognition and as- 
sociated cooperative behaviors; 4) plasticity 
versus tight genetic control of social behav- 
ior; 5) effects on demography (e.g., spacing 
behavior, dispersal, density dependent ag- 
gression); and 6) relationships between so- 
cial structure and genetic structure of pop- 
ulations. Based on a 1980 conference, the 
ASM published an influential review of 
mammalian behavioral research (Eisenberg 
and Kleiman, 1983). 

5) Density regulation. The subject of how 
population densities are regulated continues 
to be an important, exciting, and contro- 
versial area up to the present time. Past de- 
bates about “density dependent” versus 
“density independent” factors and intrinsic 
versus extrinsic regulation have abated. It 
is now widely appreciated that densities are 
influenced by a variety of factors operating 
in a variety of ways, but that eventually 
there must be a net increase in the rate at 
which negative forces act as density increas- 
es (regulation) or the Earth would be filled 
with infinite populations. Such negative 
forces impose either an upper limit for den- 
sity or result in an equilibrium level (K) 
toward which densities tend. Similarly, the 
intrinsic-extrinsic dichotomy is now gen- 
erally accepted as a non-issue. The density 
regulating machinery consists of the organ- 
ism-environment axis, and not with either 
component alone (Lidicker, 1978). Prop- 
erties of the organism and properties of its 
environment interact to result in a given 
density with the relative contribution of each 


338 LIDICKER 


varying, but with both being always in- 
volved. 

With these contentious issues behind us, 
much of importance remains. What is the 
actual regulating mechanism for a given 
population? How much does this vary spa- 
tially and temporally? Are there general pat- 
terns for certain taxonomic groups, habitat 
assemblages, trophic levels, and life styles? 
Moreover, we need to discover if one or a 
few factors are consistently of overriding 
importance for specific populations, with 
other forces being clearly secondary or con- 
tributing only to the variance of densities. 
How important are time lags and age-sex 
structure? Finally, can we learn to predict 
population trajectories accurately, and if not, 
why not? 

A surprising development has been the 
emergence of sex ratios as important de- 
mographic variables. Not only can they vary 
greatly by microhabitat (Ostfeld et al., 1985), 
be biased by dispersal (Lidicker and Sten- 
seth, 1992), and influenced by density (Clut- 
ton-Brock, 1991; Fredga et al., 1977; van 
Schaik and Hrdy, 1991), but in some cir- 
cumstances can be influenced by litter size 
and maternal social status and condition 
(Austad and Sunquist, 1986; Clutton-Brock 
and Albon, 1982; Clutton-Brock and Iason, 
1986; Clutton-Brock et al., 1977, 1982: 
Cockburn et al., 1985; Frank, 1992; Sy- 
mington, 1987; Verme, 1969). I expect fur- 
ther significant discoveries in this area. 

One important trend has been the redis- 
covery of multi-factorial models of popu- 
lation regulation. In the early part of this 
century, ecologists and wildlife biologists 
routinely accepted that populations were 
subject to a multiplicity of positive and neg- 
ative forces. Then, as the field became more 
quantitative, along with the success of re- 
ductionist and experimental approaches to 
research, pressures became intense for find- 
ing general and simple explanations for how 
things worked. Complex and especially id- 
losyncratic explanations were viewed sus- 
piciously as non-scientific. In recent de- 
cades, ecologists have become more 


comfortable with holistic views and partic- 
ularly with a research protocol that balances 
reductionist and holistic aspects (Lidicker, 
19885, 1991; Macfadyen, 1975, 1978; Mc- 
Intosh, 1980; Odum, 1977). This new per- 
spective has encouraged viewing density 
regulation in a systems context with nu- 
merous intrinsic and extrinsic factors inter- 
acting together, a multi-factor perspective 
(Finerty, 1980; Lidicker, 197359978 
1988a). Such a perspective is only the start- 
ing point, however, as the quantitative re- 
lationships among the factors remains to be 
determined. We need to know the temporal 
and spatial stability of the patterns ob- 
served, and finally we must search for gen- 
eralities in pattern. This knowledge will al- 
low us to manipulate (manage) population 
numbers effectively and to make predic- 
tions of future density changes, or at least 
to know when predictions are reliable and 
when they are not. It will also give us the 
data to look afresh at some old questions 
such as the extent to which carrying capac- 
ities of habitats and equilibrium densities 
(K) coincide. 

With such a huge agenda ahead of us, it 
is encouraging that some mammalian ecol- 
ogists are exploring effectively the realities 
of this complex world. Pioneering research 
based on multi-factorial hypotheses has been 
reported by Wagner and Stoddart (1972), 
Keith and Windberg (1978), Taitt and Krebs 
(1983), Sinclair (1986), Hansson and Hen- 
tonnen (1988), Desy and Batzli (1989), and 
others. The approach remains controver- 
sial, however (Gaines et al., 1991; Krebs, 
1979b; Tamarin, 1978a); and the future is 
as unpredictable for this field as it is for 
many mammalian population densities. 

6) Conservation. Conservation biology 
is the extension of wildlife management from 
concern for economically important species 
to the biota as a whole. As such, it was for 
many decades a legitimate part of biology. 
Then in the rush and push for ““modern sci- 
ence” that swept through biology in the 
1960s, conservation became relegated to its 
political and moral aspects, and was shunned 


POPULATIONS 339 


by the scientific establishment. However, 
with the accelerating deterioration of the 
Earth in the 1980s, along with the prospects 
for massive losses in biodiversity, and with 
the help of significant pressure from uni- 
versity students, conservation biology re- 
emerged as an important field of scientific 
inquiry. Even staid academic units began to 
offer courses, and even major programs, in 
this area. Helping to legitimatize the field 
was the establishment of two high quality 
journals, Biological Conservation in 1968 
and Conservation Biology in 1987. Coinci- 
dent with the latter event was the initiation 
of the Society for Conservation Biology, 
which was an instant success. 

Now conservation biologists are applying 
frontline basic research in population, com- 
munity, and landscape ecology, as well as 
evolutionary biology and population genet- 
ics to address the mega-threats to humanity 
caused by losses of biodiversity and the un- 
controlled growth of our own species. As 
they operate from an increasingly firm foun- 
dation in basic science, they can and are 
moving with confidence to embrace politi- 
cal, social, and even moral aspects of the 
human predicament. Thus, the realistically 
interdisciplinary nature of the problems are 
being acknowledged and addressed, but this 
time, hopefully, without losing a solid foot- 
ing in the basic sciences. At this writing, 
society at large is beginning to show a glim- 
mer of recognition for where it is headed, 
but support for research in the relevant ar- 
eas remains a tiny fraction of that provided 
for activities that tend to exacerbate the 
problems. Whether or not human society at 
large recognizes its dilemma in time to deal 
with it humanely is the mega-question for 
the future. 


Future Perspectives 


Even a cursory overview of how popu- 
lation ecology has changed during the past 
75 years reveals a dramatic ontogeny. Lan- 


guage has changed, new concepts have ap- 
peared, and the empirical base and number 
of scientists have grown enormously. All 
these facts signal that the field has not yet 
reached maturity, and so should have a long 
future. A seedling has indeed grown into a 
young tree. In this development, mammal- 
ogists have played critical and constructive 
roles. 

Setting aside this developmental meta- 
phor, one can predict with confidence that 
mammalian population ecology will not fade 
away. Just as the structure and function of 
organisms and of cells is fundamental to any 
overview of biology, so too is the structure 
and function of populations. Populations, 
moreover, are the parts (holons) for com- 
munities and landscapes that in turn cannot 
be understood without knowledge of these 
constituents. Besides, as outlined in the six 
preceding vignettes about the current status 
of subdisciplines within population ecology, 
there is much to be learned at this level as 
well. 

Trying to be as subjective as possible, I 
suggest that the following topics will receive 
increasing attention in the near future: 


1. Relating genetic structure to demograph- 
ic and social structure, giving new in- 
sights to all three areas, and tending to 
blur the traditional distinction between 
ecological and evolutionary time scales; 

Focusing on landscape-level issues, both 

for their intrinsic interest and because 

community-types are being increasingly 
fragmented; 

3. Understanding of dispersal as critical in- 
puts and outputs to population systems 
and a major connector and information 
link within meta-populations; 

4. Recognizing parasitic and cooperative 
coactions as important community or- 
ganizers; 

5. Exploring the interplay of social behav- 
ior and other aspects of population bi- 
ology, with the emphasis being on mu- 
tual effects, and on a comparative 
approach; 


N 


340 LIDICKER 


6. Appreciating the local complexity and 
global simplicity of density regulating 
mechanisms, and reconciling this ap- 
parent paradox through multi-factor 
models; and 

7. Giving all the support we can to arresting 
the decline in our life-support system 
through conservation biology and relat- 
ed efforts. 


Where do mammals and mammalogists 
fit into all of this relating, focusing, under- 
standing, recognizing, exploring, appreci- 
ating, and giving? Right at the front lines. 
Mammals are among the more complex in- 
habitants of this planet; so if we can un- 
derstand them, we can provide guidelines 
for the rest. Also, being larger and cleverer 
than most creatures, they often represent 
keystone species (strong interactors) in their 
communities. As such, they often can serve 
as indicator species for the status and sta- 
bility of intractably complex chunks of the 
biosphere. Finally, mammals include the 
species Homo sapiens. Thus for us, mam- 
mals are our closest kin, and no wonder 
many are loved, feared, admired, or reviled. 
When we study life, we learn about our 
planet and ourselves, but when we study 
mammals we come even closer to intimate 
understanding. 


Acknowledgments 


History abhors any attempt to define bound- 
aries around those who can be credited or blamed 
for any effort at reconstruction. All one’s expe- 
riences contribute in intangible ways. Neverthe- 
less, I gratefully acknowledge R. H. Tamarin who 
contributed to early discussions regarding the 
scope and content of this chapter, C. W. Wozen- 
craft who kindly trusted me with seven programs 
of ASM annual meetings from the society’s ar- 
chives, E. P. Odum for helpful discussions, and 
the editors of this volume for choosing me for 
this assignment. Both Tamarin and an anony- 
mous reviewer made many helpful suggestions 
for improving the manuscript. L. N. Lidicker 
gave logistic and other support throughout the 
project. 


Literature Cited 


ALEXANDER, R. D. 1974. The evolution of social be- 
havior. Annual Review of Ecology and Systematics, 
5:325-383. 

ALLEE, W. C., A. E. EMERSON, O. PARK, T. PARK, AND 
K. P. ScHmipt. 1949. Principles of animal ecology. 
W.B. Saunders, Philadelphia, Pennsylvania, 837 pp. 

ALLEN, T. F. H., AND T. B. STARR. 1982. Hierarchy: 
perspectives for ecological complexity. University of 
Chicago Press, Chicago, 310 pp. 

AMLANER, C. J., AND D. W. MACDONALD (EDs.). 1980. 
A handbook on telemetry and radio tracking. Per- 
gamon Press, Elmsford, New York, 804 pp. 

ANDERSON, P. K. 1970. Ecological structure and gene 
flow in small mammals. Symposia of the Zoological 
Society of London, 26:299-325. 

ANDERSON, R. M. 1982. Fox rabies. Pp. 242-261, in 
Population dynamics of infectious diseases: theory 
and applications (R. M. Anderson, ed.). Chapman 
and Hall, London, 368 pp. 

ANDERSON, R. M., ANDR. M. May. 1979. Population 
biology of infectious diseases: Part I. Nature, 280: 
361-367. 

ANDERSON, R. M., H. C. JACKSON, R. M. MAy, AND 
A.D.M. SmitH. 1981. Population dynamics of fox 
rabies in Europe. Nature, 289:765-61. 

ANDRZEJEWSKI, R., A. KAJAK, AND E. PIECZYNSKA. 
1963. Efecty migracji. Ekologia Polska, Seria B, 
9:161-172. 

ARMITAGE, K. B. 1988. Resources and social orga- 
nization of ground-dwelling squirrels. Pp. 131-155, 
in The ecology of social behavior (C. N. Slobodchi- 
koff, ed.). Academic Press, San Diego, 429 pp. 

AUSTAD, S., AND M. E. Sunquist. 1986. Sex-ratio 
manipulation in the common opossum. Nature, 324: 
58-60. 

BarasH, D. P. 1974. The evolution of marmot so- 
cieties: a general theory. Science, 185:415-—420. 

BARNETT, R. J. 1982. Chinese sages and western bi- 
ologists. Asianist (California State University, Chi- 
co), 7:3-13. 

Bates, M. 1950. The nature of natural history. Charles 
Scribner, New York, 309 pp. 

BAUCHAU, V., AND E. LEBOULENGE. 1991. Population 
biology of woodland rodents 1n a patchy landscape. 
Pp. 275-283, in Le rongeur et l’espace (M. LeBerre 
and L. Le Guelte, eds.). R. Chabaud, Paris, 362 pp. 

BerGeR, J. 1986. Wild horses of the Great Basin: 
social competition and population size. University 
of Chicago Press, Chicago, 326 pp. 

1988. Social systems, resources, and phylo- 
genetic inertia: an experimental test and its limita- 
tions. Pp. 157-186, in The ecology of social behavior 
(C. N. Slobodchikoff, ed.). Academic Press, San Di- 
ego, 429 pp. 

Berry, R. J. 1987. Population studies of mammals. 
Symposia of the Zoological Society of London, 58: 
1-5. 

BONDRUP-NIELSEN, S., AND R. A. IMs. 1988. Pre- 
dicting stable and cyclic populations of Clethriono- 
mys. Oikos, 52:178-185. 

BOURLIERE, F. 1951. Vie et moeurs des mammiferes. 
Payot, Paris, 249 pp. 


POPULATIONS 341 


Boyce, M. S. 1977. Population growth with sto- 
chastic fluctuations in the life table. Theoretical Pop- 
ulation Biology, 12:366-373. 

Boyce, M. S. (EpD.). 1988. Evolution of life histories 
of mammals. Yale University Press, New Haven, 
Connecticut, 373 pp. 

BRANDON, R. N., AND R. M. BurIAn (EDS.). 1984. 
Genes, organisms, populations: controversies over 
the units of selection. Massachusetts Institute of 
Technology Press, Cambridge, Massachusetts, 329 
pp. 

Brown, J. H. 1971. Mechanisms of competitive ex- 
clusion between two species of chipmunks. Ecology, 
52:305-311. 

Brown J. H., AND D. W. Davipson. 1977. Compe- 
tition between seed-eating rodents and ants in desert 
ecosystems. Science, 196:880-882. 

Brown, J. H., AND D. E. Witson. 1994. Natural his- 
tory and evolutionary ecology. Pp. 377-397, in Sev- 
enty-five years of mammalogy (1919-1994) (E. C. 
Birney and J. R. Choate, eds.). Special Publication, 
The American Society of Mammalogists, | 1:1-433. 

Brown, J. H., D. W. DAvipsoNn, AND O. J. REICHMAN. 
1979. An experimental study of competition be- 
tween seed-eating desert rodents and ants. American 
Zoologist, 19:1129-1143. 

CaBRERA, A. 1922. Manual de Mastozoologia. Com- 
pania Anonima de Libreria, Madrid, 440 pp. 

CaLHoun, J. B. 1949. A method for self-control of 
population growth among mammals living in the 
wild. Science, 109:333-335. 

1962. The ecology and sociology of the Nor- 
way rat. United States Department of Health, Ed- 
ucation, and Welfare, Public Health Service Publi- 
cation, 1008, Bethesda, Maryland, 288 pp. 

CAUGHLEY, G. 1970. Eruption of ungulate popula- 
tions, with emphasis on Himalayan thar in New Zea- 
land. Ecology, 51:53-72. 

CHAPMAN, R. N. 1931. Animal ecology. McGraw- 
Hill Book Company, New York, 464 pp. 

Cuitty, D. 1952. Mortality among voles (Microtus 
agrestis) at Lake Vyrnwy, Montgomeryshire in 1936- 
9. Philosophical Transactions of the Royal Society 
of London, Series B, Biological Science, 236:505- 
552: 


1955. Adverse effects of population density 
upon the viability of later generations. Pp. 57-67, in 
The numbers of man and animals (J. B. Cragg and 
N. W. Pirie, eds.). Institute of Biology Symposia, 4, 
Oliver and Boyd, Edinburgh, 152 pp. 

1958. Self-regulation of numbers through 

changes in viability. Cold Spring Harbor Symposia 

on Quantitative Biology, 22:277-280. 

1960. Population processes in the vole and 

their relevance to general theory. Canadian Journal 

of Zoology, 36:99-113. 

. 1967. The natural selection of self-regulatory 
behaviour in natural populations. Proceedings of the 
Ecological Society of Australia, 2:51-78. 

CuRISTIAN, J. J. 1950. The adreno-pituitary system 
and population cycles in mammals. Journal of Mam- 
malogy, 31:247-259. 

1955a. The effect of population size on the 

weights of the reproductive organs of white mice. 

Naval Medical Research Institute Research Report, 

13:81-88. 


1955b. Effect of population size on the ad- 

renal glands and reproductive organs of male mice 

in populations of fixed size. American Journal of 

Physiology, 182:292-300. 

1959. The roles of endocrine and behavioral 

factors in the growth of mammalian populations. Pp. 

71-97, in Comparative endocrinology (A. Gorbman, 

ed.). John Wiley & Sons, Incorporated, New York, 

746 pp. 

1961. Phenomena associated with popula- 
tion density. Proceedings of the National Academy 
of Sciences, USA, 47:428-449. 

CHRISTIAN, J. J., AND D. E. Davis. 1955. Reduction 
of adrenal weight in rodents by reducing population 
size. Transactions of the North American Wildlife 
Conference, 20:177-189. 

CLEMENTS, F. E. 1905. Research methods in ecology. 
University Publishing Company, Lincoln, Nebras- 
ka, 334 pp. 

1916. Plant succession: an analysis of the 

development of vegetation. Carnegie Institute of 

Washington, Publication, 242, Washington, D.C., 512 


CLEMENTS, F. E., AND V. E. SHELFORD. 1939. Bio- 
ecology. John Wiley & Sons, Incorporated, New York, 
425 pp. 

CLutTTon-Brock, T. H. 1991. The evolution of pa- 
rental care. Princeton University Press, Princeton, 
352 pp. 

CLuTTON-Brock, T. H., ANDS. D. ALBON. 1982. Pa- 
rental investment in male and female offspring in 
mammals. Pp. 223-258, in Growing points in ethol- 
ogy (P. P. G. Bateson and R. A. Hinde, eds.). Cam- 
bridge University Press, Cambridge, 548 pp. 

CLuTTON-Brock, T. H., AND G. R. IASON. 1986. Sex 
ratio variation in mammals. The Quarterly Review 
of Biology, 61:339-374. 

CLuTTON-Brock, T. H., F. E. GUINNESS, AND S. D. 
ALBON. 1982. Red deer: the behavior and ecology 
of two sexes. Chicago University Press, Chicago, 378 


CLUTTON-Brock, T. H., P. H. HARVEY, AND B. RUDDER. 
1977. Sexual dimorphism, socionomic sex ratio and 
body weight in primates. Nature, 269:797-800. 

Cockburn, A. 1988. Social behaviour in fluctuating 
populations. Croom Helm, Sydney, 239 pp. 

CocksBurwn, A., M. P. Scott, AND D. J. Scotts. 1985. 
Sex ratio and intra-sexual kin competition in mam- 
mals. Oecologia, 66:427-429. 

Coxe, L. C. 1948. Population phenomena and com- 
mon knowledge. Scientific Monthly, 67:338-345. 
Coney, W., AND J. D. NicHors. 1978. The use of 
models in small mammal population studies. Pp. 
14-35, in Populations of small mammals under nat- 
ural conditions (D. P. Snyder, ed.). Pymatuning Lab- 
oratory of Ecology, Special Publication, 5, 237 pp. 

Crowcrort, P. 1991. Elton’s ecologists: a history of 
the Bureau of Animal Population. University of Chi- 
cago Press, Chicago, 177 pp. 

Crowcrort, P., AND F. P. Rowe. 1957. The growth 
of confined colonies of the wild house-mouse (Mus 
msuculus L.). Proceedings of the Zoological Society 
of London, 129:359-370. 

DaAsMANN, R. F. 1964. Wildlife biology. John Wiley 
& Sons, Incorporated, New York, 231 pp. 

Davipson, D. W., AND J. H. Brown. 1980. Com- 


342 LIDICKER 


petition and the structure of granivore communities. 
BioScience, 30:233-238. 

Davis, D. E. 1949. The weight of wild brown rats at 
sexual maturity. Journal of Mammalogy, 30:125- 
130. 

Deevey, E. S., Jr. 1947. Life tables for natural pop- 
ulations of animals. The Quarterly Review of Biol- 
ogy, 22:283-314. 

Dekker, H. 1975. A simple mathematical model of 
rodent population cycles. Journal of Mathematical 
Biology, 2:57-67. 

Desy, E. A., AND G. O. BATzLt. 1989. Effects of food 
availability and predation on prairie vole demog- 
raphy: a field experiment. Ecology, 70:412-421. 

Dietz, K., AND D. SCHENZLE. 1985. Mathematical 
models for infectious disease statistics. Pp. 167-204, 
in A celebration of statistics: the ISI centenary vol- 
ume (A. C. Atkinson and S. E. Fienberg, eds.). 
Springer-Verlag, New York, 606 pp. 

Dun ap, T. R. 1981. DDT: scientists, citizens, and 
public policy. Princeton University Press, Princeton, 
318 pp. 

EISENBERG, J. F. 1967. A comparative study of rodent 
ethology with emphasis on evolution of social be- 
havior, I. Proceedings of the United States National 
Museum, 122 (3597):1-35. 

EISENBERG, J. F., AND D. G. KLEIMAN (EDsS.). 1983. 
Advances in the study of mammalian behavior. Spe- 
cial Publication, The American Society of Mam- 
malogists, 7:1-753. 

EISENBERG, J. F., AND J. O. WoLFF. 1994. Behavior. 
Pp. 398-420, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 1 1:1-433. 

E._pripGE, N. 1985. Unfinished synthesis: biological 
hierarchies and modern evolutionary thought. Ox- 
ford University Press, New York, 237 pp. 

ELton, C.S. 1927. Animal ecology. MacMillan, New 
York, 207 pp. 

. 1930. Animal ecology and evolution. Oxford 

University Press, 96 pp. 

1942. Voles, mice, and lemmings: problems 
in population dynamics. Clarendon Press, Oxford, 
496 pp. 

ErRINGTON, P. L. 1946. Predation and vertebrate 
populations. Quarterly Review of Biology, 21:144- 
177, 221-245. 

. 1963. The phenomenon of predation. Amer- 
ican Scientist, 51:180-192. 

FENNER, F. 1976. Myxoma virus and Oryctolagus 
cuniculus: two colonizing species. Pp. 485-499, in 
The genetics of colonizing species (H. G. Baker and 
G. L. Stebbins, eds.). Academic Press, New York, 
588 pp. 

Fenton, M. B., AND D. W. THomas. 1985. Migra- 
tions and dispersal of bats (Chiroptera). Pp. 409- 
424, in Migration: mechanisms and adaptive signif- 
icance (M. A. Rankin, ed.). Contributions in Marine 
Science, Supplement Volume 27, 868 pp. 

Finerty, J.P. 1980. The population ecology of cycles 
in small mammals. Yale University Press, New Ha- 
ven, Connecticut, 234 pp. 

Forbes, S. A. 1880. On some interactions of organ- 


isms. Bulletin of the Illinois State Laboratory of Nat- 

ural History, 1:3-17. 

1887. The lake as a microcosm. Bulletin of 
Science A (Peoria). Reprinted Illinois Natural His- 
tory Survey Bulletin, 15:537-550, 1925. 

ForMan, R. T. T., AND M. Gopron. 1986. Land- 
scape ecology. John Wiley & Sons, Incorporated, 
New York, 620 pp. 

FRANK, L. 1992. Spotted hyenas may use neonatal 
siblicide to bias sex ratios. Abstract of the IV Inter- 
national Behavioral Ecological Congress, Princeton 
University. 

FReEDGA, K., A. Gropp, I. WINKLING, AND F. FRANK, 
1977. A hypothesis explaining the exceptional sex 
ratio of the wood lemming (Myopus shisticolor). Her- 
editas, 85:101-104. 

FRIEDERICHS, K. 1927. Grundsatzliches tiber die Le- 
benseinheiten hoherer Ordnung und den 6kologisch- 
en Einheitsfaktor. Naturwissenschaften, 15:153-157. 

. 1930. Die Grundfragen und Gesetzmassig- 
keiten der Land und Forst-wirtschaftlichen ZoGlogie. 
Paul Parey, Berlin, 2 vols. 

GABRIELSON, I. N. 1951. Wildlife management. Mac- 
Millan, New York, 274 pp. 

Gaines, M.S. 1981. Importance of genetics to pop- 
ulation dynamics. Pp. 1-27, in Mammalian popu- 
lation genetics (M. H. Smith and J. Joule, eds.). Uni- 
versity of Georgia Press, Athens, 380 pp. 

Gaines, M. S., N. C. STENSETH, M. L. JoHNsoN, R. A. 
IMs, AND S. BONDRUP-NIELSEN. 1991. A response 
to solving the enigma of population cycles with a 
multifactorial perspective. Journal of Mammalogy, 
72:627-631. 

Gaus, G. F. 1934. The struggle for existence. Wil- 
liams & Wilkins, Baltimore, 163 pp. 

GiLL, A. E., AND W. C. WozENcRAFT. 1994. Com- 
mittees and annual meetings. Pp. 155-170, in Sev- 
enty-five years of mammalogy (1919-1994) (E. C. 
Birney and J. R. Choate, eds.). Special Publication, 
The American Society of Mammalogists, 11:1-433. 

GRANT, P. R. 1969. Experimental studies of com- 
petitive interaction in a two-species system. 1. Mi- 
crotus and Clethrionomys species in enclosures. Ca- 
nadian Journal of Zoology, 47:1059-1082. 

1972. Interspecific competition among ro- 

dents. Annual Review of Ecology and Systematics, 

3:79-106. 

1978. Competition between species of small 
mammals. Pp. 38-51, in Populations of small mam- 
mals under natural conditions (D. P. Snyder, ed.). 
Pymatuning Laboratory of Ecology, Special Publi- 
cation, 5, 237 pp. 

Grecory, R. D. 1991. Parasite epidemiology and 
host population growth: Heligmosomoides polygyrus 
(Nematoda) in enclosed wood mouse populations. 
The Journal of Animal Ecology, 60:805-821. 

HaAmILton, W. D. 1971. Geometry for the selfish 
herd. Journal of Theoretical Biology, 31:295-311. 

HamMILton, W. J., Jk. 1939. American mammals. 
McGraw-Hill Book Company, New York, 434 pp. 

1953. Migrants and emigrants. New York 

State Conservationist, June-July: 10-11. 

. 1955. Mammalogy in North America. Pp. 

661-688, in A century of progress in the natural 


POPULATIONS O45 


sciences 1853-1953. California Academy of Science, 
San Francisco, 807 pp. 

HAmMmonp, P.S. 1987. Techniques for estimating the 
size of whale populations. Symposia of the Zoolog- 
ical Society of London, 58:225-245. 

Hansson, L. 1977. Spatial dynamics of field voles 
Microtus agrestis in heterogeneous landscapes. Oi- 
kos, 29:539-544. 

Hansson, L., AND H. HENTTONEN. 1988. Rodent dy- 
namics as community processes. Trends in Ecology 
& Evolution, 3:195-200. 

Hansson, L., AND N. C. STENSETH (EDS.). 1988. Mod- 
elling small rodent population dynamics. Oikos, 52: 
137-229. 

HASKELL, E. F. 1949. A clarification of social science. 
Main Currents in Modern Thought, 7:45-51. 

HestBEck, J. B. 1988. Population regulation of cyclic 
mammals: a model of the social fence hypothesis. 
Oikos, 52:156-168. 

Hisy, A. R., AND J. S. JEFFERY. 1987. Census tech- 
niques for small populations, with special reference 
to the Mediterranean monk seal. Symposia of the 
Zoological Society of London, 58:193-210. 

HILBorn, R., AND S. C. STEARNS. 1982. On inference 
in ecology and evolutionary biology: the problem of 
multiple causes. Acta Biotheoretica, 31:145-164. 

Howarp, L. O., AND W. F. Fiske. 1911. The im- 
portation into the United States of the parasites of 
the gipsy-moth and the brown-tail moth. United 
States Department of Agriculture, Bureau of Ento- 
mology Bulletin, 91. 

Howarp, W.E. 1953. Rodent control on California 
ranges. Journal of Range Management, 6:423-434. 

1960. Innate and environmental dispersal of 

individual vertebrates. The American Midland Nat- 

uralist, 63:152-161. 

1965. Interaction of behavior, ecology, and 
genetics of introduced mammals. Pp. 461-480, in 
The genetics of colonizing species (H. G. Baker and 
G. L. Stebbins, eds.). Academic Press, New York, 
588 pp. 

Howe i, A. B., 1923. Periodic fluctuations in the 
numbers of small mammals. Journal of Mammal- 
ogy, 4:149-155. 

KALELA, O. 1961. Seasonal change of habitat in the 
Norwegian lemming, Lemmus lemmus (L.). Annuals 
of the Academy of Science Fennicae Series A, IV, 
Biology, 55:1-72. 

KAUFMAN, G. A. 1989. Use of fluorescent pigments 
to study social interactions in a small nocturnal ro- 
dent, Peromyscus maniculatus. Journal of Mam- 
malogy, 70:171-174. 

KeiTH, L. B., AND L. A. WINDBERG. 1978. A demo- 
graphic analysis of the snowshoe hare cycle. Wildlife 
Monographs, 58:1-70. 

Kina, J. A. 1955. Social behavior, social organiza- 
tion, and population dynamics in a black-tailed prai- 
rie dog town in the Black Hills of South Dakota. 
Contributions from the Laboratory of Vertebrate Bi- 
ology, University of Michigan, 67:1-123. 

KLEIMAN, D., AND J. F. EISENBERG. 1973. Compar- 
ison of canid and felid social systems from an evo- 
lutionary perspective. Animal Behaviour, 21:637- 
659. 


KonisH! M., AND Y. ito. 1973. Early entomology in 
East Asia. Pp. 1-20, in History of entomology (R. 
F. Smith, T. E. Mittler, and C. N. Smith, eds.). An- 
nual Reviews Incorporated, Palo Alto, California, 
517 pp. 

Kress, C. J. 1964. The lemming cycle at Baker Lake, 
Northwest Territories, during 1959-62. Arctic In- 
stitute of North America Technical Paper, 15, 104 
pp. 


. 1971. Genetic and behavioral studies on fluc- 
tuating vole populations. Pp. 243-256 in Dynamics 
of populations (P. J. den Boer and G. R. Gradwell, 
eds.). Proceedings of the Advanced Study Institute 
on ‘Dynamics of Numbers in Populations,’ Ooster- 
beek. Centre for Agricultural Publications and Doc- 
uments, Wageningen, The Netherlands, 611 pp. 

. 1979a. Dispersal, spacing behaviour, and ge- 

netics in relation to population fluctuations in the 

vole Microtus townsendii. Fortschrift fiir Zodlogie, 

25:61-77. 

. 19796. Small mammal ecology. Science, 203: 
350-351. 

Kress, C. J., M. S. Gaines, B. L. KELLER, J. H. Myers, 
AND R. H. TAMARIN. 1973. Population cycles in 
small rodents. Science, 179:35-41. 

Kress, J. R., AND N. B. Davies (EDs.). 1984. Behav- 
ioural ecology: an evolutionary approach, Second 
ed., Sinauer Associates, Incorporated, Sunderland, 
Massachusetts, 493 pp. 

Lack, D. 1954. The natural regulation of animal 
numbers. Oxford University Press, New York, 343 
pp. 


. 1966. Population studies of birds. Oxford 
University Press, New York, 341 pp. 

Leary, R. A. 1985. Interaction theory in forest ecol- 
ogy and management. Junk Publishers, Boston, 219 
pp. 

Leopotp, A. 1933. Game management. Charles 
Scribner, New York, 481 pp. 

Lesuigz, P.H. 1945. On the use of matrices in certain 
population mathematics. Biometrika, 33:183-212. 
Levins, R. 1968. Evolution in changing environ- 
ments: some theoretical explorations. Princeton 

University Press, Princeton, New Jersey, 120 pp. 

LEWONTIN, R.C. 1970. The units of selection. Annual 
Review of Ecology and Systematics, 1:1-18. 

Lipicker, W. Z., JR. 1962. Emigration as a possible 
mechanism permitting the regulation of population 
density below carrying capacity. The American Nat- 
uralist, 96:29-33. 

1965. Comparative study of density regula- 

tion in confined populations of four species of ro- 

dents. Researches on Population Ecology, 7:57-72. 

1973. Regulation of numbers in an island 

population of the California vole, a problem in com- 

munity dynamics. Ecological Monographs, 43:27 1- 

302. 


1975. The role of dispersal in the demogra- 
phy of small mammals. Pp. 103-128, in Small mam- 
mals: their productivity and population dynamics 
(F. B. Golley, K. Petrusewicz, and L. Ryszkowski, 
eds.). Cambridge University Press, London, 251 pp. 
. 1978. Regulation of numbers in small mam- 
mal populations—historical reflections and a syn- 


344 


thesis. Pp. 122-141, in Populations of small mam- 

mals under natural conditions (D. P. Snyder, ed.). 

Pymatuning Laboratory of Ecology, Special Publi- 

cation, 5, 237 pp. 

1979a. Analysis of two freely-growing en- 

closed populations of the California vole. Journal of 

Mammalogy, 60:447-466. 

1979b. A clarification of interactions in eco- 

logical systems. BioScience, 29:475-477. 

1984. Kazimierz Petrusewicz, 1906-1982. 

Journal of Mammalogy, 65:168-170. 

1985a. Population structuring as a factor in 

understanding microtine cycles. Acta Zoologica Fen- 

nica, 173:23-27. 

1985b. An overview of dispersal in non-vo- 

lant small mammals. Pp. 369-385 in Migration: 

mechanisms and adaptive significance (M. A. Ran- 
kin, ed.). Contributions in Marine Science, Supple- 
ment Volume 27, University of Texas, 868 pp. 

. 1988a. Solving the enigma of microtine “‘cy- 

cles.”” Journal of Mammalogy, 69:225-235. 

. 1988. The synergistic effects of reductionist 

and holistic approaches in animal ecology. Oikos, 

53:278-281. 

. 1991. In defense of a multifactor perspective 
in population ecology. Journal of Mammalogy, 72: 
631-635. 

LipIckER, W. Z., JR., AND N. C. STENSETH. 1992. To 
disperse or not to disperse: who does it and why? 
Pp. 21-36, in Animal dispersal: small mammals as 
a model (N. C. Stenseth and W. Z. Lidicker, Jr., eds.). 
Chapman and Hall, London, 365 pp. 

LipickER, W. Z., Jr., J. O. Woirr, L. N. Lipicker, 
AND M. H. SmitH. 1991. Utilization of a habitat 
mosaic by cotton rats during a population decline. 
Landscape Ecology, 6:259-268. 

Lomnicki, A. 1980. Regulation of population density 
due to individual differences and patchy environ- 
ment. Oikos, 35:184-193. 

LotKxa, A. J. 1907. Relation between birth rates and 
death rates. Science, 26:21-22. 

. 1925. Elements of physical biology. Williams 
and Wilkins, Baltimore, 460 pp. 

MACFADYEN, A. 1975. Some thoughts on the behav- 
iour of ecologists. The Journal of Animal Ecology, 
44:351-363. 

. 1978. The ecologist’s role in the international 
scientific community, Oikos, 31:1-2. 

MacLega, S. F., Jr., B. M. FitzGERALD, AND F. A. 
PITELKA. 1974. Population cycles in arctic lem- 
mings: winter reproduction and predation by wea- 
sels. Arctic and Alpine Research, 6:1—12. 

Mauer, W.J. 1967. Predation by weasels on a winter 
population of lemmings, Banks Island, Northwest 
Territories. The Canadian Field Naturalist, 81:248- 
250. 

MALtTHus, T. R. 1798. An essay on the principle of 
population. Johnson, London. (reprinted with intro- 
duction and bibliography. E. A. Wrigley and D. Sou- 
den [eds.]. 1988. W. Pickering, London, 139 pp.) 

Mares, M. A., AND G. N. CAMERON. 1994. Com- 
munity and ecosystem ecology. Pp. 348-376, in Sev- 
enty-five years of mammalogy (1919-1994) (E. C. 
Birney and J. R. Choate, eds.). Special Publication, 
The American Society of Mammalogists, 11:1-433. 


LIDICKER 


Massey, D.R., AND J. JOULE. 1981. Spatial-temporal 
changes 1n genetic composition of deer mouse pop- 
ulations. Pp. 180-201, in Mammalian population 
genetics (M. H. Smith and J. Joule, eds.). University 
of Georgia Press, Athens, 380 pp. 

May, R. M., ANDR. M. ANDERSON. 1979. Population 
biology of infectious diseases. II. Nature, 280:455- 
461. 

McAtTEE, W.L. 1936. Malthusian principle in nature. 
Scientific Monthly, 42:444-456. 

McCuttouGu, D.R. 1979. The George Reserve deer 
herd: population ecology of a K-selected species. 
University of Michigan Press, Ann Arbor, 271 pp. 

1985. Long range movements of large ter- 
restrial mammals. Pp. 444-465, in Migration: mech- 
anisms and adaptive significance (M. A. Rankin, ed.). 
Contributions in Marine Science, Supplement Vol- 
ume 27, 868 pp. 

McInrosu, R. P. 1980. The background and some 
current problems in theoretical ecology. Synthese, 
43:195-225. 

. 1985. The background of ecology: concept 
and theory. Cambridge University Press, 383 pp. 
McSHEA, W. J., AND D. M. Mapison. 1992. Alter- 
native approaches to the study of small mammal 
dispersal: insights from radiotelemetry. Pp. 319-332, 
in Animal dispersal: small mammals as a model (N. 
C. Stenseth and W. Z. Lidicker, Jr., eds.). Chapman 

and Hall, London, 365 pp. 

Mecu, L. D. 1987. Age, season, distance, direction, 
and social aspects of wolf dispersal from a Minnesota 
pack. Pp. 55-74, in Mammalian dispersal patterns: 
the effects of social structure on population genetics 
(B. D. Chepko-Sade and Z. T. Halpin, eds.). Uni- 
versity of Chicago Press, Chicago, 342 pp. 

MENDALL, H. L. 1944. Food of hawks and owls in 
Maine. The Journal of Wildlife Management, 8:198- 
208. 

MERRIAM, G. 1990. Ecological processes in the time 
and space of farmland mosaics. Pp. 121-133, in 
Changing landscapes: an ecological perspective (I. S. 
Zonnefeld and R. T. T. Forman, eds.). Springer-Ver- 
lag, New York, 286 pp. 

1991. Corridors and connectivity: animal 
populations in heterogeneous environments. Pp. 133- 
142, in Nature conservation 2: the role of corridors 
(D. A. Sauners and R. J. Hobbs, eds.). Surrey Beatty, 
Chipping Norton, New South Wales, 442 pp. 

MILter, G.S., JR. 1929. Mammalogy and the Smith- 
sonian Institution. Annual Report (1928) Smithson- 
ian Institution Publication, 2981, pp. 391-411. 

Mo .tison, D. 1977. Spatial contact models for eco- 
logical and epidemic spread. Journal of the Royal 
Statistical Society, B39:283-326. 

1987. Population dynamics of mammalian 
diseases. Symposia of the Zoological Society of Lon- 
don, 58:329-342. 

Montcomery, W. I. 1987. The application of cap- 
ture-mark-recapture methods to the enumeration of 
small mammal populations. Symposia of the Zoo- 
logical Society of London, 58:25-57. 

Morais, R. D., AND P. R. GRANT. 1972. Experimen- 
tal studies of competitive interaction in a two-species 
system. IV. Microtus and Clethrionomys species in 


POPULATIONS 345 


a single enclosure. The Journal of Animal Ecology, 
41:275-290. 

Mouncer, J.C., AND J. H. Brown. 1981. Competition 
in desert rodents: an experiment with semipermeable 
enclosures. Science, 211:510-512. 

Newsome, A. E., AND L. K. Corsett. 1975. Out- 
breaks of rodents in semi-arid and arid Australia: 
causes, preventions, and evolutionary considera- 
tions. Pp. 117-153, im Rodents in desert environ- 
ments (I. Prakash and P. K. Ghosh, eds.). Junk Pub- 
lications, The Hague, 624 pp. 

Ovum, E. P. 1953. Fundamentals of ecology. W. B. 
Saunders Company, Philadelphia, 384 pp. 

1977. The emergence of ecology as a new 
integrative discipline. Science, 195:1289-1293. 

O’NEILL, R. V., D. L. DEANGELIS, J. B. WAIDE, AND 
T. F. H. Atten. 1986. A hierarchical concept of 
ecosystems. Princeton University Press, Princeton, 
254 pp. 

OsTFELD, R. S., W. Z. LipicKER, JR., AND E. J. HESKE. 
1985. The relationship between habitat heteroge- 
neity, space use, and demography in a population of 
California voles. Oikos, 45:433-442. 

Park, T. 1946. Some observations on the history and 
scope of population ecology. Ecological Mono- 
graphs, 16:313-320. 

PEARL, R. 1922. The biology of death. Lippincott, 
Philadelphia, 275 pp. 

1925. The biology of population growth. 

Knopf, New York, 260 pp. 

. 1927. The growth of populations. The Quar- 
terly Review of Biology, 2:532-548. 

PEARSON, O. P. 1966. The prey of carnivores during 
one cycle of mouse abundance. The Journal of An- 
imal Ecology, 35:217-233. 

1971. Additional measurements of the im- 

pact of carnivores on California voles (Microtus cal- 

ifornicus). Journal of Mammalogy, 52:41-49. 

. 1985. Predation. Pp. 535-566, in Biology of 
New World Microtus (R. H. Tamarin, ed.). Special 
Publication, The American Society of Mammalo- 
gists, 8:1-893. 

PETERSON, R. O., ANDR.E. PAGE. 1983. Wolf-moose 
fluctuations at Isle Royale National Park, Michigan, 
U.S.A. Acta Zoologica Fennica, 174:251-253. 

PETRUSEWICZ, K. 1957. Investigation of experimen- 
tally induced population growth. Ekologia Polska, 
Seria A, 5:281-301. 

PITELKA, F. A., AND A. M. ScHuULTz. 1964. The nu- 
trient-recovery hypothesis for Arctic microtine cy- 
cles. Pp. 55-68 in Grazing in terrestrial and marine 
environments (D. J. Crisp, ed.). British Ecological 
Society Symposium, 4, Blackwell Scientific Publish- 
ers, Oxford, 322 pp. 

PLOWRIGHT, W. 1982. The effects of rinderpest and 
rinderpest control on wildlife in Africa. Symposia of 
the Zoological Society of London, 50:1-28. 

RiBB_eE, D. O., AND M. SALvIONI. 1990. Social or- 
ganization and nest co-occupancy in Peromyscus cal- 
ifornicus, a monogamous rodent. Behavioral Ecology 
and Sociobiology, 26:9-15. 

RosENZwEIG, M. L. 1966. Community structure in 
sympatric Carnivora. Journal of Mammalogy, 47: 
602-612. 

1973. Habitat selection experiments with a 


pair of coexisting heteromyid rodent species. Ecol- 
ogy, 54:111-117. 

Ross, J. 1982. Myxomatosis: the natural evolution 
of the disease. Symposia of the Zoological Society 
of London, 50:77-95. 

SALTHE, S. N. 1985. Evolving hierarchial systems: 
their structure and representation. Columbia Uni- 
versity Press, New York, 343 pp. 

SCHAFFER, W. M. 1974. Selection for optimal life 
histories: the effects of age structure. Ecology, 53: 
291-303. 

SCHNELL, G. D., AND R. K. SELANDER. 1981. Envi- 
ronmental and morphological correlates of genetic 
variation in mammals. Pp. 60-99, in Mammalian 
population genetics (M. H. Smith and J. Joule, eds.). 
University of Georgia Press, Athens, 380 pp. 

SCHRODER, G. D., AND M. L. ROSENzweIG. 1975. Per- 
turbation analysis of competition and overlap in 
habitat utilization between Dipodomys ordii and Di- 
podomys merriami. Oecologia, 29:9-28. 

SCHWERDTFEGER, F. 1941. Uber die Ursachen des 
Massenwechsels der Insekten. Zeitschrift fiir Ange- 
wandte Entomologie, 28:254—-303. 

Scott, M. E. 1988. The impact of infection and dis- 
ease on animal populations: implications for con- 
servation biology. Conservation Biology, 2:40-56. 

SEMPER, K. 1881. Animal life as affected by the nat- 
ural conditions of existence. Appleton, New York, 
472 pp. 

SETON, E.T. 1929. Lives of game animals. 4 volumes, 
Doubleday, Doran & Company, New York. 

SHELFORD, V.E. 1913. Animal communities in tem- 
perate America. University of Chicago Press, Chi- 
cago, 362 pp. 

. 1929. Laboratory and field ecology. Williams 
and Wilkins, Baltimore, 608 pp. 

SHERMAN, P. W., J. U. M. JARVIS, AND R. D. ALEXANDER 
(eps.). 1991. The biology of the naked mole-rat. 
Princeton University Press, Princeton, New Jersey, 
518 pp. 

SHvarts, S.S. 1969. Evolyutsionnaya ekologiya zhi- 
votnykh. Proceedings of the Institute of Plant and 
Animal Ecology, Sverdlovsk, 65:1-100. 

1977. The evolutionary ecology of animals 
(translated by A. E. Gill). Consultants Bureau, New 
York, 292 pp. 

SincLair, A. R. E. 1986. Testing multi-factor causes 
of population limitation: an illustration using snow- 
shoe hares. Oikos, 47:360-364. 

SLOBODCHIKOFF, C. H. (ED.). 1988. The ecology of 
social behavior. Academic Press, San Diego, Cali- 
fornia, 429 pp. 

SmitH, A. T., AND B. L. Ivens. 1984. Spatial rela- 
tionships and social organization in adult pikas: a 
facultatively monogamous mammal. Zeitschrift fiir 
Tierpsychologie, 66:37-47. 

SmituH, F. E. 1952. Experimental methods in popu- 
lation dynamics: a critique. Ecology, 33:441-450. 
SmitH, H. S. 1935. The role of biotic factors in the 
determination of population densities. Journal of 

Economic Entomology, 28:873-898. 

SmitH, M.H., AND J. JoULE(EDs.). 1981. Mammalian 
population genetics. University of Georgia Press, 
Athens, 380 pp. 

SmitH, M. H., M. N. MANLOVE, AND J. JouLE. 1975. 


346 LIDICKER 


Spatial and temporal dynamics of the genetic orga- 
nization of small mammal populations. Pp. 99-113, 
in Populations of small mammals under natural con- 
ditions (D. P. Snyder, ed.). Pymatuning Laboratory 
for Ecology, Special Publication, 5, 237 pp. 

Smuts, J.C. 1926. Holism and evolution. MacMillan 
Company, New York, 362 pp. 

SoLtomon, M.E. 1949. The natural control of animal 
populations. Journal of Animal Ecology, 18:1-35. 
1958. Meaning of density-dependence and 
related terms in population dynamics. Nature, 181: 

1778-1780. 

SoutHwick, C. H. 1955. The population dynamics 
of confined house mice supplied with unlimited food. 
Ecology, 36:212-225. 

STENSETH, N. C. 1978. Demographic strategies in 
fluctuating populations of small rodents. Oecologia, 
33:149-172. 

. 1981. On Chitty’s theory for fluctuating pop- 

ulations: the importance of genetic polymorphism 

in the generation of regular density cycles. Journal 

of Theoretical Biology, 90:9-36. 

1983. Causes and consequences of dispersal 

in small mammals. Pp. 63-101, in The ecology of 

animal movement (I. R. Swingland and P. J. Green- 
wood, eds.). Clarendon Press, Oxford, 311 pp. 

. 1986. On the interactions between stabilizing 
social factors and destabilizing trophic factors in small 
rodent populations. Theoretical Population Biology, 
29:365-385. 

STENSETH, N. C., AND W. Z. LIpICKER, JR. 1992a. The 
use of radioisotopes in the study of dispersal: with 
acase study. Pp. 333-352, in Animal dispersal: small 
mammals as a model (N. C. Stenseth and W. Z. 
Lidicker, Jr., eds.). Chapman and Hall, London, 365 


. 19926. Presaturation and saturation dispers- 
al 15 years later: some theoretical considerations. 
Pp. 201-223, in Animal dispersal: small mammals 
as a model (N. C. Stenseth and W. Z Lidicker, Jr., 
eds.). Chapman and Hall, London, 365 pp. 

(EDs.). 1992c. Animal dispersal: small mam- 
mals as a model. Chapman and Hall, London, 365 
pp. 

SyMINGTON, M. M. 1987. Sex ratio and maternal 
rank in wild spider monkeys: when daughters dis- 
perse. Behavioral Ecology and Sociobiology, 20:421- 
425. 

SZACKI, J., AND A. Liro. 1991. Movements of small 
mammals in the heterogeneous landscape. Land- 
scape Ecology, 5:219-224. 

TaitT, M.J., ANDC. J. Kress. 1983. Predation, cover 
and food manipulations during a spring decline of 
Microtus townsendii. Journal of Animal Ecology, 52: 
837-848. 

TAMARIN, R. H. 1978a. A defense of single-factor 
models of population regulation. Pp. 159-162, in 
Populations of small mammals under natural con- 
ditions (D. P. Snyder, ed.). Pymatuning Laboratory 
of Ecology, Special Publication, 5, 237 pp. 

. 19786. Dispersal, population regulation, and 
K-selection in field mice. The American Naturalist, 
112:545-555. 

TAMARIN, R. H., R. S. OSTFELD, S. R. PUGH, AND G. 
BUJALSKA (EDS.). 1990. Social systems and popu- 


lation cycles in voles. Birkhauser Verlag, Basel, 229 


TAYLOR, J. M., AND D. A. SCHLITTER. 1994. Award- 
ees. Pp. 71-109, in Seventy-five years of mammal- 
ogy (1919-1994) (E. C. Birney and J. R. Choate, 
eds.). Special Publication, The American Society of 
Mammalogists, 1 1:1-433. 

THIENEMANN, A. 1939. Grundziige einer allgemeinen 
Oekologie. Archives of Hydrobiology, 35:267—285. 

THompsoN, W. R. 1939. Biological control and the 
theories of the interactions of populations. Parasi- 
tology, 31:299-388. 

TRIPPENSEE, R. E. 1948. Wildlife management. Mc- 
Graw-Hill Book Company, New York, 479 pp. 

Trivers, R. L. 1971. The evolution of reciprocal al- 
truism. The Quarterly Review of Biology, 46:35—57. 

TuRNER, N. 1960. The effect of inbreeding and cross- 
breeding on numbers of insects. Annals of the En- 
tomological Society of America, 53:686-688. 

Uvarov, B. P. 1931. Insects and climate. Transac- 
tions of the Entomological Society of London, 79: 
1-247. 

VAN SCHAIK, C. P., AND S. B. Hrpy. 1991. Intensity 
of local mate competition shapes the relationship 
between maternal rank and sex ratios at birth in 
cercopithecine primates. The American Naturalist, 
138:1556-1562. 

VERME, L. H. J. 1969. Reproductive patterns of white- 
tailed deer related to nutritional plane. The Journal 
of Wildlife Management, 33:88 1-887. 

VOLTERRA, V. 1926. Fluctuations in the abundance 
of a species considered mathematically. Nature, 118: 
558-560. 

1931. Lecons sur la theorie mathematique 
que de la lutte pour la vie. Gauthiers-Villars, Paris, 
214 pp. 

Vrsa, E. S., AND N. ELDREDGE. 1984. Individuals, 
hierarchies and processes: towards a more complete 
evolutionary theory. Paleobiology, 10:146-171. 

WAGNER, F.H., ANDL.C.SToppART. 1972. Influence 
of coyote predation on black-tailed jackrabbit pop- 
ulations in Utah. The Journal of Wildlife Manage- 
ment, 36:329-342. 

Warp, A. J., D. THOMPSON, AND A. R. Hipy. 1987. 
Census techniques for grey seal populations. Sym- 
posia of the Zoological Society of London, 58:181- 
191. 

WEAVER, J. L. 1979. Wolf predation upon elk in the 
Rocky Mountain parks of North America: a review. 
Pp. 29-33, in North American elk: ecology, behavior 
and management (M. S. Boyce and L. D. Hayden- 
Wing, eds.). University of Wyoming Press, Laramie, 
294 pp. 

WEGNER, J., AND G. MERRIAM. 1990. Use of spatial 
elements in a farmland mosaic by a woodland ro- 
dent. Biological Conservation, 54:236-276. 

WELLINGTON, W. G. 1960. Qualitative changes in 
natural populations during changes in abundance. 
Canadian Journal of Zoology, 38:289-314. 

WiLBerT, H. 1963. K6nnen Insekten populationen 
durch Selektionprozesse reguliert werden? Ziet- 
schrift fiir Angewandte Entomologie, 52:185-204. 

Witson, E.O. 1975. Sociobiology, the new synthesis. 
Harvard University Press, Cambridge, 697 pp. 

Wot rr, J.O. 1980. The role of habitat patchiness in 


POPULATIONS 347 


the population dynamics of snowshoe hares. Eco- 1965. Self-regulating systems in populations 
logical Monographs, 50:111-130. of animals. Science, 147:1543-1548. 

Wynn_e-Epwarps, V. C. 1962. Animal dispersal in 1986. Evolution through group selection. 
relation to social behaviour. Hafner Publishing Blackwell Scientific Publishers, Oxford, 386 pp. 
Company, New York, 653 pp. 


COMMUNITY AND ECOSYSTEM ECOLOGY 


MICHAEL A. MARES AND Guy N. CAMERON 


Introduction 


biotic community is defined by Odum 
(1971:140) as “*. . . any assemblage of 
populations living in a prescribed area or 
physical habitat; it is an organized unit to 
the extent that it has characteristics addi- 
tional to its individual and population com- 
ponents.” Organisms forming a community 
interact in some manner with one another, 
whether through coevolutionary adapta- 
tions, as links in food chains, or any of in- 
numerable other potential biotic nexuses. 
Thus, a community may include all of the 
tree species in a particular forest, or all of 
the trees plus their associated plant and an- 
imal species, including detritus-feeding or- 
ganisms. Ecosystems, on the other hand, in- 
clude all of the organisms composing a 
community plus the abiotic components of 
the environment. Organization and inter- 
action among trophic levels, in addition to 
energy flow or nutrient cycling between the 
living and non-living parts of the system, 1s 
implied in this definition. 

While inclusion of several trophic levels 
within a single community is common, re- 
search on mammals seldom deals with an 
entire community. It is important to un- 
derstand these terms as they were classically 
employed because they are frequently mis- 
used. For example, Jaksic (1981) cited sev- 


348 


eral studies of mammals that ostensibly dealt 
with communities, but actually dealt either 
with a partial guild [e.g., a guild being (Root, 
1967:335) “a group of species that exploit 
the same class of environmental resources 
in a similar way ... without regard to tax- 
onomic positions’] or with simple taxo- 
nomic assemblages. An example of the for- 
mer might be the seed-eating rodents in a 
desert, which are a part of the granivore 
guild—the complete guild would include 
birds, ants, and other consumers of seeds. 
An example of the latter is research con- 
ducted on a “rodent community,” when in 
fact a study may have been done at the pop- 
ulation level—the community would in- 
clude all of the mammals and other organ- 
isms that interact in some important manner 
within a particular habitat or defined region 
(see, for example, the discussion of Slobod- 
kin, 1987). As May (1984:15) stated: °... 
any attempt to elucidate patterns of com- 
munity structure must deal with the ques- 
tion of how to delimit the community. Much 
academic research restricts itself to a par- 
ticular taxonomic group . . . instead of first 
consciously deciding which groups of spe- 
cies comprise a coherent and irreducible 
community.” In this context, however, it is 
important to emphasize that entire com- 


COMMUNITIES AND ECOSYSTEMS 349 


munities do not have to be studied in a 
community ecology study as long as inves- 
tigations are undertaken within a commu- 
nity-based framework. 

Our goal in this chapter is to examine how 
research on mammals has influenced, or has 
been influenced by, ideas of community and 
ecosystem organization. Mammals perform 
important functions at and above the com- 
munity level, whether through pathways of 
energy flow (e.g., mammals are trophically 
diverse and may be primary, secondary, or 
tertiary consumers), through widespread 
coevolutionary adaptations with plants and 
other organisms (e.g., pollination activities 
of tropical bats or dispersal of seeds by trop- 
ical rodents and ungulates), by affecting 
standing biomass and production, or by 
dramatic impacts on a particular habitat, 
such as elephants and ungulates in the Af- 
rican savanna community. The effects of 
mammals on each other and on other or- 
ganisms, as well as on the abiotic portions 
of the ecosystem, are extensive. A good deal 
of effort has been dedicated to understand- 
ing interactions at levels of biological or- 
ganization above the individual and the 
population. It is this area of investigation — 
research examining the place and the im- 
portance of mammals in biological com- 
munities and ecosystems—that will be re- 
viewed in this chapter. 

When the ASM was founded in 1919, in- 
formation on community and ecosystem re- 
lationships of mammals was negligible. Re- 
search at this time focused on questions 
dealing with individual and population 
ecology, championed by such giants as Jo- 
seph Grinnell; however, many of the guid- 
ing principles in community and ecosystem 
ecology were being formed (see below for 
work by Merriam, Shelford, and Elton) and 
were rapidly incorporated into studies deal- 
ing with mammals. 


Historical Overview 


Background research on communities and 
ecosystems.—Odum (1971:Chapters 1-2) 


and Kendeigh (1974) reviewed the history 
of the conceptualization of the terms “‘eco- 
system” and “community,” and McIntosh 
(1985) provided an overview of the history 
of ecology. The idea that plants and animals 
occur together in some type of non-random 
pattern is quite old. Kendeigh (1974), for 
example, mentioned a reference to species 
assemblages by Theophrastus at the time of 
Aristotle in the 4th century BC (see McIn- 
tosh, 1985; Ramalay, 1940). As early as 
1807, Humboldt and Bonpland referred to 
plant associations which could be identified 
by physiognomy and which were related to 
both latitudinal and vertical zonation. In 
1815, Humboldt devised a grid system for 
recording presence or absence of plant spe- 
cies between different landscapes (MclIn- 
tosh, 1985). The German botanist, A. 
Grisebach, in 1838, described animals and 
plants occurring together in interrelated as- 
sociations. Seventy years after Humboldt’s 
ground-breaking work, in 1877, another 
German, Karl Md6bius, discussed oyster 
communities on a coral reef; M6bius used 
the term biocoenosis, which subsequently 
became the European term for biotic com- 
munities. When Mobius’ work was trans- 
lated into English in 1883, biocoenosis be- 
came community (e.g., Allee et al., 1949). 
C. SchrGter, a Swiss botanist, working 1n the 
late 1800s and early 1900s, was one of the 
first biologists to use the concept of plant 
community consistently for describing veg- 
etation (Gigon et al., 1981). 

S. A. Forbes (1887) also used the term 
community in his classic work on lake ecol- 
ogy, and it became the term generally used 
in North America to describe interrelated 
biotic associations. Forbes, who has been 
called the complete ecologist (e.g., McIn- 
tosh, 1985), was curator of the Illinois Nat- 
ural History Museum and director of the 
State Laboratory of Natural History (=Il- 
linois Biological Survey). As McIntosh 
(1985) noted, Forbes’ influence on ecology 
was enormous, with his 1887 paper found- 
ing the science of limnology and other pa- 
pers anticipating such modern ecological 
concepts as competitive exclusion. Cur- 


350 MARES AND CAMERON 


ously, competitive exclusion was first more 
specifically defined, if in a qualitative man- 
ner, by two mammalogists, Joseph Grinnell 
(1904, 1908), the father of academic mam- 
malogy in North America (Jones, 1991), a 
charter member of ASM and president of 
the society in 1937, and Angel Cabrera 
(1932), a Spaniard, who was named an hon- 
orary member of the ASM (see Hutchinson, 
1978). 

One of the first animal ecologists, Victor 
Shelford, wrote that ecology was the science 
of communities (Shelford, 1913). Com- 
munity theory primarily developed by plant 
ecologists (e.g., A. G. Tansley, F. E. Clem- 
ents, H. C. Cowles) in the early part of this 
century initially was exemplified by the or- 
ganismic dynamic theory of Clements that 
predicted a stable, climax stage. This early 
view, widely accepted by plant ecologists 
and more or less by animal ecologists, was 
challenged from the 1930s to the 1950s by 
plant ecologists espousing an individua- 
listic theory (Gleason, 1917, 1939; McIn- 
tosh, 1975, 1980; Whittaker, 1951). Where- 
as these studies challenged the idea of the 
plant community, animal ecologists adopt- 
ed the concept of the community as an en- 
tity composed of species at equilibrium. 
Such an idea, associated with the work of 
Robert MacArthur, derived largely from the 
belief that many patterns in nature were a 
consequence of competition to promote 
niche separation (Cody and Diamond, 1975; 
Connell, 1980; Diamond, 1978). Mammal- 
ogists contributed substantially to uncov- 
ering the role of competition in structuring 
natural communities (see below). However, 
another mammalogist (Brown, 1981) ar- 
gued that theoretical population ecology 
largely failed to produce a quantitative the- 
ory applicable to community ecology. The 
largest oversight, he argued, was a failure to 
emphasize energy flow as a coalescing pat- 
tern (see also Hall et al., 1992). 

The idea of the ecosystem is more recent 
than that of the community. A botanist, A. 
G. Tansley (1935), in a review of botanical 
concepts, coined the term ecosystem, which 
expanded the concept of the biotic com- 


munity to include the interactions of the 
organisms comprising the community with 
the abiotic parts of the environment. The 
term biocoenosis was enlarged to geobio- 
coenosis by a Russian, V. N. Sukachev 
(Odum, 1971; see Sukachev, 1958), thus be- 
coming the equivalent of ecosystem. Al- 
though the terminology used in the New and 
Old World differed, and different underly- 
ing ecological philosophies influenced re- 
search within these two regions (e.g., Gigon 
et al., 1981), there was a general apprecia- 
tion of supra-individual and supra-popu- 
lation effects in ecology, especially in con- 
tributing to community stability and system 
cohesiveness. 

Inherent in the work of some ecologists 
was the idea that communities, and later, 
ecosystems, were superorganisms, respond- 
ing as unified units to experimental and evo- 
jutionary perturbations (e.g., Clements, 
1905, 1916; Semper, 1881). Tansley (1935), 
however, argued strongly that neither the 
community nor the ecosystem should be 
viewed as some type of superorganism. The 
concept of the ecosystem as a super entity 
largely has been discounted by most ecol- 
ogists. However, the idea has arisen again 
in recent years under the guise of a bio- 
spheric entity called Gaia (see Barlow, 1991). 
This mystical super life form is almost sen- 
tiently responsive to deviations from “‘nor- 
mal’? environmental parameters that are 
conducive to maintaining the life to which 
it (Gaia) 1s presently adapted. 

Mammalogists, communities, and eco- 
systems. —Despite the long history of Eu- 
ropean botanists and invertebrate biologists 
who developed community-based studies, 
a number of North American biologists, who 
also conducted important research on 
mammals, were intimately involved with 
the foundations of community and ecosys- 
tem ecology. As early as the late nineteenth 
century, C. Hart Merriam, the father of 
modern mammalogy (e.g., Osgood, 1943; 
Sterling, 1977), was the first North Ameri- 
can to develop research interests relating to 
communities and ecosystems. Merriam de- 
veloped the team method of conducting sur- 


COMMUNITIES AND ECOSYSTEMS eeu 


vey research in particular regions. This in- 
volved sending groups of researchers into 
the field to study botany, geology, and most 
aspects of vertebrate biology (systematics, 
distribution, natural history, and ecology of 
both birds and mammals), either for specific 
localities or for broader regions (e.g., Mer- 
riam, 1890, 1892, 1894, 1898). 

Merriam was among the earliest propo- 
nents in North America of a unified view 
of natural communities. The biological sur- 
veys that were conducted in a broad-based 
manner across taxa, and that included ex- 
tensive geological investigations and data 
on climate, amassed a great deal of infor- 
mation on how the biota of a region reflect- 
ed abiotic factors in the environment. In 
examining such data for the San Francisco 
Mountains region of northern Arizona, 
Merriam formulated the concept of life zones 
(Merriam, 1894, 1898). The life zone con- 
cept was the first attempt to include dom- 
inant animals in a community classification 
scheme. This concept warrants additional 
discussion because it led to early consider- 
ation of the interactions among taxonomi- 
cally diverse organisms (i.e., community in- 
teractions) and with their abiotic 
environment (i.e., ecosystems). 

Merriam attempted to explain the distri- 
bution of animals in relation to life zones 
that were themselves defined by tempera- 
ture laws that he formulated. The resultant 
zones formed altitudinal and _ latitudinal 
bands that stretched across the North 
American continent. The life zone concept 
worked effectively in the mountainous areas 
of the western United States where it was 
derived, partially because the temperature 
limits defining the faunal zones coincided 
with vegetation regions. There was a good 
deal of criticism of Merriam’s life zones (see 
Odum, 1945, for a review), and the sugges- 
tion that there were definable life zones was 
replaced by the biome concept (see below) 
which is still widely used today. 

Merriam’s revolutionary techniques of 
field research and broadly based field sur- 
veys assisted in the development of a ho- 
listic view of entire biotas as organized and 


interrelated units responding to abiotic in- 
fluences. Subsequently, several other mam- 
malogists helped lay the foundations of 
modern community and ecosystem ecology. 
Charles C. Adams, for example, who ini- 
tially worked for S. A. Forbes in the Illinois 
Natural History Survey, published some of 
the earliest work in community ecology 
when he described a number of animal com- 
munities while conducting a biological sur- 
vey of Michigan (Adams, 1905, 1909; see 
Kendeigh, 1974; and McIntosh, 1985). Ad- 
ams was a charter member of ASM, was 
nominated by president Merriam to chair 
the ASM Committee on Life Histories of 
Mammals (Hollister, 1920), and published 
the first manual on animal ecology (Adams, 
1915). 

Another landmark in the development of 
community ecology was also produced by a 
mammalian ecologist in North America. 
The first book ever published on animal or 
plant communities was by Victor Shelford 
(1913), who was the first president of the 
Ecological Society of America (in 1915) and 
who joined ASM in 1923. Some of his work 
had a physiological orientation and led to 
initial ideas about how environmental ex- 
tremes limited species (and community) 
ranges. Although he did not formalize the 
concept, Shelford’s work outlined food 
chains and made initial conceptual linkages 
between communities and ecosystems, de- 
scribing them as dynamic units responding 
to changing environmental parameters. 
[Ideas concerning food chains and the con- 
cept of the pyramid of numbers were first 
set forth by K. Semper, a North American 
zoologist, who published a book on animals 
and their relationship to their natural en- 
vironments (Semper, 1881, see McIntosh, 
1985). Semper’s work was an early zoology 
text that applied Darwin’s ideas of natural 
selection to a wide array of organisms and 
included discussion of such topics as cryp- 
sis, warning coloration, and competition be- 
tween similar species.] Shelford realized the 
importance of biological surveys (e.g., Shel- 
ford, 1926) and conducted detailed research 
on lemming populations (e.g., Shelford and 


jDZ MARES AND CAMERON 


Twomey, 1941). Shelford’s landmark work 
was the development of the biome concept 
in conjunction with the plant ecologist, F. 
Clements (Clements and Shelford, 1939). 
Shortly after these contributions of North 
American ecologists were published, semi- 
nal research on how organisms functioned 
was conducted in Great Britain. Perhaps the 
preeminent work contributing to the de- 
velopment of community and ecosystem 
theory (and to the development of ecology 
in general) was that of the mammalian ecol- 
ogist, Charles Elton (Elton joined the ASM 
in 1931), who formulated or developed in 
detail four important ecological concepts: 
the niche; differences in food particle size 
as a mechanism to reduce competition; the 
food web; and the pyramid of numbers (Duff 
and Lowe, 1981; Elton, 1927, 1933). These 
ideas became paradigms of ecological the- 
ory and contributed greatly to an appreci- 
ation of the functional relationships of or- 
ganisms in communities and ecosystems. 
Elton’s work, which built directly upon 
the research of Adams and Shelford, was 
fundamental to understanding the com- 
plexities of nature. With the pyramid of 
numbers, Elton showed that there was a 
structure to nature—organisms in a com- 
munity were not randomly organized so far 
as their abundance was concerned; rather, 
different trophic levels showed specific nu- 
merical relationships to one another (e.g., 
herbivores were more abundant than car- 
nivores). Similarly, pyramids of biomass and 
energy illustrated non-random organiza- 
tions with both biomass and energy content 
decreasing in a pyramidal fashion toward 
higher trophic levels. Even though we now 
know that only the pyramid of energy can- 
not be inverted, these descriptions of nat- 
ural communities were pivotal to the de- 
velopment of the modern underpinnings of 
ecosystem research. With the description of 
food chains and webs, Elton clearly showed 
how energy linked component species in an 
ecosystem in often unexpectedly complex 
pathways. This was a profound description 
of nature that continues to impact current 
ideas of community structure (e.g., Pimm 


et al., 1991). Elton was also responsible for 
quantitative research on mammal popula- 
tion ecology, particularly with his bench- 
mark publication on 10-year population cy- 
cles of the lynx (Elton and Nicholson, 1942), 
his classic book on population ecology of 
mice, lemmings, and voles (Elton, 1942), 
and other contributions (e.g., Elton, 1958, 
1966). 

Although the original concept of niche 
was not necessarily associated with com- 
munity studies, it has had an important im- 
pact on modern ecological theory (e.g., Ehr- 
lich and Roughgarden, 1987). It is worth 
noting that Grinnell (1914, 1917a, 19175) 
was among the earliest individuals to de- 
velop the idea of the niche. Indeed, until 
Gaffney (1973) reviewed the history of the 
niche concept and found that it was coined 
by Robert Johnson in 1910, the origin of 
the term had been attributed to Grinnell 
(Cox, 1980). 

Clearly, Adams, Shelford, Grinnell, and 
Elton utilized their ecological expertise, es- 
pecially that developed from working on 
mammals, to influence the foundations of 
ecology, particularly at the higher levels of 
biological organization. By the early 20th 
Century, mammalogists were among the 
leading ecologists in conducting studies and 
developing theories bearing on the devel- 
opment of community and ecosystem ecol- 
ogy. Their work, along with the burgeoning 
disciplines of limnology and plant com- 
munity ecology, helped drive the field into 
the modern age. Mammalogists have con- 
tinued to play a role in the development of 
community and ecosystem studies, not only 
in the field and the laboratory but, at least 
in the case of modern ecosystem research, 
in the biopolitical arena as well. 


Approaches to Community and 
Ecosystem Ecology 


Early studies in mammalian ecology mir- 
rored the natural history approach exem- 
plified by Grinnell’s work. This descriptive 
approach was reflected in biotic surveys that 


COMMUNITIES AND ECOSYSTEMS 353 


encompassed a variety of techniques to 
sample both plants and animals through the 
1940s in the United States (i.e., Fautin, 
1946). The 1940s and 1950s were a period 
during which studies were designed to de- 
scribe community processes, in particular 
trophic dynamics and energy flow (Linde- 
man, 1942; Odum, 1957; Teal, 1957). Ini- 
tial emphasis was on aquatic habitats, but 
subsequent studies in terrestrial ecosystems 
included small mammals as major consum- 
ers (e.g., Golley, 1960). 

The International Biological Program 
(1969-1974; IBP) was an important factor 
in the development of community and eco- 
system ecology because it bridged the earlier 
descriptive approach and the current em- 
phasis on empiricism. One thrust of IBP 
was to organize groups of specialists to study 
major terrestrial biomes and to integrate the 
findings with models used as predictive 
tools. This international effort at under- 
standing the structure and function of eco- 
systems on a global scale was in large part 
developed and administered by another 
mammalogist, W. Frank Blair. Many mam- 
malogists active today participated in IBP 
(IBP will be discussed in detail below). 

One of the criticisms about IBP was the 
lack of hypothesis testing. Ecological studies 
since the mid-1970s have become increas- 
ingly grounded in the scientific method, thus 
completing the transition from the descrip- 
tive approach that was begun at the turn of 
the century. To facilitate experimental stud- 
ies at appropriate ecological scales (both 
spatial and temporal), a variety of ecological 
research areas have been established, in- 
cluding Biosphere Reserves, Experimental 
Ecological Reserves, and Long-term Ex- 
perimental Research areas (Franklin et al., 
1990). Ecological experiments are conduct- 
ed in the laboratory and field, use natural 
or experimentally controlled perturbations, 
and consider factors that influence organ- 
isms over the short- or long-term (Dia- 
mond, 1986). Mammalogists have been at 
the forefront of development of empirical 
studies conducted in the field (see citations 
below) and have argued for the develop- 


ment of facilities where long-term experi- 
mental research could be undertaken. 
Mammalogists also have argued that nat- 
ural history should continue to play a crit- 
ical role in empirical studies by providing 
the crucial knowledge to design appropriate 
experiments (Bartholomew, 1986; Brown, 
1986; Mares and Braun, 1986). Finally, 
mammalogists have played a role in devel- 
oping methods to conduct and analyze field 
experiments, such as taking into account the 
effect of scale, both spatial (J. S. Brown, 
1989: Morris, 1987, 1989; Price and Kra- 
mer, 1984) and temporal (Brown and Heske, 
1990; Brown and Kurzius, 1989). 


Community Ecology 


The concept of niche. —The development 
of the concept of the niche began with sev- 
eral mammalogists. Joseph Grinnell wrote 
that ‘“‘As with zones and faunas, associa- 
tions are often capable of subdivision; in 
fact such splitting may be carried logically 
to the point where but one species occupies 
each its own niche” (Grinnell and Swarth, 
1913:218), and “A concurrent axiom 1s that 
if associational analysis 1s carried far enough, 
no two species of birds or mammals will be 
found to occupy precisely the same ecologic 
niche, although they may apparently do so 
where their respective associations are rep- 
resented fragmentarily and in intermixture” 
(Grinnell, 1914:91). Grinnell defined the 
niche as “the concept of the ultimate dis- 
tributional unit, within which each species 
is held by its structural and instinctive 
limitations...” (Grinnell, 1928/1943:192- 
194). This view of the niche as a distribu- 
tional entity was complemented by Charles 
Elton’s (1927:64) idea that the ‘“‘niche of an 
animal means its place in the biotic envi- 
ronment, its relations to food and ene- 
mies’’—the so-called functional niche. Dice 
(1952) suggested that the niche represented 
a coalescence of both functional and distri- 
butional attributes of a species. 

The current concept of the niche was for- 
malized mathematically as an n-dimen- 


354 MARES AND CAMERON 


sional hyperspace by an aquatic biologist, 
G. Evelyn Hutchinson (1957). Mammalo- 
gists have contributed to refining the niche 
concept. For example, MacMahon et al. 
(1981) discussed how the niche reflects the 
actual or potential state of an organism at 
an instant in time. They concluded that an 
organism’s niche is bounded by tolerance 
limits set by heredity, maturity, and accli- 
matization, and that changes in tolerances 
during an organism’s life cycle create on- 
togenetic bottlenecks in the niche. 
Mammalogists have contributed to our 
knowledge of the niche concept with re- 
search measuring niche parameters (Carnes 
and Slade, 1982; Churchfield, 1991; Dueser 
and Shugart, 1979, 1982; Montgomery, 
1989: Slobodchikoff and Schultz, 1980; 
Smartt, 1978; Van Horne and Ford, 1982). 
In addition, mammalogists have conducted 
empirical studies that illustrated increases 
in niche breadth with intraspecific compe- 
tition (Smartt and Lemen, 1980; Van Horne 
and Ford, 1982), variation in genetic and 
morphological measurements with niche 
breadth (i.e., the niche variation hypothesis; 
Smith, 1981), a correlation of niche breadth 
with species abundance (Brown, 1984; Sea- 
gle and McCraken, 1986), body size (Bar- 
clay and Brigham, 1991; Willig and Moul- 
ton, 1989), and partitioning of resources 
(Brown, 1973, 1975; Cameron, 1971; Em- 
mons, 1980; Mares and Williams, 1977; 
McKenzie and Start, 1989; M’Closkey, 
1980; Meserve, 1981; Owen-Smith, 1989; 
Price et al., 1991; Willig et al., 1993). 
Interspecific interactions. —In addition to 
the niche concept, mammalogists have con- 
tributed substantially to another basic con- 
cept of community and ecosystem ecology, 
that of interspecific interactions, including 
competition, predation, and mutualism. 
Again, Joseph Grinnell laid the framework 
for this concept when he wrote “‘these var- 
ious circumstances, which emphasize de- 
pendence upon cover, and adaptation in 
physical structure and temperament there- 
to, go to demonstrate the nature of the ul- 
timate associational niche occupied by the 
California thrasher. ... It is, of course, ax- 


iomatic that no two species regularly estab- 
lished in a singie fauna have precisely the 
same niche relationships” (Grinnell, 1917a: 
433), and that “‘no two species in the same 
general territory can occupy for long iden- 
tically the same ecological niche ... com- 
petitive displacement of one of the species 
by the other is bound to take place” (Grin- 
nell, 1928/1943:192-194). The great Span- 
ish mammalogist, A. Cabrera, who spent 
most of his professional life in Argentina 
and was the preeminent force in the history 
of South American mammalogy, also pub- 
lished an important paper on competitive 
exclusion that described the concept as a 
biological law (Cabrera, 1932). 

Interspecific competition was first de- 
scribed mathematically by Lotka and Vol- 
terra (see Slobodkin, 1961). Over the years, 
mammalogists have contributed to the 
modification of these models to overcome 
some of the limiting assumptions (Fryxell 
et al., 1991). Mammalogists have also de- 
vised statistical methods to measure com- 
petition in the field (Hallett and Pimm, 1970; 
Rosenzweig et al., 1984). Other mammal- 
ogists were instrumental in beginning the 
classification of this process into what is now 
known as interference and exploitation 
competition (Elton and Miller, 1954; Mil- 
ler, 1967) and in describing the relative im- 
portance of these processes (King and 
Moors, 1979). Mammalogists have com- 
pleted numerous other studies on the pro- 
cess of interspecific competition (e.g., 
Brown, 1971; Brown et al., 1979; Dickman, 
1989: Fox, 1989; Holbrook, 1979; Kirk- 
land, 1991; Pulliam and Brand, 1975; Ro- 
senzweig, 1966; Smith and Balda, 1979; 
Willig and Moulton, 1989; see below for 
role of competition in community struc- 
ture), but data gathered across entire mam- 
mal faunas to clarify competitive or other 
mechanisms that are important in structur- 
ing temperate and tropical faunas are still 
rudimentary (Lacher and Mares, 1986; Wil- 
lig, 1986). 

The niche overlap hypothesis states that 
maximum tolerable niche overlap decreases 
as the intensity of competition increases 


COMMUNITIES AND ECOSYSTEMS bey) 


(Pianka, 1974). Studies on several mam- 
malian systems offer support for this hy- 
pothesis (Fox, 1981; Lacher and Alho, 1989; 
M’Closkey, 1978; Porter and Dueser, 1981; 
but the multivariate technique used by Por- 
ter and Dueser has been questioned by 
Carnes and Slade, 1982). However, Brown 
(1975) found that niche overlap increased 
when number of species increased for North 
American desert rodents. He attributed this 
response to the fact that the Mohave desert 
communities he studied may be composed 
of more generalist species than those ex- 
amined in the other studies. 

A second interspecific interaction to which 
mammalogists have contributed is the pro- 
cess of predation. As with competition, ba- 
sic models for this process were developed 
by Lotka and Volterra. Mammalogists were 
instrumental in refining these models (Ro- 
senzweilg, 1969, 1973; Rosenzweig and 
MacArthur, 1963). Much of the subsequent 
development of this aspect of community 
ecology relied on studies of mammals; for 
example, functional and numerical re- 
sponses were described with responses be- 
tween Sorex, Blarina, and Peromyscus and 
their sawfly larva prey (Holling, 1959), and 
differences in susceptibility of age groups to 
predation were described in the moose-wolf 
system (Mech, 1966). 

Mammalogists have conducted many 
studies on the basic nature of predator-prey 
relations (e.g., Hornocker, 1970; Pearson, 
1971; Schnell, 1968; Wagner and Stoddart, 
1972). Two views on the role of predators 
arose earlier in this century. One, champi- 
oned by the mammalogist Paul Errington 
(1946), held that predators only took sur- 
plus prey above the carrying capacity, a view 
without current support. The other view 
arose in the entomological literature and 
concluded that predators regulated their 
prey. Demonstration of this phenomenon 
has been elusive largely because of the myr- 
iad of definitions given to this process (Var- 
ley, 1975); population regulation, however, 
is a density-dependent feedback of either 
increasing mortality or decreasing fecundity 
proportional with increasing predation. Er- 


linge and his colleagues (Erlinge et al., 1983, 
1984) analyzed population density of field 
voles and rabbits, as well as food habits of 
their major avian and mammalian preda- 
tors, in Sweden. They recorded both func- 
tional and numerical responses by predators 
to changes in prey numbers and concluded 
that the functional response, combined with 
switching by predators from voles to rabbits 
and vice versa when numbers of prey be- 
came low, produced a density-dependent ef- 
fect during the period of highest vole density 
(autumn). These findings were challenged 
by Kidd and Lewis (1987), who argued that 
Erlinge and his colleagues had not demon- 
strated density-dependent predation; Er- 
linge et al. (1988) responded that predator 
switching among alternative prey affected 
regulation. Korpimaki (1993), however, 
presented evidence that Microtus sp. in Fin- 
land are regulated by density-dependent 
avian predation and delayed density-depen- 
dent mammalian predation. Sinclair et al. 
(1990) concluded that house mice in Aus- 
tralia were regulated by delayed density-de- 
pendent predation at low-moderate mouse 
densities, but by inverse density-depen- 
dence at high mouse densities. Trostel et al. 
(1987) found that avian and mammalian 
predators may affect the 10-year cycle of 
snowshoe hares in a delayed density-depen- 
dent fashion. 

Mutualism has been studied much less 
intensively than either competition or pre- 
dation, but research on mammals has again 
provided perspectives on the mechanics and 
pervasiveness of this process. Mutualism can 
be a direct or indirect process. Mammal- 
plant interactions, such as seed dispersal 
(Carpenter, 1978; Sazima and Sazima, 1978; 
Simpson and Neff, 1981; Sussman and Ra- 
ven, 1978) or pollination (Fleming, 1981; 
Howe, 1980; Smith, 1970; Stapanian and 
Smith, 1978) are direct processes. In indi- 
rect mutualism, a positive interaction is 
achieved even though there is no direct con- 
tact between the species. For example, al- 
though Thompson gazelles, zebras, and wil- 
debeests eat different foods on the Serengeti, 
the gazelles prefer to feed in areas where 


356 MARES AND CAMERON 


wildebeests have grazed a month earlier, 
since such areas contain greater plant bio- 
mass (McNaughton, 1976). Brown et al. 
(1986), building upon an evolutionary hy- 
pothesis developed by Mares and Rosen- 
zwelg (1978), demonstrated that rodents in 
the Mohave desert eat large seeds, whereas 
ants prefer smaller seeds. When rodents were 
removed, large-seeded plants increased in 
abundance, reduced the abundance of small- 
seeded plants and, consequently, the small 
seed resources of ants. Thus, rodents acted 
as indirect mutualists on ants (Davidson et 
al., 1984). 

Other examples of indirect mutualism in- 
clude the observation that the progress of 
plant succession may be positively affected 
when pocket gophers alter soil characteris- 
tics and thereby affect the resultant plant 
species composition (Andersen and Mac- 
Mahon, 1985; Huntly and Inouye, 1988; 
Tilman, 1983). In a similar fashion, food 
availability for granivorous birds is affected 
positively by desert rodents that forage pref- 
erentially upon those plant species that 
compete with plant species eaten by the birds 
and that maintain areas of bare soil which 
serve as germination sites for those plants 
eaten by the birds (Mitchell et al., 1990). 
Coppock et al. (19835) also discovered that 
bison preferentially grazed in areas where 
prairie dogs had reduced the occurrence of 
less preferred plants, thereby allowing 
growth of more preferred plants. Finally, 
dispersal of seeds from parent plants re- 
duced seed predation by desert rodents and 
thereby enhanced seed germination 
(O’Dowd and Hay, 1980). 

Community structure. —Structure within 
a community is determined by both com- 
position and relative abundance of species. 
Many studies have addressed Elton’s (1927) 
concept of limited membership: Why is it 
that what does occur together constitutes a 
limited subset of what might occur together? 
One avenue of research has been to inves- 
tigate whether structure exists for subsets of 
a community [i.e., within community struc- 
ture, termed guild structure by Root (1967) 


to refer to groups of species exploiting re- 
sources in a similar way; the multiple mean- 
ings of guild, however, have been discussed 
by Hawkins and MacMahon (1989) and 
Simberloff and Dayan (1991)]. Most of this 
work has centered on insects and lower ver- 
tebrates. In one of the few studies with 
mammals, MacMahon (1976) concluded 
that similarities in guild structure of small 
mammals among sites in the deserts of the 
western United States resulted from inter- 
actions of evolutionary events and site char- 
acteristics. Fox (1989) and others (e.g., 
Findley, 1989; Humphrey et al., 1983; 
McKenzie and Start, 1989; Rosenzweig, 
1989; Smythe, 1986; Willig and Moulton, 
1989) have also examined the mechanisms 
affecting community (or guild) assembly in 
mammals. Fox (1989) used a taxonomical- 
ly-based rule for species assembly of small 
mammals in Australian heathlands that 
stipulated there was a higher probability that 
species comprising a community will have 
been drawn from a genus, guild, or taxo- 
nomically-related group of species with 
similar diets. Fox and Brown (1993) applied 
an assembly rule based upon functional 
groups to suggest that interspecific compe- 
tition was an important mechanism struc- 
turing desert rodent communities in North 
America. Willig and Moulton (1989), on the 
other hand, found that ecomorphology in 
bat communities was not different from that 
expected by a stochastic model; Willig et al. 
(1993) reported that dietary differences 
among Brazilian bats did not order com- 
munity structure, but suggested that com- 
petition for some other resource could be 
more important. 

Other research has centered on the role 
of competition in determining community 
structure. This research can be divided into 
observational and empirical evidence. Here 
again, mammalogists have played promi- 
nent roles. Several sorts of observational ev- 
idence have been used to conclude that in- 
terspecific competition has been important 
in determining community structure. Re- 
source partitioning, comparative species 


COMMUNITIES AND ECOSYSTEMS Sey) 


distributions, and character displacement 
will be considered. 

Resource partitioning, the subdivision of 
resources by two or more species, is one 
outcome of the Lotka-Volterra model of 
competition, whereby niche dimensions of 
competing species are modified such that 
niche overlap decreases (see reviews by 
Schoener, 1974, 1986a). Numerous studies 
have demonstrated resource partitioning in 
mammals (e.g., Belk et al., 1989; Brown, 
1989; Dueser and Hallett, 1980; Fleming et 
al., 1972; Hallett et al., 1983; Heithaus et 
al., 1975; McNab, 1971; McNaughton and 
Georgiadis, 1986; Meserve, 1981). 

The negative correlation between spatial 
distributions of species is another way that 
the effect of competition on community 
structure has been inferred. There are many 
examples of this effect from the literature 
on mammals. For example, mammalogists 
have noted such spatial partitioning be- 
tween Sigmodon hispidus, S. fulviventer, and 
S. ochrognathus in Durango, Mexico (Pe- 
tersen, 1970, 1973); Sigmodon leucotis and 
Microtus mexicanus in Durango, Mexico 
(Baker, 1969); among seven species of Mi- 
crotus in western North America (Ander- 
son, 1959); and among desert rodents in 
the southwestern United States (Whitford 
and Steinberger, 1989). Similarly, the 
northward withdrawal of Microtus coinci- 
dent with a gradual northward advance of 
S. hispidus is also viewed as an indication 
of competition (Baker, 1969). Other mam- 
malogists have devised methods of detect- 
ing the effects of competition by analysis of 
captures at trap stations (Hallett and Pimm, 
1970; Rosenzweig et al., 1984). 

Character displacement is the change un- 
der natural selection of morphological, 
physiological, or behavioral characteristics 
in one or more ecologically similar species 
whose ranges overlap in sympatry. Such 
evolved differences reduce competition. 
Malmquist (1985) demonstrated that Sorex 
minutus had significantly smaller jaws when 
it occurred in sympatry with S. araneus 
(Sweden) than when it occurred allopatri- 


cally (Ireland). Similarly, Dayan et al. (1989, 
1990) analyzed cranial characteristics of 
weasels in North America and Israel, and 
feline carnivores in Israel, and concluded 
that past competition for food led to pres- 
ent-day cranial differences. 

Although these studies suggest an impor- 
tant role for competition in community 
structure, they are not conclusive. Empirical 
evidence demonstrating a change in niche 
breadth in response to a change in abun- 
dance of a potential competitor is necessary. 
Such evidence can be gathered from natural 
experiments or from perturbation experi- 
ments. Natural experiments involve com- 
paring an area where a species 1s allopatric 
with a similar area where it occurs sympat- 
rically with a potential competitor; differ- 
ences in niche dimensions between the two 
areas are taken to indicate the effect of com- 
petition. For example, Glass and Slade 
(1980) reported that when S. hispidus de- 
clined locally in Kansas, Microtus ochro- 
gaster expanded its spacial use of habitats; 
there was spatial separation when both spe- 
cies were present. The greatest problem with 
such natural experiments is that the sites 
compared may differ in ways other than the 
presence or absence of the species under 
consideration. 

A perturbation experiment is arguably the 
best way to demonstrate whether compe- 
tition affects community structure. This type 
of experiment, where one species is re- 
moved or reduced in density by the inves- 
tigator, and the effect upon the remaining 
species 1s documented, avoids problems of 
possible differences between study sites. 
Such field experiments have demonstrated 
that interspecific competition affects com- 
munity structure in a wide variety of sys- 
tems (Busch and Kravetz, 1992; Connell, 
1983; Schoener, 1983, 1985; Underwood, 
1986). The inclusion in these general re- 
views of certain field experiments on mam- 
mals in which experimental flaws had been 
detected were criticized (i.e., enclosures 
smaller than home ranges; Galindo and 
Krebs, 1986; Schoener, 19866). However, 


358 MARES AND CAMERON 


Dueser et al. (1989) reaffirmed the role of 
competition in structuring rodent commu- 
nities. Details of these effects can be found 
in the numerous studies cited in the above 
reviews, such as Grant (1972), Crowell and 
Pimm (1976), and Dickman (1988). 

One of the major criticisms to the con- 
clusion that competition affects community 
structure was that many empirical studies 
were biased and that null models (1.e., mod- 
els assuming no biological effects) could ex- 
plain observed patterns of community 
structure (see Harvey et al., 1983; Strong et 
al., 1984). Community patterns of neotrop- 
ical bats seem to be affected by factors other 
than simple competitive interactions (e.g., 
Willig and Mares, 1989). While problems 
certainly existed with empirical studies, 
analyses and reanalyses of data with null 
models have reconfirmed the importance of 
competition in general, and among mam- 
mals in particular, in structuring some com- 
munities (Bowers and Brown, 1982; Brown 
and Bowers, 1984; Dayan et al., 1990; Find- 
ley, 1989). However, Owen-Smith (1989), 
studying African ungulates in savanna 
grasslands, concluded that competition had 
little effect on community structure. Simi- 
larly, Findley (1993), in a comprehensive 
analysis of data on bat communities from 
throughout the world, concluded that com- 
petitive interactions had little or no part in 
structuring the communities; rather, their 
structure had a great deal to do with sto- 
chastic processes. 

Predation also has been shown to be an 
important determinant of community 
structure (Sih et al., 1985). Removal of sea 
otters from nearshore communities along 
the coast of the western United States in- 
creased abundance of a major prey item (sea 
urchins). Abundant sea urchins decimated 
nearshore kelp communities, both in terms 
of abundance and diversity; simplification 
of the kelp community caused loss of many 
associated marine organisms. Thus, the sea 
otter can be classified as a keystone species 
in this system (Duggins, 1980; Estes and 
Palmisano, 1974; Estes et al., 1978; Simen- 


stad et al., 1978). Similarly, Brown and 
Heske (1990) classified a guild of three spe- 
cies of kangaroo rats in the Mohave Desert 
as keystone species because their removal 
decreased the abundance of bare areas (ger- 
mination sites for plants), changed the spe- 
cies composition of the plants, and favored 
invasion of the desert area by grassland spe- 
cies of mammals. Such effects were noted 
also in areas where species were introduced. 
Case and Bolger (1991) observed that in- 
troduction of mongoose, domestic dogs and 
cats, and Rattus on islands in various parts 
of the world constrains the distribution, col- 
onization, and abundance of reptiles. Pred- 
ators also affect microhabitat distribution 
of small mammals (Brown et al., 1988; 
Longland and Price, 1991). Kotler dem- 
onstrated that desert rodents forage in mi- 
crohabitats offering shelter from predators 
and that the effects of predation risk, in 
combination with resource availability, in- 
fluence structure of desert rodent assem- 
blages (Kotler, 1984, 1989; Kotler and Holt, 
1989; Kotler et al., 1988). 

Community patterns. —Community pat- 
tern was defined by Elton (1966:22) as “‘the 
repetition of certain component shapes to 
form a connected or interspersed design.” 
Here we consider patterns in species rich- 
ness, abundance, and diversity. The num- 
ber of species (species richness) of mammals 
increases with area (the well-known species 
area curve; Brown, 1971; Brown and Ni- 
coletto, 1991; Connor and McCoy, 1979; 
Dritschilo et al., 1975), although Lomolino 
(1989) warned of statistical considerations 
when interpreting the slope of the species- 
area curve (see Coleman et al., 1982). The 
distributional extent and density of mam- 
mals are also related (Brown, 1984). 

Several taxa of mammals exhibit hyper- 
diversity (Dial and Marzluff, 1989), that is, 
their biodiversity is greater than what would 
be expected by chance alone. Latitudinal 
patterns in species diversity of mammals 
are well known (Fleming, 1973; Heaney, 
1991; Harrison et al., 1992; McCoy and 
Connor, 1980; Owen, 1990a, 19906, Pagel 


COMMUNITIES AND ECOSYSTEMS 359 


et al., 1991; Rosenzweig, 1993; Schum, 
1984: Simpson, 1964; Willig and Sandlin, 
1991; Willig and Selcer, 1989), but not all 
groups of mammals respond to latitude in 
the same way. Indeed, quadrupedal mam- 
mals (as opposed to bats) do not fit the clas- 
sic pattern of increasing the diversity of spe- 
cies aS one moves toward the equator 
(Lacher and Mares, 1986; Mares, 1992; 
Mares and Ojeda, 1982). Many reasons for 
this gradient in species diversity have been 
advanced, including the supposition of a 
longer, uninterrupted time for evolution in 
the tropics [although Dritschilo et al. (1975) 
showed that rodent species introduced to 
North America within the past 2,000 years 
do not have fewer mite species than species 
that arose in the Pleistocene as predicted 
by the time hypothesis], spatial heterogene- 
ity (Hafner, 1977; Kotler and Brown, 
1988; M’Closkey, 1978), primary productiv- 
ity (Abramsky, 1989; Abramsky and Ro- 
senzweig, 1984: Brown, 1973; Brown and 
Davidson, 1977; Owen, 1988), potential 
evapotranspiration (Currie, 1991; Rosen- 
zweig, 1968), and disturbances (Fuentes and 
Jaksic, 1988). Bowers (1993) demonstrated 
that plant communities with high and low 
intensity of herbivory have lower diversity 
than when herbivory was at an intermediate 
intensity. Rosenzweig (1993) reviewed ev- 
idence from mammals and other taxa that 
reveals a productivity-diversity pattern with 
highest diversity at intermediate productiv- 
ities and suggests hypotheses to explain it, 
particularly the decline at high productivi- 
ties. 

Control of species diversity has been 
linked to the theory of limiting similarity, 
whereby the number of species in a com- 
munity may be limited by their niche over- 
lap (often measured as size ratios; Hutch- 
inson, 1959). Most data on size ratios, 
including that from mammals, do not sup- 
port limiting similarity (Brown and Lieber- 
man, 1973; Willig, 1986). In fact, the pres- 
ence of vacant niches in mammalian 
communities may facilitate invasions (Da- 
vis and Ward, 1988). 


Finally, the study of several other pat- 
terns provides insight into mammalian 
community dynamics. Differences in pat- 
terns of body mass of North American land 
mammals seen at different measurement 
scales have been attributed to diverse eco- 
logical and evolutionary processes oper- 
ating at those scales (i.e., competition, 
extinction, and allometric energetic con- 
straints; Brown and Nicoletto, 1991). Stage 
of succession affects mammalian diversity 
(Buckner and Shure, 1985; Foster and 
Gaines, 1991; Fox, 1982; Sly, 1976) and, in 
turn, mammals have a profound effect on 
patterns of plant succession by the processes 
of herbivory and disturbance; mammals 
usually facilitate the entrance of later suc- 
cessional (plant) species into a successional 
sere (Anderson et al., 1980; Huntly and In- 
ouye, 1987, 1988; Pearst, 1989; Platt, 1975; 
Tilman, 1983). Most recently, mammalian 
ecologists have begun to focus attention on 
patterns at the landscape scale. In particu- 
lar, current work is revealing the effect of 
sizes of habitat patches (particularly result- 
ing from habitat fragmentation) and corri- 
dors on dynamics of small mammal pop- 
ulations (Foster and Gaines, 1991; 
Henderson et al., 1985; Henein and Mer- 
riam, 1990; Laurance, 1991; Merriam and 
Lanoue, 1990). 

Community function. — Community 
function involves relationships among con- 
stituent species whereby energy and nutri- 
ents are exchanged among these species. 
However, other sorts of interactions among 
species affect the community. For example, 
the study of mammalian communities has 
contributed to our knowledge of ecological 
stability. McNaughton (1977, 1985) consid- 
ered how grazing mammals affected the re- 
lation among stability, diversity, and func- 
tional properties in grasslands of the 
Serengeti, concluding that the effect on 
grassland plant diversity may be different 
from the effect on grassland function (mea- 
sured as primary production). 

Trophic interactions among species are 
discussed in the section on Ecosystem Ecol- 


360 MARES AND CAMERON 


ogy below. Here we consider the impact of 
such trophic interactions and address the 
question as to the effects mammalian con- 
sumers might have on ecological commu- 
nities. Hairston et al. (1960; hereafter HSS) 
concluded that herbivores were seldom 
food-limited and unlikely to compete for 
resources, whereas producers, carnivores, 
and decomposers competed in a density- 
dependent fashion for resources. This land- 
mark study stimulated much research into 
consumer effects in various taxa, including 
mammals. Mammals usually consume 2- 
8% of available net production, but may eat 
as much as 30% under some conditions (P1- 
mentel, 1988), tending to support HSS. 
However, the addition of food results in in- 
creased population density, growth rate, and 
survival, and smaller home ranges, coun- 
tering predictions of HSS (Boutin, 1990; 
Desy et al., 1990; Dobson and Kjelgaard, 
1985; Klenner and Krebs, 1991; Mares et 
al., 1976, 1982: Sullivan et al., 1983; Taitt 
and Krebs, 1983). The conclusion that not 
all plants are edible and that food is limiting 
has been strengthened by studies demon- 
strating that dietary intake by mammalian 
consumers is restricted by the nutrient and 
secondary plant compound content of their 
food (Batzli, 1986; Batzli et al., 1980; Ber- 
geron and Jodoin, 1987; Bryant et al., 1991; 
Bucyanayandi and Bergeron, 1990; Eshel- 
man and Jenkins, 1989; Hanley, 1982; Jon- 
asson et al., 1986; Jung and Batzli, 1981; 
Kerley and Erasmus, 1991; Kuropat and 
Bryant, 1983; Marquis and Batzh, 1989; 
Randolph et al., 1991; Schultz, 1964; Seagle 
and McNaughton, 1992; Sinclair et al., 1982, 
1988; Snyder, 1992; Willig and Lacher, 
1991). 

Mammalian consumers have a variety of 
other effects on community function (Hunt- 
ly, 1991; Huntly and Inouye, 1988; Paige, 
1992; Whicker and Detling, 1988). In sum- 
mary, mammals affect plant production 
(Detling et al., 1980; Grant and French, 
1980; Reichman and Smith, 1991), fitness 
(Belsky, 1986; Edwards, 1985; Maschinsk1 
and Whitham, 1989; McNaughton, 1986; 


Paige and Whitham, 1987), pollination and 
seed dispersal (Borchert and Jain, 1978: 
Fleming, 1982; Golley et al., 1975; Howell 
and Roth, 1981), vegetative diversity 
(Archer et al., 1987; Batzli and Pitelka, 1970; 
Borchert and Jain, 1978; Bryant, 1987; Cof- 
fin and Lauenroth, 1988; Fox and Bryant, 
1984; Fuentes et al., 1983; Grant et al., 1982; 
Lidicker, 1989; Reichman and Smith, 1985; 
Reichman et al., 1993; Spatz and Mueller- 
Dombois, 1973; Stapanian and Smith, 1986; 
Truszkowski, 1982), and nutrient content 
(Coppock et al., 1983a). The complexity of 
biotic and abiotic interactions can be pro- 
nounced. For example, Grant et al. (1977) 
demonstrated that addition of nitrogen and 
water affected composition and density of 
a short-grass prairie and, concomitantly, af- 
fected structure of the mammalian com- 
munity (see also Grant et al., 1980, for the 
effects of burrowing by fossorial mammals 
on plant production). 

Convergent evolution and the develop- 
ment of communities. —Community ecol- 
ogists have paid a good deal of attention to 
determining if communities develop over 
evolutionary time in a predictable manner. 
Because all species within a community re- 
spond to complex stimuli in an evolution- 
ary manner, it might appear that popula- 
tions evolving under broadly similar 
climatic regimes would develop suites of 
similar adaptations. Certainly it has long 
been known that several mammals are re- 
markably convergent, and this general mor- 
phological similarity is particularly preva- 
lent among desert rodents, perhaps because 
they inhabit areas that are especially chal- 
lenging to the physiology and ecology of 
small mammals (Eisenberg, 1975; Hatt, 
1932: Schmidt-Nielsen, 1964). 

Pianka (1969, 1973, 1975, 1985, 1986) 
and Cody (1970, 1973, 1974, 1975) were 
among the first evolutionary ecologists to 
examine community convergence. Pianka 
conducted research on lizard communities 
in the United States, Australia, and Africa. 
Cody studied birds occurring on different 
continents in similar habitats (Mediterra- 


COMMUNITIES AND ECOSYSTEMS 361 


nean chaparral-scrubland birds of Califor- 
nia and Chile). Both examined various as- 
pects of ecology and community structure, 
and devised quantitative methods for com- 
paring niche parameters of faunas. Broadly 
speaking, birds were more convergent than 
lizards, although in each area there were 
striking examples of ecologically and mor- 
phologically convergent pairs, as well as re- 
markably different species. Karr and James 
(1975) studied the bird faunas of forested 
habitats of North and Central America and 
of Africa. Utilizing multivariate techniques, 
they concluded that convergence was pro- 
nounced among some species that differed 
phylogenetically, whereas divergence was 
evident among some species with similar 
phylogenetic backgrounds. 

At about this same time, Mares (1975, 
1976), for desert rodents, and Findley 
(1976), for bats, used multivariate analyses 
of morphoecological data to assess similar- 
ities and differences between faunas occur- 
ring on different continents. Both concluded 
that convergence was pronounced; mor- 
phology (and ecology) had evolved in many 
members of each fauna in a similar manner. 
Nevo (1979) demonstrated that fossorial 
rodents on many continents converged in 
ecological, morphological, behavioral, 
physiological, genetic, and many other char- 
acteristics in response to the subterranean 
environment. 

Mares (1980, 1993a, 19935) later extend- 
ed his original analysis, which had been lim- 
ited to an examination of desert and non- 
desert rodents in North and South America, 
to small mammals inhabiting all of the 
world’s deserts. His results showed that 
community-wide convergence of morphol- 
ogy and ecology generally was detectable 
when species with widely different phylog- 
enies were compared. Similar results were 
found by Berman (1985) in a rigorous mor- 
phological analysis of the evolution of bi- 
pedality among small mammals in deserts. 
Mares (1983:37-38) noted: “If one were to 
go into an unknown desert region, there are 
many predictions that could be made con- 


cerning the small mammal fauna... of the 
area... [AJ]t least some rodents ... would 
exhibit the following adaptations: special- 
ized kidneys .. . a counter-current heat ex- 
change system in the nasal region; modified 
brain cells responsible for ADH secretion; 
lowered metabolic rate; facultative torpor; 
ability to exist without free water; mini- 
mization of water loss through respiratory, 
excretory, and defecatory pathways; inflated 
tympanic bullae or elongate pinnae; bipe- 
dality ... [which]... could occur in all tro- 
phic categories except the completely fos- 
sorial niche ... [and] coexisting species 
might exhibit regular patterns of body size 
differences.” These comments about the 
pervasiveness of convergent evolution on 
the biology of organisms were in broad 
agreement with Nevo (1979). 

The International Biological Program 
dedicated a great deal of effort to assess the 
pervasiveness and predictability of conver- 
gent evolution between communities (Ma- 
bry et al., 1977; Orians and Solbrig, 1977; 
Simpson, 1977). The results of these exten- 
sive studies indicated that, differences in 
history, phylogeny, and climate notwith- 
standing, ecosystematic convergence can be 
quite pronounced, especially for some of the 
components of the ecosystem. 

Recent research on convergent evolution 
indicated that similar evolutionary adap- 
tations to similar physical environments 
may not only be striking, but may extend 
beyond morphological traits to complex be- 
havioral and ecological attributes. For ex- 
ample, Mares and Lacher (1987) showed 
that mammals that are specialized for life 
on isolated piles of boulders in different parts 
of the world can develop strongly conver- 
gent suites of characteristics that are asso- 
ciated with life in this rocky environment. 
These similarities will override phylogenet- 
ic similarities to such an extent that, for the 
traits examined, animals in different orders 
that inhabited very similar microenviron- 
ments (e.g., hyraxes, Cavia and Procavia of 
Africa, and the rock cavy, Kerodon, of the 
Brazilian Caatinga), were more similar to 


362 MARES AND CAMERON 


one another than they were to their own 
confamilials. 

Curiously, when the entire mammal fau- 
na of the Brazilian Caatinga was examined, 
there was little or no faunal convergence 
evidence between the Caatinga’s fauna and 
those of other semiarid areas in the world 
(Mares et al., 1985). The Caatinga, although 
an extensive tropical dry area, has had a 
special history of isolation from grasslands 
where pre-adaptations for aridity might have 
developed over time, as they did for the 
other deserts and semideserts of the world. 
Rather, the Caatinga is a tropical dryland 
surrounded by moist forests, an unusual 
zone that undergoes periodic and cata- 
strophic droughts (perhaps every two de- 
cades). Mares et al. (1985) showed that the 
largely tropically adapted fauna of the Caa- 
tinga was unable to adapt to aridity because 
droughts likely functioned as a frequent bot- 
tleneck that regularly eliminated most small 
mammals from the region. This research 
made clear the role of history, climate, and 
surrounding habitats on the evolution of 
convergent assemblages of mammals. 

Research on convergent evolution is con- 
tinuing for many groups of organisms (e.g., 
Luke, 1986; Schluter, 1986, 1990). Many 
questions remain to be answered. What is 
the influence of history on the evolution of 
similar species in similar areas? How chal- 
lenging must an environment be to limit the 
evolutionary responses of organisms and 
thus make convergence likely? To what ex- 
tent can phylogeny be overridden by natural 
selection? At the higher levels of organiza- 
tion (e.g., alpha and beta diversity, coexis- 
tence, competitive interactions, predation 
effects), what are the factors that cause con- 
vergence to be manifested, and can con- 
vergence be measured in some meaningful 
manner when entire faunas are compared 
(see Mares, 1993a)? 


Ecosystem Ecology 


Energetics. —With the publication of 
Tansley’s (1935) classic paper on plant ecol- 


ogy, 1t was possible to begin formulating 
experiments that would describe the func- 
tional relationships of organisms in a de- 
fined area. Perhaps because it is difficult to 
define the boundaries of an ecosystem [Col- 
invaux (1973:296) noted: ““Ecosystems are 
in the eye of the beholder .. .”’], it follows 
that the breakthrough in ecosystem ecology 
was made by an investigator studying lakes, 
which by their nature have distinct bound- 
aries. The landmark paper on ecosystem 
ecology was Lindeman’s (1942) report on 
the energetics of organisms in Cedar Bog 
Lake in Minnesota. Lindeman determined 
the standing crop of the various trophic lev- 
els in the lake and then assigned caloric val- 
ues to the productivity at each level. Thus, 
the currency of systems ecology (energy) was 
defined, quantified, and applied. Addition- 
ally, it subsequently became possible to have 
at least a frame of comparison for param- 
eters of standing crop, turnover, productiv- 
ity, and so forth. 

After Lindeman, ecosystems were con- 
sidered a basic unit of ecology (e.g., Evans, 
1956; Odum, 1953), and many ecologists, 
particularly those working in aquatic sys- 
tems, began conducting research on either 
natural systems or systems constructed in 
the laboratory (e.g., Slobodkin, 1962). If 
lakes have relatively well-defined bound- 
aries, and test tube communities even more 
so, terrestrial communities are notoriously 
difficult to control or even to obtain mea- 
surements of their component species. As 
Engelmann (1966) observed, it is a daunting 
task to apply a systems approach to a ter- 
restrial ecosystem. It was almost surely this 
difficulty in capturing, observing, and quan- 
tifying population sizes (standing crop), de- 
termining the energetics of respiration and 
of daily activities, estimating turnover rates, 
and obtaining the myriad of other data re- 
quired to understand how the system func- 
tioned, that delayed the application of 
Lindeman’s ideas (and those that had ex- 
panded systems theory in the intervening 
period) to a terrestrial system. It would be 
18 years before a trophic dynamic study of 
a terrestrial community would be conduct- 


COMMUNITIES AND ECOSYSTEMS 363 


ed. That classic paper would be provided 
by a mammalogist, Frank Golley (1960). 

Golley, who joined ASM in 1955 and 
would later publish an important text in 
mammalogy with David E. Davis (Davis 
and Golley, 1963) and field guides to the 
mammals of Georgia and South Carolina 
(Golley 1962, 1966), began a study of an 
old field terrestrial ecosystem whose vege- 
tation consisted largely of grasses and herbs. 
The main herbivore was a vole (Microtus 
pennsylvanicus) and the major predator was 
the least weasel (Mustela nivalis). As might 
be expected, Golley had to census plants, 
determine their energy content and the pro- 
portion of energy that the plants devoted to 
respiration, and estimate their productivity. 
Similar measurements (e.g., standing crop 
biomass and energy content, population dy- 
namics, growth, reproduction, assimilation 
efficiency, energy consumption) had to be 
made for Microtus and Mustela. Clearly, 
Golley’s study required a prodigious effort, 
yet it remains one of the few examples of 
energy flow through a simple terrestrial sys- 
tem (e.g., “Even the work of Golley ... is 
not very comprehensive,” Collier et al., 
1973:420). This criticism notwithstanding, 
Golley’s work established the field of ter- 
restrial energetics in vertebrate populations 
(see also Golley, 1961, 1967, 1968, 1983; 
Golley and Golley, 1972; Golley et al., 
LOTS): 

Shortly after Golley published his paper, 
Odum et al. (1962) expanded the scope of 
research on energy flow in another old field 
ecosystem. They examined energy flow 
through more components of the food chain 
than Golley did, including grasshoppers, a 
cricket, a sparrow, and the old field mouse, 
Peromyscus polionotus. Their research al- 
lowed them to tease apart differences in en- 
ergy flow between vertebrates and inverte- 
brates, as well as between herbivores and 
granivores. 

Much research into the energetics of 
mammals was devoted to determining the 
energy costs associated with various daily 
activities for mammals. This was generally 
carried out in the laboratory on resting an- 


imals, or utilized physiological instrumen- 
tation to compare resting and active rates 
of metabolism. These investigations cen- 
tered on single species and the results often 
were compared to energetic assumptions and 
determinations made by Golley (e.g., Chew 
et al., 1965; Gessaman, 1973; Golley et al., 
1965; Gorecki, 1965; Grodzinski and Go- 
recki, 1967; McNab, 1963, 1991; McNab 
and Morrison, 1963; Pearson, 1960). 

Terrestrial ecosystems were as difficult to 
study after Golley’s research had been pub- 
lished as they were before, but the publi- 
cation of his study on energy flow showed 
that, in principle, terrestrial systems, albeit 
extremely complex, were amenable to field 
research. Investigators thus began the dif- 
ficult task of examining energy flow through 
other systems. One of the first to publish on 
this topic was a mammalogist, Oliver Pear- 
son, who examined populations of several 
species of rodents and various carnivores 
(including feral house cats) in a large park 
in California (Pearson, 1964). Pearson cen- 
sused rodent populations to determine den- 
sity, then deduced the impact of carnivores 
on rodents by intensively collecting feces of 
predators. He also measured plant standing 
crop and estimated energetics of the organ- 
isms involved in energy flow through the 
system. This study was important because 
it dealt with a system which was more com- 
plex than that studied by Golley, although 
it was done over a much shorter time, ne- 
cessitating more assumptions than did Gol- 
ley’s work. 

Several studies dealing with one or an- 
other aspect of secondary productivity in 
ecosystems were published by Petrusewicz 
(1967), but it was another mammalogist who 
directed the research that would provide the 
next major energy flow study in a complex 
field situation. Robert Chew and his wife, 
Alice Eastlake Chew (Chew and Chew, 1965, 
1970), conducted an extensive study on the 
energetics of a desert scrub community, in- 
cluding its mammals. Working in a creosote 
bush (Larrea tridentata) scrubland, the 
Chews determined bioenergetics of plants, 
including density, productivity, and stand- 


364 MARES AND CAMERON 


ing crop, and gathered the same information 
on 13 species of small- and medium-sized 
mammals that occurred on the area. Their 
work remains one of the finest studies of 
energy flow in mammals ever conducted and 
provided important data to understand the 
pathways of energy flow through a desert 
system, ecological efficiencies of herbivores 
and granivores, and the net energy flow 
through various links in the food chain. 
Their research described the minor role 
played by small mammals (herbivores, 
granivores) in energy transfer in a com- 
munity, converting only 0.016% of the pri- 
mary above-ground production to mammal 
tissue that was then available as a food re- 
source to carnivores in higher trophic levels. 
This work provided dramatic quantitative 
data on the shape of the pyramids of energy 
and biomass. 

Following these early seminal studies, 
other investigators began to refine our un- 
derstanding of energy flow through mam- 
mal species and communities (e.g., Collier 
et al., 1975; Collins and Smith, 1976; Fle- 
harty and Choate, 1973; French et al., 1976; 
Gebczynska, 1970; Gebczynski et al., 1972; 
Grodzinski, 1971; Grodzinski and French, 
1983; Kenagy, 1973; McNaughton, 1976; 
Merritt and Merritt, 1978; Montgomery and 
Sunquist, 1975; Myrcha, 1975; Soholt, 
1973). These studies were conducted in 
temperate and tropical areas, and on both 
small and large mammals. 

The International Biological Program 
(IBP).—Undoubtedly, the major research 
stimulus to work on bioenergetics, and a 
continuing factor throughout the world on 
current interest in community dynamics, 
was the establishment of the IBP in the 
1960s. Because of its importance to research 
on ecosystems, some background on 
IBP is provided. 

In 1962 Ledyard Stebbins, a plant genet- 
icist at the University of California, Davis, 
published a paper on the activities of the 
International Union of Biological Sciences, 
of which he was Secretary-General (Steb- 
bins, 1962; see Blair, 1977). In that report, 


he outlined the International Biological 
Program, a program of global ecological re- 
search. W. Frank Blair, who was then Pres- 
ident of the Ecological Society of America, 
and who had been one of the leading mam- 
malian ecologists in ASM before dedicating 
his research program to the evolutionary 
ecology of amphibians and reptiles (cf., Blair, 
1939, 1941, 1953, 1955), became intimate- 
ly involved in the complex planning that 
ultimately resulted in the establishment of 
an internationally organized and funded 
program of comparative ecosystem research 
in 1967. The initial program had limited 
funding; broad-based financial support pro- 
vided by congressional action did not be- 
come available until Blair, in his role as 
Chairman of the US/IBP, led the fight to 
push funding bills through committees of 
both the House and Senate between 1967 
and 1970. 

IBP was dedicated to elucidating the 
structure and function of the earth’s major 
ecosystems. The methodologies employed 
were those of population ecology, energet- 
ics, community structure, mathematical 
modeling, and elemental cycling, among 
others. At the heart of this multi-country 
research effort were the biome programs. 
These included programs focusing on the 
major terrestrial biomes (Tundra Biome, 
Grassland Biome, Desert Biome, Conifer- 
ous Forest Biome, Deciduous Forest Bi- 
ome, Tropical Forest Biome), as well as pro- 
grams dealing with the Conservation of 
Ecosystems, Man in the Andes, Circum- 
polar Peoples, Upwelling Areas, and Origin 
and Structure of Ecosystems, which exam- 
ined the role of convergent evolution in 
structuring communities of organisms in 
North and South America. 

IBP was big science in all of its glory, and 
with all of its problems (Blair, 1977). Be- 
cause it cut across disciplines and countries, 
IBP was an extremely difficult undertaking 
and was widely criticized by scientists who 
were not involved in the programs or who 
felt that this type of coordinated research 
was not the way to do science (Michell et 


COMMUNITIES AND ECOSYSTEMS 365 


al., 1976). It was viewed negatively by some 
(e.g., Boffey, 1976), but time has provided 
a more balanced historical perspective. As 
McIntosh (1985:215) noted, “the status of 
ecology and ecologists at the inception of 
IBP was clearly ‘minor’, but IBP changed 
“the way ecology was done and the way 
ecologists thought about ecology” (McIn- 
tosh, 1985:219). IBP was a maturing force 
in the development of ecosystem ecology; 
it pushed this type of investigation into the 
forefront of organismal biology, giving it a 
high public profile and underscoring the im- 
portance of developing an understanding of 
how the natural environment functions. At- 
tempts to devise mathematical models of 
ecosystems were clearly less than successful 
(e.g., Berlinski, 1976), but ecosystem ecol- 
ogy has continued to develop, both concep- 
tually and methodologically (McIntosh, 
1985). 

The effect of IBP on world ecology was 
pronounced (Kormondy and McCormick, 
1981). In reviewing country after country, 
it is clear that field research flourished where 
IBP sites had been located. The program 
functioned as a training ground for students 
in the various fields of ecology, including 
mammalian community structure, energet- 
ics, population dynamics, and evolution. 
Literally thousands of papers on mammals 
have been published from work that was 
funded by, or related to, the IBP’s many 
foci. In Poland, for example, Kajak and 
Pieczynska (1981:287-—288) reported: ‘Four 
major periods can be distinguished in the 
development of Polish ecology after 1945: 
... [f]rom 1969 to 1975 was a period of 
intense studies on ecological productivity 
and of ecosystem studies connected with. . . 
IBP ... [including especially s]tudies on 
smail mammals.” For Sweden, Sjors (1981: 
305-306) noted, “The ... (IBP) meant in- 
creased contacts among ecologists all over 
the world. ... Thanks to the IBP [produc- 
tion and biomass studies] became highlight- 
ed in basic research.”’ In most countries 
where IBP research was conducted, mam- 
mal investigations were extensive. 


The results of the efforts of the IBP are 
still being witnessed today in mammalogi- 
cal research. There are ecosystem-oriented 
studies and research based in energetics that 
are currently providing important infor- 
mation on the ecology of populations and 
communities of mammals. Research stim- 
ulated by the projects or scientists who par- 
ticipated in IBP is still being conducted in 
a wide array of habitats throughout the 
world. Even as we approach two decades 
since the termination of IBP, there has 
probably been insufficient time to assess ob- 
jectively the impacts and contributions of 
the entire program on a global basis. How- 
ever, scientists involved with IBP not only 
conducted research on ecosystem function, 
community evolution, and ecosystem de- 
velopment, but were also instrumental in 
carrying on empirical research on the effects 
of abiotic factors on the structure of mam- 
malian communities. The Structure of Eco- 
systems Program was dedicated to this goal. 
IBP and ecosystem studies will be inti- 
mately associated in the future. 


Conclusions 


Mammalogists have contributed greatly 
to the development of community and eco- 
system ecology. Their influence has been 
pervasive and continuous, and extends from 
the very foundations of these fields of re- 
search. Present trends indicate that impor- 
tant empirical and theoretical contributions 
to elucidating patterns of community and 
ecosystem structure and function will con- 
tinue to be made by mammalogists. There 
is no doubt that members of the ASM have 
been, and will continue to be, at the fore- 
front of this research. 


Acknowledgments 


We thank O. J. Reichman and M. R. Willig 
for critical reviews of the manuscript. We also 


366 MARES AND CAMERON 


thank J. K. Braun, R. B. Channell, and R. Hum- 
phrey for assistance in preparing the manuscript. 


Literature Cited 


ABRAMSKY, Z. 1989. Communities of gerbilline ro- 
dents in sand dunes of Israel. Pp. 205-217, in Pat- 
terns in the structure of mammalian communities 
(D. W. Morris, Z. Abramsky, B. J. Fox, and M. R. 
Willig, eds.). Texas Tech University Press, Lubbock, 
266 pp. 

ABRAMSKY, Z., AND M. L. RosENzweic. 1984. Til- 
man’s predicted productivity-diversity relationship 
shown by desert rodents. Nature, 309:150-151. 

Apams, C. C. 1905. The postglacial dispersal of the 
North American biota. The Biological Bulletin, 9:53- 
71. 


1909. An ecological survey of Isle Royal, 
Lake Superior. Report of the Board of the Geological 
Survey of 1908, Lansing, Michigan, 468 pp. 

. 1913. Guide to the study of animal ecology. 
Macmillan, New York, 183 pp. 

ALLEE, W. C., A. E. EMERSON, O. PARK, T. PARK, AND 
K. P. Scumipt. 1949. Principles of animal ecology. 
W. B. Saunders Company, Philadelphia, Pennsyl- 
vania, 837 pp. 

ANDERSEN, D.C., ANDJ. A. MACMAHON. 1985. Plant 
succession following the Mount St. Helens volcanic 
eruption: facilitation by a burrowing rodent, Tho- 
momys talpoides. The American Midland Natural- 
ist, 114:62-69. 

ANDERSEN, D. C., J. A. MACMAHON, AND M. L. WOLFE. 
1980. Herbivorous mammals along a montane sere: 
community structure and energetics. Journal of 
Mammalogy, 61:500-519. 

ANDERSON, S. 1959. Mammals of the Grand Mesa, 
Colorado. University of Kansas Publications, Mu- 
seum of Natural History, 9:405-414. 

ARCHER, S., M. G. GARRETT, AND J. K. DETLING. 1987. 
Rates of vegetation change associated with prairie 
dog (Cynomys ludovicianus) grazing in North Amer- 
ican mixed-grass prairie. Vegetatio, 72:159-166. 

BAKER, R. H. 1969. Cotton rats of the Sigmodon 
fulviventer group. Pp. 177-232, in Contributions in 
Mammalogy (J. K. Jones, ed.). Miscellaneous Pub- 
lications, Museum of Natural History, University of 
Kansas, 51:1-428. 

BARCLAY, R. M. R., AND R. M. BRIGHAM. 1991. Prey 
detection, dietary niche breadth, and body size in 
bats: why are aerial insectivorous bats so small? The 
American Naturalist, 137:693-703. 

BarLow, C. 1991. From Gaia to selfish genes. MIT 
Press, Cambridge, Massachusetts, 273 pp. 

BARTHOLOMEW, G. A. 1986. The role of natural his- 
tory in contemporary biology. BioScience, 36:324- 
329. 

Batzu, G. O. 1986. Nutritional ecology of the Cal- 
ifornia vole: effects of food quality on reproduction. 
Ecology, 67:406-412. 

Batzu, G. O., AND F. A. PireckaA. 1970. Influence 
of meadow mouse populations on California grass- 
land. Ecology, 51:1027-1039. 

Batzu, G. O., R. G. Wuite, S. F. MACLEAN, F. A. 
PITELKA, AND B. D. CoLiier. 1980. The herbivore- 
based trophic system. Pp. 335-410, im An Arctic 


ecosystem: the coastal tundra at Barrow, Alaska (J. 
Brown, P. C. Miller, L. L. Tieszen, and F. L. Bunnell, 
eds.). Dowden, Hutchinson & Ross, Incorporated, 
Stroudsburg, Pennsylvania, 571 pp. 

BELK, M. C., H. D. SMitH, AND J. Lawson. 1989. Use 
and partitioning of montane habitat by small mam- 
mals. Journal of Mammalogy, 69:688-695. 

Betsky, A. J. 1986. Does herbivory benefit plants? 
A review of the evidence. The American Naturalist, 
127:870-892. 

BERGERON, J. M., AND L. Joporn. 1987. Defining 
“high quality” food resources of herbivores: the case 
for meadow voles (Microtus pennsylvanicus). Oec- 
ologia, 71:510-517. 

BERLINSKI, D. 1976. On systems analysis: an essay 
concerning the limitations of some mathematical 
methods in the social, political, and biological sci- 
ences. MIT Press, Cambridge, Massachusetts. pp. 


BERMAN, S. L. 1985. Convergent evolution in the 
hindlimb of bipedal rodents. Zeischrift fiir Zoolo- 
gische Systematik und Evolutionsforschung, 23:59- 
Vile 

Biair, W. F. 1939. Faunal relationships and geo- 
graphic distribution of mammals in Oklahoma. The 
American Midland Naturalist, 22:85-133. 

1941. Techniques for the study of small 

mammal populations. Journal of Mammalogy, 22: 

148-157. 

1953. Population dynamics of rodents and 

other small mammals. Advances in Genetics, 5:1- 

41. 


1955. Size differences as a possible isolation 
mechanism in Microhyla. The American Naturalist, 
89:297-302. 

. 1977. Big biology. Dowden, Hutchinson and 
Ross, Incorporated, Stroudsburg, Pennsylvania, 261 
pp. 

Borrey, P. M. 1976. International Biological Pro- 
gram: was it worth the cost and effort? Science, 193: 
866-868. 

Borcuert, J. I., AND S. K. JAIN. 1978. The effect of 
rodent seed predation on four species of California 
annual grasses. Oecologia, 33:101-113. 

Boutin, S. 1990. Food supplementation experiments 
with terrestrial vertebrates: patterns, problems, and 
the future. Canadian Journal of Zoology, 68:203- 
220. 

Bowers, M.A. 1993. Influence of herbivorous mam- 
mals on an old-field plant community: years 14 
after disturbance. Oikos, 67:129-141. 

Bowers, M., AND J. H. Brown. 1982. Body size and 
coexistence in desert rodents: chance or community 
structure? Ecology, 63:39 1-400. 

Brown, J. H. 1971. Species richness of boreal mam- 
mals living on the montane islands of the Great Ba- 
sin. The American Naturalist, 105:467-478. 

. 1973. Species diversity of seed-eating desert 

rodents in sand dune habitats. Ecology, 54:775-787. 

. 1975. Geographical ecology of desert rodents. 

Pp. 315-341, in Ecology and evolution of commu- 

nities (M. L. Cody and J. M. Diamond, eds.). Har- 

vard University Press, Cambridge, Massachusetts, 

543 pp. 

. 1981. Two decades of homage to Santa Rosa- 

lia: toward a general theory of diversity. The Amer- 

ican Zoologist, 21:877-888. 


COMMUNITIES AND ECOSYSTEMS 367 


. 1984. On the relationship between abun- 
dance and distribution of species. The American 
Naturalist, 124:255-279. 

. 1986. Two decades of interaction between 
the MacArthur-Wilson model and the complexities 
of mammalian distributions. Biological Journal of 
the Linnean Society, 28:231-251. 

Brown, J. H., AND M. A. Bowers. 1984. Patterns 
and processes in three guilds of terrestrial verte- 
brates. Pp. 282-296, in Ecological communities. 
Conceptual issues and the evidence (D. R. Strong, 
Jr., D. Simberloff, L. G. Abele, and A. B. Thistle, 
eds.). Princeton University Press, Princeton, New 
Jersey, 613 pp. 

Brown, J. H., AND D. W. Davipson. 1977. Com- 
petition between seed-eating rodents and ants in des- 
ert ecosystems. Science, 196:880-882. 

Brown, J. H., AND E. J. Heske. 1990. Control of a 
desert-grassland transition by a keystone rodent guild. 
Science, 250:1705-1707. 

Brown, J. H., AND M. Kurzius. 1989. Spatial and 
temporal variation in guilds of North American gra- 
nivorous desert rodents. Pp. 71-90, in Patterns in 
the structure of mammalian communities (D. W. 
Morris, Z. Abramsky, B. J. Fox, and M. R. Willig, 
eds.). Texas Tech University Press, Lubbock, 266 
pp. 

Brown, J. H., AND G. A. LIEBERMAN. 1973. Resource 
utilization and coexistence of seed-eating desert ro- 
dents in sand dune habitats. Ecology, 54:788-797. 

Brown, J. H., AND P. F. NicoLetto. 1991. Spatial 
scaling of species composition: body masses of North 
American land mammals. The American Naturalist, 
138:1478-1512. 

Brown, J. H., D. W. DAvIDSON, AND O. J. REICHMAN. 
1979. An experimental study of competition be- 
tween seed-eating rodents and ants. The American 
Zoologist, 19:1129-1143. 

Brown, J. H., D. W. Davipson, J. C. MUNGER, AND 
R. S. INouye. 1986. Experimental community 
ecology: the desert granivore system. Pp. 41-62, in 
Community ecology (J. Diamond and T. Case, eds.). 
Harper and Row, New York, 665 pp. 

Brown, J. S. 1989. The role of resource variability 
in structuring desert rodent communities. Pp. 141- 
154, in Patterns in the structure of mammalian com- 
munities (D. W. Morris, Z. Abramsky, B. J. Fox, 
and M. R. Willig, eds.). Texas Tech University Press, 
Lubbock, 266 pp. 

Brown, J. S., B. P. Kotter, R. J. SMITH, AND W. O. 
Wirtz, II. 1988. The effects of owl predation on 
the foraging behavior of heteromyid rodents. Oec- 
ologia, 76:408-415. 

BRYANT, J. P. 1987. Feltleaf willow-snowshoe hare 
interactions: plant carbon/nutrient balance and 
floodplain succession. Ecology, 68:1319-1327. 

BRYANT, J. P., F. D. PROVENzA, J. PAsTor, P. B. REI- 
CHARDT, T. P. CLAUSEN, AND J. T. puTorr. 1991. 
Interactions between woody plants and browsing 
mammals mediated by secondary metabolites. An- 
nual Review of Ecology and Systematics, 22:431- 
446. 

BUCKNER, C. Z., AND D. J. SHURE. 1985. The response 
of Peromyscus to forest opening size in the southern 
Appalachian Mountains. Journal of Mammalogy, 66: 
299-307. 


BUCYANAYANDI, J. F., AND J. M. BERGERON. 1990. 
Effects of food quality on feeding patterns of meadow 
voles (Microtus pennsylvanicus) along a community 
gradient. Journal of Mammalogy, 71:390-396. 

Buscu, M., AND F. O. KrAvetz. 1992. Competitive 
interactions among rodents (Akodon azarae, Calo- 
mys laucha, Calomys musculinus and Oligoryzomys 
flavescens) in a two-habitat system. II. Effect of spe- 
cies removal. Mammalia, 56:540-554. 

CaprerRA, A. 1932. La incompatabilidad ecoldogica: 
una ley biolégica interesante. Anales Sociedad Cien- 
tifica de Argentina, 114:243-260. 

CAMERON, G. N. 1971. Niche overlap and compe- 
tition in woodrats. Journal of Mammalogy, 52:288- 
296. 

Carnes, B. A., AND N. A. SLADE. 1982. Some com- 
ments on niche analysis in canonical space. Ecology, 
63:888-893. 

CARPENTER, F. L. 1978. Hooks for mammal polli- 
nation. Oecologia, 35:123-132. 

Case, T. J., AND D. T. BotGer. 1991. The role of 
interspecific competition in the biogeography of is- 
land lizards. Trends in Ecology and Evolution, 6:135- 
139. 

Cuew, R. M., AND A. E. CHEw. 1965. The primary 
productivity of a desert scrub (Larrea tridentata) 
community. Ecological Monographs, 35:355-375. 

1970. Energy relationships of the mammals 
of a desert scrub (Larrea tridentata) community. 
Ecological Monographs, 40:1-21. 

CHeEw, R. M., R. G. LINDBERG, AND P. HAYDEN. 1965. 
Circadian rhythm of metabolic rate in pocket mice. 
Journal of Mammalogy, 46:477-494. 

CHURCHFIELD, S. 1991. Niche dynamics, food re- 
sources, and feeding strategies in multispecies com- 
munities of shrews. Pp. 23-34, in The biology of the 
Soricidae (J. S. Findley and T. L. Yates, eds.). Special 
Publication, The Museum of Southwestern Biology, 
1:1-91. 

CLEMENTS, F. E. 1905. Research methods in ecology. 
University Publishing Company, Lincoln, Nebras- 
ka. 

1916. Plant succession. An analysis of the 
development of vegetation. Carnegie Institute of 
Washington Publication, 242:1-512. 

CLEMENTS, F. E. AND V. E. SHELFORD. 1939. Bio- 
ecology. John Wiley & Sons, Incorporated, New York, 
425 pp. 

Copy, M. L. 1970. Chilean bird distributions. Ecol- 
ogy, 51:455-464. 

1973. Parallel evolution and bird niches. Pp. 

307-338, in Mediterranean-type ecosystems: origin 

and structure (F. di Castri and H. A. Mooney, eds.). 

Springer-Verlag, New York, 405 pp. 

1974. Competition and the structure of bird 

communities. Princeton University Press, Prince- 

ton, New Jersey, 318 pp. 

1975. Towards a theory of continental spe- 
cies diversity: bird distributions over Mediterranean 
habitat gradients. Pp. 214-257, in Ecology and evo- 
lution of communities (M. Cody and J. Diamond, 
eds.). Harvard University Press, Cambridge, Mas- 
sachusetts, 543 pp. 

Copy, M. L., AND J. M. DIAMOND (eEDs.). 1975. Ecol- 
ogy and evolution of communities. Harvard Uni- 
versity Press, Cambridge, Massachusetts, 543 pp. 


368 MARES AND CAMERON 


CorFin, D. P., AND W. K. LAUENROTH. 1988. The 
effects of disturbance size and frequency on a short- 
grass plant community. Ecology, 69:1609-1617. 

COLEMAN, B. D., M. A. Mares, M. R. WILLIG, AND Y. 
H. Hsrew. 1982. Randomness, area, and species 
richness. Ecology, 63:1121-1133. 

CoLINvAUx, P. 1973. Introduction to ecology. John 
Wiley & Sons, Incorporated, New York, 621 pp. 
Co.uigrR, B. D., G. W. Cox, A. W. JOHNSON, AND P. 
C. MILLER. 1973. Dynamic ecology. Prentice-Hall, 
Incorporated, Englewood Cliffs, New Jersey, 563 pp. 

Co ier, B. D., N. C. STENSETH, S. BARKLEY, AND R. 
Osporn. 1975. A simulation model of energy ac- 
quisition and utilization by the brown lemming 
Lemmus trimucronatus at Barrow, Alaska. Oikos, 
26:276-294. 

Co.iins, V. R., AND M. H. SmitH. 1976. Field de- 
termination of energy flow in a small nocturnal 
mammal. Journal of Mammalogy, 57:149-158. 

CONNELL, J. H. 1980. Diversity and the coevolution 
of competitors, or the ghost of competition past. 
Oikos, 35:131-138. 

1983. On the prevalence and relative im- 
portance of interspecific competition: evidence from 
field experiments. The American Naturalist, 122:661- 
696. 

Connor, E. F., AND E. D. McCoy. 1979. The statis- 
tics and biology of the species-area relationship. The 
American Naturalist, 113:791-833. 

Coppock, D. L., J. K. DETLING, J. E. ELLIs, AND M. I. 
Dyer. 1983a. Plant-herbivore interactions in a 
North American mixed-grass prairie. I. Effects of 
black-tailed prairie dogs on intraseasonal above- 
ground plant biomass and nutrient dynamics and 
plant species diversity. Oecologia, 56:1-9. 

Coppock, D. L., J. E. ELuis, J. K. DETLING, AND M. I. 
Dyer. 1983. Plant-herbivore interactions in a 
North American mixed-grass prairie. II. Responses 
of bison to modification of vegetation by prairie dogs. 
Oecologia, 56:10-15. 

Cox, D. 1980. A note on the queer history of “niche.” 
Bulletin of the Ecological Society of America, 64: 
201-202. 

CROWELL, K. L., AND S. L. Pimm. 1976. Competition 
and niche shifts of mice introduced onto small is- 
lands. Oikos, 27:251-258. 

Currig, D. J. 1991. Energy and large-scale patterns 
of animal- and plant-species richness. The American 
Naturalist, 137:27-49. 

Davipson, D. W., R. S. INoUYE, AND J. H. BRown. 
1984. Granivory in a desert ecosystem: experimen- 
tal evidence for indirect facilitation of ants by ro- 
dents. Ecology, 65:1780-1786. 

Davis, D. E., AND F. B. GOLLEY. 1963. Principles in 
mammalogy. Reinhold Publishing Company, New 
York, 335 pp. 

Davis, R., AND O. G. WARD. 1988. A vacant ‘‘Mi- 
crotus niche” now occupied by the yellow-nosed cot- 
ton rat (Sigmodon ochrognathus) on an isolated 
mountain in southeastern Arizona. Journal of Mam- 
malogy, 69:362-365. 

Dayan, T., D. SIMBERLOFF, E. TCHERNOV, AND Y. 
Yom-Tov. 1989. Inter- and intraspecific character 
displacement in mustelids. Ecology, 70:1526-1539. 

. 1990. Feline canines: community-wide char- 


acter displacement among the small cats of Israel. 
The American Naturalist, 136:39-60. 

Desy, E. A., G. O. BATZLI, AND J. Lru. 1990. Effects 
of food and predation on behaviour of prairie voles. 
Oikos, 58:159-168. 

DETLING, J. K., M. I. Dyer, C. PROCTER-GREGG, AND 
D. T. Winn. 1980. Plant-herbivore interactions: 
examination of potential effects of bison saliva on 
regrowth of Bouteloua gracilis. Oecologia, 45:26-31. 

DIAL, K. P., AND J. M. MARZLUFF. 1989. Nonrandom 
diversification within taxonomic assemblages. Sys- 
tematic Zoology, 38:26—37. 

DiamMonpD, J. M. 1978. Niche shifts and the redis- 
covery of interspecific competition. American Sci- 
entist, 66:322-331. 

. 1986. Overview: laboratory experiments, field 
experiments, and natural experiments. Pp. 3-22, in 
Community ecology (J. M. Diamond and T. Case, 
eds.). Harper and Row, New York, 665 pp. 

Dice, L. 1952. Natural communities. University of 
Michigan Press, Ann Arbor, 547 pp. 

DICKMAN, C.R. 1988. Body size, prey size, and com- 
munity structure in insectivorous mammals. Ecol- 
ogy, 69:569-580. 

1989. Patterns in the structure and diversity 
of marsupial carnivore communities. Pp. 241-251, 
in Patterns in the structure of mammalian com- 
munities (D. W. Morris, Z. Abramsky, B. J. Fox, 
and M. R. Willig, eds.). Texas Tech University Press, 
Lubbock, 266 pp. 

Dosson, F. S., AND J. D. KJELGAARD. 1985. The 
influence of food resources on life history in Colum- 
bian ground squirrels. Canadian Journal of Zoology, 
63:2105-2109. 

DrRITSCHILO, W., H. CoRNELL, D. NAFus, AND B. 
O’Connor. 1975. Insular biogeography of mice 
and mites. Science, 190:467—-469. 

Dueser, R. D., AND J. G. HALLETT. 1980. Compe- 
tition and habitat selection in a forest-floor small 
mammal fauna. Oikos, 35:293-297. 

DueserR, R. D., AND H. H. SHuGART. 1979. Niche 
pattern in a forest-floor small-mammal fauna. Ecol- 
ogy, 60:108-118. 

—. 1982. Reply to comments by Van Horne and 
Ford and by Carnes and Slade. Ecology, 63:1174— 
1175. 

Dueser, R. D., J. H. PoRTER, AND J. L. J. DOOLEY. 
1989. Direct tests for competition in North Amer- 
ican rodent communities: synthesis and prognosis. 
Pp. 105-126, in Patterns in the structure of mam- 
malian communities (D. W. Morris, Z. Abramsky, 
B. J. Fox, and M. R. Willig, eds.). Texas Tech Uni- 
versity Press, Lubbock, 266 pp. 

Durr, A. G., AND P. D. Lowe. 1981. Great Britain. 
Pp. 141-156, in Handbook of contemporary devel- 
opments in world ecology (E. J. Kormondy and J. 
F. McCormick, eds.). Greenwood Press, Westport, 
Connecticut, 776 pp. 

Duaains, D. O. 1980. Kelp beds and sea otters: an 
experimental approach. Ecology, 61:447-453. 

Epwarps, J. 1985. Effects of herbivory by moose on 
flower and fruit production of Aralia nudicaulis. The 
Journal of Ecology, 73:861-868. 

Eur icu, P. R., AND J. ROUGHGARDEN. 1987. The 
science of ecology. Macmillan, New York, 710 pp. 

EISENBERG, J. F. 1975. The behavior patterns of des- 


COMMUNITIES AND ECOSYSTEMS 369 


ert rodents. Pp. 189-224, in Rodents in desert en- 
vironments (I. Prakash and P. K. Ghosh, eds.). Junk, 
b.v., The Hague, The Netherlands, 624 pp. 

ELTon, C. 1927. Animal ecology. Sidgwick & Jack- 
son, London, 207 pp. 

1933. The ecology of animals. Methuen and 

Company Limited, London, 97 pp. 

1942. Voles, mice, and lemmings: problems 

in population dynamics. Clarendon Press, Oxford, 

496 pp. 

1958. The ecology of invasions by animals 

and plants. Methuen & Company Limited, London, 

181 pp. 

1966. The pattern of animal communities. 
Methuen & Company Limited, London, 432 pp. 
ELTON, C., AND R. S. MILLER. 1954. The ecological 
survey of animal communities with a practical sys- 
tem of classifying habitats by structural characters. 

The Journal of Ecology, 42:460-496. 

ELTON, C., AND M. NIcHOLSON. 1942. The ten-year 
cycle in numbers of the lynx in Canada. The Journal 
of Animal Ecology, 11:215-244. 

Emmons, L. H. 1980. Ecology and resource parti- 
tioning among nine species of African rain forest 
squirrels. Ecological Monographs, 50:31-54. 

ENGELMANN, M.D. 1966. Energetics, terrestrial field 
studies, and animal productivity. Advances in Eco- 
logical Research, 3:73-115. 

ERLINGE, S., ET AL. 1983. Predation as a regulation 
factor on small rodent populations in southern Swe- 
den. Oikos, 40:36-52. 

. 1984. Can vertebrate predators regulate their 

prey? The American Naturalist, 123:125-133. 

1988. More thoughts on vertebrate predator 
regulation of prey. The American Naturalist, 132: 
148-154. 

ERRINGTON, P. L. 1946. Predation and vertebrate 
populations. The Quarterly Review of Biology, 21: 
144-177. 

ESHELMAN, R. D., AND S. H. JENKINS. 1989. Food 
selection by Belding’s ground squirrels in relation to 
plant nutritional features. Journal of Mammalogy, 
70:846-851. 

Estes, J. A., AND J. F. PALMISANO. 1974. Sea otters: 
their role in structuring nearshore communities. Sci- 
ence, 185:1058-1060. 

Estes, J. A., N.S. SMITH, AND J. F. PALMISANO., 1978. 
Sea otter predation and community structure. Ecol- 
ogy, 59:822-833. 

Evans, F. C. 1956. Ecosystems as the basic unit in 
ecology. Science, 123:1127-1128. 

FAUTIN, R. 1946. Biotic communities of the northern 
desert shrub biome in western Utah. Ecological 
Monographs, 16:251-310. 

Finb.ey, J. S. 1976. The structure of bat commu- 
nities. The American Naturalist, 110:129-139. 

. 1989. Morphological patterns in rodent com- 

munities of southwestern North America. Pp. 253- 

263, in Patterns in the structure of mammalian com- 

munities (D. W. Morris, Z. Abramsky, B. J. Fox, 

and M. R. Willig, eds.). Texas Tech University Press, 

Lubbock, 266 pp. 

. 1993. Bats. A community perspective. Cam- 
bridge University Press, New York, 200 pp. 

FLEHARTY, E. D., AND J. R. CHOATE. 1973. Bioen- 


ergetic strategies of the cotton rat, Sigmodon hispi- 
dus. Journal of Mammalogy, 54:680-692. 

FLEMING, T. H. 1973. Numbers of mammal species 
in North and Central American forest communities. 
Ecology 54:555-563. 

1981. Fecundity, fruiting pattern, and seed 

dispersal in Piper amalago (Piperaceae), a bat-dis- 

persed tropical shrub. Oecologia, 51:42-46. 

1982. Foraging strategies of plant visiting 
bats. Pp. 287-326, in Ecology of bats (T. H. Kunz, 
ed.). Plenum Press, New York, 425 pp. 

FLEMING, T. H., E. T. Hooper, AND D. E. WILSON. 
1972. Three Central American bat communities: 
structure, reproductive cycles, and movement pat- 
terns. Ecology, 53:555-569. 

Forses, S.A. 1887. The lake asa microcosm. Bulletin 
Science Association of Peoria, Illinois, 1887:77-87. 

Foster, J., AND M. S. Gaines. 1991. The effects of 
a successional habitat mosaic on a small mammal 
community. Ecology, 72:1358-1373. 

Fox, B. J. 1981. Niche parameters and species rich- 
ness. Ecology, 62:1415-1425. 

1982. Fire and mammalian secondary suc- 

cession in an Australian coastal heath. Ecology, 63: 

1332-1341. 

. 1989. Small-mammal community pattern in 
Australian heathland: a taxonomically-based rule for 
species assembly. Pp. 91-103, in Patterns in the 
structure of mammalian communities (D. W. Mor- 
ris, Z. Abramsky, B. J. Fox, and M. R. Willig, eds.). 
Texas Tech University Press, Lubbock, 266 pp. 

Fox, B. J., AND J. H. Brown. 1993. Assembly rules 
for functional groups in North American desert ro- 
dent communities. Oikos, 67:358-370. 

Fox, J. F., AND J. P. BRYANT. 1984. Instability of the 
snowshoe hare and woody plant interaction. Oec- 
ologia, 63:128-135. 

FRANKLIN, J. F., C. S. BLEDSOE, AND J. T. CALLAHAN. 
1990. Contributions of the long-term ecological re- 
search program. BioScience, 40:509-523. 

FRENCH, N. R., W. E. GRANT, W. GRODZINSKI, AND 
D. M. Swirr. 1976. Small mammal energetics in 
grassland ecosystems. Ecological Monographs, 46: 
201-220. 

FRYXELL, J. M., D. J. T. HUsse.i, A. B. LAMBERT, AND 
P.C. SmitH. 1991. Time lags and population fluc- 
tuations in white-tailed deer. The Journal of Wildlife 
Management, 55:377-385. 

FUENTES, E. R., AND F. M. JAksic. 1988. The hump- 
backed species diversity curve: why has it not been 
found among land animals? Oikos, 53:139-143. 

FUENTES, E. R., F. M. JAKsitc, AND J. A. SIMONETTI. 
1983. European rabbits versus native rodents in 
central Chile: effects on shrub seedlings. Oecologia, 
58:411-414. 

GAFENEY, P. M. 1973. Roots of the niche concept. 
The American Naturalist, 109:490. 

GALINDO, C., AND C. J. Kress. 1986. Evidence for 
competition in small rodents. Oikos, 46:116-118. 
GeEBCZYNSKA, Z. 1970. Bioenergetics of a root vole 

population. Acta Theriologica, 15:33-66. 

GeBcCZYNSKI, M., A. GORECKI, AND A. Drozpz. 1972. 
Metabolism, food assimilation and bioenergetics of 
three species of dormice (Gliridae). Acta Theriolo- 
gica, 17:271-294. 


370 MARES AND CAMERON 


GESSAMAN, J. A. (ED.). 1973. Ecological energetics of 
homeotherms. A view compatible with ecological 
modeling. Monograph Series, Utah State University 
Press, Logan, 20:1-155. 

Gicon, A., F. KL6TzL1, AND U. RAHM. 1981. Swit- 
zerland. Pp. 313-329, in Handbook of contemporary 
developments in world ecology (E. J. Kormondy and 
J. F. McCormick, eds.). Greenwood Press, Westport, 
Connecticut, 776 pp. 

Giass, G. E., AND N. A. SLADE. 1980. The effect of 
Sigmodon hispidus on spatial and temporal activity 
of Microtus ochrogaster: evidence for competition. 
Ecology, 61:358-370. 

Gieason, H. A. 1917. The structure and develop- 
ment of the plant association. Bulletin of the Torrey 
Botanical Club, 43:463-481. 

. 1939. The individualistic concept of the plant 
association. The American Midland Naturalist, 21: 
92-110. 

Got..ey, F.B. 1960. Energy dynamics ofa food chain 
of an old-field community. Ecological Monographs, 
30:187-206. 

. 1961. The energy value of ecological mate- 

rials. Ecology, 42:581-584. 

1962. The mammals of Georgia: a study of 

their distribution and functional role in the ecosys- 

tem. University of Georgia Press, Athens, Georgia, 

218 pp. 

. 1966. South Carolina mammals. Contribu- 

tions of the Charleston Museum, No. 15. 181 pp. 

1967. Methods of measuring secondary pro- 

ductivity in terrestrial vertebrate populations. Pp. 

99-124, in Secondary productivity of terrestrial eco- 

systems (K. Petrusewicz, ed.). 2 vol. Panstwowe Wy- 

dawnictwo Naukowe, Warsaw, Poland, 879 pp. 

1968. Secondary productivity in terrestrial 

ecosystems. American Zoologist, 8:53-59. 

(ED.). 1983. Tropical rainforest ecosystems: 
structure and function. Ecosystems of the world, 14A, 
Elsevier Scientific Publishing Company, Amster- 
dam, 381 pp. 

GoLLey, F. B., K. PETRUSEWICz, AND L. RySZKOWSKI 
(EDs.). 1975. Small mammals: their productivity and 
population dynamics. Cambridge University Press, 
Cambridge, 451 pp. 

GoL Ley, F. B., G. A. PETRIpDEs, E. L. RAUBER, AND J. 
H. JENKINS. 1965. Food intake and assimilation by 
bobcats under laboratory conditions. The Journal of 
Wildlife Management, 29:442-447. 

Go. ey, F. B., J. T. McGINnnis, R. G. CLEMENTs, G. 
I. CHILD, AND M. J. DuEver. 1975. Mineral cycling 
in a tropical moist ecosystem. University of Georgia, 
Athens, 248 pp. 

GoLLEY, P. M., AND F. B. GOLLEy (EDs.). 1972. Trop- 
ical ecology with an emphasis on organic production. 
University of Georgia, Athens. 

Gorecki, A. 1965. Energy values of body in small 
mammals. Acta Theriologica, 10:333-352. 

GRANT, P. R. 1972. Interspecific competition among 
rodents. Annual Review of Ecology and Systematics, 
3:79-196. 

GRANT, W. E., AND N. R. FRENCH. 1980. Evaluation 
of the role of small mammals in grassland ecosys- 
tems: a modelling approach. Ecological Modelling, 
8:15-37. 


GRANT, W. E., N. R. FRENCH, AND L. J. Forse, Jr. 
1980. Effects of pocket gopher mounds on plant 
production in shortgrass prairie ecosystems. The 
Southwestern Naturalist, 25:215-224. 

GRANT, W.E., N. R. FRENCH, AND D. M. Swirr. 1977. 
Response of a small mammal community to water 
and nitrogen treatments in a short-grass prairie eco- 
system. Journal of Mammalogy, 58:637-652. 

GRANT, W. E., E. C. Birney, N. R. FRENCH, AND D. 
M. Swirr. 1982. Structure and productivity of 
grassland small mammal communities related to 
grazing-induced changes in vegetative cover. Journal 
of Mammalogy, 63:248-260. 

GRINNELL, J. 1904. The origin and distribution of the 
chestnut-backed chickadee. The Auk, 21:364—382. 
1908. The biota of the San Bernardino 
Mountains. University of California Publications in 

Zoology, 5:1-170. 

. 1914. An account of the mammals and birds 

of the lower Colorado Valley. University of Califor- 

nia Publications in Zoology, 12:51-294. 

1917a. The niche-relationships of the Cali- 

fornia thrasher. The Auk, 34:427-433. 

1917b. Field test of theories concerning dis- 

tributional control. The American Naturalist, 51:115- 

128. 


1928. The presence and absence of animals. 
University of California Chronicle, 30:429-450 (Re- 
printed in: Joseph Grinnell’s Philosophy of Nature, 
Berkeley, 1943. Pp. 187-208). 

GRINNELL, J., AND H. S. SwArtH. 1913. An account 
of the birds and mammals of the San Jacinto area 
of southern California, with remarks upon the be- 
havior of geographic races on the margins of their 
habitats. University of California Publications in Zo- 
ology, 10:197-406. 

GrRoDZINSKI, W. 1971. Energy flow through popu- 
lations of small mammals in the Alaska taiga forest. 
Acta Theriologica, 16:231-276. 

GRODZINSKI, W., AND N. R. FRENCH. 1983. Produc- 
tion efficiency in small mammal populations. Oec- 
ologia, 56:41-49. 

GRODZINSKI, W., AND A. GORECKI. 1967. Daily en- 
ergy budgets of small rodents. Pp. 295-314, in Sec- 
ondary productivity of terestrial ecosystems (K. Pe- 
trusewicz, ed.). 2 vol. Panstwowe Wydawnictwo 
Naukowe, Warsaw, Poland, 879 pp. 

Harner, M.S. 1977. Density and diversity in Mojave 
Desert rodent and shrub communities. The Journal 
of Animal Ecology, 46:925-938. 

Hairston, N. G., F. E. SMITH, AND L. B. SLOBODKIN. 
1960. Community structure, population control and 
competition. The American Naturalist, 94:421-425. 

Hatt, A. S., J. A. STANFORD, AND F. R. HAuver. 1992. 
The distribution and abundance of organisms as a 
consequence of energy balances along multiple en- 
vironmental gradients. Oikos, 65:377-390. 

HALLETT, J. G., AND S. L. Primm. 1970. Direct esti- 
mation of competition. The American Naturalist, 
113:593-600. 

HALLETT, J. G., M. A. O’CONNELL, AND R. L. 
Honeycutt. 1983. Competition and habitat se- 
lection: test ofa theory using small mammals. Oikos, 
40:175-181. 

Han.ey, T. A. 1982. The nutritional basis for food 


COMMUNITIES AND ECOSYSTEMS ST 


selection by ungulates. Journal of Range Manage- 
ment, 35:146-151. 

HarrIson, S., S. J. Ross, AND J. H. LAwron. 1992. 
Beta diversity on geographic gradients in Britain. 
The Journal of Animal Ecology, 61:151-158. 

Harvey, P. H., R. K. COLWELL, J. W. SILVERTON, AND 
R. M. May. 1983. Null models in ecology. Annual 
Review of Ecology and Systematics, 14:189-212. 

Hatt, R.T. 1932. The vertebral column of ricochetal 
rodents. Bulletin American Museum of Natural His- 
tory, 63:599-738. 

Hawkins, C. P., AND J. A. MACMAHON. 1989. Guilds: 
the multiple meanings of a concept. Annual Review 
of Entomology, 34:423-451. 

Heaney, L. R. 1991. An analysis of patterns of dis- 
tribution and species richness among Philippine fruit 
bats (Pteropodidae). Bulletin American Museum of 
Natural History, 206:145-167. 

HEITHAUuS, E. R., T. H. FLEMING, AND P. A. OPLER. 
1975. Foraging patterns and resource utilization in 
seven species of bats in a seasonal tropical forest. 
Ecology, 56:841-854. 

HENDERSON, M. T., G. MERRIAM, AND J. WEGNER. 
1985. Patchy environments and species survival: 
chipmunks in an agricultural mosaic. Biological 
Conservation, 31:95-105. 

HENEIN, K., AND G. MERRIAM. 1990. The elements 
of connectivity where corridor quality is variable. 
Landscape Ecology, 4:157-170. 

Hosrook, S. J. 1979. Habitat utilization, compet- 
itive interactions and coexistence of three species of 
cricetine rodents in east-central Arizona. Ecology, 
60:758-769. 

Ho Luna, C. S. 1959. The components of predation 
as revealed by a study of small mammal predation 
in the European pine sawfly. Canadian Entomolo- 
gist, 91:293-320. 

Ho uister, N. 1920. Editorial comment. Journal of 
Mammalogy, 1:108-110. 

Hornocker, M. G. 1970. An analysis of mountain 
lion predation upon mule deer and elk in the Idaho 
Primitive Area. Wildlife Monographs, 21:1-39. 

Howe, H. F. 1980. Monkey dispersal and waste ofa 
neotropical fruit. Ecology, 61:944-959. 

Howe tL, D. J., AND B.S. RotH. 1981. Sexual repro- 
duction in agaves: the benefits of bats; the cost of 
semelparous advertising. Ecology, 62:1-7. 

Humpnurey, S. R., F. J. BONACCORSO, AND T. L. ZINN. 
1983. Guild structure of surface-gleaning bats in 
Panama. Ecology, 64:284—294. 

Hunt ty, N. 1991. Herbivores and the dynamics of 
communities and ecosystems. Annual Review of 
Ecology and Systematics, 22:477-504. 

Hunt y, N., ANDR.S.INouyeE. 1987. Small mammal 
populations of an old-field chronosequence: succes- 
sional patterns and association with vegetation. 
Journal of Mammalogy, 68:739-745. 

. 1988. Pocket gophers in ecosystems: patterns 
and mechanisms. BioScience, 38:786-793. 

HutTcuinson, G. E. 1957. Concluding remarks. Cold 
Spring Harbor Symposium on Quantitative Biology, 
22:415-427. 

1959. Homage to Snata Rosalia or why are 

there so many kinds of animals? The American Nat- 

uralist, 93:145-159. 


. 1978. An introduction to population ecology. 
Yale University Press, New Haven, Connecticut, 260 
pp. 

Jaxsic, F. M. 1981. Abuse and misuse of the term 
“guild” in ecological studies. Oikos, 37:397—400. 
Jonasson, S., J. P. BRYANT, F. S. I. CHAPIN, AND M. 
ANDERSSON. 1986. Plant phenols and nutrients in 
relation to variations in climate and rodent grazing. 

The American Naturalist, 128:379-408. 

Jones, J. K., JR. 1991. Genealogy of twentieth-cen- 
tury systematic mammalogists in North America: 
the descendants of Joseph Grinnell. Pp. 48-56, in 
Latin American mammalogy: history, biodiversity, 
and conservation (M. A. Mares and D. J. Schmidly, 
eds.). University of Oklahoma Press, Norman, 468 


pp. 

JunGc, H. J. G., AND G. O. Batzut. 1981. Nutritional 
ecology of microtine rodents: effects of plant extracts 
on the growth of Arctic microtines. Journal of Mam- 
malogy, 62:286-292. 

KAJAK, Z., AND E. PlEcZYNSKA. 1981. Poland. Pp. 
287-302, in Handbook of contemporary develop- 
ments in world ecology (E. J. Kormondy and J. F. 
McCormick, eds.). Greenwood Press, Westport, 
Connecticut, 776 pp. 

Karr, J. R., AND F. C. JAMes. 1975. Eco-morpho- 
logical configurations and convergent evolution in 
species and communities. Pp. 258-291, in Ecology 
and evolution of communities (M. Cody and J. Di- 
amond, eds.). Harvard University Press, Cambridge, 
Massachusetts, 543 pp. 

KENAGY, G. J. 1973. Daily and seasonal patterns of 
activity and energetics in a heteromyid rodent com- 
munity. Ecology, 54:1201-1219. 

KENDEIGH, S.C. 1974. Ecology, with special reference 
to animals and man. Prentice-Hall, Inc., Englewood 
Cliffs, New Jersey, 474 pp. 

KERLEY, G. I. H., AND T. ErAsmus. 1991. What do 
mice select for in seeds? Oecologia, 86:26 1-267. 
Kipp, N. A. C., AND G. B. Lewis. 1987. Can verte- 
brate predators regulate their prey? A reply. The 

American Naturalist, 130:448-453. 

Kina, C. M., AND P. J. Moors. 1979. On co-exis- 
tence, foraging strategy and the biogeography of wea- 
sels and stoats (Mustela nivalis and M. erminea) in 
Britain. Oecologia, 39:131-150. 

KIRKLAND, G. L., JR. 1991. Competition and coex- 
istence in shrews (Insectivora: Soricidae). Pp. 15-22, 
in The biology of the Soricidae (J. S. Findley and T. 
L. Yates, eds.). Special Publication, The Museum of 
Southwestern Biology, 1:1-91. 

KLENNER, W., AND C. J. Kress. 1991. Red squirrel 
population dynamics. I. The effect of supplemental 
food on demography. The Journal of Animal Ecol- 
ogy, 60:961-978. 

KORMONDY, E. J., AND J. F. MCCorMIckK (EDS.). 1981. 
Handbook of contemporary developments in world 
ecology. Greenwood Press, Westport, Connecticut, 
776 pp. 

KorpiMAKI, E. 1993. Regulation of multiannual vole 
cycles by density-dependent avian and mammalian 
predation? Oikos, 66:359-363. 

Kot Ler, B. P. 1984. Risk of predation and the struc- 
ture of desert rodent communities. Ecology, 65:689- 
701. 


ay 2 MARES AND CAMERON 


1989. Temporal variation in the structure of 
a desert rodent community. Pp. 127-139, in Patterns 
in the structure of mammalian communities (D. W. 
Morris, Z. Abramsky, B. J. Fox, and M. R. Willig, 
eds.). Texas Tech University Press, Lubbock, 266 


KoTLer, B. P., AND J. S. BRown. 1988. Environ- 
mental heterogeneity and the coexistence of desert 
rodents. Annual Review of Ecology and Systematics, 
19:281-308. 

Kot Ler, B. P., ANDR. D. Hott. 1989. Predation and 
competition: the interaction of two types of species 
interactions. Oikos, 54:256-260. 

Kot er, B. P., J. S. Brown, R. J. SMitH, AND W. O. 
Wirtz, II. 1988. The effects of morphology and 
body size on rates of owl predation on desert rodents. 
Oikos, 53:145-152. 

Kuropat, P., AND J. P. BRYANT. 1983. Digestibility 
of caribou summer forage in Arctic Alaska in relation 
to nutrient, fiber, and phenolic constituents. Acta 
Zoologica Fennica, 175:51-52. 

LACHER, T. E., JR., AND C. J. R. ALHo. 1989. The 
structure of a small mammal community in a central 
Brazilian savanna. Pp. 137-162, in Advances in neo- 
tropical mammalogy (K. H. Redford and J. F. Ei- 
senberg, eds.). Sandhill Crane Press, Gainesville, 
Florida, 614 pp. 

LACHER, T. E., JR., AND M. A. Mares. 1986. Neo- 
tropical mammal communities: an appraisal. Re- 
vista Chilena de Historia Natural, 59:121-134. 

LAURANCE, W. F. 1991. Edge effects in tropical forest 
fragments: application of a model for the design of 
nature reserves. Biological Conservation, 57:205-219. 

LipIcKER, W. Z., Jk. 1989. Impacts of non-domes- 
ticated vertebrates on California grasslands. Pp. 135- 
150, in Grassland structure and function: California 
annual grassland (L. F. Huenneke and H. Mooney, 
eds.). Kluwer Academic Publishers, Dordrecht, 
Netherlands. 

LINDEMAN, R. L. 1942. The trophic-dynamic aspect 
of ecology. Ecology, 23:399-418. 

LoMoLino, M. V. 1989. Interpretations and com- 
parisons of constants in the species-area relationship: 
an additional caution. The American Naturalist, 133: 
277-280. 

LONGLAND, W. S., AND M. V. Price. 1991. Direct 
observations of owls and heteromyid rodents: can 
predation risk explain microhabitat use? Ecology, 
72:2261-2273. 

LuKE, C. 1986. Convergent evolution of lizard toe 
fringes. Biological Journal of the Linnean Society, 
27:1-16. 

Mapry, T., J. HUNZIKER, AND D. R. DIFEo, Jr. (EDs.). 
1977. Creosote bush: biology and chemistry of Lar- 
rea in New World deserts. Dowden, Hutchinson and 
Ross, East Stroudsburg, Pennsylvania, 284 pp. 

MacMauon, J. A. 1976. Species and guild similarity 
of North American desert mammal faunas: a func- 
tional analysis of communities. Pp. 133-148, in Evo- 
lution of desert biota (D. W. Goddall, ed.). Univer- 
sity of Texas, Austin, 250 pp. 

MacManuon, J. A., D. J. ScHmmpr, D. C. ANDERSON, 
K. G. SMITH, AND R. L. BAyn, JR. 1981. An or- 
ganism-centered approach to some community and 
ecosystem concepts. Journal of Theoretical Biology, 
88:287-307. 


Ma.mautsT, M. G. 1985. Character displacement and 
biogeography of the pygmy shrew in Northern Eu- 
rope. Ecology, 66:372-377. 

Mares, M. A. 1975. South American mammal zoo- 
geography: evidence from convergent evolution in 
desert rodents. Proceedings of the National Acade- 
my of Science, USA, 72:1702-1706. 

1976. Convergent evolution of desert ro- 

dents: multivariate analysis and zoogeographic im- 

plications. Paleobiology, 2:39-64. 

1980. Convergent evolution among desert 

rodents: a global perspective. Bulletin of Carnegie 

Museum of Natural History, 16:1-51. 

1983. Desert rodent adaptation and com- 

munity structure. Great Basin Naturalist Memoirs, 

7:30-43. 

1992. Neotropical mammals and the myth 

of Amazonian biodiversity. Science, 255:976-979. 

1993a. Desert rodents, seed consumption, 

and convergence: the evolutionary shuffling of ad- 

aptations. BioScience, 43:372-379. 

1993b. Heteromyids and their ecological 
counterparts: a pandesertic view of rodent ecology 
and evolution. Pp. 652-714, in Biology of the Het- 
eromyidae (H. H. Genoways and J. H. Brown, eds.). 
Special Publication, The American Society of Mam- 
malogists, 10:1-719. 

Mares, M. A., AND J. K. BRAUN. 1986. An inter- 
national survey of the popular and technical litera- 
ture of mammalogy. Annals of Carnegie Museum, 
55:149-209. 

Mares, M. A., AND T. E. LACHER, JR. 1987. Ecolog- 
ical, morphological and behavioral convergence in 
rock-dwelling mammals. Current Mammalogy, 
1:307-348. 

Mares, M. A., AND R. A. OJEDA. 1982. Patterns of 
diversity and adaptation in South American hystri- 
cognath rodents. Pp. 393-432, in Mammalian bi- 
ology in South America (M. A. Mares and H. H. 
Genoways, eds.). Pymatuning Laboratory of Ecol- 
ogy, Special Publication 6, Linesville, Pennsylvania, 
539 pp. 

Mares, M. A., AND M. L. RosENzweIG. 1978. Gra- 
nivory in North and South American deserts: ro- 
dents, birds and ants. Ecology, 59:235-241. 

Mares, M. A., AND D. F. WitiiaAms. 1977. Experi- 
mental support for food particle size resource allo- 
cation in heteromyid rodents. Ecology, 58:1186- 
1190. 

Mares, M. A., M. WATSON, AND T. E. LACHER, JR. 
1976. Home range perturbations in 7amias striatus: 
food supply as a determinant of home range and 
density. Oecologia, 25:1-12. 

Mares, M. A., M. R. WILLIG, AND T. E. LACHER, JR. 
1985. The Brazilian Caatinga in South American 
zoogeography: tropical mammals in a dry region. 
Journal of Biogeography, 12:57-69. 

Mares, M. A., T. E. LACHER, Jr., M. R. WILLIG, N. 
A. BiTrar, R. ADAmMs, A. KLINGER, AND D. TAZIK. 
1982. An experimental analysis of social spacing in 
Tamias striatus. Ecology, 63:267-273. 

Marauts, R. J., AND G. O. Batzui. 1989. Influence 
of chemical factors on palatability of forage to voles. 
Journal of Mammalogy, 70:503-511. 

MASCHINSKI, J., AND T. G. WHITHAM. 1989. Thecon- 
tinuum of plant responses to herbivory: the influence 


COMMUNITIES AND ECOSYSTEMS 373 


of plant association, nutrient availability, and tim- 
ing. The American Naturalist, 134:1-19. 

May, R. M. 1984. An overview: real and apparent 
patterns in community structure. Pp. 3-16, in Eco- 
logical communities: conceptual issues and the ev- 
idence (D. R. Strong, Jr., et al., eds.). Princeton Uni- 
versity Press, Princeton, New Jersey, 613 pp. 

McCoy, E. D., AND E. F. CONNor. 1980. Latitudinal 
gradients in the species diversity of North American 
mammals. Evolution, 34:193-203. 

MclIntosn, R. P. 1975. H. A. Gleason, “‘individu- 
alistic ecologist,” 1882-1975: his contributions to 
ecological theory. Bulletin of the Torrey Botanical 
Club, 102:263-273. 

1980. The background and some current 

problems of theoretical ecology. Synthese, 43:195- 

255. 


1985. The background of ecology: concept 
and theory. Cambridge University Press, Cam- 
bridge. 383 pp. 

McKenzie, N. L., AND A. N. Start. 1989. Structure 
of bat guilds in mangroves: environmental distur- 
bance and determinism. Pp. 167-178, in Patterns in 
the structure of mammalian communities (D. W. 
Morris, Z. Abramsky, B. J. Fox, and M. R. Willig, 
eds.). Texas Tech University Press, Lubbock, 266 


pp. 

M’CLoskEy, R. T. 1978. Niche separation and as- 
sembly in four species of Sonoran desert rodents. 
The American Naturalist, 1 12:683-694. 

1980. Spatial patterns in sizes of seeds col- 
lected by four species of heteromyid rodents. Ecol- 
ogy, 61:486-489. 

McNas, B. K. 1963. A model of the energy budget 
of a wild mouse. Ecology, 44:521-532. 

. 1971. The structure of tropical bat faunas. 

Ecology, 52:352-358. 

. 1991. The energy expenditure of shrews. Pp. 
35-45, in The biology of the Soricidae (J. S. Findley 
and T. L. Yates, eds.). Special Publication, The Mu- 
seum of Southwestern Biology, 1:1-91. 

McNas, B. K., AND P. Morrison. 1963. Body tem- 
perature and metabolism in subspecies of Peromys- 
cus from arid and mesic environments. Ecological 
Monographs, 33:63-82. 

McNauauton, S. J. 1976. Serengeti migratory wil- 
debeest: facilitation of energy flow by grazing. Sci- 
ence, 191:92-94, 

1977. Diversity and stability of ecological 

communities: a comment on the role of empiricism 

in ecology. The American Naturalist, 111:515-525. 

1985. Ecology of a grazing ecosystem: the 

Serengeti. Ecological Monographs, 55:259-294. 

1986. On plants and herbivores. The Amer- 
ican Naturalist, 128:765-770. 

McNauGutTon, S. J., AND N. J. Georaiapis. 1986. 
Ecology of African grazing and browsing mammals. 
Annual Review of Ecology and Systematics, 17:39- 
65. 

Mecu, L. D. 1966. The wolves of Isle Royale. Fauna 
of the National Parks of the United States. Fauna 
Series, 7:1-210. 

MERRIAM, C. H. 1890. Results of a biological survey 
of the San Francisco Mountain region and the desert 
of the Little Colorado, Arizona. North American 
Fauna, 3:1-136. 


1892. The geographic distribution of life in 
North America. Proceedings of the Biological So- 
ciety of Washington, 7:1-64. 

1894. Laws of temperature control of the 

geographical distribution of terrestrial animals and 

plants. National Geographic Magazine, 6:229-238. 

. 1898. Life zones and crop zones. United States 
Department of Agriculture Bulletin 10, 79 pp. 

MERRIAM, G., AND A. LANOuE. 1990. Corridor use 
by small mammals: field measurement for three ex- 
perimental types of Peromyscus leucopus. Landscape 
Ecology, 4:123-131. 

Merritt, J. F.,AND J. M. Merritr. 1978. Population 
ecology and energy relationships of Clethrionomys 
gapperi in a Colorado subalpine forest. Journal of 
Mammalogy, 59:576-598. 

MEseRVE, P.L. 1981. Resource partitioning in a Chil- 
ean semi-arid small mammal community. The Jour- 
nal of Animal Ecology, 50:745-758. 

MICHELL,R.,R.A.MAYER,ANDJ. DowNHOwWER. 1976.An 
evaluation of three biome programs. Science, 192: 
859-865. 

Miiier, R. S.. 1967. Pattern and process in compe- 
tition. Advances in Ecological Research, 4:1-74. 
MITCHELL, W. A., Z. ABRAMSKY, B. P. KoTLer, B. 
PINSHOw, AND J. S. BRown. 1990. The effect of 
competition on foraging activity in desert rodents: 

theory and experiments. Ecology, 71:844-854. 

Montcomery, G. G. 1989. Population parameters, 
spatial division, and niche breadth in two Apodemus 
species sharing a woodland habitat. Pp. 45-58, in 
Patterns in the structure of mammalian communi- 
ties (D. W. Morris, Z. Abramsky, B. J. Fox, and M. 
R. Willig, eds.). Texas Tech University Press, Lub- 
bock, 266 pp. 

MontTGOMERY, G. G., AND M. E. SunquistT. 1975. 
Impact of sloths on neotropical forest energy flow 
and nutrient cycling. Pp. 69-98, in Tropical ecolog- 
ical systems (F. B. Golley and E. Medina, eds.). 
Springer-Verlag, New York, 398 pp. 

Morris, D. W. 1987. Spatial scale and the cost of 
density-dependent habitat selection. Evolutionary 
Ecology, 1:379-388. 

1989. The effect of spatial scale on patterns 
of habitat use: red-backed voles as an empirical mod- 
el of local abundance for northern mammals. Pp. 
23-34, in Patterns in the structure of mammalian 
communities (D. W. Morris, Z. Abramsky, B. J. Fox, 
and M. R. Willig, eds.). Texas Tech University Press, 
Lubbock, 266 pp. 

Myrcua, A. 1975. Bioenergetics of an experimental 
population and individual laboratory mice. Acta 
Theriologica, 20:175-226. 

Nevo, E. 1979. Adaptive convergence and diver- 
gence of subterranean mammals. Annual Review 
Ecology and Systematics, 10:269-308. 

O’Dowp, D. J., AND M. E. Hay. 1980. Mutualism 
between harvester ants and a desert ephemeral: seed 
escape from rodents. Ecology, 61:531—540. 

Opum, E. P. 1945. The concept of the biome as ap- 
plied to the distribution of North American birds. 
The Wilson Bulletin, 57:191-201. 

. 1953. Fundamentals of ecology. W. B. Saun- 

ders Company, Philadelphia. 384 pp. 

. 1971. Fundamentals of ecology. Third ed. 

W. B. Saunders Company, Philadelphia, 574 pp. 


374 MARES AND CAMERON 


OpuM, E. P., C. E. CONNELL, AND L. B. DAVENPORT. 
1962. Population energy flow of three primary con- 
sumer components of old-field ecosystems. Ecology, 
43:88-96. 

Opum, H. T. 1957. Trophic structure and produc- 
tivity of Silver Springs, Florida. Ecological Mono- 
graphs, 27:55-115. 

OrRIANS, G. H., ANDO. T. SOLBRIG. 1977. Convergent 
evolution in warm deserts. Dowden, Hutchinson and 
Ross, East Stroudsburg, Pennsylvania, 333 pp. 

Oscoop, W.H. 1943. Clinton Hart Merriam— 1855- 
1942. Journal of Mammalogy, 24:421-436. 

Owen, J.G. 1988. On productivity as a predictor of 
rodent and carnivore diversity. Ecology, 69:1161- 
1165. 


1990a. An analysis of the spatial structure 

of mammalian distribution patterns in Texas. Ecol- 

ogy, 71:1823-1832. 

. 19906. Patterns of mammalian species rich- 
ness in relation to temperature, productivity, and 
variance in elevation. Journal of Mammalogy, 71: 
1-13. 

OweEN-SMITH, N. 1989. Morphological factors and 
their consequences for resource partitioning among 
African savanna ungulates: a simulation modelling 
approach. Pp. 155-165, im Patterns in the structure 
of mammalian communites (D. W. Morris, Z. 
Abramsky, B. J. Fox, and M. R. Willig, eds.). Texas 
Tech University Press, Lubbock, 266 pp. 

PAGEL, M. D., R. M. May, AND A. R. CoL.iz. 1991. 
Ecological aspects of the geographic distribution and 
diversity of mammalian species. The American Nat- 
uralist, 137:791-815. 

Paice, K. N. 1992. Overcompensation in response 
to mammalian herbivory: from mutualistic to an- 
tagonistic interactions. Ecology, 73:2076-2085. 

PaiGE, K. N., AND T. G. WHITHAM. 1987. Overcom- 
pensation in response to mammalian herbivory: the 
advantage of being eaten. The American Naturalist, 
129:407-416. 

PEARSON, O. P. 1960. The oxygen consumption and 
bioenergetics of harvest mice. Physiological Zoolo- 
gy, 33:152-160. 

. 1964. Carnivore-mouse predation: an ex- 
ample of its intensity and bioenergetics. Journal of 
Mammalogy, 45:177-188. 

1971. Additional measurements of the im- 
pact of carnivores on California voles (Microtus cal- 
ifornicus). Journal of Mammalogy, 52:41-49. 

PearsT, D. R. 1989. Species interactions in a suc- 
cessional grassland. III. Effect of canopy gaps, gopher 
mounds, and grazing on colonization. Journal of 
Ecology, 77:267-289. 

PETERSEN, M. K. 1970. Competition between the cot- 
ton rats, Sigmodon fulviventer and S. hispidus. Ph.D. 
dissert., Michigan State University, East Lansing, 
110 pp. 

1973. Interactions between the cotton rats, 
Sigmodon fulviventer and S. hispidus. The American 
Midland Naturalist, 90:319-333. 

PETRUSEWICZ, K. (ED.). 1967. Secondary productivity 
of terrestrial ecosystems, 2 vol. Panstwowe Wy- 
dawnictwo Naukowe, Warsaw, Poland, 879 pp. 

PIANKA, E. R. 1969. Habitat specificity, speciation, 
and species density in Australian desert lizards. Ecol- 
ogy, 50:498-502. 


1973. The structure of lizard communities. 
Annual Review of Ecology and Systematics, 4:53- 
74, 


. 1974. Niche overlap and diffuse competition. 
Proceedings of the National Academy of Sciences, 
USA, 71:2141-2145. 

1975. Niche relations of desert lizards. Pp. 
292-314, in Ecology and evolution of communities 
(M. Cody and J. Diamond, eds.). Harvard Univer- 
sity Press, Cambridge, Massachusetts, 543 pp. 
1985. Some intercontinental comparisons of 
desert lizards. National Geographic Research, 1:490- 
504. 


1986. Ecology and natural history of desert 
lizards. Princeton University Press, Princeton, New 
Jersey, 208 pp. 

PIMENTEL, D. 1988. Herbivore population feeding 
pressure on plant hosts: feedback evolution and host 
conservation. Oikos, 53:289-302. 

Pim, S. L., J. H. LAwTon, AND J. E. COHEN. 1991. 
Food web patterns and their consequences. Nature, 
350:669-674. 

Piatt, W. J. 1975. The colonization and formation 
of equilibrium plant species associations on badger 
disturbances in a tall-grass prairie. Ecological Mono- 
graphs, 45:285-305. 

Porter, J. H., AND R. D. DUEsER. 1981. Niche over- 
lap and competition in an insular small mammal 
fauna: a test of the niche overlap hypothesis. Oikos, 
39:228-236. 

Price, M. V., AND K. A. KRAMER. 1984. On mea- 
suring microhabitat affinities with special reference 
to small mammals. Oikos, 42:349-354. 

Price, M. V., W. S. LONGLAND, AND R. L. GOLDINGAY. 
1991. Niche relationship of Dipodomys agilis and 
D. stephensi: two sympatric kangaroo rats of similar 
size. The American Midland Naturalist, 126:172- 
186. 

PULLIAM, H. R., AND M. R. BRAND. 1975. The pro- 
duction and utilization of seeds in plains grassland 
of southeastern Arizona. Ecology, 56:1158-1166. 

RaMALAy, R. 1940. The growth of a science. Uni- 
versity of Colorado Studies, General Series A, 26:3- 
14. 

RANDOLPH, J. C., G. N. CAMERON, AND J. A. WRAZEN. 
1991. Dietary choice of generalist grassland herbi- 
vore, Sigmodon hispidus. Journal of Mammalogy, 
72:300-3 13. 

REICHMAN, O. J., AND S. C. SmitH. 1985. Impact of 
pocket gopher burrows on overlying vegetation. 
Journal of Mammalogy, 66:720-725. 

REICHMAN, O. J., AND S. C. SMitH. 1991. Responses 
to simulated leaf and root herbivory by a biennial, 
Tragopogon dubius. Ecology, 72:116-124. 

REICHMAN, O. J., J. H. BENEDIX, JR., AND T. R. SEA- 
STEDT. 1993. Distinct animal-generated edge ef- 
fects in a tallgrass prairie community. Ecology, 74: 
1281-1284. 

Root, R. B. 1967. The niche exploitation pattern of 
the blue-gray gnatcatcher. Ecological Monographs, 
37:31 7=350) 

RosENzweIG, M. L. 1966. Community structure in 
sympatric Carnivora. Journal of Mammalogy, 47: 
602-612. 

. 1968. Net primary productivity of terrestrial 


COMMUNITIES AND ECOSYSTEMS a15 


communities: prediction from climatological data. 

The American Naturalist, 102:67-74. 

1969. Why the prey curve has a hump. The 

American Naturalist, 103:81-87. 

. 1973. Evolution of the predator isocline. 

Evolution, 27:84-94. 

1989. Habitat selection, community orga- 

nization, and small mammal studies. Pp. 5-21, in 

Patterns in the structure of mammalian communi- 

ties (D. W. Morris, Z. Abramsky, B. J. Fox, and M. 

R. Willig, eds.). Texas Tech University Press, Lub- 

bock, 266 pp. 

1993. Species diversity gradients: we know 
more and less than we thought. Journal of Mam- 
malogy, 73:715-730. 

ROSENZWEIG, M. L., AND R. H. MACARTHUR. 1963. 
Graphical representation and stability conditions of 
predator-prey interactions. The American Natural- 
ist, 97:209-223. 

ROSENZWEIG, M. L., Z. ABRAMSKY, AND S. BRAND. 
1984. Estimating species interactions in heteroge- 
neous environments. Oikos, 43:329-340. 

SAzIMA, M., AND I. SAzIMA. 1978. Bat pollination of 
the passion flower, Passiflora mucronata, in south- 
eastern Brazil. Biotropica, 10:100-109. 

SCHLUTER, D. 1986. Tests for similarity and conver- 
gence of finch communities. Ecology, 67:1073-1083. 

1990. Species-for-species marching. The 
American Naturalist, 136:560-568. 

SCHMIDT-NIELSEN, K. 1964. Desert animals. Clar- 
endon Press, Oxford, England, 277 pp. 

SCHNELL, J. H. 1968. The limiting effects of natural 
predation on experimental cotton rat populations. 
The Journal of Wildlife Management, 32:698-711. 

SCHOENER, T. W. 1974. Competition and the form 
of habitat shift. Theoretical Population Biology, 
6:265-307. 

1983. Field experiments on interspecific 

competition. The American Naturalist, 122:240-285. 

1985. Some comments on Connell’s and my 

reviews of field experiments on interspecific com- 

petition. The American Naturalist, 125:730-740. 

1986a. Mechanistic approaches to commu- 

nity ecology: a new reductionism? American Zool- 

ogist, 26:81-106. 

1986b. When should a field experiment be 
counted? A reply to Galindo and Krebs. Oikos, 46: 
119-121. 

ScHuLtTz, A. M. 1964. The nutrient-recovery hy- 
pothesis for Arctic microtine cycles. Pp. 57-68, in 
Grazing in terrestrial and marine environments (D. 
Crisp, ed.). British Ecological Society Symposium 4. 
Blackwell Scientific Publishers, Oxford, 322 pp. 

ScHuM, M. 1984. Phenetic structure and species rich- 
ness in North and Central American bat faunas. 
Ecology, 65:1315-1324. 

SEAGLE, S. W., AND G. F. MCCRAKEN. 1986. Species 
abundance, niche position, and niche breadth for five 
terrestrial animal assemblages. Ecology, 67:816-818. 

SEAGLE, S. W., ANDS. J. MCNAUGHTON. 1992. Spatial 
variation in forage nutrient concentrations and the 
distribution of Serengeti grazing ungulates. Land- 
scape Ecology, 7:229-241. 

Semper, K. 1881. Animal life as affected by the nat- 
ural conditions of existence. Appleton, New York. 

SHELFORD, V.E. 1913. Animal communities in tem- 


perate North America. University of Chicago Press, 

Chicago, 368 pp. 

. 1926. Naturalists’ guide to the Americas. 
Williams & Wilkins Company, Baltimore, Mary- 
land, 761 pp. 

SHELFORD, V. E., AND A. C. Twomey. 1941. Tundra 
animal communities in the vicinity of Churchill, 
Manitoba. Ecology, 22:47-69. 

Sin, A., P. CRowLey, M. McPEEk, J. PETRANKA, AND 
K.STROHMEIER. 1985. Predation, competition, and 
prey communities: a review of field experiments. 
Annual Review of Ecology and Systematics, 16:269- 
S12 

SIMBERLOFF, D., AND T. DAYAN. 1991. The guild con- 
cept and the structure of ecological communities. 
Annual Review of Ecology and Systematics, 22:1 15- 
144. 

SIMENSTAD, C. A., J. A. Estes, AND K. W. KENYON. 
1978. Aleuts, sea otters, and alternate stable-state 
communities. Science, 200:403-411. 

Simpson, B. B. 1977. Mesquite: its biology in two 
desert ecosystems. Dowden, Hutchinson and Ross, 
East Stroudsburg, Pennsylvania, 250 pp. 

Simpson, B. B., AND J. L. NEFF. 1981. Floral rewards: 
alternatives to pollen and nectar. Annals of the Mis- 
souri Botanical Garden, 68:301-322. 

Simpson, G.G. 1964. Species density of North Amer- 
ican recent mammals. Systematic Zoology, 13:57- 
73% 

SmncLarr, A. R. E., C. J. KREBS, AND M. SmitH. 1982. 
Diet quality and food limitation in herbivores: the 
case of the snowshoe hare. Canadian Journal of Zo- 
ology, 60:889-897. 

Sincuair, A. R. E., P. D. OLSEN, AND T. D. REDHEAD. 
1990. Can predators regulate small mammal pop- 
ulations? Evidence from house mouse outbreaks in 
Australia. Oikos, 59:382-392. 

Sinciair, A. R. E., C. J. Kress, J. N. M. SMITH, AND 
S. BouTin. 1988. Population biology of snowshoe 
hares. III. Nutrition, plant secondary compounds and 
food limitation. The Journal of Animal Ecology, 57: 
787-806. 

SyOrs, H. 1981. Sweden. Pp. 303-312, in Handbook 
of contemporary developments in world ecology (E. 
J. Kormondy and J. F. McCormick, eds.). Green- 
wood Press, Westpoprt, Connecticut, 776 pp. 

SLOBODCHIKOFF, C. N., AND W. C. SCHULTz. 1980. 
Measures of niche overlap. Ecology, 61:1051-1055. 

SLOBODKIN, L. B. 1961. Growth and regulation of 
animal populations. Holt, Rinehart, and Winston, 
New York, 184 pp. 

. 1962. Energy in animal ecology. Advances 

in Ecological Research, 4:69-101. 

. 1987. How to be objective in community 
studies. Pp. 93-108, in Neutral models in biology 
(M. H. Nitecki and A. Hoffman, eds.). Oxford Uni- 
versity Press, Oxford, 166 pp. 

Sty, G. R. 1976. Small mammal succession on strip- 
mined land in Vigo County, Indiana. The American 
Midland Naturalist, 95:257-267. 

SMARTT, R. A. 1978. A comparison of ecological and 
morphological overlap in a Peromyscus community. 
Ecology, 59:216-220. 

Smartt, R. A., ANDC. LEMEN. 1980. Intrapopulation 
morphological variation as a predictor of feeding 


376 MARES AND CAMERON 


behavior in deermice. The American Naturalist, 116: 
891-894. 

Smitu, C. C. 1970. The coevolution of pine squirrels 
(Tamiasciurus) and conifers. Ecological Mono- 
graphs, 40:349-371. 

SmiTH, C. C., AND R. P. BALDA. 1979. Competition 
among insects, birds and mammals for conifer seeds. 
The American Zoologist, 19:1965-1083. 

SmitH, M. F. 1981. Relationships between genetic 
variability and niche dimensions among coexisting 
species of Peromyscus. Journal of Mammalogy, 62: 
273-285. 

SMYTHE, N. 1986. Competition and resource parti- 
tioning in the guild of neotropical terrestrial frugiv- 
orous mammals. Annual Review of Ecology and Sys- 
tematics, 17:169-188. 

SnypDER, M. A. 1992. Selective herbivory by Abert’s 
squirrel mediated by chemical variability in pon- 
derosa pine. Ecology, 73:1730-1741. 

SoHOLT, L. 1973. Consumption of primary produc- 
tion by a population of kangaroo rats (Dipodomys 
merriami) in the Mojave Desert. Ecological Mono- 
graphs, 43:357-376. 

Spatz, G., AND D. MUELLER-DomBols. 1973. The 
influence of feral goats on Koa tree reproduction in 
Hawaii Volcanoes National Park. Ecology, 54:870- 
877. 

STAPANIAN, M. A., AND C. C. SmitH. 1978. A model 
for seed scatter-hoarding: coevolution of fox squir- 
rels and black walnuts. Ecology, 59:884-896. 

1986. How fox squirrels influence the inva- 
sion of prairies by nut-bearing tress. Journal of 
Mammalogy, 67:326-332. 

STEBBINS, G. L. 1962. Toward better international 
cooperation in the life sciences. Plant Science Bul- 
letin, 8:1-10. 

STERLING, K. B. 1977. Last of the naturalists: the 
career of C. Hart Merriam. Arno Press, New York, 
472 pp. 

STRONG, D. R., D. SIMBERLOFF, L. G. ABELE, AND A. 
B. THIsTLe. 1984. Ecological communities: con- 
ceptual issues and the evidence. Princeton Univer- 
sity Press, Princeton, New Jersey, 613 pp. 

SUKACHEV, V. N. 1958. On the principles of genetic 
classification in biocenology. Ecology, 39:364—-367. 

SULLIVAN, T. P., D. S. SULLIVAN, AND C. J. KREBS. 
1983. Demographic responses of a chipmunk (Eu- 
tamias townsendi) population with supplemental 
food. The Journal of Animal Ecology, 52:743-756. 

SUSSMAN, R. W., AND P. H. RAVEN. 1978. Pollination 
by lemurs and marsupials: an archaic coevolutionary 
system. Science, 200:73 1-736. 

Taitt, M. J., AND C. J. Kress. 1983. Predation, cov- 
er, and food manipulations during a spring decline 
of Microtus townsendi. Journal of Animal Ecology, 
52:837-848. 

TANSLEY, A. G. 1935. The use and abuse of vegeta- 
tional concepts and terms. Ecology, 16:284-307. 
TEAL, J. M. 1957. Community metabolism in a tem- 
perate cold spring. Ecological Monographs, 27:283- 

302. 

TILMAN, D. 1983. Plant succession and gopher dis- 
turbance along an experimental gradient. Oecologia, 
60:285-292. 


TROSTEL, K., A. SINCLAIR, C. WALTERS, AND C. KREBS. 
1987. Can predation cause the 10-yr hare cycle? 
Oecologia, 74:185-192. 

TRUSZKOWSKI, J. 1982. The impact of the common 
vole on the vegetation of agroecosystems. Acta The- 
riologica, 27:305-345. 

UNDERWOOD, T. 1986. The analysis of competition 
by field experiments. Pp. 240-268, in Community 
ecology: pattern and process (J. Kikkawa and D. 
Anderson, ed.). Blackwell Scientific Publishers, Ox- 
ford, England, 432 pp. 

VARLEY, G. C. 1975. Should we control the use of 
the word control? Bulletin of the British Ecological 
Society, 6:7. 

VAN Horne, B., ANDR. G. Forp. 1982. Niche breadth 
calculation based on discriminant analysis. Ecology, 
63:1172-1174. 

WAGNER, F.H., ANDL.C.SToDDART. 1972. Influence 
of coyote predation on black-tailed jackrabbit pop- 
ulations in Utah. The Journal of Wildlife Manage- 
ment, 36:329-342. 

Wuicker, A. D., ANDJ.K. DETLING. 1988. Ecological 
consequences of prairie dog disturbances. Bio- 
Science, 38:778-785. 

WHITFORD, W. G., AND Y. STEINBERGER. 1989. The 
long-term effects of habitat modification on a desert 
rodent community. Pp. 33-43, in Patterns in the 
structure of mammalian communities (_D. W. Mor- 
ris, Z. Abramsky, B. J. Fox, and M. R. Willig, eds.). 
Texas Tech University Press, Lubbock, 266 pp. 

WHITTAKER, R. H. 1951. A criticism of the plant 
association and climatic climax concepts. Northwest 
Science, 26:17-31. 

Wiiuic, M. R. 1986. Bat community structure in 
South America: a tenacious chimera. Revista Chi- 
lena de Historia Natural, 59:151-168. 

WILLIG, M. R., AND T. E. LACHER, JR. 1991. Food 
selection of a tropical folivore in relation to leaf- 
nutrient content. Journal of Mammalogy, 72:314—- 
321. 

Wiiuic, M. R., AND M. A. Mares. 1989. A compar- 
ison of bat assemblages from phytogeographic zones 
of Venezuela. Pp. 59-67, in Patterns in the structure 
of mammalian communities (D. W. Morris, Z. 
Abramsky, B. J. Fox, and M. R. Willig, eds.). Texas 
Tech University Press, Lubbock, 266 pp. 

WILLIG, M. R., AND M. P. MouLTon. 1989. The role 
of stochastic and deterministic processes in struc- 
turing Neotropical bat communities. Journal of 
Mammalogy, 70:323-329. 

WILLIc, M. R., AND E. A. SANDLIN. 1991. Gradients 
of species density and turnover in New World bats: 
a comparison of quadrat and band methodologies. 
Pp. 81-96, in Latin American mammalogy: history, 
biodiversity, and conservation (M. A. Mares and D. 
J. Schmidly, eds.). University of Oklahoma Press, 
Norman, 468 pp. 

WILLIG, M. R., AND K. W. SELceR. 1989. Bat species 
density gradients in the New World: a statistical as- 
sessment. Journal of Biogeography, 16:189-195. 

WILLIG, M. R., G. R. CAMILO, AND S. J. NoBLe. 1993. 
Dietary overlap in frugivorous and insectivorous bats 
from edaphic cerrado habitats of Brazil. Journal of 
Mammalogy, 74:117-128. 


NATURAL HISTORY AND EVOLUTIONARY 


ECOLOGY 


JAMES H. BROWN AND DON E. WILSON 


Introduction 


he last 75 years have seen dramatic 
changes in both the theoretical con- 
cepts of ecology and evolution, and in the 
field and laboratory studies that provide the 
empirical basis for theoretical advances. On 
the one hand, there has been a trend toward 
increasing conceptual specialization as the 
broad field of natural history has been sup- 
planted by the specialized study of life his- 
tories, population dynamics, community 
organization, and morphological, physio- 
logical, and behavioral adaptation. On the 
other hand, there has been a corresponding 
trend toward decreasing taxonomic spe- 
cialization as the practitioners of these dis- 
ciplines have chosen to study organisms on 
the basis of their suitability for testing eco- 
logical theory. 

This chapter attempts to describe how 
these trends have influenced the develop- 
ment of North American mammalogy, as 
well as how studies of mammals have con- 
tributed to the theory and data of modern 
ecology and evolutionary biology. When the 
ASM was founded in 1919, many of its 
charter members and earliest recruits in- 
cluded the leading natural historians of the 
early 20th Century (Merriam, Bailey, Jack- 


ag, 


son, Allen, Osgood, Nelson, Goldman). 
Over the 75-year history of the society, 
studies of mammals have continued to play 
key roles as classical natural history has 
evolved into modern evolutionary ecology. 

The history of changes in ecological and 
evolutionary studies of mammals reflect 
more fundamental changes in the devel- 
opment of modern science. As we shall see, 
many of the questions posed by early nat- 
ural historians have not yet been completely 
answered and are still the subject of major 
research programs today. This is not to say 
that there has been no progress. A great deal 
has been learned about how wild mammals 
survive, reproduce, and coexist in diverse 
habitats, and this knowledge frequently 
raises more questions than it answers. The 
questions have become more focused and 
the standards for acceptable answers have 
become more rigorous. New tools, such as 
mathematical models, field experiments, 
and statistical analyses, have been devel- 
oped to facilitate the interplay of theory and 
data. Broad syntheses have been attempted. 
In all of these developments, studies of 
mammals have played major roles. 

For convenience, this history can be di- 


378 BROWN AND WILSON 


vided into several phases. The first, the dis- 
covery phase, began with the earliest studies 
of mammals in the Americas; the last, the 
evolutionary ecology phase, is a major theme 
of contemporary research. 


Discovery Phase 


Humans have always been curious about 
the plants and animals that share their world, 
and they have always had a special interest 
in their nearest relatives, other mammals. 
The earliest humans studied the ecology and 
behavior of mammals out of necessity, be- 
cause different mammal species were im- 
portant food sources, deadly predators, se- 
rious competitors, helpful mutualists, and 
objects of admiration and worship. As mod- 
ern human civilizations developed, they re- 
tained their fascination with the natural 
world and with the lives of their wild mam- 
malian relatives. Mammals figure promi- 
nently in the art and writing of ancient Ori- 
ental, Mediterranean, African, European, 
and American civilizations. As western civ- 
ilization emerged from the middle ages, Eu- 
ropean naturalists such as Linneaus, Cuvier, 
and Buffon began to describe, classify, and 
study the lives of their native mammals. 
These studies received added impetus when 
the voyages of discovery returned from 
around the world bearing specimens of 
amazing new kinds of mammals and other 
living things. 

North American mammalogy began in 
earnest when the newly arrived European 
colonists began to explore the continent and 
assess its natural resources. As with so many 
other human endeavors, the initial impetus 
for this exploration was economic. Beaver 
and other furbearers were among the ear- 
liest of North America’s vast natural re- 
sources to be exploited by Europeans. De- 
mand for beaver pelts drove fur trappers 
and mountain men into parts of the conti- 
nent that previously had been accessible only 
to indigenous tribes (Chittenden, 1954). The 
Hudson Bay Company and the Pacific Fur 


Company kept meticulous records of their 
annual trade in pelts that provided long- 
term records of population fluctuations and 
predator-prey dynamics. These data have 
been analyzed by several generations of 
ecologists, beginning with Elton (1942). Al- 
though the immediate influence of the fur 
trade on studies of natural history was lim- 
ited, one major contribution was a land- 
mark study of beaver by Lewis H. Morgan 
(1868), one of America’s first ethologists. 
The fur trappers did much to stimulate in- 
terest in wildlife and exploration when they 
returned to the outposts of civilization with 
tales of a vast continent inhabited by ani- 
mals unknown to Europeans. 

The early part of the 19th Century saw 
several exploring expeditions that contrib- 
uted importantly to our knowledge of mam- 
mals. When the newly independent United 
States had acquired the immense Louisiana 
Purchase from France in 1803, President 
Thomas Jefferson dispatched an expedition 
under command of Captains Merriweather 
Lewis and William Clark to survey and map 
the Missouri and Columbia rivers, to study 
the natural history and natural resources of 
the area, and to provide a detailed report of 
all Indian tribes and how to deal with them 
peacefully (Thwaites, 1904). Lewis and 
Clark’s journals provided the first descrip- 
tions of many North American mammals, 
and specimens were also brought back. Un- 
fortunately, the United States had no na- 
tional museum at the time, and all of the 
specimens ultimately were lost. Lewis and 
Clark’s collection was deposited in Peale’s 
Museum in Philadelphia; subsequently, 
most of it was purchased by P.T. Barnum 
and destroyed by fire in 1865 (Gunderson, 
1976). Other notable early expeditions that 
obtained valuable information and speci- 
mens of mammals were those of J. J. Au- 
dubon and J. Bachman, and of T. Say. 

Mammalian natural history also benefit- 
ed from expeditions directed towards the 
discovery of a Northwest Passage that would 
provide access between the Atlantic and Pa- 
cific. Beginning in 1819, several expeditions 


EVOLUTIONARY ECOLOGY a7 


to northern Canada were led by Sir John 
Franklin and Sir William Edward Parry. Sir 
John Richardson was surgeon-naturalist on 
the earlier Franklin expeditions and he also 
described birds and mammals collected on 
the Parry expeditions. Richardson’s (1829) 
Fauna Boreali-Americana contains a com- 
plete volume on mammals. All three of these 
early explorers have been honored with pat- 
ronyms proposed for ground squirrels: 
Spermophilus franklinii, S. parryi, and S. 
richardsonii. 

North American mammalogy, like other 
branches of natural history, is indebted to 
another set of explorations, begun in the 
1850s to seek routes for a transcontinental 
railroad (Miller, 1929). After passage of the 
Railroad Surveys Bill in 1853, the Federal 
Government set out surveying parties that 
were accompanied by physician-naturalists 
from the U.S. Army Medical Corps. They 
made mammal collections of enormous 
breadth and value that were deposited in 
the newly founded (1846) Smithsonian In- 
stitution. These were first studied and de- 
scribed by Professor Spencer Fullerton 
Baird, whose Mammals of North America 
(1859, which appeared as Volume VIII of 
the Pacific Railway Survey Reports) pro- 
vided a state-of-the-art synopsis of the then 
758 known species of North American 
mammals. 

Another physician-naturalist of the U.S. 
Army Medical Corps, Dr. Edgar Alexander 
Mearns, accompanied the survey of the 
U.S.-Mexican International Boundary. 
From his field work in 1892-1894, Mearns 
contributed over 30,000 specimens of plants 
and animals, including over 7,000 mam- 
mals, to the National Museum (Mearns, 
1907). Earlier, Mearns had contributed to 
the specimens that established the verte- 
brate collections of the American Museum 
of Natural History in New York. Later, dur- 
ing two tours of duty in the Philippines, then 
accompanying President Theodore Roose- 
velt’s African expedition, and finally as a 
collaborator with Childs Frick on two ad- 
ditional African expeditions, Mearns con- 


tinued his field studies and collected many 
additional specimens. 

The discovery phase of American mam- 
malogy owes much to the physician-natu- 
ralists of the U.S. Army Medical Corps. In 
addition to Mearns and others associated 
with the Pacific Railroad and Mexican 
Boundary Surveys, a major contributor was 
Dr. Elliot Coues. Coues published his first 
scientific paper at 19 and received his M.D. 
2 years later. In 1864 he joined the Army 
Medical Corps and spent the next 20 years 
doing field work, collecting specimens, and 
publishing extensively on mammals, birds, 
and other vertebrates. He served as the first 
curator of mammals after Baird had orga- 
nized the National Museum in 1879. He 
also served as secretary and naturalist to the 
Geological and Geographical Survey of the 
Territories under F. V. Hayden. Among 
other contributions, Coues wrote five 
monographs on rodents, which form Vol- 
ume 4 of the Hayden Survey Monographs 
(Coues, 1877a), and a classic revision of the 
family Mustelidae (Coues, 18775). 

Owing largely to the contributions of these 
explorer-physician-naturalists, most of the 
North American continent had been sur- 
veyed, and most, but by no means all, of 
the native mammals had been collected and 
classified, by the beginning of the 20th Cen- 
tury. Increasingly, naturalists were con- 
cerned, not with describing new species, but 
with understanding what determines the 
distribution and abundance of the species 
that they now knew about. They began to 
study the lives of wild mammals directly by 
observation and indirectly by trapping and 
tracking. The study of North American 
mammals had begun to pass from the dis- 
covery phase to a natural history phase. 


Natural History Phase 


When the ASM was founded in 1919, its 
charter members included some of the most 
prominent North American biologists. The 
majority of these, including Hartley H. T. 


380 


Jackson, C. Hart Merriam, Edward W. Nel- 
son, Wilfred H. Osgood, Marcus Ward Lyon, 
Jr., T. S. Palmer, Edward A. Preble, Glover 
M. Allen, Joseph Grinnell, Gerrit S. Miller, 
Jr., Angel Cabrera, A. H. Howell, Ned Hol- 
lister, Harold E. Anthony, Vernon Bailey, 
Edgar Alonzo Goldman, Laurance M. Huey, 
Remington Kellogg, Nagamichi Kuroda, 
Austin Roberts, Waldo L. Schmitt, Arthur 
deC. Sowerby, Witmer Stone, Oldfield 
Thomas, Alexander Wetmore, A. H. Winge, 
and Joel Asaph Allen (the first Honorary 
Member), were primarily taxonomists, still 
actively engaged in classifying species and 
documenting their distributions over the 
continent. Even though their systematic 
work represented the culmination ofthe dis- 
covery phase, their studies were becoming 
increasingly synthetic, analytical, and con- 
ceptual. It is no accident that these taxon- 
omists also made some of the most impor- 
tant contributions to natural history. 
Perhaps the most seminal of these early 
contributions that bridged the gap between 
taxonomy and natural history were those of 
C. Hart Merriam. Gerrit S. Miller, Jr., him- 
selfa talented taxonomist, argued that while 
the writings of Darwin had aroused initial 
interest in mammalian natural history, it 
was Merriam who developed the techniques 
for the systematic study of mammals (Mil- 
ler, 1929). Merriam, yet another medical 
doctor influenced by Baird, had an early 
interest in ornithology and an impressive 
combination of intellect, energy, and fore- 
sight that enabled him to establish in 1885 
an organization that began as the Section of 
Ornithology in the Division of Entomology 
under the Commissioner of Agriculture. 
Within a year the Division of Ornithology 
had attained independence, and by 1888 it 
had expanded to the Division of Economic 
Ornithology and Mammalogy. Merriam’s 
later predilection for mammals was illus- 
trated by his staff's reference to the “Divi- 
sion of Economic Ornithology and Extray- 
agant Mammalogy” (Osgood, 1943). This 
unit, which became the Bureau of Biological 


BROWN AND WILSON 


Survey in 1905, and the cadre of distin- 
guished field and museum personnel assem- 
bled by Merriam between 1885 and 1910, 
were the major reasons for the rapid ad- 
vance in our knowledge of North American 
mammals early in this century. 

Merriam’s efforts were facilitated by the 
otherwise unremarkable decision by a man- 
ufacturing company to turn its attention 
from making clothes wringers to producing 
and marketing a truly better mousetrap, the 
“Cyclone” trap, which made its appearance 
in the 1880s. Merriam knew from his stud- 
ies of birds that a key to advancing the sys- 
tematics of mammals was to accumulate and 
study large series of uniformly prepared 
specimens from throughout the range. The 
new cyclone trap made this possible for small 
mammals (Miller, 1929). 

Merriam’s monumental contributions to 
mammalogy were made possible by a com- 
bination of personal science, inspired lead- 
ership, and ability to recruit outstanding sci- 
entists (Osgood, 1943). Merriam personally 
described 660 new species of mammals and 
published more than 600 papers (Grinnell, 
1943). Perhaps his most important paper 
was the one that used the observed eleva- 
tional and latitudinal zonation of flora and 
fauna to develop the life zone concept (Mer- 
riam, 1890). In addition, Merriam empha- 
sized the use of cranial characters in clas- 
sification, and he perfected field and 
museum methods that are still in use today. 
He initiated a new publication series, North 
American Fauna, and wrote the first 11 vol- 
umes himself. From its first volume in 1889 
to its 75th and most recent one (Timm et 
al., 1989), this series has been extremely 
influential; several volumes represent mile- 
stones in the transition from the discovery 
to the natural history phase of North Amer- 
ican mammalogy. Under Merriam’s lead- 
ership the Biological Survey became the pri- 
mary center of mammalogical research. 
Much of this was owing to his genius for 
picking exceptional colleagues. 

The group assembled at the Biological 


EVOLUTIONARY ECOLOGY 381 


Survey comprised an extraordinary group 
of field and museum biologists. Beginning 
about 1883, Merriam had communicated 
with a Minnesota farm boy named Vernon 
Bailey, who had supplied him with difficult 
to obtain specimens, such as shrews. Soon 
after accepting the position in Washington, 
Merriam hired Bailey, thus beginning a close 
and productive friendship between two gi- 
ants of American mammalogy. Bailey and 
his wife, Florence, who was Merriam’s sis- 
ter, crisscrossed the continent collecting 
mammals and birds and described their 
studies in a series of volumes on the fauna 
of various states and geographic regions. 

Two other remarkable members of the 
Biological Survey were Edward W. Nelson 
and Edward Alphonso Goldman. Beginning 
in 1892, Nelson, who later became Chief of 
the Survey, undertook a 14-year biological 
survey of Mexico. He hired Goldman, then 
an 18-year-old California youth, to accom- 
pany him. The results of their collaborative 
study are undoubtedly the most important 
ever achieved by two individuals for a single 
country. They obtained 12,400 specimens 
of birds and 17,400 specimens (including 
354 new species and subspecies) of mam- 
mals. In addition, they collected reptiles, 
amphibians, and plants, and their field re- 
ports contained a wealth of information on 
the vegetation and climate of Mexico (Gold- 
man, 1951). 

If Merriam’s life zone concept was the 
first important ecological principle that sig- 
naled the shift from the discovery phase to 
the natural history phase, Joseph Grinnell’s 
(1917a, 19175) niche concept was the sec- 
ond. Grinnell’s concept, which emphasized 
the role of environmental conditions in lim- 
iting the distribution of a species, was later 
redefined and formalized by Hutchinson 
(1957). Although Grinnell used a bird spe- 
cies, the California thrasher, to illustrate his 
idea of the niche, he made enormous con- 
tributions to both mammalogy and orni- 
thology. Grinnell not only published 554 
papers between 1893 and 1939, he also 


started a mammalogical dynasty by training 
an exceptional cadre of students at the Uni- 
versity of California at Berkeley (Jones, 
1991; Whitaker, 1994). 

Although the majority of the classic de- 
scriptive studies that marked the transition 
from the discovery phase to the natural his- 
tory phase were done on rodents, many were 
performed on other groups that have more 
unusual or conspicuous lifestyles, such as 
bats, ungulates, carnivores, and marine 
mammals. Pioneering studies on bats in- 
cluded Glover Merrill Allen’s (1939) classic 
treatise and two important papers by A. B. 
Howell (1920a, 19205) in the first volume 
of the Journal of Mammalogy. A subse- 
quent volume by Griffin (1958) emphasized 
behavioral and physiological studies of 
echolocation, but also summarized much of 
the information on natural history. These 
early studies were limited to insights that 
could be obtained from studies at roosts, 
direct observations of flying bats, and lab- 
oratory experiments, until the use of Japa- 
nese mist nets around the middle of the cen- 
tury. 

Carnivores and ungulates were the sub- 
jects of important early studies, especially 
those of E. T. Seton (e.g., 1909, 1923), which 
included a multivolume work on the lives 
of game animals (1929). More recent classic 
studies were Murie’s (1944) and Young and 
Goldman’s (1944) on wolves, Hall’s (1951; 
which included taxonomy as well as natural 
history) on weasels, Taylor’s (1956) on deer, 
and Altmann’s (1952) on elk. Important 
early studies of marine mammals included 
papers by Evermann (1921) and Kellogg 
(1921), both in the second volume of the 
Journal of Mammalogy. The more recent 
tradition of natural history studies is illus- 
trated by Bartholomew and Peterson’s 
(1967; the first Special Publication of the 
American Society of Mammalogists) mono- 
graph on the California sea lion and Le Boeuf 
and colleagues’ studies of the northern el- 
ephant seal (e.g., Le Boeuf and Reiter, 1988). 

Natural history studies gathered momen- 


382 BROWN AND WILSON 


tum in the 1920s and 1930s, and they con- 
tinued to dominate ecological mammalogy 
until the 1960s. Some of these, such as those 
by Grinnell and his students on different 
areas in California (Grinnell and Storer, 
1924: Grinnell et al., 1930, 1937), focused 
on particular geographic regions. Others 
were restricted to single species or a few 
related species. Natural history investiga- 
tions reached their epitome in monographic 
studies of various kinds of rodents. These 
included major works on woodrats (Finley, 
1958; Linsdale and Tevis, 1951; Vorhies 
and Taylor, 1940), ground squirrels (Lins- 
dale, 1946), deer mice (McCabe and Blan- 
chard, 1950), microtines (Elton, 1942; Er- 
rington, 1963), and heteromyid rodents 
(Eisenberg, 1963; Reynolds, 1958, 1960). In 
contrast to these large, integrated studies of 
particular species or genera, the contribu- 
tions of two of the most influential mam- 
malian natural historians, W. J. Hamilton, 
Jr., and W. H. Burt, consisted primarily of 
a combination of books on all mammals 
and shorter papers on particular kinds (e.g., 
Burt, 1940, 1946; Hamilton, 1939; Layne 
and Whitaker, 1992; Muul, 1990). 

The natural history phase of research in 
mammalogy also saw the beginnings of the 
conservation movement. W. T. Hornaday 
(1899) detailed the life history and near ex- 
tinction of the North American bison, and 
Volume 2 of the Journal of Mammalogy 
contained a paper on the status of the Eu- 
ropean bison (Ahrens, 1921). The works of 
Seton (1909, 1923, 1929) are filled with ac- 
counts of the relentless killing, declining 
abundances, and contracting ranges of car- 
nivores and ungulates. The Biological Sur- 
vey monitored the status of furbearers and 
the fur trade (Ashbrook, 1922). Lang (1923) 
called attention to the plight of the white 
rhinoceros in Volume 4 of the Journal of 
Mammalogy. Aldo Leopold (e.g., 1933) 
emerged as an eloquent advocate for con- 
servation and developed wildlife manage- 
ment as an applied science based on the 
principles of natural history and ecology. E. 
P. Walker worked diligently to stimulate in- 


terest in mammalian conservation during 
his years at the National Zoological Park, 
and culminated his career with his opus on 
mammals of the world (Walker, 1964). 

Natural history is still a significant com- 
ponent of contemporary American mam- 
malogy. This is apparent from the success 
of the ASM’s Mammalian Species series of 
publications and from the large number of 
““descriptive”’ papers appearing in the Jour- 
nal of Mammalogy and other journals. 


Mammalogy and the New Synthesis 


By the 1930s the study of ecology and 
evolution was already beginning to enter a 
new phase. The new synthesis was laying a 
theoretical and genetical foundation for the 
study of evolution. Fisher, Wright, and Hal- 
dane introduced mathematical models to 
characterize the genetic mechanisms of evo- 
lutionary change, as well as experimental 
and statistical techniques to test rigorously 
the predictions of these models. Simpson, 
Dobzhansky, and Mayr developed a broad 
view of evolution that incorporated not only 
genetic mechanisms, but also systematics, 
biogeography, paleontology, and ecology. 

In North American mammalogy, the in- 
fluence of the new synthesis is seen most 
clearly in two research programs. One 1s the 
work on the genetics of Peromyscus by F. 
B. Sumner and L. R. Dice. These studies 
rivaled those of Drosophila, if not for their 
elucidation of genetic mechanisms per se, 
then for their insights into the adaptive con- 
text of genetic variation. Sumner (e.g., 1932) 
showed that coat color and other traits of 
Peromyscus were heritable, and Dice and 
his students at Michigan, P. M. Blossom, 
W. F. Blair, and B. E. Horner, went on to 
explain geographic variation in coat color 
and morphology in terms of natural selec- 
tion by predators in environments that dif- 
fer in background coloration and vegetation 
structure (e.g., Dice, 1947; Dice and Blos- 
som, 1937; see also Benson, 1933). This 
research program is notable for its use of 


EVOLUTIONARY ECOLOGY 383 


Peromyscus as an empirical “model sys- 
tem” for addressing general conceptual 
questions, for its combination of controlled 
experiments in the laboratory to test mech- 
anisms and comparative field observations 
to place the experimental results in a real- 
istic natural context, and for its use of rig- 
orous experimental designs and statistical 
analyses. 

The other major contribution of North 
American mammalogy to the new synthesis 
was G. G. Simpson’s interpretation of the 
historical record of evolution, based on his 
studies of fossil and Recent mammals. 
Simpson (1940, 1943, 1944, 1947a, 19475, 
1950, 1953) focused on the evolution of the 
North and South American faunas, and on 
the effects of the interchange of species across 
the Interamerican and Bering land bridges. 
He also brought new quantitative approach- 
es to paleontology and comparative biology 
by developing mathematical techniques for 
assessing similarity among faunas, quanti- 
fying diversity, and measuring rates of evo- 
lutionary change. Simpson can be credited 
with primary responsibility for giving the 
new synthesis an historical and biogeo- 
graphic dimension. 

With a few conspicuous exceptions, such 
as Dice and Simpson, descriptive natural 
history studies predominated in North 
American mammalogy until after World 
War II. 


Evolutionary Ecology Phase 


In the late 1950s and 1960s, a major em- 
phasis on science in the U.S. and Canada 
was stimulated by military and scientific 
competition with the U.S.S.R. This period 
saw the emergence of modern evolutionary 
ecology. The seminal event was the Cold 
Spring Harbor Symposium in Quantitative 
Biology in 1957. This symposium is note- 
worthy for three things. First, it had several 
papers on the dynamics of small mammal 
populations (Chitty, 1957; Pitelka, 1957). 
These signaled a shift to North America of 


the research on the dramatic fluctuations in 
rodent populations that had been pioneered 
in Europe by Elton (1927, 1942). Second, 
the mix of genetics, ecology, and evolution 
indicated an effort to expand the new syn- 
thesis to include ecology. Here and in the 
symposium on the genetics of colonizing 
species held in Syracuse in the mid-1960s 
(Lewontin, 1968), the foundations of evo- 
lutionary ecology were laid. Finally, Hutch- 
inson (1957) in his “concluding remarks,” 
capped the symposium by presenting his 
theory of the multidimensional niche. This 
was by no means the first use of mathe- 
matical models in ecology, but it took the- 
oretical ecology beyond the problems of 
population growth and regulation that had 
preoccupied ecologists prior to that time. It 
provided a new conceptual framework to 
address questions about limiting factors, in- 
terspecific interactions, species diversity, 
and adaptation. 

Population dynamics. -Mammalian 
ecologists were well represented at the Cold 
Spring Harbor Symposium. Attendees in- 
cluded Frank Pitelka, Dennis Chitty, Paul 
Errington, John B. Calhoun, John J. Chris- 
tian, and David E. Davis. Charles Elton, 
perhaps the most eminent of all British ecol- 
ogists, had attracted much interest to the 
population fluctuations of microtines. Elton 
had begun field work in the Scandinavian 
arctic in the 1920s, and had summarized 
much of this work in his Voles, Mice and 
Lemmings (1942). Errington (1946, 1963) 
had been working in Iowa since the 1930s 
on the role of predation, disease, and other 
factors in limiting muskrat populations. In- 
fluential papers in the Cold Spring Harbor 
Symposium by Pitelka (1957) on lemming 
cycles at Point Barrow, Alaska, and by Chit- 
ty (1957) on the genetics and behavioral 
components of microtine population regu- 
lation signaled the seminal roles that these 
two newcomers would play in North Amer- 
ican mammalian ecology. 

The challenge that microtines pose to 
ecologists is to explain the dramatic mul- 
tiannual fluctuations in populations. 


384 BROWN AND WILSON 


Whether microtine populations “‘cycle”’ and 
what causes the fluctuations are the two 
questions that have preoccupied microtine 
ecologists since Elton (1942) and Errington 
(1946, 1963). The chapter by Lidicker (1994) 
documents the history and accomplish- 
ments of the enormous research program 
that developed in both North America and 
Europe to address these questions (see also 
Gaines et al., in press; Henttonen et al., 1984; 
Krebs and Myers, 1974; Krohne, 1982; Lid- 
icker, 1988, in press; Taitt and Krebs, 1985; 
Tamarin, 1985). 

Although much attention has been de- 
voted to microtines, important investiga- 
tions of population dynamics have been 
performed on other mammals. Many stud- 
ies have focused on other rodents, because 
of their small size, ease of trapping, and 
occurrence in a wide variety of habitats (e.g., 
Adler and Tamarin, 1984; Brown and 
Heske, 1990; Brown and Zeng, 1989; Pet- 
ticrew and Sadler, 1974; Stickel and War- 
bach, 1960; Whitford, 1976). These have 
often implicated temporal variation in cli- 
mate and food supply as the primary cause 
of population fluctuations. Yet another per- 
spective is offered by large mammals, whose 
population dynamics often appear to be 
controlled by complex relationships be- 
tween food supply and susceptibility to pre- 
dation (e.g., Fowler and Smith, 1981; 
McCullough, 1979). Thus, mammals con- 
tinue to offer a wealth of different patterns 
of population fluctuations, of different 
mechanisms of population regulation, and 
of different kinds of populations for study. 

Species diversity and community struc- 
ture.—After formulating the multidimen- 
sional ecological niche in his ““Concluding 
remarks” at the Cold Spring Harbor Sym- 
posium, Hutchinson (1959) gave a presi- 
dental address to the American Society of 
Naturalists entitled ““Homage to Santa Ro- 
salia, or Why are there so many kinds of 
animals?” By explicitly focusing on patterns 
of species diversity, resource utilization, and 
coexistence, and on processes of population 
regulation, interspecific interaction, and ad- 


aptation, these two papers laid much of the 
foundation for modern community ecology. 
Although David Lack had addressed some 
of these problems in his Darwin’s Finches 
in 1947, they were not pursued vigorously 
until the late 1950s. Other important con- 
tributions at this time included Brown and 
Wilson’s (1956) treatise on character dis- 
placement and MacArthur’s (1958, 1960, 
1965, 1970, 1972) empirical and theoretical 
studies. 

Data from mammals figured prominently 
in these studies in community ecology. 
Hutchinson (1959) used weasels as exam- 
ples of the regular ratios in the body sizes 
or trophic appendages that can be observed 
among coexisting species and that were hy- 
pothesized to reflect the influence of inter- 
specific competition on community struc- 
ture. Hutchinson and MacArthur (1959) 
used the frequency distribution of body sizes 
among all species of North American mam- 
mals to develop models of niche relation- 
ships and coexistence. 

Others were quick to exploit the advan- 
tages of mammals for studies in evolution- 
ary ecology. In 1959, Hall and Kelson pub- 
lished a major taxonomic treatise, The 
Mammals of North America, which con- 
tained, among other information, detailed 
range maps of every species. Simpson (1964) 
used this data base to quantify patterns of 
species diversity across the continent. Thus 
began a long tradition of using these range 
maps to address Hutchinson’s question 
about the ecological processes causing geo- 
graphic variation in species diversity 
(Brown, 1981; Hagmeir and Stults, 1964; 
MacArthur, 1972; Owen, 1990; Rapoport, 
1982; Wilson, 1974; see also Fleming, 1973). 
Unfortunately, despite a great deal of re- 
search, the question remains largely unan- 
swered. The major geographic gradients in 
species richness, including the dramatic in- 
crease in diversity from poles to equator, 
have been increasingly well documented in 
mammals and other organisms, but inves- 
tigators have had only limited success in 
evaluating the contributions of several, and 


EVOLUTIONARY ECOLOGY 385 


not necessarily exclusive, mechanisms that 
may have caused these patterns (e.g., Brown, 
1988; MacArthur, 1972). 

One ecological legacy of Dice’s genetic 
research on Peromyscus was two elegant ex- 
perimental studies of habitat selection. Har- 
ris (1952) showed that forest and grassland 
races of P. maniculatus preferred artificial 
habitats of different structure in the labo- 
ratory. Wecker (1963, 1964) took this ap- 
proach to the field, where he showed not 
only that young mice exhibited a strong 
preference for appropriate habitat, but also 
that there were both inherited and learned 
components of this behavior. Rosenzweig, 
Dueser, M’Closkey, Price, and Morris (see 
references below) continued to investigate 
habitat selection, using it as a vehicle to 
understand population dynamics and com- 
munity structure. 

MacArthur’s student, Rosenzweig, hav- 
ing analyzed geographic variation in body 
size in North American mammals for his 
doctoral dissertation (Rosenzweig, 1966, 
1968), began to study habitat selection, re- 
source utilization, and coexistence in desert 
rodents. Rosenzweig’s studies (e.g., Rosenz- 
weig, 1973; Rosenzweig and Sterner, 1970; 
Rosenzweig and Winakur, 1969; Rosenz- 
weig et al., 1975; Schroder and Rosenzweig, 
1975) were the first of many (see Brown and 
Harney, 1993) that used the desert rodent 
system to address fundamental questions in 
community ecology. From these and other 
studies we have learned that species diver- 
sity and composition vary on geographic 
scales with precipitation and productivity 
(Brown, 1973, 1975), and on local to re- 
gional scales with soil and vegetation type 
(M’Closkey, 1976, 1978; Rosenzweig and 
Winakur, 1969; Rosenzweig et al., 1975). 
Coexisting species tend to be more different 
in body size, body shape, and other attri- 
butes than expected by random community 
assembly (Bowers and Brown, 1982; Brown, 
1973; Dayan and Simberloff, in press; Fin- 
dley, 1989; Hopf and Brown, 1986), and 
they tend to use different microhabitats 
(Brown and Liebermen, 1973; Lemen and 


Rosenzweig, 1978; M’Closkey, 1981; Price, 
1978; Rosenzweig, 1973; Rosenzweig and 
Winakur, 1969). These observations sug- 
gest that interspecific competition plays a 
major role in the organization of these com- 
munities. Field experiments in which some 
species increased in abundance or shifted 
their microhabitat use in response to re- 
moval of other species have provided ad- 
ditional direct evidence for interspecific 
competition (Bowers et al., 1987; Brown and 
Munger, 1985; Freeman and Lemen, 1983; 
Munger and Brown, 1981; Price, 1978; see 
also Larsen, 1986). Clever experiments that 
have altered the risk of predation have 
shown that it influences foraging behavior 
and microhabitat use and probably interacts 
with competition to affect community 
structure (Brown etal., 1987; Kotler, 1984a, 
1984b, 1985: Thompson, 1982a, 1982b). 
Although studies of desert rodents rival 
those of Darwin’s finches and Anolis lizards 
for their contributions to community ecol- 
ogy, many questions remain unanswered and 
several research programs are pursuing 
them. Populations appear to be limited 
largely by food supplies and to fluctuate with 
climatic conditions that determine the 
availability of seeds, insects, and foliage (e.g., 
Beatley, 1976), but the coupling between the 
abiotic environment and population dy- 
namics varies among species and 1s poorly 
understood (Brown and Heske, 1990). There 
has been widespread agreement that differ- 
ences in microhabitat use promote coexis- 
tence, but the extent and significance of food 
resource partitioning has been much more 
controversial (Brown, 1975; Brown and 
Lieberman, 1973; Dayan and Simberloff, in 
press; Lemen, 1978; Mares and Williams, 
1977; Rosenzweig and Sterner, 1970; Smi- 
gel and Rosenzweig, 1974). Although the 
importance of predation and interspecific 
competition no longer seems to be in doubt, 
the way that these processes separately and 
jointly affect population dynamics and 
community structure requires further study. 
Finally, the importance of character dis- 
placement and other kinds of coevolution- 


386 


ary responses to biotic interactions is re- 
ceiving considerable study, but remains 
largely unresolved. 

By no means were all of the important 
community-level studies were of desert ro- 
dents. Miller (1967), Dueser and Shugart 
(1978), Dueser and Hallett (1980), Morris 
(1984), Kirkland (1985), and others inves- 
tigated habitat selection and interspecific in- 
teractions of rodents and shrews in forest 
and grassland habitats. Competitive inter- 
actions among chipmunk species were stud- 
ied in coniferous forest habitats in several 
places in western North America (Brown, 
1971; Chappell, 1978; Heller, 1971; Shep- 
pard, 1971). Findley (1973, 1976, 1993) used 
bats for pioneering studies of ecomorphol- 
ogy, the relationships between patterns of 
morphological variation among species and 
the composition of ecological communities. 
Moors (1984), Ralls and Harvey (1985), and 
Dayan et al. (1989) performed more de- 
tailed morphological and field studies to re- 
examine Hutchinson’s and Rosenzweig’s 
inferences about sexual dimorphism, re- 
source partitioning, and character displace- 
ment in mustelids. Fleming (1971, 1973, 
1988; Fleming et al., 1972) and Wilson 
(1971, 1973; Wilson and Findley, 1970) pi- 
oneered studies of tropical communities of 
both terrestrial mammals and bats, and these 
were followed by others (August, 1983; Hei- 
thaus et al., 1975; Sanchez-Cordero and 
Fleming, 1993). 

Life history studies. —In terms of their use 
of direct observations in the field to learn 
about important events in the lives of in- 
dividual free-living mammals, the most di- 
rect descendants of the classical natural his- 
tory studies of the early 1900s were the life 
history studies of the latter half of the cen- 
tury. Because of their high densities and di- 
urnal habits, colonial sciurid rodents were 
frequently chosen for longitudinal studies 
of life histories. J. A. King’s (1955) work on 
black-tailed prairie dogs (Cynomys ludovi- 
cianus) was probably the most influential, 
if not the first, of the detailed field studies 


BROWN AND WILSON 


of a single population of marked individ- 
uals. This was followed by Armitage’s (1962) 
work on marmots, and then by many other 
studies using different species of ground 
squirrels (reviewed in Murie and Michener, 
1984). 

These studies have been much more than 
descriptive natural history; they have been 
instrumental in gathering data to build and 
test theories of social behavior and life his- 
tory tactics. Together with studies of the 
wolf by Mech (1966, 1970), of Scottish red 
deer (Cervus elaphus) by Clutton-Brock et 
al. (1982), of African carnivores (e.g., Kruuk, 
1972; Packer, 1986; Packer et al., 1988), 
and of primates (e.g., Altmann and Alt- 
mann, 1970; Cheney et al., 1988), the body 
of work on North American sciurids has 
been instrumental in the development of 
our ideas about the roles of environmental 
conditions and social interactions in deter- 
mination of individual reproductive success 
and in evolution of social systems. Perhaps 
the two most extreme and _ spectacular 
mammalian life histories—and ones that 
have far-reaching theoretical implica- 
tions—are the eusocial systems of naked 
mole rats and the semelparous life histories 
of some dasyurid marsupials. Mole-rats 
(Heterocephalus glaber) resemble social bees 
and ants, living in large colonies in which 
a single dominant female effectively cas- 
trates and enslaves her relatives (Jarvis, 
1981). Marsupial mice (Genus Antechinus) 
resemble salmon and certain plants in that 
the males of several species are semelpa- 
rous; they put all of their resources into a 
single reproductive effort and then die after 
just one breeding season (Lee and Cock- 
burn, 1985). 

Another approach to studying the evo- 
lution of life histories and social systems 
that was pioneered by North American 
mammalogists involved allometric rela- 
tionships—patterns of variation with re- 
spect to body size. In 1963, McNab pub- 
lished an influential paper on the correlation 
between home range size and body size (see 


EVOLUTIONARY ECOLOGY 307 


also Schoener, 1968). This was followed by 
several studies of the allometry of life his- 
tory traits, such as litter size, gestation length, 
and maternal investment in offspring (e.g., 
Calder, 1984; Clutton-Brock and Harvey, 
1983; Eisenberg, 1981; Eisenberg and Wil- 
son, 1979; Peters, 1983). These studies have 
not only demonstrated correlates of body 
size that are expressed in allometric rela- 
tionships across large samples of mammal 
species, they have also pointed out devia- 
tions from these relationships that can be 
attributed to evolutionary constraints or 
to adaptations to special ecological condi- 
tions, or both. For other evolutionary and 
ecological approaches to the study of mam- 
malian life histories see Millar (1977) and 
Millar and Zammuto (1983). 

Coevolution. —Several early naturalists 
noted that mammals play potentially im- 
portant roles as dispersers, as well as con- 
sumers, of seeds. Smith (1970) put these 
kinds of interactions in a modern perspec- 
tive with a classic study of coevolution be- 
tween red squirrels (Tamiasciurus hudsoni- 
us and T. douglasii) and conifers. He showed 
that the two squirrel species have different 
morphological and behavioral traits that re- 
flect adaptations to the different kinds of 
conifers that predominate in their geograph- 
ic ranges, and the trees also exhibit adap- 
tations to promote dispersal and to limit 
consumption by the squirrels. Small forest- 
dwelling mammals, such as deer mice and 
voles, have been shown to play a major role 
in dispering the mutualistic mycorrhizal 
fungi that are obligately associated with the 
roots of many tree species (e.g., Maser et 
al., 1978). Howell (1979) found that a bat, 
Leptonycteris sanbornii, 1s the principal pol- 
linator of several century plant and cactus 
species in the Sonoran and Chihuahuan des- 
erts. 

Subsequently, much of the attention 
turned to the tropics, where both rodents 
and bats were shown to be important dis- 
persers of seeds of fleshy-fruited trees (e.g., 
Fleming, 1988; Janzen, 1983; Smythe, 


1970). These studies have for the most part 
supported Janzen’s (1970) suggestion that 
animals, especially frugivorous and graniv- 
orous mammals and birds, play a major role 
in the structure and dynamics of tropical 
forests. Rodents and bats are particularly 
important in carrying seeds away from the 
parent tree, where they are subject to heavy 
predation from insect consumers and mi- 
crobial pathogens, to distant sites that may 
be more favorable for survival and germi- 
nation. Janzen’s (1981) discovery that in- 
troduced horses are important agents of seed 
dispersal for some tropical tree species led 
to the suggestion that the extinction of the 
Pleistocene megafauna and the extirpation 
of modern species of large mammals, such 
as tapirs and peccaries, may be causing sub- 
stantial changes in tropical forests (Janzen 
and Martin, 1982). 

Recently, evolutionary ecologists have 
speculated about coevolutionary relation- 
ships between parasitic or symbiotic organ- 
isms and their hosts (e.g., Holmes and Price, 
1986; Price, 1980). Studies of mammals 
have supported suggestions that “parasites” 
may not always have significant negative 
effects on their hosts; in fact, some apparent 
parasites might even benefit their hosts 
(Munger and Holmes, 1988). Other fasci- 
nating symbiotic relationships have been 
discovered. Several tropical mammals have 
symbiotic insects that live in their fur, their 
nests, or both, and prey on lice, fleas, and 
other ectoparasites (e.g., Ashe and Timm, 
1987: Timm and Ashe, 1988). In desert and 
arid grassland habitats bannertailed kan- 
garoo rats (Dipodomys spectabilis) and 
woodrats (Neotoma sp.) construct large dens 
that provide refuges for many kinds of in- 
vertebrates and small vertebrates (Monson 
and Kessler, 1940). In addition, the seed 
stores of the bannertailed kangaroo rats are 
inhabited by many kinds of fungi that have 
been suggested to have beneficial effects on 
their rodent hosts by enhancing the nutri- 
tional value of infested seeds (e.g., Hawkins, 
1992; Reichman et al., 1985). 


388 BROWN AND WILSON 


The Transition from Natural 
History to Evolutionary 
Ecology 


The transition. —The period of active re- 
search in mammalogy in North America, 
from about 1850 to the present, marked the 
transition from studies that emphasized de- 
scriptive taxonomy, morphology, distribu- 
tion, paleontology, and natural history to 
investigations that were motivated by the 
theoretical questions of modern disciplines 
such as biomechanics, physiology, behav- 
ior, genetics, evolution, systematics, ecol- 
ogy, and biogeography. The 19th-Century 
naturalists were generalists. The greatest of 
them, such as Cuvier, Darwin, Wallace, 
Bates, von Humboldt, and Prinz Maximil- 
ian zu Wied, were knowledgeable about 
plants, invertebrates, and vertebrates, stud- 
ied geology and paleontology as well as bi- 
ology, and developed concepts and theories 
to explain their empirical observations. Even 
the early 20th-Century mammalogists were 
amazingly diverse scientists. For example, 
Merriam published in geography and an- 
thropology as well as mammalogy (Grin- 
nell, 1943; Osgood, 1943), and Grinnell 
wrote influential papers on the behavior, 
ecology, biogeography, and systematics of 
both birds and mammals (Miller, 1943). 

The natural historians of the first half of 
the 20th Century represented a transition 
from the 19th-Century naturalists to mod- 
ern evolutionary ecologists. These natural 
historians, best represented by individuals 
such as Linsdale, Murie, Vorhies, and Ham- 
ilton, made detailed, descriptive studies of 
particular species that emphasized behav- 
ior, reproductive biology, and distribution 
with respect to habitat. Today their work 
may seem quaint, descriptive, and lacking 
theoretical motivation. It is important how- 
ever, to recognize the extent to which the 
natural historians laid the foundations for 
the more conceptual approach of contem- 
porary mammalogy. Taxonomic mammal- 
ogists were still describing new species and 


mapping their geographic ranges well into 
the present century. It was necessary to doc- 
ument the basic biology of these mammals 
before it was apparent which ones were well 
suited for addressing ecological and evolu- 
tionary questions of theoretical interest. 

The dependence of modern evolutionary 
ecologists on the work of their more de- 
scriptive antecedents is illustrated by two 
observations. First, many of the evolution- 
ary ecologists were trained either by natural 
historians or by taxonomists. Note, for ex- 
ample, the academic histories of Findley, 
Krebs, Lidicker, Eisenberg, Wilson, Brown, 
Fleming, and other mammalian evolution- 
ary ecologists (Jones, 1991; Whitaker, 1994). 
Second, the influence of the natural histo- 
rians is illustrated by the frequency with 
which the studies of Linsdale, Grinnell, Hall, 
Vorhies and Taylor, Findley, and others are 
cited in recent publications. The descriptive 
observations of the natural historians often 
provide the inspiration for the modern ex- 
perimental studies of evolutionary ecolo- 
gists. 

The role of theory.—The transition from 
natural history to evolutionary ecology can 
be attributed largely to the influence of 
mathematical theory and the seminal con- 
tributions of Hutchinson, MacArthur, and 
others. The foundations of the new synthe- 
sis were laid by the mathematical models 
of genetic evolutionary change of Fisher, 
Wright, and Haldane. Although this work 
was largely completed before World War II, 
the consolidation of the new synthesis did 
not come until the major works of Dobz- 
hansky (1937), Simpson (1944, 1953) and 
Mayr (1942, 1963). These major advances 
in understanding the evolutionary process 
demonstrated the power of mathematical 
models to motivate important experimental 
and synthetic empirical studies. 

The incorporation ofan evolutionary per- 
spective into studies of ecology and life his- 
tory can be attributed largely to the influ- 
ence of Hutchinson and his student, 
MacArthur. As mentioned above, Hutch- 
inson (1957, 1959) set much of the agenda 


EVOLUTIONARY ECOLOGY 389 


for the next several decades with his papers 
on the niche and the diversity of species. 
MacArthur (e.g., 1960, 1970, 1972) fol- 
lowed with mathematical treatments of spe- 
cies abundance and diversity, competition 
and resource utilization, coexistence and 
coevolution, life history theory, optimal 
foraging, and island biogeography. There 
was hardly a topic in modern evolutionary 
ecology that MacArthur did not address. He 
was almost certainly the most influential 
ecologist who ever lived, an assessment that 
is borne out by the total number of times 
his papers have been cited (see Science Ci- 
tation Index). 

The specific mathematical models devel- 
oped by Hutchinson, MacArthur, and oth- 
ers have had mixed success. Some, such as 
the broken stick distribution of niches and 
the idea that complexity promotes stability, 
were misguided or just plain wrong, and 
have been abandoned. Others, such as r and 
K reproductive strategies and the limiting 
similarity of species were too simplistic; they 
represented important advances, but were 
eventually replaced by more complex and 
realistic theory. Still others, such as re- 
source-based competition equations and the 
theory of island biogeography are still wide- 
ly used to motivate both theoretical and em- 
pirical studies. Despite the mixed success of 
these models, their influence on the devel- 
opment of modern evolutionary ecology is 
enormous. Almost every influential empir- 
ical paper since 1960 cites theoretical lit- 
erature and attempts to evaluate predictions 
of mathematical theory. 

Even more important than its success in 
explaining evolutionary and ecological phe- 
nomena, however, was the way that math- 
ematical theory revolutionized the science. 
It led to more conceptual, question-asking, 
quantitative, analytical, experimental, and 
statistical approaches to both theoretical and 
empirical studies. To produce mathemati- 
cal theory requires conceptual innovation, 
quantitative skills, and analytical rigor. To 
test empirically the predictions of theory 
requires understanding the theory, choice of 


an appropriate system for study, design and 
execution of appropriate experiments or 
comparative observations, and rigorous sta- 
tistical analysis and inference. 

Mammalian systems for testing theory. — 
The appearance of compelling mathemati- 
cal models called for empirical tests of their 
assumptions and predictions in appropriate 
natural systems. Beginning with Hutchin- 
son and MacArthur’s (1959) paper on the 
distribution of body sizes among species, 
North American mammals have played a 
major role in the interaction between theory 
and data. Some of this was largely seren- 
dipitous. Thanks to the efforts of the natural 
historians, mammals had already been rel- 
atively well studied and young scientists 
trained in more descriptive mammalogy 
soon became interested in testing the excit- 
ing new theories. 

Furthermore, certain kinds of mammals 
possess combinations of characteristics that 
have made them excellent systems for quan- 
titative and experimental field studies. The 
influential roles of sciurid rodents, pri- 
mates, and ungulates in life history studies, 
of microtine rodents in studies of popula- 
tion dynamics, and of desert rodents in in- 
vestigations of coexistence and interactions 
of species are no accident. Each of these 
groups has specific traits that facilitate ob- 
servation, quantification, and experimental 
manipulation to obtain definitive tests of 
hypotheses and theoretical predictions. No 
organism is ideal for all kinds of studies, 
and some groups of birds, lizards, insects, 
plants, and intertidal organisms, have 
rivaled mammals as empirical systems for 
studies in evolutionary ecology. Neverthe- 
less, mammals have played and will con- 
tinue to play an influential role in the de- 
velopment of evolutionary ecology (see 
citations in the previous section). 

Increasing scientific rigor. —-As men- 
tioned above, the development of mathe- 
matical theory had a profound effect on the 
way that empirical studies of mammals were 
conducted. The emphasis shifted from qual- 
itative description motivated by economic 


390 BROWN AND WILSON 


concerns or investigator fancy, to statisti- 
cally rigorous, experimental hypothesis- 
testing motivated by theoretical issues. 

Once empirical studies shifted from de- 
scribing the natural history of mammal spe- 
cies to evaluating the assumptions and pre- 
dictions of particular theories they then 
needed to provide more definitive answers. 
This required formulating and testing hy- 
potheses. Usually the goals of natural his- 
tory studies were essentially similar to Lins- 
dale’s (1946:vii): ““The need for an extensive 
study of the life of the California Ground 
Squirrel has grown with increasing rapidity 
as more and more questions have been raised 
about this animal, its habitat, distribution, 
and characteristics.”” There is no way to 
frame this objective in the form of a single 
hypothesis, or to satisfy this need except by 
doing the kind of broad, descriptive study 
that Linsdale did. This changed when the 
goal became to learn whether habitat het- 
erogeneity affects the dynamics of a micro- 
tine population or whether two desert ro- 
dent species are competing. Each of these 
questions can be cast as a specific hypoth- 
esis, and answered definitively with a single 
set of observations or experimental manip- 
ulations. 

Another impact of theory, then, was that 
it led to an increased emphasis on the design 
and execution of controlled experiments to 
give definitive tests of hypotheses. Connell 
(1961) brought to modern evolutionary 
ecology the approach, long practiced by 
British plant ecologists, of doing replicated 
manipulative experiments in the field. It did 
not take long for field experiments to be 
applied to mammalian ecology, first and 
most notable in manipulations of microtine 
populations by Krebs and colleagues (Krebs 
et al., 1969) and in Rosenzweig’s (1973) 
“habitat tailoring’ experiments on desert 
rodents. Now a large proportion of field 
studies in mammalian ecology are well-de- 
signed experiments, with appropriate con- 
trols, adequate replication, and standard- 
ized data collection. Of course, a number of 
conceptually or practically important ques- 
tions simply cannot be answered by manip- 


ulative experiments. It is either impractical 
to experiment on the spatial or temporal 
scale required to test the hypothesis (e.g., to 
address biogeographic questions), or it is 
illegal or unethical to perturb the natural 
system (e.g., in the case of endangered spe- 
cies). This does not diminish the need to 
adopt a rigorous, hypothesis-testing ap- 
proach, but it requires that carefully de- 
signed comparative observations be substi- 
tuted for artificial manipulations (e.g., 
Brooks and McLennan, 1991; Harvey and 
Pagel, 1991). 

Finally, the emphasis on evaluating the- 
ory, testing hypotheses, and doing experi- 
ments has led to the development of an in- 
creasingly powerful battery of statistical 
techniques. In fact, many of the analyses 
were developed by theoreticians, including 
Fisher and Wright, for testing empirically 
the predictions of their models. Statistical 
analyses are virtually absent from most of 
the natural history literature before World 
War II, although means and occasionally 
some measure of variance were sometimes 
reported. Now ecological papers contain 
such sophisticated experimental designs and 
statistical analyses that constant updating 
of biometric skills is required to interpret 
the results, let alone to do state-of-the-art 
research (e.g., see Dueser et al., 1989). 

It is hard to overestimate how much our 
science has changed since World War II. In 
just a few decades traditional descriptive 
natural history has fallen into disfavor in 
the classroom and the journals. It has been 
eclipsed by an evolutionary approach to 
ecology that asks theoretical questions, and 
uses sophisticated experimental and statis- 
tical techniques to answer them. Mammal- 
ogists have not simply responded to this 
revolution, they have often been in the fore- 
front, using the special advantages of mam- 
mals to make important conceptual and 
empirical advances. 


Summary 


The history of North American mam- 
malogy began with the exploration of the 


EVOLUTIONARY ECOLOGY 391 


continent by Europeans. There was added 
incentive to study mammals, because one 
of the resources most valuable to the early 
colonists was fur, especially beaver pelts. 
After the fur trade slackened, official voy- 
ages to explore and survey the remote parts 
of the continent were usually accompanied 
by scientists, many of them physician-nat- 
uralists with particular interests in mam- 
mals. During this discovery phase, the early 
naturalists were concerned with describing 
and classifying the different kinds of mam- 
mals and beginning to accumulate infor- 
mation on their distributions and habits. 

By the beginning of the 20th Century, 
most of the species of North American 
mammals had been described and mam- 
malogists were beginning to specialize. One 
of the specialties was natural history, which 
encompassed all aspects of ecology and be- 
havior. In contrast to modern disciplines, 
natural history was a descriptive science. Its 
goal was to describe the environmental re- 
lationships and lives of particular species, 
groups of related species, or entire assem- 
blages of coexisting species. 

The new evolutionary synthesis began the 
transformation of the field sciences into 
modern theory-testing, experimental disci- 
plines. Early studies of mammalian genet- 
ics, adaptations, paleontology, and bioge- 
ography contributed importantly to the data 
and concepts of the new synthesis. 

After World War II, the interjection of 
evolutionary concepts and mathematical 
modeling was instrumental in the transfor- 
mation of traditional, descriptive natural 
history into the modern discipline of evo- 
lutionary ecology. Mammals were used to 
make important empirical and theoretical 
contributions to our understanding of pop- 
ulation dynamics, community organiza- 
tion, foraging, habitat selection, life history 
traits, and coevolution. 


Acknowledgments 


We thank W. L. Gannon, L. Hawkins, E. J. 
Heske, B. D. Patterson, and R. M. Timm for 


their comments on this manuscript. Brown’s re- 
search has been supported by the National Sci- 
ence Foundation, most recently by Grant BSR- 
8718139. 


Literature Cited 


ADLER, G. H., AND R. H. TAMARIN. 1984. Demog- 
raphy and reproduction in island and mainland white- 
footed mice (Peromyscus leucopus) in southeastern 
Massachusetts. Canadian Journal of Zoology, 62:58- 
64. 

AHRENS, T. G. 1921. The present status of the Eu- 
ropean bison or wisent. Journal of Mammalogy, 2:58— 
62. 

ALLEN, G. M. 1939. Bats. Harvard University Press, 
Cambridge, 368 pp. 

ALTMANN, M. 1952. Social behavior of elk, Cervus 
canadensis nelsoni, in the Jackson Hole area of Wy- 
oming. Behaviour, 4:116-143. 

ALTMANN, S. A., AND J. ALTMANN. 1970. Baboon 
ecology: African field research. University of Chi- 
cago Press, Chicago, 220 pp. 

ARMITAGE, K. B. 1962. Social behavior of a colony 
of the yellow-bellied marmot (Marmota flaviventris). 
Animal Behaviour, 10:319-331. 

ASHBROOK, F. G. 1922. The fur trade and the fur 
supply. Journal of Mammalogy, 3:1-7. 

ASHE, J. S., AND R. M. Timm. 1987. Predation by 
and activity patterns of ‘parasitic’ beetles of the genus 
Amblyopinus (Coleoptera: Staphylinidae). Journal of 
Zoology (London), 212:429-437. 

AucustT, P. V. 1983. The role of habitat complexity 
and heterogeneity in structuring tropical mammal 
communities. Ecology, 64:1495-1507. 

BairD, S. F. 1859. Mammals: general report upon 
the zoology of the several Pacific railroads routes. 
Reports, explorations and surveys for railroad route 
from Mississippi River to Pacific Ocean. Volume 8 
(part 1), Washington, D.C., 757 pp. 

BARTHOLOMEW, G. A., AND R. S. PETERSON. 1967. 
The natural history and behavior of the California 
sea lion. Special Publications, The American Society 
of Mammalogists, 1:1-79. 

BEATLEY, J.C. 1976. Environments of kangaroo rats 
(Dipodomys) and effects of environmental change on 
populations in southern Nevada. Journal of Mam- 
malogy, 57:67-93. 

Benson, S. B. 1933. Concealing coloration among 
some desert rodents of the southwestern United 
States. University of California Publication of Zo- 
ology, 40:1-70. 

Bowers, M. A., AND J. H. BRown. 1982. Body size 
and coexistence in desert rodents: chance or com- 
munity structure. Ecology, 63:391-400. 

Bowers, M. A., D. B. THompson, AND J. H. BRown. 
1987. Foraging and microhabitat use in desert ro- 
dents: the role of a dominant competitor. Oecologia, 
72:77-82. 

Brooks, D. L., AND D. A. MCLENNAN. 1991. Phy- 
logeny, ecology, and behavior. University of Chicago 
Press, Chicago, 434 pp. 

Brown, J. H. 1971. Mechanisms of competitive ex- 


392 BROWN AND WILSON 


clusion between two species of chipmunks. Ecology, 

52:305-311. 

1973. Species diversity of seed-eating desert 

rodents in sand dune habitats. Ecology, 54:775-787. 

. 1975. Geographical ecology of desert rodents. 

Pp. 315-341, in Ecology and evolution of commu- 

nities (M. L. Cody and J. M. Diamond, eds.). Bel- 

knap Press, Cambridge, Massachusetts, 545 pp. 

. 1981. Two decades of homage to Santa Rosa- 

lia: toward a general theory of diversity. American 

Zoologist, 21:877-888. 

. 1988. Species diversity. Pp. 57-90, in Ana- 
lytical biogeography (A. A. Myers and P. S. Giller, 
eds.). Chapman and Hall, London, 578 pp. 

Brown, J. H., AND B. A. HARNEY. 1993. Population 
and community ecology of heteromyid rodents in 
temperate habitats. Pp. 618-651, in Biology of the 
Heteromyidae (H. H. Genoways and J. H. Brown, 
eds.). Special Publications, The American Society of 
Mammalogists, 10:1-719. 

Brown, J. H., AND E. J. Heske. 1990. Temporal 
changes in a Chihuahuan Desert rodent community. 
Oikos, 59:290-302. 

Brown, J. H., AND G. A. LIEBERMAN. 1973. Resource 
utilization and coexistence of seed-eating desert ro- 
dents in sand dune habitats. Ecology, 54:788-797. 

Brown, J. H., AND J. C. MuNGER. 1985. Experi- 
mental manipulation of a desert rodent community: 
food addition and species removal. Ecology, 66:1545- 
1563. 

Brown, J. H., AND Z. ZENG. 1989. Comparative life 
population ecology of eleven species of rodents in 
the Chihuahuan Desert. Ecology, 70:1507-1525. 

Brown, J. S., B. P. KoTLer, R. J. SMITH, AND W. O. 
Wirtz, II. 1987. The effects of owl predation on 
the foraging of heteromyid rodents. Oecologia, 76: 
408-415. 

Brown, W. L., JR., AND E. O. WiLson. 1956. Char- 
acter displacement. Systematic Zoology, 5:49-64. 
Burt, W. F. 1940. Territorial behavior and popu- 
lations of some small mammals in southern Mich- 
igan. Miscellaneous Publication of the Museum of 

Zoology, University of Michigan, 45:1—58. 

. 1946. The mammals of Michigan. University 
of Michigan Press, Ann Arbor, 288 pp. 

CaALpeR, W. A., III. 1984. Size, function, and life 
history. Harvard University Press, Cambridge, Mas- 
sachusetts, 431 pp. 

CHAPPELL, M. A. 1978. Behavioral factors in the al- 
titudinal zonation of chipmunks (Eutamias). Ecol- 
ogy, 59:565-579. 

CHENEY, D. L., R. M. SEYFARTH, S. J. ANDELMAN, AND 
P. C. Lee. 1988. Reproductive success in vervet 
monkeys. Pp. 384-402, in Reproductive success (T. 
H. Clutton-Brock, ed.). University of Chicago Press, 
Chicago, 538 pp. 

CHITTENDEN, H. 1954. The American fur trade. Stan- 
ford University Press, Stanford, California, 2 vol- 
umes, 1,029 pp. 

Cuitry, D. 1957. Self-regulation of numbers through 
changes in viability. Cold Spring Harbor Symposium 
in Quantitative Biology, 22:277—280. 

CLutton-Brock, T. H., AND P. H. HARvey. 1983. 
The functional significance of variation in body size 
in mammals. Pp. 632-663, in Advances in the study 


of mammalian behavior (J. F. Eisenberg and D. G. 
Kleiman, eds.). Special Publications, The American 
Society of Mammalogists, 7:1—753. 
CLuTTON-Brock, T. H., F. E. GUINNEss, AND S. D. 
ALBON. 1982. Red deer: behavior and ecology of 
two sexes. University of Chicago Press, Chicago, 378 


CONNELL, J. H. 1961. The influence of interspecific 
competition and other factors on the distribution of 
the barnacle, Chthamalus stellatus. Ecology, 42:710- 
723; 

Coues, E. 1877a. Monographs of North American 
rodents. United States Geological Survey of the Ter- 
ritories, Washington, D.C., 1,091 pp. 

1877b. Fur-bearing animals. Miscellaneous 
Publication No. 8, United States Geological Survey 
of the Territories, Washington, 349 pp. 

DayYANn, T., AND D. SIMBERLOFF. In press. Morpho- 
logical relationships among coexisting heteromyids: 
an incisive dental character. The American Natu- 
ralist. 

Dayan, T., D. SIMBERLOFF, E. TCHERNOV, AND Y. 
Yom-Tov. 1989. Inter- and intraspecific character 
displacement in mustelids. Ecology, 70:1526-1539. 

Dice, L. R. 1947. Effectiveness of selection by owls 
of deer mice (Peromyscus maniculatus) which con- 
trast in color with their background. Contributions 
from the Laboratory of Vertebrate Biology, Univer- 
sity of Michigan, 50:1-15. 

Dice, L. R., AND P. M. BLossom. 1937. Studies of 
mammalian ecology in southwestern North America 
with special attention to the colors of desert mam- 
mals. Carnegie Institution of Washington, Publica- 
tion, 485:1-129. 

DoBzsHANSKY, T. 1937. Genetics and the origin of 
species. Columbia University Press, New York, 364 
pp. 

Dueser, R. D., AND J. G. HALLETT. 1980. Compe- 
tition and habitat selection in a forest-floor small 
mammal fauna. Oikos, 35:293-297. 

Dueser, R. D., J. H. PORTER, AND J. L. DOOLEY, JR. 
1989. Direct tests for competition in North Amer- 
ican rodent communities: synthesis and prognosis. 
Pp. 105-125, in Patterns in the structure of mam- 
malian communities (D. W. Morris, Z. Abramsky, 
B. J. Fox, and M. R. Willig, eds.). Special Publica- 
tions, The Museum, Texas Tech University, 28:1- 
XXX. 

Dueser, R. D., AND H. H. SHUGART. 1978. Micro- 
habitats in a forest-floor small mammal fauna. Ecol- 
ogy, 59:89-98. 

EISENBERG, J. F. 1963. The behavior of heteromyid 
rodents. University of California Publications in Zo- 
ology, 69:1-114. 

1981. The mammalian radiations. Univer- 
sity of Chicago Press, Chicago, 610 pp. 

EISENBERG, J. F., AND D. E. Witson. 1979. Relative 
brain size and feeding strategies in the Chiroptera. 
Evolution, 32:740-751. 

1981. Relative brain size and demographic 
strategies in didelphid marsupials. The American 
Naturalist, 118:1-15. 

ELton, C. 1927. Animal ecology. The Macmillan 
Company, New York, 207 pp. 


EVOLUTIONARY ECOLOGY ope! 


1942. Voles, mice and lemmings. Clarendon 
Press, Oxford, 496 pp. 

ERRINGTON, P. L. 1946. Predation and vertebrate 
populations. The Quarterly Review of Biology, 21: 
144-177, 221-245. 

1963. Muskrat populations. Iowa State Uni- 
versity Press, Ames, 665 pp. 

EVERMANN, B. W. 1921. The Ano Nuevo Steller sea 
lion rookery. Journal of Mammalogy, 2:16-19. 

FINDLEY, J. S. 1973. Phenetic packing as a measure 
of faunal diversity. The American Naturalist, 107: 
580-584. 

. 1976. The structure of bat communities. The 

American Naturalist, 110:129-139. 

. 1989. Morphological patterns in rodent com- 

munities of southwestern North America. Pp. 253- 

264, in Patterns in the structure of mammalian com- 

munities (D. W. Morris, Z. Abramsky, B. J. Fox, 

and M. R. Willig, eds.). Special Publications, The 

Museum, Texas Tech University, 28:1—-xxx. 

. 1993. Bats: a community perspective. Cam- 
bridge University Press, 167 pp. 

Fintey, R. B., Jk. 1958. The wood rats of Colorado: 
distribution and ecology. University of Kansas Pub- 
lications, Museum of Natural History, 10:213-552. 

FLEMING, T. H. 1971. Population ecology of three 
species of neotropical rodents. Miscellaneous Pub- 
lications of the Museum of Zoology, University of 
Michigan, 143:1-77. 

. 1973. Numbers of mammal species in North 

and Central American forest communities. Ecology, 

54:555-563. 

1988. The short-tailed fruit bat. University 
of Chicago Press, Chicago, 363 pp. 

FLEMING, T. H., E. T. Hooper, AND D. E. WILSON. 
1972. Three Central American bat communities: 
structure, reproductive cycles and movement pat- 
terns. Ecology, 53:555-569. 

Fow er, C. W., AND T. D. SMITH (EDs.). 1981. Dy- 
namics of large mammal populations. Wiley, New 
York, 477 pp. 

FREEMAN, P. W., ANDC. LEMEN. 1983. Quantification 
of competition among coexisting heteromyids in the 
Southwest. The Southwestern Naturalist, 28:41—-46. 

Gaines, M. S., N. C. STENSETH, M. L. JOHNSON, R. A. 
IMs, AND S. BONDRUP-NIELSEN. In press. A re- 
sponse to solving the enigma of population cycles 
with a multifactorial perspective. Journal of Mam- 
malogy. 

GoLpMAN, E. A. 1951. Biological investigations in 
Mexico. Smithsonian Miscellaneous Collections, 1 15: 
1-476. 

GRIFFIN, D. R. 1958. Listening in the dark. Yale 
University Press, New Haven, 413 pp. 

GRINNELL, H. W. 1943. Bibliography of Clinton Hart 
Merriam. Journal of Mammalogy, 24:436-457. 

GRINNELL, J. 1917a. Field tests of theories concern- 
ing distributional controls. The American Natural- 
ist, 51:115-128. 

. 1917b. The niche-relationship of the Cali- 
fornia thrasher. Auk, 34:427-433. 

GRINNELL, J., J. Drxon, AND J. M. LINSDALE. 1930. 
Vertebrate natural history of Lassen Peak. Univer- 
sity of California Publications in Zoology, 35:1-594. 


1937. Fur-bearing mammals of California. 
University of California Press, Berkeley, 2 volumes. 

GRINNELL, J., AND T. I. Storer. 1924. Animal life 
in the Yosemite. University of California Press, 
Berkeley, California, 752 pp. 

GuNDERSON, H.L. 1976. Mammalogy. McGraw-Hill 
Book Company, New York, 483 pp. 

Hacmeirr, E. M., AND C. D. Stutts. 1964. A nu- 
merical analysis of the distributional patterns of North 
American mammals. Systematic Zoology, 13:125- 
155: 

Hatt, E. R. 1951. American weasels. University of 
Kansas Publications, Museum of Natural History, 
4:1-466. 

Ha i, E. R., AND K. R. KELson. 1959. The mammals 
of North America. The Ronald Press Company, New 
York, 2 volumes, 1,083 pp. 

HamiLton, W. J. 1939. American mammals. Mc- 
Graw-Hill Book Company, New York, 434 pp. 

Harris, V.T. 1952. Anexperimental study of habitat 
selection by prairie and forest races of the deer mouse, 
Peromyscus maniculatus. University of Michigan 
Contributions from the Laboratory of Vertebrate Bi- 
ology, 56:1-53. 

Harvey, P. H., AND M. D. PAGEL. 1991. The com- 
parative method in evolutionary biology. Oxford 
University Press, Oxford, 239 pp. 

Hawkins, L. K. 1992. Banner-tailed kangaroo rats 
and molds: a possible vertebrate-fungus mutualism. 
Ph.D. dissert., University of New Mexico, Albu- 
querque, XXX pp. 

HerrHaus, E. R., T. H. FLEMING, AND P. A. OPLER. 
1975. Foraging patterns and resource utilization in 
seven species of bats in a seasonal tropical forest. 
Ecology, 56:841-854. 

Hetter, H. C. 1971. Altitudinal zonation of chip- 
munks (Eutamias): interspecific aggression. Ecology, 
52:312-319. 

HENTTONEN, H., J. TAst, J. ViITALA, AND A. KAIKUSA- 
Lo. 1984. Ecology of cyclic rodents in northern 
Finland. Society Flora Fauna Fennici, Memoirs, 60: 
84-92. 

Howes, J. C., AND P. W. Price. 1986. Communities 
of parasites. Pp. 187-213, in Community ecology: 
patterns and process (D. J. Anderson and J. Kik- 
kawa, eds.). Blackwell Scientific Publications, Ox- 
ford, xxx pp. 

Hopr, F. A., AND J. H. Brown. 1986. The bull’s-eye 
method for testing randomness in ecological com- 
munities. Ecology, 67:1139-1155. 

Hornapay, W. T. 1899. The extermination of the 
American bison with a sketch of its discovery and 
life history. Smithsonian Institution Annual Report, 
1887, Washington, D.C., pp. 367-548. 

Howe Lt, A. B. 1920a. Contribution to the life-his- 
tory of the California mastiff bat. Journal of Mam- 
malogy, 1:111-117. 

1920b. Some Californian experiences with 
bat roosts. Journal of Mammalogy, 1:169-177. 

Howe Lt, D. J. 1979. Flock foraging in nectar-feeding 
bats: advantages to the bats and to the host plants. 
The American Naturalist, 1 13:23-49. 

Hutcuinson, G. E. 1957. Concluding remarks. Cold 
Spring Harbor Symposium in Quantitative Biology, 
22:415-427. 


394 BROWN AND WILSON 


1959. Homage to Santa Rosalia or why are 
there so many kinds of animals? The American Nat- 
uralist, 93:145-159. 

HuTCHINSON, G. E., AND R. H. MACARTHUR. 1959. 
A theoretical ecological model of size distributions 
among species of animals. The American Naturalist, 
93:117-126. 

JANZEN, D. H. 1970. Herbivores and the number of 
tree species in tropical forests. The American Nat- 
uralist, 104:501-528. 

. 1981. Guanacaste tree seed-swallowing by 

Costa Rican range horses. Ecology, 62:587-592. 

1983. Dispersal of seeds by vertebrate guts. 

Pp. 232-262, in Coevolution (D. J. Futuyma and M. 

Slatkin, eds.). Sinaur, Sunderland, Massachusetts, xxx 


pp. 

JANZEN, D. H., AND P. S. MARTIN. 1982. Neotropical 
anachronisms: the fruits the gomphotheres ate. Sci- 
ence, 215:19-27. 

Jarvis, J. M. 1981. Eusociality in a mammal: co- 
operative breeding in naked mole-rat colonies. Sci- 
ence, 212:571-573. 

Jones, J. K., Jk. 1991. Genealogy of Twentieth-cen- 
tury systematic mammalogists in North America: 
the descendants of Joseph Grinnell. Pp. 48-56, in 
Latin American mammalogy: history, biodiversity, 
and conservation (M. A. Mares and D. J. Schmidly, 
eds.). University of Oklahoma Press, Norman, 468 


pp. 

KeELLoGG, R. 1921. A new pinniped from the Upper 
Pliocene of California. Journal of Mammalogy, 
2:221-226. 

Kina, J. A. 1955. Social behavior, social organiza- 
tion, and population dynamics in a black-tailed prai- 
rie dog town in the Black Hills of South Dakota. 
University of Michigan Contributions from the Lab- 
oratory of Vertebrate Zoology, 67:1-123. 

KIRKLAND, G. L., JR. 1985. Small mammal com- 
munities in temperate North American forests. Aus- 
tralian Mammalogy, 8:137-144. 

Kotter, B. P. 1984a. Predation risk and the structure 
of desert rodent communities. Ecology, 65:689-701. 

19845. Harvesting rates and predatory risk 

in desert rodents: a comparison of two communities 

on different continents. Journal of Mammalogy, 65: 

91-96. 

. 1985. Owl predation on desert rodents which 
differ in morphology and behavior. Journal of Mam- 
malogy, 66:824—-828. 

Kress, C. J., B. L. KELLER, AND R. H. TAMARIN. 1969. 
Microtus population biology: demographic changes 
in fluctuating populations of Microtus ochrogaster 
and M. pennsylvanicus in southern Indiana. Ecology, 
50:587-607. 

Kress, C. J., AND J. H. Myers. 1974. Population 
cycles in small mammals. Advances in Ecological 
Research, 8:267-399. 

KROHNE, D. T. 1982. Demography of low-litter-size 
populations of Microtus californicus. Canadian Jour- 
nal of Zoology, 60:368-374. 

Kruuk, H. 1972. The spotted hyena. University of 
Chicago Press, Chicago, 335 pp. 

Lack, D. 1947. Darwin’s finches. Cambridge Uni- 
versity Press, Cambridge, 204 pp. 


LanGc, H. 1923. Recent and historical notes on the 
square-lipped rhinoceros (Ceratotherium simum). 
Journal of Mammalogy, 4:155-163. 

Larsen, E. C. 1986. Competitive release in micro- 
habitat use among coexisting desert rodents: a nat- 
ural experiment. Oecologia, 69:231-237. 

Layne, J. N., AND J.O. WHITAKER, JR. 1992. William 
J. Hamilton. Jr. 1902-1990. Journal of Mammalogy, 
73:693-706. 

LE BoeuF, B. J., AND J. REITER. 1988. Lifetime re- 
productive success in northern elephant seals. Pp. 
344-362, in Reproductive success (T. H. Clutton- 
Brock, ed.). University of Chicago Press, Chicago, 
538 pp. 

Lee, A. K., AND A. CockKBuRN. 1985. The evolu- 
tionary ecology of marsupials. Cambridge Univer- 
sity Press, Cambridge, 274 pp. 

Lemen, C. A. 1978. Seed size selection in hetero- 
myids. Oecologia, 35:13-19. 

LEMEN, C. A., AND M. L. ROSENZWEIG. 1978. Micro- 
habitat selection in two species of heteromyid ro- 
dents. Oecologia, 33:127-135. 

LeopoLp, A. 1933. Game management. Scribners, 
New York, 481 pp. 

LEWONTIN, R. C. 1968. Population biology and evo- 
lution. Syracuse University Press, Syracuse, New 
York, 205 pp. 

Lipicker, W. Z., JR. 1988. Solving the enigma of 
microtine “cycles.”’ Journal of Mammalogy, 69:693- 
704. 


1994. Population Ecology. Pp 323-347, in 

seventy-five years of mammalogy (1919-1994) (E. 

C. Birney and J. R. Choate, eds.). Special Publica- 

tion, American Society of Mammalogists, | 1:1-433. 

In press. In defense of a multifactor per- 
spective in population ecology: a response to Gaines, 
Stenseth, Johnson, Ims, and Bondrup-Nielsen. Jour- 
nal of Mammalogy. 

LInSDALE, J. M. 1946. The California ground squir- 
rel. University of California Press, Berkeley, 475 pp. 

LINSDALE, J. M., AND L. P. TEvis. 1951. The dusky- 
footed wood rat. University of California Press, 
Berkeley, 664 pp. 

M’CLoskey, R. T. 1976. Community structure in 
sympatric rodents. Ecology, 57:728-739. 

. 1978. Niche separation and assembly in four 

species of Sonoran Desert rodents. The American 

Naturalist, 1 12:683-694. 

1981. Microhabitat use in coexisting desert 
rodents—the role of population density. Oecologia, 
50:310-315. 

MacArTHvur, R.H. 1958. Population ecology of some 
warblers in northeastern coniferous forests. Ecology, 
39:599-619. 

. 1960. On the relative abundance of species. 

The American Naturalist, 94:25-36. 

. 1965. Patterns of species diversity. Biological 

Reviews, 40:510-533. 

. 1970. Species packing and competitive equi- 

librium for many species. Theoretical Population Bi- 

ology, 1:1-11. 

1972. Geographical ecology. Patterns in the 

distribution of species. Harper and Row, New York, 

269 pp. 


EVOLUTIONARY ECOLOGY Gis) 


Mares, M. A., AND D. F. WiLiiAms. 1977. Experi- 
mental support for food particle size resource allo- 
cation in heteromyid rodents. Ecology, 58:1186- 
1190. 

Maser, C., J. M. TRAPPE, AND R. A. NussBAuM. 1978. 
Fungal-small mammal interrelationships with em- 
phasis on Oregon coniferous forests. Ecology, 59: 
799-809. 

Mayr, E. 1942. Systematics and the origin of species. 
Columbia University Press, New York, 334 pp. 

1963. Animal species and evolution. Har- 
vard University Press, Cambridge, Massachusetts, 
813 pp. 

McCasg, T. T., AND B. D. BLANCHARD. 1950. Three 
species of Peromyscus. Rood Associates, Santa Bar- 
bara, California, 136 pp. 

McCu.L.LouGu, D.R. 1979. The George Reserve deer 
herd. University of Michigan Press, Ann Arbor, 271 
pp. 

McNa~s, B. K. 1963. Bioenergetics and the deter- 
mination of home range size. The American Natu- 
ralist, 97:133-140. 

Mearns, E. A. 1907. The mammals of the Mexican 
boundary of the United States. Bulletin of the United 
States National Museum, 56:1-530. 

Mecu, L. D. 1966. The wolves of Isle Royale. U.S. 
National Park Service, Fauna ser. 7, 210 pp. 

1970. The wolf: the ecology and behavior of 
an endangered species. American Museum of Nat- 
ural History, Natural History Press, Garden City, 
New York, 384 pp. 

ME_ERRIAM, C. H. 1890. Results of a biological survey 
of the San Francisco Mountain Region and desert of 
the Little Colorado, Arizona. North American Fau- 
na, 3:1-136. 

MILiar, J.S. 1977. Adaptive features of mammalian 
reproduction. Evolution, 31:370-386. 

MILLAR, J. S., AND R. M. ZAMMuTO. 1983. Life his- 
tories of mammals: an analysis of life tables. Ecology, 
64:63 1-635. 

Miter, A. H. 1943. Joseph Grinnell’s philosophy 
of nature. University of California Press, Berkeley, 
California, 237 pp. 

MILter, G.S. 1929. Mammalogy and the Smithsoni- 
an Institution. Smithsonian Report for 1928, Pub- 
lication 2995, United States Government Printing 
Office, pp. 391-411. 

Miiter, R. S. 1967. Pattern and process in compe- 
tition. Advanced Ecology Research, 4:1-74. 

Monson, G., AND W. KessLeR. 1940. Life history 
notes on the banner-tailed kangaroo rat, Merriam’s 
kangaroo rat, and the white-footed wood rat in Ar- 
izona and New Mexico. The Journal of Wildlife 
Management, 4:37-43. 

Moors, P. J. 1984. Coexistence and interspecific 
competition in the carnivore genus Mustela. Acta 
Zoologica Fennica, 172:37-40. 

Moraan, L. H. 1868. The American beaver and his 
works. J. B. Lippincott, Philadelphia, 330 pp. 

Morris, D. W. 1984. Patterns and scale of habitat 
use in two temperate-zone small mammal faunas. 
Canadian Journal of Zoology, 62:1517-1523. 

Muncer, J.C., AND J. H. BRown. 1981. Competition 


in desert rodents: an experiment with semipermeable 
exclosures. Science, 211:510-512. 

Muncer, J. C., AND J.C. Hotmes. 1988. Benefits of 
parasitic infection: a test using a ground squirrel sys- 
tem. Canadian Journal of Zoology, 66:222-227. 

Morte, A. 1944. The wolves of Mt. McKinley. Unit- 
ed States Department of the Interior, National Park 
Service Fauna series, 5:1—238. 

MuRriIe_, J. O., AND G. R. MICHENER (EDS.). 1984. The 
biology of ground-dwelling squirrels. University of 
Nebraska Press, Lincoln, 464 pp. 

Muut, I. 1990. William Henry Burt: 1903-1987. 
Journal of Mammalogy, 71:113-116. 

Oscoop, W.H. 1943. Clinton Hart Merriam— 1855- 
1942. Journal of Mammalogy, 24:421-436. 

Owen, J.G. 1990. An analysis of the spatial structure 
of mammalian distribution patterns in Texas. Ecol- 
ogy, 71:1823-1832. 

Packer, C. 1986. The ecology of sociality in felids. 
Pp. 429-451, in Ecological aspects of social evolu- 
tion: birds and mammals (D. I. Rubenstein and R. 
W. Wrangham, eds.). Princeton University Press, 
Princeton, New Jersey, xxx pp. 

PACKER, C., ET AL. 1988. Reproductive success of 
lions. Pp. 363-383, in Reproductive success (T. H. 
Clutton-Brock, ed.). University of Chicago Press, 
Chicago, 538 pp. 

Peters, R. H. 1983. Ecological implications of body 
size. Cambridge University Press, Cambridge, 329 


Peterson, R. S., AND G. A. BARTHOLOMEW. 1967. 
The natural history and behavior of the California 
sea lion. Special Publications, The American Society 
of Mammalogists, 1:1-79. 

PETTICREW, B. G., AND R. M. F. S. SADLER. 1974. 
The ecology of the deer mouse Peromyscus mani- 
curist in a coastal coniferous forest. I. Population 
dynamics. Canadian Journal of Zoology, 52:107-118. 

PireL_KA, F. A. 1957. Some aspects of population 
structure in the short-term cycles of the brown lem- 
ming in northern Alaska. Cold Spring Harbor Sym- 


Price, M. V. 1978. The role of microhabitat in struc- 
turing desert rodent communities. Ecology, 59:910- 
921. 

Price, P. W. 1980. Evolutionary biology of parasites. 
Princeton University Press, Princeton, New Jersey, 
237 pp. 

RALLs, K., AND P. H. Harvey. 1985. Geographic 
variation in size and sexual dimorphism of North 
American weasels. Biological Journal of the Linnean 
Society, 25:119-167. 

Rapoport, E.H. 1982. Areography. Pergamon Press, 
New York, 269 pp. 

REICHMAN, O. J., D. T. WICKLOW, AND C. REBAR. 1985. 
Ecological and mycological characteristics of the 
caches in the mounds of Dipodomys spectabilis. Jour- 
nal of Mammalogy, 66:643-651. 

Reyno_ps, H. G. 1958. Ecology of Merriam’s kan- 
garoo rat on the grazing lands of southern Arizona. 
Ecological Monographs, 28:111-127. 

1960. Life history notes on Merriam’s kan- 

garoo rat in southern Arizona. Journal of Mam- 

malogy, 41:48-58. 


396 BROWN AND WILSON 


RICHARDSON, J. 1829. Fauna boreali-Americana; or 
the zoology of the northern parts of British America: 
containing descriptions of the objects of natural his- 
tory collected on the late northern land expedition 
under command of Captain Sir John Franklin. J. 
Murray, London, 4 volumes. 

Rosenzweic, M. L. 1966. Community structure in 
sympatric Carnivora. Journal of Mammalogy, 47: 
602-612. 

1968. The strategy of body size in mam- 

malian carnivores. The American Midland Natu- 

ralist, 80:299-315. 

1973. Habitat selection experiments with a 
pair of coexisting heteromyid rodent species. Ecol- 
ogy, 54:111-117. 

ROSENZWEIG, M. L., B. SMIGEL, AND A. KRAFT. 1975. 
Patterns of food, space and diversity. Pp. 241-268, 
in Rodents in desert environments (I. Prakash and 
P. Gosh, eds.). W. Junk, The Hague, The Nether- 
lands. 

ROSENZWEIG, M. L., AND P. W. STERNER. 1970. Pop- 
ulation ecology of desert rodent communities: body 
size and seed husking as bases for heteromyid co- 
existence. Ecology, 51:217-224. 

ROSENZWEIG, M. L., AND J. WINAKUR. 1969. Popu- 
lation ecology of desert rodent communities: habi- 
tats and environmental complexity. Ecology, 50:558- 
S72. 

SANCHEZ-CORDERO, V., AND T. H. FLEMING. 1993. 
Ecology of tropical heteromyids. Pp. 596-617, in 
Biology of heteromyid rodents (H. H. Genoways and 
J. H. Brown, eds.). Special Publications, The Amer- 
ican Society of Mammalogists, 10:1-719. 

SCHOENER, T. W. 1968. Sizes of feeding territories 
among birds. Ecology, 49:123-131. 

SCHRODER, G. D., AND M. L. RosENzweEIG. 1975. Per- 
turbation analysis of competition and overlap in 
habitat utilization between Dipodomys ordii and Di- 
podomys merriami. Oecologia, 19:9-28. 

Seton, E.T. 1909. Life histories of northern animals. 
Charles Scribner’s Sons, New York, 2 volumes, 299 


1923. The arctic prairies. Charles Scribner’s 

Sons, New York, 308 pp. 

1929. Lives of game animals. Doubleday, 
Doran and Company, Inc., Garden City, New York, 
4 volumes. 

SHEPPARD, D. H. 1971. Competition between two 
chipmunk species. Ecology, 52:320-329. 

Stmpson, G. G. 1940. Mammals and land bridges. 
Journal of the Washington Academy of Sciences, 30: 
137-163. 

. 1943. Mammalsand the nature of continents. 

American Journal of Science, 241:1-31. 

1944. Tempo and mode in evolution. Co- 

lumbia University Press, New York, 237 pp. 

1947a. Evolution, interchange and resem- 
blance of the North American and Eurasian Ceno- 

zoic mammalian fauna. Evolution, 1:218-220. 

1947b. Holarctic mammalian faunas and 

continental relationships during the Cenozoic. Bul- 

letin of the Geological Society of America, 58:61 3- 

688. 


. 1950. History of the fauna of Latin America. 
American Scientist, 38:36 1-389. 


1953, The major features of evolution. Co- 

lumbia University Press, New York, 424 pp. 

. 1964. Species density of North American Re- 
cent mammals. Systematic Zoology, 13:57-73. 

SMIGEL, B. W., AND M. L. RoSENzweIG. 1974. Seed 
selection in Dipodomys merriami and Perognathus 
penicillatus. Ecology, 55:329-339. 

SmituH, C. C. 1970. The coevolution of pine squirrels 
(Tamiasciurus) and conifers. Ecological Mono- 
graphs, 40:349-371. 

SMYTHE, N. 1970. Relationships between fruiting 
seasons and seed dispersal methods in a Neotropical 
forest. The American Naturalist, 104:25-35. 

STICKEL, L. F., ANDO. WARBACH. 1960. Small-mam- 
mal populations ofa Maryland woodlot, 1949-1954. 
Ecology, 41:269-286. 

SUMNER, F. B. 1932. Genetic, distributional, and evo- 
lutionary studies of the subspecies of deer mice 
(Peromyscus maniculatus). Bibliography of Genet- 
ics, 9:1-106. 

Taitt, M. J., AND C. J. Kress. 1985. Population 
dynamics and cycles. Pp. 567-620, in Biology of 
New World Microtus (R. H. Tamarin, ed.). Special 
Publications, The American Society of Mammalo- 
gists, 8:1-893. 

TAMARIN, R. H. (ED.). 1985. Biology of New World 
Microtus. Special Publications, The American So- 
ciety of Mammalogists, 8:1-893. 

TAYLOR, W. P. (ED.). 1956. The deer of North Amer- 
ica. The Stackpole Company, Harrisburg, Pennsyl- 
vania, and the Wildlife Management Institute, 
Washington, D.C., 668 pp. 

TuHompson, S. D. 1982a. Microhabitat utilization and 
foraging behavior of bipedal and quadrupedal het- 
eromyid rodents. Ecology, 63:1303-1312. 

1982b. Structure and species composition of 
desert heteromyid rodent species assemblages: ef- 
fects of a simple habitat manipulation. Ecology, 63: 
1313-1321. 

Tuwalites, R. G. 1904. The original journals of the 
Lewis and Clark expeditions. Dodd, Mead and Com- 
pany, Inc., New York, 8 volumes, 

Timm, R. M., AND J. S. AsHE. 1988. The mystery of 
the gracious hosts. Natural History, 9:6-9. 

Timm, R. M., D. E. Witson, B. L. CLAuson, R. K. 
LAVAL, AND C. S. VAUGHAN. 1989. Mammals of 
the La Selva-Braulio Carrillo Complex, Costa Rica. 
North American Fauna, 75:1-162. 

Voruies, C. T., AND W. P. TAyYLor. 1940. Life history 
and ecology of the white-throated wood rat, Neot- 
oma albigula Hartley, in relation to grazing in Ari- 
zona. University of Arizona College of Agriculture, 
Agricultural Experiment Station Technical Bulletin, 
86:453-529. 

WALKER, E. P. 1964. Mammals of the world. The 
Johns Hopkins University Press, Baltimore, Mary- 
land, 3 volumes. 

WECKER, S. C. 1963. The role of early experience in 
habitat selection by the prairie deer mouse, Pero- 
myscus maniculatus bairdii. Ecological Monographs, 
33:307-325. 

. 1964. Habitat selection. Scientific American, 
211:109-116. 

WuitaKker, J. O., JR. 1994. Academic propinquity, 
Pp. 121-138, in Seventy-five years of mammalogy 


EVOLUTIONARY ECOLOGY ao7 


(1919-1994). Special Publications, The American 
Society of Mammalogists, 11:1-433. 

WuHiItTForRD, W. G. 1976. Temporal fluctuations in 
density and diversity of desert rodent populations. 
Journal of Mammalogy, 57:351-369. 

Witson, D. E. 1971. Ecology of Myotis nigricans 
(Mammalia: Chiroptera) on Barro Colorado Island, 
Panama Canal Zone. Journal of Zoology, 163:1-13. 

. 1973. Bat faunas: a trophic comparison. Sys- 

tematic Zoology, 22:14-29. 


Witson, D. E., AND J. S. FINDLEY. 1970. Reproduc- 
tive cycle of a Neotropical insectivorous bat, Myotis 
nigricans. Nature, 225:1155. 

Witson, J. W., II. 1974. Analytical zoogeography of 
North American mammals. Evolution, 28:124-140. 

Younsa, S. P., ANDE. A.GOLDMAN. 1944. The wolves 
of North America. American Wildlife Institute, 
Washington, D.C., 636 pp. 


BEHAVIOR 


JOHN F. EISENBERG AND JERRY O. WOLFF 


Introduction 


umankind is a product of organic evo- 
lution, as are all living organisms on 
the planet Earth. Although anthropologists 
may disagree as to the date of transition 
from Homo erectus to H. sapiens, the cul- 
tural transition of some 100 to 70 x 103 
years B.P. by H. sapiens was profound. 
Modern humans developed the capacity for 
rapid cultural evolution and, in conjunction 
with a very large brain, began to set about 
taming the environment in ways we can 
scarcely comprehend except through the 
minds and evidence of archaeologists. Our 
early ancestors were gifted naturalists and 
keen observers of nature. The manner of 
life styles exhibited by organisms of concern 
to the early economic systems were well 
known and communicated by direct partic- 
ipation in hunting, gathering, and also pre- 
sumably by an oral tradition. Thus, a 
knowledge of the behavior and life history 
of animals and plants has been part and 
parcel to our cultural heritage as human be- 
ings (Count, 1973). 

By the middle of the 19th Century, Eu- 
ropean naturalists were beginning to move 
away from the naming and describing of 
floras and faunas and attempting to grapple 
with the more intangible aspects of biology. 
One aspect that preoccupied attention was 


398 


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4 
4, 


animal behavior. The behavior of organ- 
isms has always held a special fascination. 
Consider the admonition of King Solomon 
“Behold the ant, thy sluggard and consider 
her ways” (Proverbs 6:6, King James ver- 
sion of the Bible). By the middle of the 19th 
Century two major schools of thought had 
developed. On the one hand, the empiri- 
cists, following René Descartes, tried to an- 
alyze the behavior of non-human mammals 
in terms of a mechanistic model. Questions 
were posed concerning what an animal could 
perceive and thus respond to. Elaborate ex- 
periments were designed to determine the 
limits of human and non-human animal 
perceptions (Mach, 1959). On the other 
hand, a determined group of naturalists per- 
sisted in attempting to describe (in writing) 
what animals actually did in their natural 
habitat. Charles Darwin, arguably the finest 
19th-Century English speaking, objective 
observer and recorder of nature, who wrote 
and communicated his thoughts in the 
1800s, was also concerned with animal be- 
havior (Darwin, 1859, 1872). His influence 
was profound because he not only offered 
an explanation for morphological change 
through natural selection, but also suggested 
avenues for the study of behavioral change, 
ultimately controlled by natural selection. 


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BEHAVIOR 299) 


A Brief History of Ethology: 
its Origins, Reception, and 
Modification in America 


In 1973, Niko Tinbgergen, Konrad Lo- 
renz, and Karl von Frisch received the No- 
bel Prize in Physiology and Medicine. These 
three men exemplified the early 20th-Cen- 
tury fruit of the Darwinian revolution in 
terms of the analysis of animal behavior (see 
Lorenz, 1981 for a review). Lorenz empha- 
sized the close observation of behavior in- 
volving animals kept at semi-liberty, but 
habituated to human observers. The com- 
parative method was stressed. Tinbergen 
championed the observation of wild crea- 
tures and was ingenious in developing ex- 
perimental techniques applied to free-living 
populations. von Frisch was the consum- 
mate experimentalist and studied the per- 
ceptual capacities of fish and bees (von 
Frisch, 1950). The general theory that these 
three men developed was first consolidated 
in Tinbergen’s book, The Study of Instinct, 
published in English in 1951. The book out- 
lined a theoretical framework for the anal- 
ysis of behavior, but the analysis rested 
firmly on the correct description of an an- 
imal’s repertoire—the ethogram. It was not- 
ed that an animal’s behavior consisted at 
least of two major types of actions: 1) seek- 
ing an appropriate goal (appetitive behav- 
ior); and 2) satisfying a need (consumma- 
tory behavior). With this publication the 
framework was set for the next 20 years of 
research in North America and Europe. 

The theoretical framework built in Eu- 
rope on Darwinian foundations was resisted 
by many 20th-Century students of animal 
behavior in North America. This situation 
derived mainly from the fact that the North 
Americans were often associated with psy- 
chology departments that were strongly tied 
to the experimental method and Cartesian 
reductionism. Experimental design was par- 
amount and asa result two American schools 
developed: 1) those devoted to the physi- 
ological mechanisms underlying discrete 


behaviors—a reductionist position; and 2) 
those devoted to the analysis of how ani- 
mals learn. Both approaches (with a view 
toward public funding) were justified before 
the general public and elected officials on 
the grounds that animal “surrogates” could 
lead to a better understanding of human 
behavior. Thus, the study of non-human 
animals in and of themselves was subli- 
mated to utilitarian needs in terms of hu- 
man welfare. A possible third North Amer- 
ican tradition was grounded in an attempt 
to understand cognitive processes. While 
experimental designs were important, the 
concept of higher mental processes and how 
to study the phenomenon has remained elu- 
sive (Dewsbury, 1989a; Schusterman et al., 
1986). 

Strangely enough, in Europe and North 
America, many of the earlier studies of 
mammalian behavior were undertaken by 
naturalists and wildlife managers who jus- 
tified their activities in terms of understand- 
ing the life history of organisms that were 
of economic importance in terms of “har- 
vesting” or “‘control’’ by humans (Leopold, 
1933). In fact, the applied researchers in- 
vestigating vertebrate behavior were often 
considered outside the boundaries of “‘pure 
science” by many academics and such a 
dreary dichotomy was to persist for some 
time (Wilson and Eisenberg, 1990). Nev- 
ertheless, regardless of the motivation, an- 
imal behavior has held a great fascination 
for all observers across all cultures. 

The social behavior of animals has long 
preoccupied mankind. Aside from the won- 
ders of individualistic behaviors, the vari- 
ety of patterns displayed during mating, pa- 
rental care, and seemingly altruistic 
behaviors were of special concern after the 
Darwinian revolution. Darwin (1859), Kro- 
potkin (1902), and Deegener (1918) grap- 
pled with the problem. Tinbergen (1951) 
pointed out that a social system is basically 
a communication system and thus opened 
a new arena of research. A paper by W. D. 
Hamilton (1964) was revolutionary because 
it laid the groundwork for a rational analysis 


400 EISENBERG AND WOLFF 


of how societies could evolve through nat- 
ural selection. Eisenberg (1966) outlined the 
evolutionary trends of social behavior with- 
in the class Mammalia. Trivers (1971, 1972, 
1974) amplified and clarified some intricate 
problems raised by Hamilton (1964), and 
E. O. Wilson (1975) brought the most re- 
cent, comprehensive synthesis to the fore- 
front. Darwinian selection, Mendelian ge- 
netics, ecology, and behavior had been 
wedded into a system of testable hypothe- 
ses. This revolution in thought will be treat- 
ed in a later section. 


A Record of the Beginnings of 
Animal Behavior Studies in 
North America 


Given that the basis of ethology is an- 
chored in the systematic observation of an 
animal’s behavior, who can we identify as 
the first North American mammalian ethol- 
ogist? With all due respect, we must over- 
look the preliterate but viable cultures of 
hunter-gatherers that preceded European 
occupation of the continent. We suggest 
(among others) L. H. Morgan as a candi- 
date. Not only did he write a classic work 
on the beaver (Castor canadensis) (Morgan, 
1868), but he also wrote a magnificent eth- 
nography of the Iroquois Indians in New 
York and Ontario (Morgan, 1851). While 
not an ethologist in the 20th-Century sense, 
he nevertheless was an objective observer 
and dutiful recorder of his observations. 
While such naturalists as Audubon and 
Bachman (1846-1854) recorded facts con- 
cerning the habits of their subjects, L. H. 
Morgan concentrated on a single species or 
a human culture and described their behav- 
ior and social structure in astonishing detail. 

Through the late 19th and early 20th cen- 
turies, mammalian behavior patterns con- 
tinued to be described often in a fragmented 
fashion or as a series of anecdotes. Ernest 
Thompson Seton (1953) made a fine con- 
tribution in the compilation of anecdotes 


by organizing descriptions of behavior with- 
in species accounts in the form of a func- 
tional classification, e.g., mating behavior, 
parental care, feeding and foraging, and the 
like (Note: Seton did not apply these exact 
subheadings, but the sense was there.) 

The beginning of a theoretical framework 
for behavior studies grounded in Darwinian 
theory started within North America during 
the late 19th and early 20th centuries with 
the work of C. O. Whitman, who observed 
that the courtship of pigeons was composed 
of numerous stereotypic components. The 
behavioral units and their sequencing were 
often characteristic of each domestic breed. 
Could some types of behavior be compared 
among breeds or species in the manner of 
a comparative anatomist? Were the units of 
behavior as expressions of nerve-muscle re- 
lationships subject to the laws of heredity 
(Whitman, 1899, 1919)? 

Whitman’s student, Wallace Craig, chose 
bird song for comparative study and soon 
discovered that while some songs were spe- 
cies specific and relatively fixed, other spe- 
cies show plasticity and a good deal of learn- 
ing in song development (Craig, 1918). Craig 
and Whitman were pioneers in their studies. 
Parallel efforts in Europe by Oskar Heinroth 
and Konrad Lorenz led to the founding of 
European ethology, but in North America, 
behavior studies developed on many dif- 
ferent fronts with little intellectual cross fer- 
tilization (Dewsbury, 19895). Application 
of these concepts to mammalian behavior 
was to occur much later (see the next sec- 
tion). 


Mammalian Behavior Studies 
Prior to 1965 


Threads in the loom—behavior studies. — 
Ethology as a discipline did not become 
consolidated in the U.S. until the mid-1950s. 
Although a knowledge of “‘species-typical 
behavior’ was a working tool for all great 
naturalists, to presume that behavior stud- 
ies represent something new is to oversim- 


BEHAVIOR 401 


plify a very complex situation. Our prede- 
cessors and seniors of the last 70 years were 
involved with behavior studies, whether or 
not their labors were organized into a formal 
system. For example, Vernon Bailey, who 
worked with the U.S. Biological Survey, was 
intrigued by the behavior of his subjects 
(Bailey, 1931). Ned Hollister, before he took 
command of the U.S. National Zoological 
Park, wrote a classic paper concerning the 
effects of captivity and captive diets on the 
skull morphology of African lions (Hollis- 
ter, 1917). Joseph Grinnell, the spirit of the 
Museum of Vertebrate Zoology at Berkeley 
during its most formative years, was one of 
the most astute observers of vertebrate be- 
havior ever to document his observations 
(Grinnell, 1914). A. Brazier-Howell was 
deeply concerned with the problems of form 
and function, a true behaviorist by anyone’s 
definition (Howell, 1944). Shadle (1946) 
with his delightful, yet incisive, observa- 
tions on the sexual life of porcupines is also 
a case in point. While on the subject of 
mammalian reproduction, the efforts of R. 
K. Enders (1935, 1952) and O. P. Pearson 
(1944) in mammalian behavior studies stand 
out, not to diminish their other considerable 
contributions. Many others could be cited 
(Bronson, 1989 for review). 

One area of the discipline of behavior that 
has not received much attention from the 
standpoint of the “behaviorist” is that vague 
area of energetics and behavior, or “‘eco- 
physiology,” which not only has had a long 
history, but also a profound influence on 
the types of questions that behaviorists ask. 
The beginnings may go back to Claude Ber- 
nard in the 19th Century but the fact of the 
matter remains that in the 1940s mam- 
malogists began asking hard questions con- 
cerning how mammals were able to with- 
stand the rigors of adverse environments. 
Morrison and B. K. McNab began to ask 
the questions and seek the answers (Mor- 
rison and McNab, 1962), as did Bartholo- 
mew (Bartholomew and Cade, 1957). Feed- 
back between the so-called behaviorists and 
the physiologists continued (McNab, 1983). 


Another area of research with a long his- 
tory of mammals as subjects includes be- 
havioral genetics. Sumner (1932) and sub- 
sequently Lee Dice literally pioneered the 
research on the genetics of non-domesti- 
cated mammals (Dice, 1933). Peromyscus 
was their genus of choice and it was a sound 
one. With the Michigan stocks, Howard 
(1948), Harris (1952), and King (1961) were 
to shape the thinking of younger biologists 
concerning the genetics of behavior in the 
1950s (see also King, 1968). 

Population dynamics and the behavior of 
mammalian species at different densities has 
become a focus of interest since the synthe- 
sis published by Elton (Crowcroft, 1991 for 
review). The pioneers on this frontier of the 
1950s included D. E. Davis, D. Chitty, J. 
B. Calhoun, and J. Christian (Anderson, 
1989; Cockburn, 1988 for reviews). The role 
of density-dependent and density-indepen- 
dent factors on the regulation of population 
size was a “hot topic” at that time, and the 
discovery that endocrine changes could me- 
diate and be mediated by behavioral changes 
only added fuel to the fires of controversy 
(Calhoun, 1963a, 19636; Christian, 1963). 
That behavior could be linked to the genetic 
background of an individual led to a flurry 
of productive research and once again be- 
havioral studies were an integral part of the 
effort (Calhoun, 1963a; Harris, 1952). 

The unique sensory abilities of mammals 
had long been recognized, but D. R. Grif- 
fin’s publication on the echolocation of bats 
in 1958 was truly a watershed. Kellogg 
(1961) synthesized similar data for dol- 
phins. Bioacoustics became a field unto its 
own. 

At Cornell, W. J. Hamilton, Jr. and his 
colleagues initiated important studies on 
mammalian food habits. Although many 
other aspects of mammalian behavior were 
studied at Cornell, perhaps one of the most 
notable single-species monographs was 
James Layne’s contribution on the behavior 
and ecology of the red squirrel (Tamuasiurus 
hudsonicus) (Layne, 1954). 

The use of livetraps for the purpose of 


402 EISENBERG AND WOLFF 


trap, mark, and release studies opened a new 
era in the studies of how mammals use space. 
H. B. Sherman invented a successful metal 
livetrap in the late 1930s that is marketed 
to this day. Sherman and his students at the 
University of Florida developed a series of 
studies aimed at clarifying microhabitat use 
and the spacing behavior of small mammals 
utilizing the trap, mark, and release scheme. 
William Burt, utilizing a livetrap modifi- 
cation of his own at Michigan, wrote an 
influential paper in 1940 proposing that 
some species of small mammals appeared 
to show territorial behavior (Burt, 1940). 
The study of nocturnal, cryptic mammals 
and their movements received an enormous 
assist with the introduction of radiotele- 
metric techniques in the 1960s. Perhaps the 
most pioneering group was associated with 
the University of Minnesota with their mag- 
nificent setup at the Cedar Creek Natural 
History Area (Tester et al., 1964). 

As an aside, immobilization of mammals 
with a reliable series of drugs and instru- 
ments for projection was revolutionary 
(Harthoorn, 1976). Younger students will 
not appreciate fully the revolution intro- 
duced by reliable telemetry and pharma- 
ceutical systems. 

Given the advanced techniques of trap, 
mark, and release, monographic treatises 
involving these methods began to supple- 
ment direct observation. The focus was of- 
ten ecological, but behavior became more 
and more a concern regardless of technique: 
Linsdale and Tevis (1951) on the dusky- 
footed woodrat, Neotoma fuscipes; Linsdale 
(1946) on the California ground squirrel; 
Linsdale and Tomich (1953) on Odocoileus 
hemionus; Moore (1957) on Sciurus niger; 
and Layne (1954) on Tamiasciurus hud- 
sconicus all appeared in the 1940s and 1950s. 
One of the benchmark field studies of mam- 
malian social behavior was John King’s 
monograph on the black-tailed prairie dog, 
Cynomys ludovicianus (King, 1955). This 
classic study demonstrated that careful ob- 
servations of marked individuals could yield 
insight into the use of space, mode of com- 


munication, and relations among kin. King’s 
effort paved the way for field experiments 
and ever more sophisticated studies of di- 
urnal sciurids (Murie and Michener, 1984, 
1989 for review). 

Nocturnal small mammals still presented 
problems because direct observation was not 
possible. Eisenberg, following the tech- 
niques developed by Eibl-Eibesfeldt (1958) 
in Germany, developed the strategy of com- 
bining field studies with captive studies. 
With the aid of the electronic flash camera, 
behavior patterns of small, nocturnal mam- 
mals could be recorded on film for analysis 
(Eisenberg, 1962, 1963). 

Kaufmann in the late 1950s and early 
1960s carried out a classic field study on a 
diurnal carnivore, the coati (Nasua narica) 
in Panama (Kaufmann, 1962). His creative 
analysis of the social use of space by female 
bands has stood the test of time. Shortly 
thereafter, Valerius Geist produced his clas- 
sic study of Ovis dalli and O. canadensis in 
British Columbia (Geist, 1971). Ungulate 
behavior studies had come of age. Kleiman 
(1967) stimulated interest in the compara- 
tive social behavior of the Canidae. McKay 
(1973), based upon his studies in the 1960s, 
brought the Asiatic elephant to the forefront 
of attention. 

DeVore, in his studies of the baboon (Pa- 
pio cynocephalus) in Kenya, ushered in the 
new era of primate studies; Schaller’s study 
of free-ranging mountain gorillas (Gorilla 
gorilla beringei) was a true milestone in the 
art of field work (DeVore, 1965; Schaller, 
1963). They demonstrated that a field work- 
er could habituate the subjects to the pres- 
ence of an intruder. Eisenberg and Kuehn 
(1966) attempted a synthesis for neotropical 
primates. 

The pure ethological approach based on 
efforts as applied to mammals (Hediger, 
1942) was summarized in R. F. Ewer’s 
(1969) classic, The Ethology of Mammals. 
New disciplines were already forming 
around the interface between ecology and 
behavior. Suitably inspired, Smythe and 
Wemmer working in the 1960s provided 


BEHAVIOR 403 


important contributions (Smythe, 1970; 
Wemmer, 1977). With the inclusion of pop- 
ulation genetics the stage was set for the 
development of sociobiology as a synthetic 
discipline by Wilson in 1975 (see section 
From ethology to sociobiology). 

The watershed of the late 1960s.—In 1969, 
the Smithsonian Institution convened its 
public symposium under the broad title of 
Man and Beast, the results of which were 
published in 1971 (Eisenberg and Dillon, 
1971). The symposium and its published 
results were an attempt to focus public at- 
tention on the relevance of animal behavior 
studies to understanding human behavior. 
To this end, the participants in the sym- 
posium included biologists, philosophers, 
psychologists, anthropologists and sociolo- 
gists. In part, the public symposium was in 
response to the recently published work of 
Konrad Lorenz titled in English translation, 
On Aggression. The notion that some as- 
pects of human behavior could have a ge- 
netic basis was anathema to some of the 
North American social scientists. As Wat- 
son (1914) had proclaimed some years be- 
fore, the human mind could be considered 
at birth as a tabula rasa, where environ- 
mental conditioning reigns supreme in 
forming the life of the infant, juvenile, and 
subadult. 

One member of the conference, E. O. Wil- 
son, who delivered a provocative paper on 
the evolution of territoriality, was deeply 
moved by the conference. By his own ad- 
mission, it inspired him to produce his clas- 
sic Sociobiology. The raging controversy that 
accompanied the publication of Wilson’s 
synthesis remains a remarkable quirk in the 
development of the behavioral sciences. 
Many of us regarded, with dismay, the vit- 
riolic attacks, often personal, to which Wil- 
son was subjected. However, wounds heal, 
and those aspects of philosophical confron- 
tation that seemed so desperately important 
in the early 1970s diminished, and by the 
time a sequel to the volume was prepared 
via the mechanism of a conference at the 
Smithsonian in 1986, barely a flicker oc- 


curred within the halls of academe. The re- 
sults of this conference were published in 
1991 under the title: Man and Beast Revis- 
ited (Robinson and Tiger, 1991). 

A certain amount of emotional maturity 
must have occurred in the intervening 16 
to 20 years, and one might hope that the 
healing process will continue. It should be 
noted that there were no philosophical vil- 
lains leading to the first major confronta- 
tion, following Man and Beast (1969). To 
the contrary, the philosophical confronta- 
tion of the mid-1970s was long overdue and, 
sadly, somewhat protracted in the manner 
in which the participants registered their 
viewpoints. One felt at the conclusion of the 
1986 symposium in Washington, D.C., that 
the burning issues of the relevance of animal 
behavior studies to the interpretation of hu- 
man behavior had somewhat declined. This 
is not to say that the cross-fertilization dur- 
ing the intervening 20 years had not been 
useful. It simply says that rapid and facile 
generalizations forthcoming from popular- 
ists did not necessarily solve any of the cur- 
rent problems of the human race. Clearly 
the social scientists contributing to the 1986 
symposium, such as Helen Fisher and Lio- 
nel Tiger, had gleaned a great deal from the 
earlier ruminations in 1969. 


The Influence of Some Seminal 
Institutions 


The American Museum of Natural His- 
tory and relations with the New York Zoo- 
logical Society. —The American Museum of 
Natural History (AMNH) was one of the 
earliest museums in the United States to 
create a separate Department of Animal Be- 
havior. The origin of the behavior group 
was established under G. Kingsley Noble 
(See Koestler, 1971, for an account of the 
midwife toad scandal and Noble’s role.). Al- 
though best known for his work with the 
Amphibia, Noble was a pioneer in the anal- 
ysis of the relationship between hormones 
and behavior (Noble, 1931). Thus, he 


404 EISENBERG AND WOLFF 


founded an experimentally based discipline 
that was basically reductionist. After No- 
ble’s premature death, Frank A. Beach was 
appointed to head the group and created the 
Department of Animal Behavior while pur- 
suing the role of hormones and behavior 
(Beach, 1948). He recruited T. C. Schneirla 
in the late 1940s to join him. After World 
War II, the department began in earnest to 
assemble a graduate student group. Beach 
championed the hormone and behavior tra- 
dition, but also brought some of his own 
interests. Beach had been a student of Lash- 
ley, who had pioneered brain and behavior 
studies, and thus a second reductionist tra- 
dition was added. Beach left AMNH for 
Yale, and Schneirla succeeded him as chair. 
E. Tobach, L. Aronson, and D. Lehrman 
became the key players as former students. 
D. Lehrman, a contemporary, would later 
found the Institute of Animal Behavior at 
Rutgers. Aronson, pursuing brain-behavior 
relationships, would continue with fish, but 
also turned to cat behavior. Aronson be- 
came chair on the occasion of Schneirla’s 
retirement. 

Given the ties of the AMNH with the 
local New York universities and subse- 
quently with Rutgers, its influence was con- 
siderable. The research efforts were often 
grounded in attempting to understand phys- 
iological mechanisms underlying behavior 
and were often allied with colleagues in hu- 
man medicine. The application of the re- 
sults of animal-based research to human 
problems became for some a guiding ideal 
(Rosenblatt and Komisaruk, 1977). 

The New York Zoological Society (NYZS) 
maintained relations with the AMNH pri- 
marily through curators in various depart- 
ments of vertebrate zoology. Early in the 
Century, the NYZS sponsored field research 
with an aim to improve knowledge appli- 
cable to the proper captive maintenance of 
exotics. William Beebe was supported and 
his attempts to found field stations in the 
Neotropics are renowned. In the early 1900s 
Beebe had assembled groups of researchers 
in what is now Guayana. Beebe (1925) pub- 


lished the first behavioral ecology study of 
the three-toed sloth, an effort not to be 
equalled until research by Montgomery and 
Sunquist (1975). In the late 1960s, the NYZS 
established the unit that was to become 
‘“Wildlife Conservation International,” 
thereby supporting a core group of mam- 
malogists concerned with the interface be- 
tween ecology and behavior including R. 
Payne, T. Struhsaker, and G. Schaller in the 
original assemblage. 

The University of Chicago.—The Uni- 
versity of Chicago established connections 
with the Field Museum of Natural History 
at an early stage. These close ties contrib- 
uted greatly to the study of zoogeography 
and ecology. Many students of the first au- 
thor’s generation studied the classic Prin- 
ciples of Ecology by Allee, Park, Park, Em- 
erson and Schmidt. The ecologists of the 
Chicago group also had a deep concern with 
the behavior of animals. Emerson concen- 
trated on social insects and the problem of 
the evolution of social behavior. Allee shared 
many of Emerson’s interests, but his con- 
cerns were more wide ranging. Although 
neither Emerson nor Allee may be consid- 
ered mammalogists, their contribution to 
the theoretical links between behavior and 
ecology is incalculable. Indeed the highest 
student award conferred at the annual meet- 
ings of the U. S. Animal Behavior Society 
is the W. C. Allee Award. Upon leaving 
Chicago, Allee joined the University of 
Florida where he had an influence on the 
direction of behavioral research at that in- 
stitution. 

Yale and the primatologists. —Robert 
Yerkes of Yale University pioneered the 
study of primate behavior. A psychologist 
by training, he founded what was to become 
the Yerkes Primate Institute at Orange Park, 
Florida (now at Atlanta, Georgia under 
Emory University). Although Yerkes’ ef- 
forts were directed at captive, nonhuman 
primates, he actively sponsored field re- 
search with a genuine concern for objective 
descriptions of naturalistic behavior (Yer- 
kes and Yerkes, 1929). Bingham and Nissen 


BEHAVIOR 405 


were dispatched to Africa (Bingham, 1932; 
Nissen, 1931); while C. R. Carpenter was 
sent to Panama. Carpenter’s studies of Ateles 
and Alouatta stand today as classics (Car- 
penter, 1934, 1935). He went on to study 
Hylobates and Macaca in Asia (Carpenter, 
1964, for a summary). Sherwood Wash- 
burn, a graduate student during the gibbon 
project, subsequently promoted primate 
studies after World War II. His students 
(including I. DeVore) created a nexus of ac- 
tive research, first at Chicago, and then at 
Berkeley, during the late 1950s and 1960s. 

The history of the Smithsonian in the pro- 
motion of animal behavior studies. —The 
beginnings of animal behavior studies at the 
Smithsonian were rooted in the traditions 
of natural history. The collections at the Na- 
tional Zoological Park (NZP) were studied 
and sketched by artists, most notably by 
Ernest Thompson Seton, to illustrate, in 
part, his Lives of Game Animals. Although 
Ned Hollister and William Mann made nu- 
merous contributions to mammalian nat- 
ural history, behavior studies and docu- 
mentation were not systematically 
approached until E. P. Walker became As- 
sistant Director of the NZP in 1930. Walker 
was interested in photography and pio- 
neered the techniques of the use of syn- 
chronized flash bulbs, allowing bats and fly- 
ing squirrels to be photographed in mid- 
flight. He recorded primate sounds with an 
early version of the sound spectrograph, and 
attempted to describe the vocal repertoire 
of the night monkey (4otus). His arduous 
pursuit of photography eventually led to the 
publication of Mammals of the World after 
his retirement (Walker, 1964). 

The creation of a unit at the NZP with 
the mandate of studying the ethology of 
higher vertebrates was not to occur until 
1965. For the last 28 years, the NZP has 
provided leadership in the study of animal 
behavior and in the interface between be- 
havior and ecology. The full maturity of the 
Smithsonian’s role in behavioral studies 
came at two important points: 1969 when 
the symposium Man and Beast was con- 


vened; and in 1973 when a consortium 
among the University of Maryland, George 
Washington University, and the Smithsoni- 
an Institution hosted the XIth International 
Ethology Conference, marking the first time 
that this international body had convened 
in the USA. 

The University of California, Berkeley. — 
Zoologists at Berkeley had an early interest 
in animal behavior. Samuel J. Holmes pub- 
lished his Animal Intelligence in 1910, and 
W. E. Ritter published The California 
Woodpecker and I in 1938. Thereafter the 
animal behavior studies, particularly of 
higher vertebrates, mainly derived from the 
Museum of Vertebrate Zoology (MVZ). The 
emphasis at the museum was often behavior 
and ecology, or behavior and evolution, both 
approaches firmly anchored in the Darwin- 
ian tradition, and the guiding force in the 
museum was Joseph Grinnell. A student of 
David Starr Jordan, Grinnell was to found 
one of the great dynasties in American 
mammalogy (see Jones, 1991; Whitaker, 
1994). 

Mammalian behavior studies were not the 
sole domain of the MVZ. The Department 
of Psychology also had some giants in the 
field of learning studies, including E. C. Tol- 
man (1932). Tolman’s influence was pro- 
found, because he did not pursue a reduc- 
tionist approach, but rather championed the 
more holistic approach of cognition and 
“higher mental processes.’”’ A. Kroeber, in 
the Department of Anthropology, stimulat- 
ed the study of human cultures on a com- 
parative basis (Kroeber, 1925) and Karl 
Sauer, in the Department of Geography, 
championed the analysis of the role of H. 
sapiens in altering the contemporary envi- 
ronments (1969). All the elements were in 
place for the synthesis at Berkeley that would 
commence in the mid-fifties. 

A case study of synergism: Berkeley, Cal- 
ifornia— 1955—1965.—To illustrate the in- 
terdependency of behavior studies with re- 
spect to related disciplines, allow us to 
pursue a case study—Berkeley, California 
(UC), from 1955 to 1965. At the beginning 


406 EISENBERG AND WOLFF 


of the period, the great museum legacy of 
Grinnell was in place and viable. If we con- 
fine ourselves to senior staff who worked 
with mammals, F. A. Pitelka, A. Starker 
Leopold, O. P. Pearson, and S. B. Benson 
were powerful influences on the cadre of 
aspiring young mammalogists. The special- 
ties of ecology, wildlife management, phys- 
iological ecology, and systematics were well 
represented. In addition, the MVZ had close 
ties with the Department of Paleontology. 
Between 1957 and 1959, four new faculty 
were added to the biological sciences who 
had a significant impact on the ““mammal 
group”: W. Z. Lidicker, Jr., in the MVZ, P. 
V. Marler in Zoology, S. A. Washburn in 
Anthropology, and F. A. Beach in Psychol- 
ogy. 

Leopold, Beach, Washburn, and Marler 
were instrumental in developing the behav- 
ioral research station in the Berkeley hills, 
during the 1960s, but more importantly they 
actively encouraged interdisciplinary stud- 
ies at a significant crossroads in the matu- 
ration of behavioral research at the graduate 
level at UC. In addition, the long standing 
field station, ‘““The Hastings Reserve,” was 
emphasized as a place to do research. Lid- 
icker became a catalyst in promoting an in- 
terface between systematics and mamma- 
lian ecology. Those were indeed “‘heady” 
times. Washburn introduced primates as 
suitable subjects for field studies, Beach ex- 
tolled the virtues of the controlled experi- 
ment and a modified view of the reduction- 
ists’ vision of behavior, and Marler 
presented us with the philosophy of the 
ethologists. 

The original, senior faculty gave all of us 
an anchor associated with the MVZ and 
those virtues as set out by Grinnell. We may 
miss some names, but the younger mam- 
malogists who completed their Ph.D. de- 
grees in Anthropology, Psychology, Zoolo- 
gy, and Paleontology during that period 
included: W. J. Hamilton III, P. K. Ander- 
son, J. Mary Taylor, G. Heinsohn, M. Mu- 
rie, D. Isaac, J. Kaufmann, C. Thaler, T. 
Struhsaker, S. David Webb, B. LeBoeuf, T. 


Grand, S. R. Ripley, P. Jay, D. D. Thiessen, 
L. Clemens, and one of us (J. F. Eisen- 
berg)—J. O. Wolff was of the next genera- 
tion. In addition, we had many close asso- 
ciations with other vertebrate zoologists 
(pre- and postdoctoral) who went on to earn 
their “‘spurs’’ as behaviorists and systema- 
tists including: R. B. Root, D. Wilhoft, R. 
Behnke, Jerram Brown, D. Dewsbury, M. 
Konishi, J. Mulligan, K. Nelson, E. Neil, F. 
Notebaum, G. Orians, J. Nelson, and G. 
Hirsch (see also Marler, 1985). 

If we consider only the cadre of post-bac- 
calaureate ““mammalogists”’ within the pe- 
riod of that “‘magic’’ decade, 12 well-ac- 
claimed books have been produced as of 
1993, one member became the President of 
the ASM, two members became President 
of the Animal Behavior Society (ABS), one 
member won the C. Hart Merriam Award 
at the ASM, one member was President of 
the American Society of Paleontologists 
(ASP), one member became the director of 
a major US metropolitan museum, and all 
taught and mentored graduate students and 
produced numerous publications. In their 
efforts, all had influence to the far corners 
of the Earth including (exclusive of the USA) 
Australia, Canada, Botswana, Namibia, 
Panama, Mexico, Uganda, Kenya, India, Sri 
Lanka, Madagascar, Venezuela, Honduras, 
Chilé, Argentina, and Brazil. 


The Years of Consolidation and 
Subsequent Fractionation 


The Second World War interrupted all 
aspects of pure biological research. Com- 
munication with European colleagues was 
almost non-existent. Some of the ideas from 
European ethologists had begun to be ac- 
cepted by American mammalogists, often 
paradoxically via the ornithological or ich- 
thyological literature. Visits by N. Tinber- 
gen and G. Baerends to North America dur- 
ing the 1950s helped disseminate some of 
these concepts, and the hiring of European 
ethologists at North American universities 


BEHAVIOR 407 


facilitated the process (Dewsbury, 1989a, 
1992). Notable among these early “immi- 
grants” were Peter Marler at Berkeley, Franz 
Sauer at Florida, Erik Klinghammer at Pur- 
due, and Fritz Walther at Missouri and sub- 
sequently at Texas A&M. Whether called 
ethology or animal behavior, the study of 
the behavior of mammals rapidly became 
a part of the curriculum at every major uni- 
versity in North America. There were par- 
allel developments in Australia, South Af- 
rica, New Zealand, Israel, Japan, Kenya, and 
India. Thus a European tradition had taken 
root in many new locations. 

Literally hundreds of students in the 
United States during the 1960s and 1970s 
became involved in animal behavior stud- 
ies. The short period of consolidation was 
followed by the creation of new subdisci- 
plines and new societies. The Animal Be- 
havior section of the Ecological Society of 
America became a full society in 1964. 
Through an arrangement with the British 
Society for the Study of Animal Behavior, 
a newly organized journal of Animal Be- 
haviour served as a publication outlet for 
the fledgling effort. Subsequently, new so- 
cieties were formed with their own journals 
based on taxonomic lines: Chiroptera, Pri- 
mates, Cetaceans, or a “wedding” between 
ecology and behavior. 

One of us (J. F. Eisenberg) remembers at 
our meeting of the ASM in 1964 in Mexico 
City when papers dealing with behavior were 
ararity. By 1983, at our meetings in Gaines- 
ville, Florida, the behavior section was well 
represented (231 presentations). In 1983, the 
ASM also published Special Publication No. 
7, Advances in the Study of Mammalian Be- 
havior (Eisenberg and Kleiman, 1983). This 
volume marks a point of recognition, name- 
ly that behavioral studies had “come of age.” 
There were 27 participants contributing to 
the volume drawn not only from the United 
States, but also from Canada, Australia, En- 
gland, Germany, Israel, and France. In or- 
der to illustrate how behavioral studies span 
many disciplines, we will briefly outline the 
organization of this volume. 


Part one deals with the interwoven themes 
of structure, development, and function; 
obviously, the underpinnings of behavior. 
The second part of the volume deals with 
mechanisms of communication. Commu- 
nication is still the touchstone of behavioral 
studies. That is to say, whether an animal 
be solitary or social, it must have infor- 
mation concerning its conspecific neigh- 
bors, or for that matter, its competitors and 
potential predators. The third section deals 
with case studies of mammalian behavior. 
In this time when testable hypotheses seem 
to dominate as a reason for practicing sci- 
ence, we wish to remind everyone that good, 
solid descriptions are still the matrix and 
the foundation for all subsequent research. 
Part four was entitled, The adaptiveness of 
behavior: constraints, population mecha- 
nisms and evolution. Obviously, the recent 
developments and fragmentation of the 
ethological group are reflected in the eclectic 
nature of the subtitle. Clearly, behavioral 
studies have relevance to students of phys- 
iology, population ecology, genetics, and 
evolution. 


From Ethology to Sociobiology 
and Socioecology—the 
Last 25 Years 


The level of selection—the 1970s.—The 
last 25 years of research in mammalian be- 
havior still have been strongly influenced 
by Darwin’s theory of evolution by natural 
selection. Descriptions of ethograms and 
mechanistic aspects of specific behaviors 
that predominated throughout the 1960s 
were largely replaced by observational and 
empirical studies concerning the adaptive 
or evolutionary significance of behavioral 
patterns. Behavior was still looked upon as 
an adaptive strategy, but within a more re- 
fined context. Research became more ex- 
perimental and was conducted more often 
in natural environments. The “group selec- 
tion” arguments for behavior, such as alarm 
calls and other apparent altruistic behavior 


408 


(Wynne-Edwards, 1962), were largely, but 
not entirely, explained away by kin selection 
(Hamilton, 1964), individualistic selection 
(Williams, 1966), or selfish gene (Dawkins, 
1976) theories. Hamilton’s kin selection, or 
inclusive fitness theory, presented concep- 
tual and mathematical reasoning to explain 
cooperative and nepotistic behavior among 
related individuals, and antagonistic or self- 
ish behavior exhibited toward nonrelatives. 
Also during this period an emphasis was 
placed on concepts, theory, and hypothesis 
testing, rather than studying a species per 
se. The state-of-the-art of animal behavior 
in the early 1970s was reviewed by Richard 
Alexander (1974) and further summarized 
by Alexander and Tinkle (1981), and of 
course E. O. Wilson’s (1975) treatise, So- 
clobiology—the New Synthesis. 

Parental investment and the influence of 
Robert L. Trivers. — Associated with kin se- 
lection and selfish gene theory, several piv- 
otal papers were published in the early 1970s 
that strongly influenced our understanding 
of mammalian behavior. Perhaps the most 
influential paper published during this time 
was Robert L. Trivers’ (1972) theory on pa- 
rental investment and sexual selection. 
Trivers proposed that when one gender pro- 
vided greater parental investment than the 
other, competition occurred among the lat- 
ter for the former. When applied to mam- 
mals, this theory explained the intense com- 
petition observed among males, the 
significance of social organs and secondary 
sex characteristics associated with sexual di- 
morphism, and the predominance of polyg- 
ynous mating systems (Geist, 1974; Ralls, 
1977). Two other contributions by Trivers 
were his theories of reciprocal altruism 
(Trivers, 1971) and parent-offspring conflict 
(Trivers, 1974). Reciprocal altruism was 
used to explain communal nesting in bats 
(Trune and Slobodchikoff, 1978), helping 
among dolphins (Connor and Norris, 1982), 
and cooperative coalition behavior among 
male baboons (Papio anubis, Packer, 1977). 
Reciprocal altruism became an alternative 
explanation for apparent altruistic behavior 


EISENBERG AND WOLFF 


that did not have an inclusive fitness payoff. 
Supportive evidence for the parent-off- 
spring conflict theory was provided in wean- 
ing studies on bighorn sheep (Ovis cana- 
densis, Berger, 1979), red deer (Cervus 
elephas, Clutton-Brock et al., 1984), and 
Rhesus macaques (Macaca mulatta, Go- 
mendio, 1991). 

Facultative sex ratio adjustment. —In 
1973, Trivers published a provocative the- 
ory on facultative sex ratio adjustment 
(Trivers and Willard, 1973). The theory 
states that females should provide more pa- 
rental investment in the sex offspring that 
exhibits the greater variance in reproductive 
success. In mammals, this is usually con- 
sidered to be males. Trivers argued that by 
providing more maternal investment in 
male offspring, sons would be healthier and 
better competitors as adults and thus pass 
on more genes than if their mothers pro- 
vided less investment, even though the male 
cohort could be reduced in numbers at 
adulthood. Likewise, dominant or high 
ranking females or those females in “‘good”’ 
condition should produce sons, or at least 
provide more investment in them, than they 
do in daughters. Conversely, lower ranking 
and less healthy females should produce 
daughters, or at least provide more invest- 
ment in them, than in sons. Support for this 
theory was found in such diverse mammals 
as Galapagoes fur seals (Arctocephalus gal- 
apagoensis, Trillmich, 1986), red deer 
(Clutton-Brock et al., 1968), opossums (Di- 
delphis virginiana, Austad and Sunquist, 
1988; Sunquist and Eisenberg, 1993), toque 
macaques, (Macaca sinical, Dittus, 1977), 
and domestic swine (Sus scrofa, Meikle et 
al., 1993). In 1983, Joan Silk provided an 
alternative hypothesis, which stated that in 
social systems where females compete lo- 
cally for resources (referred to as the local- 
resource competition hypothesis), mothers 
should provide more investment in daugh- 
ters than in sons. Supportive evidence was 
provided for this theory in white-tailed deer 
(Odocoileus virginianus, Caley and Nudds, 
1987) and several primate species (Clark, 


BEHAVIOR 409 


1978; Silk, 1983). The relationship between 
social systems and male and female repro- 
ductive strategies with respect to facultative 
sex ratio adjustment remains an active area 
of research in mammal behavior in the 
1990s. 

Evolutionarily stable strategies (ESSs).— 
Another significant development in animal 
behavior that came out of the 1970s was 
John Maynard Smith’s concept of an evo- 
lutionarily stable strategy or ESS (Maynard 
Smith, 1974, 1982). An ESS is a strategy 
which when adopted by most members of the 
population cannot be beaten by any other 
strategy in the game. The theory attempts 
to explain the “‘best”’ or optimal behavioral 
strategy for an individual to exhibit. This 
behavior is often dependent on what other 
members of the population are doing and 
therefore is subject to frequency-dependent 
selection (Dawkins, 1980). ESS theory was 
used to explain hawk-dove strategies in an- 
imal contests (Clutton-Brock et al., 1979), 
parental investment (Maynard Smith, 1977), 
balanced sex ratios (Maynard Smith, 1981), 
cooperative mating (Packer and Pusey, 
1982), sex-biased natal dispersal (Krebs and 
Davies, 1987), and optimal foraging behav- 
ior (Belovsky, 1984). The theory was very 
helpful in demonstrating why altruism and 
group-benefit traits are not evolutionarily 
stable (Dawkins, 1976, 1980) unless they 
benefit the inclusive fitness of kin (Hamil- 
ton, 1964). ESS or optimality theory also 
contended that individuals would some- 
times be prevented from behaving opti- 
mally due to risk of predation or interfer- 
ence from better competitors and therefore 
would have to “make the best of a bad job” 
(Dawkins, 1980; Maynard Smith, 1982). 
These “‘conditional’’ ESSs (Dawkins, 1980) 
were used to explain “‘sneaky”’ mating tac- 
tics in subordinate red deer (Clutton-Brock 
et al., 1982) and reproductively-suppressed 
helpers in communal or cooperatively 
breeding mongooses (Helogale parvula), 
black-backed jackals (Canis mesomelas), 
and hunting dogs (Lycaon pictus), reviewed 
in Gittleman (1989). Since its inception in 


1974, ESS theory has been a central theme 
in developing arguments for the adaptive 
significance of behavioral patterns. 
Optimization. —Optimization models 
began achieving prominence in animal be- 
havior in the 1970s when they were applied 
to “decision-making” rules associated with 
foraging efficiency, risk sensitivity, and life 
histories (R. M. Alexander, 1982; Maynard 
Smith, 1974). Optimality theory was first 
applied to foraging behavior in birds (Mac- 
Arthur and Pianka, 1966) and later to mam- 
mals, such as forage selection in moose (Alces 
alces, Belovsky, 1978) and hoarding behav- 
ior in chipmunks (Elliot, 1978). In general, 
herbivores exhibit a trade-off between max- 
imizing energy intake and some external 
constraint such as obtaining an adequate 
mix of nutrients (Owen-Smith and Novel- 
lie, 1982) or avoiding plant secondary com- 
pounds (Freeland and Janzen, 1974). Be- 
lovsky (1978) demonstrated that moose 
tended to optimize energy intake subject to 
a sodium constraint. Habitat choice for 
white-tailed deer during winter is a trade- 
off between maximizing energy intake with- 
in a thermal constraint (Schmitz, 1991). 
Caraco and Wolf (1975) calculated that the 
mean size of African lion prides was not 
optimal for foraging efficiency, but was 
probably evolutionarily stable with respect 
to defense of carcasses, feeding territories, 
or offspring (Packer et al., 1990). Optimality 
theory has also been applied to nursing be- 
havior and reproductive success in female 
house mice (Fuchs, 1982), territorial de- 
fense (Schoener, 1987), managing range- 
lands of the western United States (Painter 
and Belsky, 1993), foraging behavior of pri- 
mates (Robinson, 1986), and harvesting 
management for whales (Horwood, 1990) 
and white-tailed deer (Leberg et al., 1987). 
Optimality theory, evolutionarily stable 
strategies, and game theory have been used 
extensively by bird and insect behavioral 
ecologists, more so by British and European 
biologists than by North American mam- 
malogists. These three behavioral concepts 
have contributed considerably to behavior- 


410 EISENBERG AND WOLFF 


al theory and should be used more by mam- 
mal behaviorists. Beware, however, that al- 
though these concepts provide a powerful 
set of tools, truly long-term studies may raise 
many more questions (Clutton-Brock, 
1988). 

Sex-biased natal dispersal. — Historically, 
dispersal was examined from ecological or 
population-level perspectives (e.g., Lidick- 
er, 1975; Stenseth and Lidicker, 1992; see 
also Chepko-Sade and Halpin, 1987). Be- 
haviorists, on the other hand, were inter- 
ested in the proximate mechanisms and ul- 
timate consequences of dispersal to the 
individual. During the 1960s and 1970s, the 
general consensus regarding dispersal of ju- 
veniles away from their natal home range 
or social group was that adults forced the 
dispersal of their offspring to reduce re- 
source or reproductive competition or both 
in the natal home range (reviewed in An- 
derson, 1989; Shields, 1982). During the late 
1970s and continuing to the present, an em- 
phasis has been placed on natal dispersal as 
being an adaptive mechanism for juveniles 
to separate from opposite-sex relatives to 
prevent inbreeding (Crockett, 1984; Harvey 
and Ralls, 1986; Pusey, 1987; Wolff, 1993). 
Packer (1979) was the first to propose that 
juvenile male baboons dispersed “‘volun- 
tarily” from their natal social group to avoid 
inbreeding with female relatives. This idea 
was criticized by Moore and Ali (1984), but 
later substantiated by Packer (1985). Since 
then, theoretical (Clutton-Brock, 1989a, 
1989b) and empirical (Wolff, 1993) argu- 
ments have been made to demonstrate that 
juvenile dispersal is correlated with the 
presence of opposite-sex parents 1n the natal 
home range and does not result from pa- 
rental aggression. Several experimental 
studies conducted in the mid-1980s and 
early 1990s confirmed that juvenile dis- 
persal functions to avoid inbreeding (e.g., 
marmots, Marmota flaviventris, Brody and 
Armitage, 1985; white-tailed deer, Odocoi- 
leus virginianus, Holzenbein and Marchin- 
ton, 1992; and white-footed mice, Pero- 


myscus leucopus, Wolff, 1992). The current 
trend is to consider juvenile dispersal as an 
adaptive, evolved behavior that benefits the 
inclusive fitness interests of both the dis- 
persing juvenile and the relatives it left be- 
hind—in short, a possible long-term com- 
promise. 

Mating systems and certainty of paterni- 
ty.—An important component of mam- 
malian behavior has been male and female 
mating strategies, especially as they pertain 
to mate guarding, pair bonding, and pater- 
nal care. Early classifications of mammalian 
mating systems included the basic monog- 
amy, polygyny, polyandry, and promiscui- 
ty. This classification system proved to be 
too simplistic and was later divided to in- 
clude, for instance: serial and permanent 
monogamy, harem-defense and territorial 
polygyny, and broadcast and arena prom- 
iscuity (Wittenberger, 1981). In the late 
1980s and 1990s, mating systems were fur- 
ther classified based on male and female 
mating bonds and defense systems that were 
ultimately based on ecological and social 
conditions (Clutton-Brock, 1989b; Eisen- 
berg, 1981). Although as many as 20 dif- 
ferent male and female bonding and defense 
systems have been described, mating sys- 
tems of over 95% of all mammal species 
reportedly are polygynous or promiscuous, 
with less than 5% being monogamous (Klei- 
man, 1977). Paternal care is extensive in 
monogamous species or even in some uni- 
male polygynous systems in which males 
are confident of paternity. As altruism is 
rarely described for mating systems in 
mammals, any type of paternal care must 
be associated with confidence of paternity. 

The use of molecular techniques such as 
electrophoresis and DNA fingerprinting that 
employ polymorphic blood proteins as ge- 
netic markers have revolutionized our 
thinking about male and female reproduc- 
tive strategies (Amos and Pemberton, 1992). 
Foltz (1981) was the first to use electropho- 
retic techniques to demonstrate that the old- 
field mouse, Peromyscus polionotus, was 


BEHAVIOR 411 


truly monogamous with males and females 
forming long-term pair bonds and all the 
young of a given female were sired by her 
mate. Conversely, Birdsall and Nash (1973) 
had earlier demonstrated that Peromyscus 
maniculatus was promiscuous. Ribble 
(1992) used DNA fingerprinting to corrob- 
orate the monogamous mating system of 
Peromyscus californicus. Similarly, in uni- 
male polygynous black-tailed prairie dogs, 
Cynomys ludovicianus, and yellow-bellied 
marmots, Marmota flaviventris, paternity 
analyses confirmed that all offspring within 
a territory were sired by the resident male 
(Foltz and Hoogland, 1981; Schwartz and 
Armitage, 1980). Pope (1991) demonstrat- 
ed that the dominant male in multi-male 
troops of A/ouatta seniculus sired most off- 
spring. 

In species where males do not defend fe- 
males and competition for estrous females 
is intense, several males may mate with the 
same female, possibly resulting in sperm 
competition and multiple paternity (see EI- 
liot, 1978, for a review). Female Belding’s 
ground squirrels, Spermophilus beldingi, and 
thirteen-lined ground squirrels, Spermoph- 
ilus tridecemlineatus, mate promiscuously 
with three to five different males and litters 
are often sired by up to three different males. 
In both species, first males sire 60-75% of 
the offspring (Foltz and Schwagmeyer, 1988; 
Hanken and Sherman, 1981), and conse- 
quently, males do not guard females, but 
leave to search for more mates as soon as 
copulation is over. In some species, how- 
ever, first males do not have a reproductive 
advantage (Dewsbury, 1984), and in those 
species males are more apt to guard females 
after copulation (Sherman, 1989). An in- 
teresting correlation of species in which fe- 
males are promiscuous and sperm compe- 
tition occurs is that males have larger testes 
than in species in which females mate with 
only one male (Harcourt et al., 1981; Heske 
and Ostfeld, 1990). 

Electrophoretic paternity analyses have 
also revealed that many species that were 


once thought to be polygynous were in fact 
promiscuous, with a relatively large portion 
of the offspring sired by nonresident males 
(Peromyscus leucopus, Xia and Millar, 1991; 
Microtus pennsylvanicus, Boonstra et al., 
1993). DNA fingerprinting has been used to 
relate reproductive success to harem mem- 
bership in red deer, Cervus elephas (Pem- 
berton et al., 1992); parentage, kinship, and 
cooperation in African lions, Panthera leo 
(Gilbert et al., 1991; Packer et al., 1991); 
demonstrate that high-ranking males sire 
most of the offspring in a troop of long- 
tailed macaques, Macaca fascicularis 
(DeRuiter et al., 1992): confirm that wolf 
(Canis lupus) packs consist of an unrelated 
pair and their related offspring (Lehman et 
al., 1992); and describe the unique mating 
systems in pilot whales (Globicephala me- 
las), in which pods consist of adult females 
and related males, but all mating occurs with 
nonpod members (Amos et al., 1991). 
Infanticide as a reproductive strategy. — 
John Calhoun was one of the first behav- 
lorists to document the killing of pups by 
adults while studying crowding behavior in 
Norway rats (Rattus norvegicus) in a semi- 
natural environment (Calhoun, 1963a). This 
early account considered infanticide an ab- 
errant pathological behavior associated with 
crowded or unnatural conditions. Infanti- 
cide was first observed in the wild in the 
early 1970s. Rudran, working with primates 
in both Sri Lanka and Venezuela, pioneered 
in the observations of infanticide and sug- 
gested a density dependent model as an ex- 
planation (Rudran, 1973). Hrdy (1977), 
while studying a naturally occurring popu- 
lation of langurs (Presbytis entellus) in Abu, 
expounded on the phenomenon. The initial 
reports of infanticide in wild populations of 
primates precipitated a series of often hasty 
observational and empirical studies on in- 
fanticide in a variety of mammal species 
(Hausfater and Hrdy, 1984). In 1979, Hrdy 
presented five hypotheses to explain the 
functional significance of infanticidal be- 
havior that had been observed in a variety 


412 EISENBERG AND WOLFF 


of species in a variety of situations: 1) off- 
spring were killed to be eaten for food; 2) 
sexual selection—where males would kill 
pups to remove genetic competitors and ter- 
minate lactation to stimulate the onset of 
estrus in the victim female; 3) competition 
for resources— where females would kill off- 
spring of other females as a mechanism of 
competing for burrows or nest sites; 4) pa- 
rental manipulation of offspring numbers or 
sex ratio; and 5) social pathology. Eisenberg 
(1981) opposed a simplistic explanation and 
advocated that several mechanisms could 
possibly be operative under natural selec- 
tion. 

Observational and experimental field and 
laboratory studies tested these hypotheses 
and provided support primarily, but not ex- 
clusively, for the sexual selection and re- 
source competition hypotheses. Killing of 
offspring by strange males to terminate lac- 
tation and stimulate the onset of estrus was 
reported in several taxa of mammals in- 
cluding lions (Panthera leo, Packer and Pu- 
sey, 1983), horses (Equus caballus, Berger, 
1983), several primate species (reviewed in 
Pusey and Packer, 1987), sciurids (e.g., Mc- 
Lean, 1983), and murids (Labov et al., 1985; 
Wolff and Cicirello, 1989). Resource-com- 
petition infanticide committed by females 
was reported in Belding’s ground squirrels 
(Spermophilus beldingi, Sherman, 1981), 
prairie dogs (Cynomys ludovicianus, Hoog- 
land, 1985), Peromyscus sp. (Wolff and Ci- 
cirello, 1991), and wild rabbits (Oryctolagus 
cuniculus, Kunkele, 1992). The current per- 
spective on infanticide in mammals 1s that 
killing of offspring by nonrelated adults 1s 
an adaptive and evolutionarily stable re- 
productive strategy. Killing of offspring as 
a social pathology, as originally proposed 
by Calhoun, seems to be not often recorded 
and certainly is not evolutionarily stable; 
however, long-term studies of long-lived 
mammals are sparse. Younger students must 
maintain an open mind (Clutton-Brock, 
1988). 

Socioecology—the contemporary synthe- 
sis.—In the last 10 years, animal behavior 


has become behavioral ecology or socio- 
ecology. Behavior now includes an animal’s 
entire behavioral repertoire, which is shaped 
largely by the distributions and abundance 
of resources, risk of predation, and com- 
petition from conspecifics. Following E. O. 
Wilson’s introduction to this synthesis in 
1975, several major contributions have been 
made to this field in the form of texts, and 
long-term case studies, and two new jour- 
nals have been produced: Behavioral Ecol- 
ogy, and Behavioral Ecology and Socio- 
biology. 

A major influence in this field in the 1980s 
has been the texts and edited volumes of 
John Krebs and Nicholas Davies (1984, with 
subsequent revisions). The editors and con- 
tributors to these books have used an evo- 
lutionary approach to synthesize published 
works derived from a variety of taxa into 
major concepts in animal behavioral ecol- 
ogy. In the most recent volume (Krebs and 
Davies, 1993), mammals have been used to 
address dispersal theory, sexual selection, 
parental investment, optimal foraging, ter- 
ritoriality, and many other evolutionary 
principles. All of these concepts should pro- 
mote future research. Other excellent and 
synthetic books include: Eisenberg and 
Kleiman’s Advances in the Study of Mam- 
malian Behavior (1983), Ecological Aspects 
of Social Evolution (Rubenstein and Wran- 
gham, 1987), Social Evolution (Trivers, 
1983), Sociobiology and Behavior (Barash, 
1982), and The Ecology of Social Behavior 
(Slobodchikoff, 1988). Comprehensive case 
studies that have made major contributions 
to the field of mammal behavioral ecology 
include Red Deer (Cervus elaphus): the Be- 
haviour and Ecology of Two Sexes (Clutton- 
Brock et al. 1982), Wild Horse (Equus cal- 
labus) of the Great Basin (Berger, 1986), as 
well as several summary texts such as Pri- 
mates in Nature (Richard, 1985), and Pri- 
mate Societies (Smuts et al., 1987), Primate 
Social Systems (Dunbar, 1988), Carnivore 
Behavior, Ecology, and Evolution (Gittle- 
man, 1989), Behavioral Ecology of Ground 
Squirrels (Michener and Murie, 1989), 


BEHAVIOR 413 


Marmots: Social Behavior and Ecology 
(Barash, 1989), and Social Systems and 
Population Cycles of Voles (Tamarin et al., 
1990). 

Applying behavioral theory to humans. — 
Comparisons of human behavior and so- 
cieties with those of nonhuman animals 
dates back to pre-Darwinian times and has 
always shadowed studies on mammals and 
behavior through time. We presented an 
historical aspect for the implications of ap- 
plying sociobiological theory to humans 
earlier in this chapter and illustrate here that 
human behavior is still a main concern of 
mammalogists as well as anthropologists and 
psychologists in the 1990s. Several promi- 
nent texts that have deservedly received at- 
tention in the last 20 years include Ortner 
(1983), Eibl-Eibesfeldt (1989), and Dissay- 
anake (1992), as well as the volumes edited 
by Napoleon Chagnon and William Irons, 
Evolutionary Biology and Human Social 
Behavior (1979), and George Barlow and 
James Silverberg’s Sociobiology: Beyond 
Nature/Nurture (1980). Martin Daly and 
Margo Wilson (1983) keep providing up- 
dated editions of their well-used undergrad- 
uate text Sex, Evolution, and Behavior. 
Donald Symons’ publication of The Evo- 
lution of Human Sexuality in 1979 sparked 
considerable controversy, primarily from the 
feminist movement, which countered with 
Hrdy’s The Woman that Never Evolved in 
1981. The journal Ethology and Sociobiol- 
ogy was Started in 1979 and is strongly ori- 
ented toward humans. A new interdisci- 
plinary society was organized in the late 
1980s, the Human Behavior and Evolution 
Society, which held its fifth annual meeting 
in 1993. The popular writing style of Rich- 
ard Dawkins’ The Selfish Gene and David 
Barash’s The Whisperings Within helped 
these books reach much of the general pub- 
lic—a laudable but perhaps futile effort. Al- 
though an integration of evolutionary the- 
ory into the social sciences has been a task 
with much resistance, mammal behavior- 
ists continue to promote the universal theme 
of evolution by natural selection applicable 


to the social systems of a// mammals. Yet, 
some caution is necessary, and a reading of 
Pepper (1958) could be of help. 


Some Advances in Sister 
Disciplines 


Form and function—paleontology.—The 
evolution of mammals and the behavior of 
early mammals has been an area of active 
research. Outstanding contributions have 
derived from the efforts of Crompton and 
his associates and students. Lillegraven and 
the Wyoming group have broadened our 
horizons with new perspectives on the eu- 
therian-marsupial dichotomy. Guthrie, in 
Alaska, has brought paleontology, behav- 
ior, and ecology to a grand synthetic treat- 
ment (1990; see Haynes, 1991). Paleocom- 
munities and their relevance for 
understanding community form and ex- 
tinction events in contemporary times has 
been pioneered also by Webb (1983), Val- 
kenburgh (1990), and Behrensmeyer et al. 
(1992). A little-appreciated area of research 
is the use by humans of animal resources as 
revealed by archaeologists (Sigler-Eisen- 
berg, 1988). 

Behavior and conservation.—The New 
York Zoological Society and the Smithsoni- 
an Institution deserve special note in this 
arena of research. In both institutions an 
emphasis on field studies with an aim to 
apply results to conservation issues has been 
overwhelming. From NYZS have come 
seminal studies on the behavior of the 
humpbacked whale (Megaptera), African 
lion (Panthera leo), African forest primates 
(Cercopithecidae), the giant panda (4//urop- 
oda), and the ungulates of the Tibetan pla- 
teau (Payne, 1983; Schaller, 1972, 1993; 
Struhsaker, 1975). From the Smithsonian- 
NZP came such studies as the behavior and 
ecology of the golden lion tamarin (Leon- 
topithecus), the ecology of the tiger (Panthe- 
ra tigris) in Nepal, the behavior and ecology 
of the Asiatic elephant (E/ephas maximus), 


414 EISENBERG AND WOLFF 


and the behavior and conservation of Pere 
David’s deer (Elaphurus davidianus), as well 
as ecosystem issues (Beck and Wemmer, 
1983: Kleiman et al., 1986; Seidensticker 
and Lumpkin, 1991; Sunquist, 1981). 

The US Fish and Wildlife Service and the 
US National Park Service, often in con- 
junction with the NZP and the US National 
Museum of Natural History, have taken a 
new turn in research emphasis by encour- 
aging important work on cetaceans, pinni- 
peds, manatees, and sea lions. Leadership 
in conservation biology is also noted (Scho- 
newald-Cox et al., 1983). This effort is often 
under-appreciated by those outside the ser- 
vice. Meanwhile, there have been wide- 
spread efforts to cope with human-animal 
conflicts as the march of human population 
growth proceeds (Redford and Padoch, 
1992; Robinson and Redford, 1991; Smythe, 
1991). 


Quo Vadis? 


Some readers may consider this docu- 
ment a rather personal account, and in many 
ways this is true, because John F. Eisenberg 
was deeply involved in the processes that 
led to the acceptance of mammalian behav- 
ior as a legitimate discipline of study in 
North America. Despite the fractionation 
following the synthesis, we firmly believe 
that, when necessary, the disparate students 
of behavior will come together for a new 
round of synthetic activity. In short, what 
comes around, will come around again. 

Let us point out that we trust some of us 
have not lost our primary mission as biol- 
ogists, which is to say that if given a tract 
of land and custodianship, then inventory, 
monitoring, and long-term population stud- 
ies are the rule. This is especially true as 
some of us begin to expand and train stu- 
dents in third world countries for roles in 
conservation. In such a situation, the infra- 
structure often does not exist without going 
back to basics, and the study of animal be- 
havior touches everything. If we choose to 


be a bit foolish, then please indulge us. We 
think of the discipline of behavior as em- 
bodied by “Tinker Bell,’ who will appear 
again and again, when necessary, to keep 
our view of nature forever young. 


Literature Cited 


ALEXANDER, R. D. 1974. The evolution of social be- 
havior. Annual Review of Ecology and Systematics, 
5:325-383. 

ALEXANDER, R. D., AND D. W. TINKLE. 1981. Natural 
selection and social behavior: recent research and 
new theory. Chiron Press, New York, 532 pp. 

ALEXANDER, R. M. 1982. Optima for animals. Ed- 
ward Arnold, London, 112 pp. 

Amos, B., J. BARRETT, AND G. A. Dover. 1991. 
Breeding behaviour of pilot whales revealed by DNA 
fingerprinting. Heredity, 67:49-55. 

Amos, B., AND J. PEMBERTON. 1992. DNA finger- 
printing in non-human populations. Current Opin- 
ion in Genetics and Development, 2:857—860. 

ANDERSON, P. K. 1989. Dispersal in rodents: a res- 
ident fitness hypothesis. Special Publications, The 
American Society of Mammalogists, 9:1-141. 

AuDUBON, J. J., AND J. BACHMAN. 1846-1854. The 
viviparous quadrupeds of North America, 3 vols. V. 
G. Audubon, New York. Reprint 1989, Wellfleet 
Press, Seacaucus, New Jersey, 440 pp. 

AusTAD, S. N., AND M. E. SuNquisT. 1988. Sex-ratio 
manipulation in the common opossum. Nature, 324: 
58-60. 

BaiLtey, V. 1931. Mammals of New Mexico. North 
American Fauna, 53:1-412. 

BaRASH, D. 1979. The whisperings within. Harper 
and Row, New York, 274 pp. 

. 1982. Sociobiology and behavior. Elsevier, 

Amsterdam, 426 pp. 

. 1989. Marmots: social behavior and ecology. 
Stanford University Press, Stanford, 360 pp. 

BaRLow, G. W., AND J. SILVERBERG. 1980. Socio- 
biology: beyond nature/nurture? AAAS Selected 
Symposia Series, Westview Press, Boulder, 627 pp. 

BARTHOLOMEW, G. A., AND T. J. CADE. 1957. Tem- 
perature regulation, hibernation and aestivation in 
the little pocket mouse. Journal of Mammalogy, 38: 
60-72. 

Beacu, F. A. 1948. Hormones and behavior. Harper 
Brothers, New York, 368 pp. 

Beck, B., AND C. WEMMER. 1983. Pere David’s deer: 
conservation of an extinct species. Garland Press, 
New York, 193 pp. 

BEEBE, W. 1925. The three-toed sloth. Zoologica, 7:1- 
67. 

BEHRENSMEYER, A., J. D. DAMUTH, W. A. DEMICHELE, 
R. Potts, H. D. Sues, AND S. L. Winc. 1992. Ter- 
restrial ecosystems through time. The University of 
Chicago Press, Chicago, 568 pp. 

BELovsky, G. E. 1978. Diet optimization in a gen- 
eralist herbivore; the moose. Theoretical Population 
Biology, 14:105-134. 


BEHAVIOR 415 


1984. Herbivore optimal foraging: a com- 
parative test of three models. The American Natu- 
ralist, 124:97-115. 

BERGER, J. 1979. Weaning conflict in desert and 
mountain bighorn sheep (Ovis canadensis): an eco- 
logical interpretation. Zeitschrist fiir Tierpsycholo- 
gie, 50:188-—200. 

. 1983. Induced abortion and social factors in 

wild horses. Nature, 303:59-61. 

1986. Wild horses of the Great Basin: social 
competition and group size. The University of Chi- 
cago Press, Chicago, 325 pp. 

BinGHAM, H. C. 1932. Gorillas in native habitat. 
Publications of Carnegie Institute (Carnegie Institute 
of Washington, Washington, D. C.), 426:1-66. 

BIRDSALL, D. A., AND D. NAsH. 1973. Occurrence of 
successful multiple insemination of females in nat- 
ural populations of deer mice (Peromyscus mani- 
culatus). Evolution, 27:106-110. 

BoonstTRA, R., X. XIA, AND L. PAVONE. 1993. Mating 
system of the meadow vole, Microtus pennsylvani- 
cus. Behavioral Ecology, 4:83-89. 

Bropy, A. K., AND K. B. ARMITAGE. 1985. The effects 
of adult removal on dispersal of yearling yellow- 
bellied marmots. Canadian Journal of Zoology, 63: 
2560-2564. 

Bronson, F. H. 1989. Mammalian reproductive bi- 
ology. The University of Chicago Press, Chicago, 
325 pp. 

Burt, W. H. 1940. Territorial behavior and popu- 
lations of some small mammals in southern Mich- 
igan. Miscellaneous Publication of the Museum of 
Zoology, The University of Michigan, 45:1-58. 

CaLey, M. J., AND T. D. Nupps. 1987. Sex-ratio 
adjustment in Odocoileus virginianus: does local re- 
source competition play a role? The American Nat- 
uralist, 129:452-457. 

CALHOUN, J. B. 1963a. The ecology and sociology of 
the Norway rat. U.S. Department of Health, Edu- 
cation, and Welfare, Public Health Service, Bethes- 
da, Maryland, 288 pp. 

1963b. The social use of space. Pp. 2-187, 
in Physiological mammalogy (Van Gelder and Maier, 
eds.). Academic Press, New York, 1:1—380. 

CaRACO, T., AND L. L. WotF. 1975. Ecological de- 
terminants of group sizes of foraging lions. The 
American Naturalist, 109:343-352. 

CARPENTER, C.R. 1934. A field study of the behavior 
and social relations of howler monkeys (Alouatta 
palliata). Comparative Psychology Monographs, 10: 
1-168. 


1935. Behavior of red spider monkeys in 

Panama. Journal of Mammalogy, 16:171-180. 

1964. Naturalistic behavior of non-human 
primates. Pennsylvania State University Press, Uni- 
versity Park, 454 pp. 

CHAGNON, N. A., AND W. IRons. 1979. Evolutionary 
biology and human social behavior: an anthropo- 
logical perspective. Duxbury Press, North Scituate, 
Massachusetts, 623 pp. 

CHEPKO-SADE, B. D., AND Z. T. HALPIN. 1987. Mam- 
malian dispersal patterns. The University of Chicago 
Press, Chicago, 342 pp. 

CHRISTIAN, J. J. 1963. Endocrine adaptive mecha- 


nisms and the physiological regulation of population 
growth. Physiological Mammalogy, 1:189-353. 

CLARK, A. B. 1978. Sex ratio and local resource com- 
petition in a prosimian primate. Science, 201:163- 
165. 

CLUTTON-BrRock, T. H. (EpD.). 1988. Reproductive 
success. The University of Chicago Press, Chicago, 
538 pp. 

1989a. Female transfer and inbreeding 

avoidance in social mammals. Nature, 337:70-72. 

1989b. Mammalian mating systems. Pro- 
ceedings of the Royal Society of London, 236:339- 
21 

CLuTTON-Brock, T. H., S. D. ALBoN, AND F. E. 
Guiness. 1984. Maternal dominance, breeding 
success, and birth sex ratios in red deer. Nature, 308: 
358-360. 

. 1986. Great expectations: dominance, breed- 
ing success and offspring sex ratios in red deer. An- 
imal Behaviour, 34:460-471. 

CLUTTON-BrRock, T. H., F. E. GurNess, AND S. D. 
ALBON. 1982. Red deer: the behaviour and ecology 
of two sexes. The University of Chicago Press, Chi- 
cago, 378 pp. 

CLuttTon-Brock, T. H., S. D. ALBon, R. M. Gipson, 
AND F. E. GuIness. 1979. The logical stag: adaptive 
aspects of fighting in red deer (Cervus elephas L.). 
Animal Behaviour, 27:211-225. 

CockBurNn, A. 1988. Social behavior in fluctuating 
populations. Croom Helm Ltd., North Ryde, New 
South Wales, 239 pp. 

Connor, R. C., AND K. S. Norris. 1982. Are dol- 
phins reciprocal altruists? The American Naturalist, 
119:358-374. 

Count, E. 1973. Being and becoming human. Van 
Nostrand, New York, 349 pp. 

Craic, W. 1918. Appetites and aversions as constit- 
uents of instincts. Biological Bulletin of Woods Hole, 
34:91-107. 

CrockeETT, C. 1984. Emigration by female red howler 
monkeys. Pp. 159-173, in Female primates (M. F. 
Small, ed.). Liss, New York, 272 pp. 

CrowcrortT, P. 1991. Elton’s ecologists. The Uni- 
versity of Chicago Press, Chicago, 177 pp. 

Day, M., AND M. Wirson. 1983. Sex, evolution, 
and behavior. Wadsworth Publishing Company, 
Belmont, California, 402 pp. 

Darwin, C. 1859. The origin of species. John Mur- 
ray, London (and many reprints), 816 pp. 

. 1872. The expression of the emotions in man 
and animals. Appleton, New York, London, 372 pp. 

Dawkins, R. D. 1976. The selfish gene. Oxford Uni- 
versity Press, Oxford, 224 pp. 

1980. Good strategy or evolutionarily stable 
strategy. Pp. 331-367, in Sociobiology: beyond na- 
ture/nurture? (G. W. Barlow and J. Silverberg, eds.). 
AAAS Selected Symposia Series. Westview Press, 
Boulder, Colorado, 627 pp. 

DEEGENER, P. 1918. Die Formen der Vergesellschaf- 
tung im Tierriche. Verlag von Vict, Leipzig, 420 pp. 

DeRutrter, S. W., Jr., G. J. J. M. TROMMELEN, A. G. 
S. W. UTTERLINDEN, R. D. MARTIN, AND J. VAN 
Hoorr. 1992. Male social rank and reproductive 
success in wild long-tailed macaques. Paternity ex- 


416 EISENBERG AND WOLFF 


clusions by blood analysis and DNA fingerprinting. 
Pp. xx—xx, in Paternity in primates: genetic tests and 
theories, implications of human DNA fingerprinting 
(R. D. Martin, A. F. Dixson, and E. J. Wickings, 
eds.). S. Karger, Basel. 

DeVore, B. I. (Ep.). 1965. Primate behavior: field 
studies of monkeys and apes. Holt, Rinehart and 
Winston, New York, 654 pp. 

Dewssury, D. A. 1984. Sperm competition in mu- 
roid rodents. Pp. 547-571, in Sperm competition 
and the evolution of animal mating systems (R. L. 
Smith, ed.). Academic Press, New York, 687 pp. 

. 1989a. A brief history of animal behavior in 

North America. Perspectives in Ethology, 8:85-122. 

. 1989b. Comparative psychology, ethology and 

animal behavior. Annual Review of Psychology, 40: 

581-602. 

. 1992. Comparative psychology and ethology: 
a reassessment. American Psychology, February:208- 
Dales: 

Dice, L.R. 1933. Fertility relationships between some 
of the species and subspecies of mice in the genus 
Peromyscus. Journal of Mammalogy, 14:298-305. 

DIsSANAYAKE, E. 1992. Homo aestheticus. Free Press, 
New York, 297 pp. 

Dittus, W. 1977. The social regulation of population 
density and age-specific distribution in the toque 
monkey. Behaviour, 63:28 1-322. 

Dunsar, R.I.M. 1988. Primate social systems. Cor- 
nell University Press, Ithaca, 378 pp. 

ErBt-EIBesFELDT, I. 1958. Das Verhalten der Naye- 
tiere. Handbuch der Zoologie, Band 8, Lieferung, 12, 
10:1-88. 

. 1989. Human ethology. Aldine de Gruyter, 
New York, 848 pp. 

EISENBERG, J. F. 1962. Studies on the behavior of 
Peromyscus maniculatus gambelii and Peromyscus 
californicus parasiticus. Behaviour, 19:177-—207. 

1963. The behavior of heteromyid rodents. 

University of California Publication in Zoology, 69: 

1-100. 


1966. The social organizations of mammals. 

Handbuch der Zoologie, Band 8, Lieferung, 39, 10: 

1-92. 

. 1981. The mammalian radiations. The Uni- 
versity of Chicago Press, Chicago, 610 pp. 

EISENBERG, J. F., AND W.S. DILLON (EDs.). 1971. Man 
and beast. Smithsonian Press, Washington, D.C., 401 


EISENBERG, J. F., AND D. G. KLEIMAN (EDs.). 1983. 
Advances in the study of mammalian behavior. Spe- 
cial Publication, The American Society of Mam- 
malogists, 7:1-753. 

EISENBERG, J. F., AND R. E. KUEHN. 1966. Behavior 
of Ateles geoffroyi and related species. Smithsonian 
Miscellaneous Collection, 151:63 pp. 

Exuiot, L. 1978. Social behavior and foraging ecology 
of the chipmunk (7Jamias striatis). Smithsonian 
Contributions to Zoology, 265:1-107. 

ENperS, R. K. 1935. Mammalian life histories from 
Barro Colorado Island, Panama. Bulletin of the Mu- 
seum of Comparative Zoology, Harvard, 78:385- 
302: 


. 1952. Reproduction in the mink. Proceedings 
of the American Philosophical Society, 96:69 1-755. 

Ewer, R. 1969. The ethology of mammals. Logos, 
London, 418 pp. 

Fottz, D. W. 1981. Genetic evidence for long-term 
monogamy in a small rodent Peromyscus polionotus. 
The American Naturalist, 117:665-675. 

Foitz, D. W., AND J. L. HOOGLAND. 1981. Analysis 
of the mating system in the black-tailed prairie dog 
(Cynomys ludovicianus) by likelihood of paternity. 
Journal of Mammalogy, 62:706-712. 

Foitz, D. W., AND P. L.SCHWAGMEYER. 1988. Sperm 
competition in the thirteen-lined ground squirrel: 
differential fertilization success under field condi- 
tions. The American Naturalist, 133:257-265. 

FREELAND, W. J., AND D. H. JANZEN. 1974. Strategies 
in herbivory by mammals: the role of plant second- 
ary compounds. The American Naturalist, 108:269- 
284. 

FriscH, K. von. 1950. Bees, their vision, chemical 
senses and language. Cornell University Press, Ith- 
aca, 157 pp. 

Fucus, S. 1982. Optimality of parental investment: 
the influence of nursing on reproductive success of 
mother and female young house mice. Behavioral 
Ecology and Sociobiology, 10:39-51. 

Geist, V. 1971. Mountain sheep. The University of 
Chicago Press, Chicago, 383 pp. 

. 1974. On the relationship of social evolution 
and ecology in ungulates. American Zoologist 14: 
205-220. 

GILBERT, D. A., C. PACKER, A. E. Pusey, J. C. STE- 
PHENS, AND S. J. O’BriEN. 1991. Analytical DNA 
fingerprinting in lions: parentage, genetic diversity, 
and kinship. Journal of Heredity, 82:378-386. 

GITTLEMAN, J. L. (ED.). 1989. Carnivore behavior, 
ecology, and evolution. Cornell University Press, 
Ithaca, 620 pp. 

GomeENDI0, M. 1991. Parent/offspring conflict during 
the weaning process in rhesus macaques. Animal 
Behaviour, 42:993-1005. 

GriFFIN, D. 1958. Listening in the dark. Yale Uni- 
versity Press, New Haven, 356 pp. 

GRINNELL, J. 1914. An account of the mammals and 
birds of the lower Colorado Valley with special ref- 
erence to the distributional problems presented. 
University of California Publications in Zoology, 12: 
51-294. 

Guturig, R. D. 1990. Frozen fauna of the mammoth 
steppe. The University of Chicago Press, Chicago, 
323 pp. 

HAMILTON, W.D. 1964. The genetical theory of social 
behavior. Journal of Theoretical Biology, 7:1-52. 
HANKEN, J., AND P. W. SHERMAN. 1981. Multiple 
paternity in Belding’s ground squirrel litters. Science, 

212:351-353. 

Harcourt, A. H., P. H. Harvey, S. G. LARSON, AND 
R. V. SHort. 1981. Testis weight and breeding 
system in primates. Nature, 293:55-57. 

Harris, V.T. 1952. Anexperimental study of habitat 
selection by prairie and forest races of the deer- 
mouse, Peromyscus maniculatus. Contributions from 
the Laboratory of Vertebrate Biology, University of 
Michigan, 56:1-53. 


BEHAVIOR 417 


HARTHOORN, A. M. 1976. The chemical capture of 
animals. Baillare, Tyndale, London, 416 pp. 

Harvey, P. H., AND K. RALLS. 1986. Do animals 
avoid incest? Nature, 320:575-576. 

HAUuUSFATER, G., AND S. B. Hrpy (eEps.). 1984. Infan- 
ticide: comparative and evolutionary perspectives. 
Aldine, New York, 598 pp. 

Haynes, G. 1991. Mammoths, mastodonts, and el- 
ephants. Cambridge University Press, 413 pp. 

Hepicer, H. 1942. Wildetiere in Gafangenschaft. 
Benno Schwabe, Basel, 205 pp. 

Heske, E. J., AND R. S. OstFELD. 1990. Sexual di- 
morphism in size, relative size of testes, and mating 
systems in North American voles. Journal of Mam- 
malogy, 71:510-519. 

Ho .uister, N. 1917. Some effects of environment 
and habitat on captive lions. Proceedings of the 
United States National Museum, 53:177-193. 

HOLZENBEIN, S., AND R. L. MARCHINTON. 1992. Spa- 
tial integration of maturing-male white-tailed deer 
in the adult population. Journal of Mammalogy, 73: 
326-334. 

HOOGLAND, J. L. 1985. Infanticide in prairie dogs: 
lactating females kill offspring of close kin. Science, 
230:1037-1040. 

Horwoop, J. W. 1990. Whale management: an op- 
timality approach. Report of the International Whal- 
ing Commission, 40:493-497. 

Howarp, W.E. 1948. Dispersal, amount of inbreed- 
ing and longevity in a local population of deer mice. 
Contributions from the Laboratory of Vertebrate Bi- 
ology, University of Michigan, 43:1—50. 

Howe Lt, A. B. 1944. Speed in animals. The Uni- 
versity of Chicago Press, Chicago, 270 pp. 

Hrpy, S. B. 1977. The langurs of Abu: female and 
male strategies of reproduction. Harvard University 
Press, Cambridge, 361 pp. 

1979. Infanticide among animals: a review, 

classification, and examination of the implications 

for reproductive strategies of females. Ethology and 

Sociobiology, 1:13—40. 

1981. The woman that never evolved. Har- 
vard University Press, Cambridge, 256 pp. 

Jones, J. K., Jk. 1991. Geneology of 20th century 
mammalogists in North America. Pp. 33-47, in Lat- 
in American mammalogy (M. Mares and D. Schmid- 
ly, eds.). University of Oklahoma Press, Norman, 
468 pp. 

KAUFMANN, J. H. 1962. Ecology and social behavior 
of the coati, Nasua narica on Barro Colorado Island, 
Panama. University of California Publications in 
Zoology, 60:95-222. 

KELLOGG, W. N. 1961. Porpoises and sonar. The 
University of Chicago Press, Chicago, 177 pp. 

Kina, J. A. 1955. Social behavior, social organiza- 
tion, and population dynamics in a black-tailed 
prairiedog town in the Black Hills of South Dakota. 
Contributions from the Laboratory of Vertebrate Bi- 
ology, University of Michigan, 67:1-123. 

1961. Development and behavioral evolu- 

tion in Peromyscus. Pp. 122-147, in Vertebrate spe- 

ciation (W. F. Blair, ed.). University of Texas Press, 

Austin, Texas, 642 pp. 

. (ED.). 1968. Biology of Peromyscus (Roden- 


tia). Special Publications, The American Society of 
Mammalogists, 2:1-593. 

KLEIMAN, D. G. 1967. Some aspects of social behav- 
ior in the Canidae. American Zoologist, 7:365—372. 

. 1977. Monogamy in mammals. Quarterly 
Review of Biology, 52:39-69. 

KLEIMAN, D. G., B. B. BEck, J. M. Dietz, L. A. Dietz, 
J. D. BALLou, AND A. J. CoImBRA-FILHO. 1986. 
Conservation program for the golden lion tamarin: 
captive research and management, ecological stud- 
ies, educational strategies, and reintroduction. Pp. 
959-979, in Primates: the road to self-sustaining 
populations (K. Benirschke, ed.). Springer-Verlag, 
New York, 1,044 pp. 

KoestLer, A. 1971. The case of the midwife toad. 
Random House, New York, 187 pp. 

Kress, J. R., AND N. B. Davies. 1984. Behavioural 
ecology. Blackwell Scientific Publications, Oxford, 
493 pp. 

. 1987. An introduction to behavioural ecol- 

ogy. Blackwell Scientific Publications, Oxford, (Sec- 

ond ed.) 389 pp. 

(EDS). 1991. Behavioural ecology: an evolu- 

tionary approach. Blackwell Scientific Publications, 

Oxford, (Third ed.) 482 pp. 

. 1993. An introduction to behavioral ecology. 
Blackwell Scientific Publications, Oxford, (Third ed.) 
420 pp. 

KroeBerR, A. 1925. Yana and Yani. Handbook of the 
Indians of California. Bulletin of American Ethnol- 
ogy, 78. Smithsonian Institution, Washington, D.C. 

KRoporkKIN, P. 1902. Mutual aid: a factor in evolu- 
tion. A. Knopf, New York, 362 pp. (Republished in 
1914.) 

KUuNKELE, J. 1992. Infanticide in wild rabbits (Orvc- 
tolagus cuniculus). Journal of Mammalogy, 73:317- 
320. 

Lasov, J. B., U. W. Huck, R. W. ELwoop, AND R. J. 
Brooks. 1985. Current problems in the study of 
infanticidal behavior of rodents. The Quarterly Re- 
view of Biology, 60:1-20. 

Layne, J. N. 1954. The biology of the red squirrel, 
Tamiasciurus hudsonicus lognax, in central New 
York. Ecology Monographs, 24:227-267. 

LeBerG, P. L., M. H. SmMiTH, AND K. T. SCRIBNER. 
1987. Optimization strategies in a white-tailed deer 
harvest program. Transaction of Congress of the In- 
ternational Union of Game Biologists, 18:102-103. 

LEHMAN, N., P. CLARKSON, L. D. MEcu, T. J. MEIER, 
R. K. WAYNE, AND S. W. Zack. 1992. A study of 
the genetic relationships within and among wolf packs 
using DNA fingerprinting and mitochondrial DNA. 
Behavioral Ecology and Sociobiology, 30:83-94. 

LeopoLp, A. 1933. Game management. Scribners, 
New York, 481 pp. 

Lipicker, W. Z., Jk. 1975. The role of dispersal in 
the demography of small mammals. Pp. 203-233, 
in Small mammals: their productivity and popula- 
tion dynamics (F. B. Golley, F. Petrusewicz, and L. 
Ryskowski, eds.). International Biological Program, 
Cambridge University Press, Cambridge, 5:1-451. 

LinspDALE, J. M. 1946. The California ground squir- 
rel. University of California Press, Berkeley, 475 pp. 

LINSDALE, J. M., AND L. P. Tevis. 1951. The dusky- 


418 EISENBERG AND WOLFF 


footed wood rat. University of California Press, 
Berkeley, 664 pp. 

LINSDALE, J. M., AND P. Q. Tomicn. 1953. A herd of 
mule deer. University of California Press, Berkeley, 
567 pp. 

Lorenz, K. Z. 1981. The foundations of ethology. 
Springer-Verlag, New York, 380 pp. 

MacArtTuur, R. H., AND E. R. PIANKA. 1966. On 
the optimal use ofa patchy environment. The Amer- 
ican Naturalist, 100:603-609. 

Macu, E. 1959. Analysis of sensations. Dover, New 
York, 310 pp. (reprint.) 

Marter, P. 1985. Hark ye to the birds. Pp. 315-346, 
in Studying animal behavior (D. A. Dewsbury, ed.). 
The University of Chicago Press, Chicago, 512 pp. 

MAYNARD SMITH, J. 1974. The theory of games and 
the evolution of animal conflicts. Journal of Theo- 
retical Biology, 47:209-221. 

1977. Parental investment—a prospective 

analysis. Animal Behaviour, 25:1-9. 

1981. Will a sexual population evolve to an 

ESS? The American Naturalist, 117:1015-1018. 

1982. Evolution and theory of games. Cam- 
bridge University Press, Cambridge, 224 pp. 

McKay, G. 1973. Behavior and ecology of the Asi- 
atic elephant in southeastern Ceylon. Smithsonian 
Contributions to Zoology, 125:113 pp. 

McLean, I. G. 1983. Paternal behaviour and killing 
of young Arctic ground squirrels. Animal Behaviour, 
31:32-44. 

McNags, B. K. 1983. Would-be ecological and be- 
havioral consequences of adaptation to various food 
resources. Pp. 664-697, in Advances in the study of 
mammalian behavior (J. F. Eisenberg and D. G. 
Kleiman, eds.). Special Publications, The American 
Society of Mammalogists, 7:1-753. 

MEIKLE, D. B., L. C. DRICKAMER, S. H. VEssEy, T. L. 
ROSENTHAL, AND K. S. FITZGERALD. 1993. Mater- 
nal dominance rank and secondary sex ratio in do- 
mestic swine. Animal Behaviour, 46:79-85. 

MontTGoMEerY, G. G., AND M. E. Sunquist. 1975. 
Impact of sloths on Neotropical forest energy flow. 
Pp. 69-98, in Tropical ecological systems (F. B. Gol- 
ley and E. Medina, eds.). Springer-Verlag, New York, 
398 pp. 

Moore, J., AND R. Aur. 1984. Are dispersal and in- 
breeding avoidance related? Animal Behaviour, 32: 
94-112. 

Moore, J. C. 1957. Natural history of Shermann’s 
fox squirrel. Bulletin of the American Museum of 
Natural History, 113:71 pp. 

MorGan, L. H. 1851. League of the Ho-De-No-Sau- 
Nee. Rochester, New York. 

1868. The American beaver and his works. 
Burt Franklin, New York, 330 pp. (reprint 1970.) 
MorrIson, P., AND B. K. McNas. 1962. Daily torpor 
in a Brazilian murine oppossum (Marmosa). Com- 
parative Biochemical Physiology, A. Comparative 

Physiology, 6:57—-68. 

Murie, J.. AND G. MICHENER. 1984. The biology of 
ground-dwelling squirrels. University of Nebraska 
Press, Lincoln, 459 pp. 

Nissen, H. W. 1931. A field study of the chimpanzee. 
Comparative Psychology Monographs, 8:1-121. 


Noste, J. K. 1931. The biology of the Amphibia. 
McGraw-Hill Book Company, Ltd. Inc., New York, 
577 pp. 

ORTNER, D. J. (ED.). 1983. How humans adapt: a 
biocultural odyssey. Smithsonian Institution, Wash- 
ington, D.C., 560 pp. 

OweN-SmMITH, N., AND P. NoveLiiz. 1982. What 
should a clever ungulate eat? The American Natu- 
ralist, 119:151-178. 

PAcKER, C. 1977. Reciprocal altruism in olive ba- 
boons. Nature, 265:441-443. 

1979. Inter-group transfer and inbreeding 

avoidance in Papio anubis. Animal Behaviour, 27: 

1-36. 


. 1985. Dispersal and inbreeding avoidance. 
Animal Behaviour, 33:676-678. 

PACKER, C., AND A. E. Pusey. 1982. Cooperation and 
competition within a coalition of male lions: kin 
selection of game theory? Nature, 296:740-742. 

1983. Adaptations of female lions to infan- 
ticide by incoming males. The American Naturalist, 
121:716-728. 

PACKER, C., D. SCHEEL, AND A. E. Pusey. 1990. Why 
lions form groups: food is not enough. The American 
Naturalist, 136:1-19. 

Packer, C., D. A. GILBERT, A. E. PUSEY, AND S. J. 
O’BriEN. 1991. Kinship and cooperation in Afri- 
can lions: a molecular genetic analysis. Nature, 351: 
562-565. 

PAINTER, E. L., AND A. J. BEtsky. 1993. Application 
of herbivore optimization theory to rangelands of 
the western United States. Ecological Applications, 
3:2-9, 

PAYNE, R. (ED.). 1983. Communication and behavior 
of whales. Westview Press, Boulder, 643 pp. 

PEARSON, O. P. 1944. Reproduction in the shrew 
(Blarina brevicauda Say). The American Journal of 
Anatomy, 75:39-93. 

PEMBERTON, J. M., S. D. ALBon, F. E. Gurness, T. H. 
CLUTTON-BROCK, AND G. A. Dover. 1992. Be- 
havioral estimates of male mating success tested by 
DNA fingerprinting in a polygynous mammal. Be- 
havioural Ecology, 3:66-75. 

Pepper, S.C. 1958. The sources of value. University 
of California Press, Berkeley, 732 pp. 

Pope, T. 1991. The reproductive consequences of 
male cooperation in red howler monkeys. Behavioral 
Ecology and Sociobiology, 27:439-446. 

Pusey, A. E. 1987. Sex-biased dispersal and inbreed- 
ing avoidance in birds and mammals. Trends in 
Ecology and Evolution, 2:295-299. 

Pusey, A. E., AND C. PACKER. 1987. Dispersal and 
philopatry. Pp. 250-266, in Primate societies (B. B. 
Smuts, D. L. Cheney, R. M. Seyfarth, R. W. Wran- 
gham, and T. T. Struhsaker, eds.). The University 
of Chicago Press, Chicago, 578 pp. 

Rais, K. 1977. Sexual dimorsphism in mammals: 
avian models and unanswered questions. The Amer- 
ican Naturalist, 111:917-938. 

REDFORD, K., AND C. PADOCH (EDs.). 1992. Conser- 
vation of Neotropical forests. Columbia University 
Press, New York, 475 pp. 

Risse, D.O. 1992. The monogamous mating system 
of Peromyscus californicus as revealed by DNA fin- 


BEHAVIOR 419 


gerprinting. Behavioral Ecology and Sociobiology, 
29:161-166. 

RICHARD, A. 1985. Primates in nature. W. H. Free- 
man, New York, 558 pp. 

Rosinson, J. G. 1986. Seasonal variation in use of 
time and space by the wedge capped capuchin mon- 
key. Smithsonian Contributions to Zoology, 431:1- 
60. 

Rosinson, J. G., AND K. H. REDFORD (EDs.). 1991. 
Neotropical use and conservation. The University 
of Chicago Press, Chicago, 520 pp. 

Rosinson, M., AND L. Tier. 1991. Man and beast 
revisited. Smithsonian Press, Washington, D.C., 386 
pp. 

ROSENBLATT, J., AND B. KoMISARUK. 1977. Repro- 
ductive behavior and evolution. Plenum Press, New 
York, 169 pp. 

RUBENSTEIN, D. I., AND R. W. WRANGHAM (EDS.). 1987. 
Ecological aspects of social evolution. Princeton 
University Press, Princeton, New Jersey, 551 pp. 

RupRAN, R. 1973. Adult male replacement in troops 
of purple faced langurs. Folia Primatologica, 19:41- 
60. 

SAUER, K. 1969. Seed, spades, hearths and herbs. 
MIT Press, Cambridge, Massachusetts, 194 pp. (re- 
print.) 

SCHALLER, G. B. 1963. The mountain gorilla: ecology 
and behavior. The University of Chicago Press, Chi- 
cago, 431 pp. 

1972. The Serengeti lion. The University of 

Chicago Press, Chicago, 480 pp. 

1993. The last panda at the zoo. The Uni- 
versity of Chicago Press, Chicago, 292 pp. 

Scumitz, O. J. 1991. Thermal constraints and opti- 
mization of winter feeding and habitat choice in 
white-tailed deer. Holarctic Ecology, 14:104-111. 

SCHOENER, T. 1987. Time budgets and territory size: 
some simultaneous optimization models for energy 
maximizers. American Zoologist, 27:259-291. 

SCHONEWALD-Cox, C. M., S. M. CHAMBERS, B. 
MAcBryDE, AND W. L. THOMAS (EDS.). 1983. Ge- 
netics and conservation. Benjamin Cummings, Lon- 
don, 174 pp. 

SCHUSTERMAN, R., J. THOMAS, AND F. Woop (EDs.). 
1986. Dolphin cognition and behavior: a compar- 
ative approach. Laurence Erlbaum Associates, Hills- 
dale, New Jersey, 393 pp. 

SCHWARTZ, O. A., AND K. B. ARMITAGE. 1980. Ge- 
netic variation in social mammals: the marmot mod- 
el. Science, 207:665-667. 

SEIDENSTICKER, J., AND S. LUMPKIN (EDS.). 1991. Asian 
elephants. Friends of the National Zoo in the Zoo- 
logical National Park, Washington D.C., 271 pp. 

SeToN, E. T. 1953. Lives of game animals, vols. 1- 
4. Charles T. Brandford, Boston, Originally pub- 
lished in 1909 and 1925, 949 pp. (reprint.) 

SHADLE, A. R. 1946. Copulation in the porcupine. 
The Journal of Wildlife Management 10:159-162. 

SHERMAN, P. W. 1981. Reproductive competition and 
infanticide in Belding ground squirrels and other an- 
imals. Pp. 311-331, in Natural selection and social 
behavior: recent research and new theory (R. D. Al- 
exander and D. W. Tinkle, eds.). Chiron Press, New 
York, 532 pp. 


1989. Mate guarding as paternity assurance 
in Idaho ground squirrels. Nature, 338:418-420. 
SHreLps, W. M. 1982. Philopatry, inbreeding and the 
evolution of sex. State University New York Press, 

Albany, 245 pp. 

SIGLER-EISENBERG, B. J. 1988. Settlement, subsis- 
tence and environment. Pp. 291-306, in Wetlands 
site archaeology (B. Purdy, ed.). Telford Press, Cald- 
well, New Jersey, 338 pp. 

Sik, J. B. 1983. Local resource competition and fac- 
ultative adjustment of sex ratios in relation to com- 
petitive ability. The American Naturalist, 121:56- 
66. 

SLOBODCHIKOFF, C. N. 1988. The ecology of social 
behavior. Academic Press, San Diego, 429 pp. 

Smuts, B. B., D. L. CHENEY, R. M. SEYFARTH, R. W. 
WRANGHAM, AND T. T. STRUSAKER (EDS.). 1987. 
Primate societies. The University of Chicago Press, 
Chicago, 578 pp. 

SmyTHE, N. 1970. Relationships between fruiting 
seasons and seed dispersal methods. The American 
Naturalist, 104:25-35. 

1991. Steps toward the domestication of the 
paca. Pp. 202-216, in Neotropical use and conser- 
vation (J. G. Robinson and K. Redford, eds.). Uni- 
versity of Chicago Press, Chicago, 520 pp. 

STENSETH, N. C., AND W. Z. LipickeR, JR. 1992. An- 
imal dispersal: small mammals as a model. Chap- 
man and Hall, London, 365 pp. 

STRUHSAKER, T. 1975. The red colobus monkey. The 
University of Chicago Press, Chicago, 311 pp. 

SuMNER, F. B. 1932. Genetic, distributional, and evo- 
lutionary studies of the subspecies of deer mice 
(Peromyscus). Bibliographica Genetica, 9:1-106. 

Sunquist, M. E. 1981. The social organization of 
tigers (Panthera tigris). Pp. xx-xxx, in Royal Chi- 
tawan National Park. Smithsonian Contributions to 
Zoology, 336:1-98. 

SunquisT, M. E., AND J. F. EISENBERG. 1993. Repro- 
ductive strategies in female Didelphis. Bulletin of the 
Florida Museum of Natural History, Biological Sci- 
ences, 36:109-140. 

Symons, D. 1979. The evolution of human sexuality. 
Oxford University Press, New York, 358 pp. 

TAMARIN, R. H., R. S. OsTFELD, S. R. PUGH, AND G. 
BusJALsKA. 1990. Social systems and population 
cycles of voles. Birkhauser, Basel, Switzerland, 229 


TeEsTER, J. R., D. W. WARNER, AND W. W. COCHRAN. 
1964. A radio tracking system for studying move- 
ments of deer. The Journal of Wildlife Management, 
28:42-45. 

TINBERGEN, N. 1951. The study of instinct. Oxford 
University Press, London, 228 pp. 

ToLMAN, E. 1932. Purposive behavior in animals and 
men. University of California Press, Berkeley, 463 


TRILLMICH, F. 1986. Maternal investment and sex 
allocation in the Galapagoes fur seal, Arctocephalus 
galapagoensis. Behavioral Ecology and Sociobiolo- 
gy, 19:157-164. 

Trivers, R. L. 1971. The evolution of reciprocal al- 
truism. The Quarterly Review of Biology, 46:35-57. 

1972. Parental investment and sexual selec- 


420 EISENBERG AND WOLFF 


tion. Pp. 136-179, in Sexual selection and the de- 

scent of man 1871-1971 (B. Campbell, ed.). Aldine, 

Chicago, 378 pp. 

. 1974. Parent-offspring conflict. American 

Zoologist, 14:249-264. 

. 1983. Social evolution. Benjamin Cum- 
mings, Menlo Park, 462 pp. 

Trivers, R. L., AND D. E. WILLARD. 1973. Natural 
selection of parental ability to vary the sex ratio of 
offspring. Science, 179:90-92. 

TRUNE, D. R., AND C. N. SLOBODCHIKOFF. 1978. Po- 
sition of immatures in pallid bat clusters: a case of 
reciprocal altruism? Journal of Mammalogy, 59:193- 
195. 

VALKENBURGH, B. VAN. 1990. Skeletal and dental 
predictors of body mass in carnivores. Pp. 181-205, 
in Body size in mammalian paleobiology (J. Damuth 
and B. MacFadden, eds.). Cambridge University 
Press, Cambridge, 397 pp. 

WALKER, E. P. 1964. Mammals of the world, 3 vols. 
Johns Hopkins University Press, Baltimore, 2:1-268. 

Watson, J. B. 1914. Behavior: an introduction to 
comparative psychology. Holt, New York, 439 pp. 

Wess, S. D. 1983. The rise and fall of the ungulate 
fauna in the Miocene. Pp. 267-306, in Coevolution 
(M. H. Nitecki, ed.). The University of Chicago Press, 
Chicago, 392 pp. 

WEMMER,C. 1977. Comparative ethology of the large 
spotted genet. Smithsonian Contributions to Zool- 
ogy, 239:1-93. 

WHITAKER, J. O., JR. 1994. Academic propinquity. 
Pp. 121-138, in Seventy-five years of mammalogy 
(1919-1994) (E. C. Birney and J. R. Choate, eds.). 
Special Publication, The American Society of Mam- 
malogists, 11:1-433. 

WHITMAN, C. O. 1899. Animal behavior. Biological 


Lectures of the Marine Biological Laboratory (Wood’s 

Hole), 6:285-338. 

. 1919. The behavior of pigeons. Publications 
of Carnegie Institute, Carnegie Institute of Wash- 
ington, Washington, D.C., 257:1-161. 

WILLIAMS, G. G. 1966. Adaptation and natural se- 
lection. Princeton, New Jersey, 307 pp. 

Wiison, D. E., AND J. F. EISENBERG. 1990. Origin 
and applications of mammalogy in North America. 
Current mammalogy (H. H. Genoways, ed.). 2:1-35. 

Witson, E. O. 1975. Sociobiology—the New Syn- 
thesis. Harvard University Press, Cambridge, Mas- 
sachusetts, 697 pp. 

WITTENBERGER, J. F. 1981. Animal social behavior. 
Duxbury Press, Boston, 722 pp. 

Wo LFF, J. O. 1992. Parents suppress reproduction 
and stimulate dispersal in opposite-sex juvenile white- 
footed mice. Nature, 359:409-410. 

1993. What is the role of adults in mam- 
malian juvenile dispersal? Oikos, 68:173-176. 

WoLrFF, J. O., AND D. M. CictRELLO. 1989. Field 
evidence for sexual selection and resource compe- 
tition infanticide in white-footed mice. Animal Be- 
haviour, 36:456-465. 

. 1991. Comparative paternal and infanticidal 
behavior of white-footed mice and deermice. Be- 
havioural Ecology, 2:38-45. 

WYNNE-EDWARDS, V. C. 1962. Animal dispersion in 
relation to social behaviour. Oliver and Boyd, Ed- 
inburgh, 653 pp. 

YERKES, R. M., AND A. W. YERKES. 1929. The great 
apes. Yale University Press, New Haven, Connect- 
icut, 652 pp. 

XIA, X., AND J. S. MILLAR. 1991. Genetic evidence 
of promiscuity in Peromyscus leucopus. Behavioral 
Ecology and Sociobiology, 28:171-178. 


CONSERVATION AND MANAGEMENT 


JAMES H. SHAW AND DAvipD J. SCHMIDLY 


Introduction 


f the space allocated them in zoos 
throughout the world is an accurate 
gauge, the public considers mammals to be 
the most popular class of vertebrates. This 
popularity is confirmed by the history of 
conservation, in which wild species of 
mammals, from American bison (Bison bi- 
son) in the 19th Century to the giant panda 
(Ailuropoda melanoleuca) in the late 20th 
Century have been prominently featured. 
Yet popularity alone is not enough to en- 
sure survival. Some characteristics of mam- 
mals, including thick, luxurious coats of hair, 
have prompted commercial exploitation, 
depletion and, in some cases extinction, 
within historical times. Higher energy de- 
mands imposed by homeothermy require 
larger areas of natural habitat to sustain 
populations of mammals, as compared with 
reptiles of similar body size and food habits. 
The large brains of mammals, together with 
lengthy periods of lactation and parental 
protection, generally correlate with rela- 
tively low reproductive rates. Animals with 
low reproductive rates are slow to recover 
from population reductions and fare poorly 
in unstable environments. 
Wild mammals are thus esthetically pop- 
ular, commercially valuable, and biologi- 
cally vulnerable. In a world increasingly 


421 


U.S. FISH AND WILDLIFE SERVICE 
DEPARTMENT OF THE INTERIOR 


NATIONAL WILDLIFE REFUGE 


dominated by human activities, political 
clashes over the fate of wild mammals will 
increase. The early successes of North 
American conservation stemmed more from 
shifts in public attitudes than from the sci- 
ence of mammalogy. Indeed, direct legal 
protection, popularly supported and mgor- 
ously enforced, remains a cornerstone of 
conservation. 

But the problems faced by mammalian 
species worldwide are now far more com- 
plex and subtle than direct overharvesting. 
These include habitat destruction, isolation 
through fragmentation, assorted effects of 
scale, genetic depletion, introduced organ- 
isms, and the prospects of global climatic 
changes. Since its inception, the ASM has 
actively promoted the conservation of wild 
mammals, but today’s more pervasive and 
complicated threats require greater involve- 
ment by mammalogists and other scientists. 
Thus, a major theme featured here is the 
increasing role of science in the formulation 
and evaluation of conservation. 


Before 1919 


From the establishment of the first col- 
onies through the 19th Century, Americans 


422 SHAW AND SCHMIDLY 


of European descent viewed wild animals 
as obstacles to progress that would, like the 
American Indian, vanish before the ad- 
vance of civilization. Wild mammals were, 
at best. perceived as temporary resources 
for uses ranging from subsistence by early 
settlers to a means of enriching speculators 
through the fur trade. 

Given such attitudes and conditions, game 
abundance around settlements declined. The 
Massachusetts Bay Colony, for example, first 
closed the season on deer in 1694 (Mat- 
thiessen, 1987). 

Subsistence hunting by settlers and mar- 
ket hunting by native Americans for trade 
with whites had begun to take its toll by the 
time of American independence. Principal- 
ly through analysis of early trade records, 
McCabe and McCabe (1984) estimated that 
white-tailed deer (Odocoileus virginianus) 
numbered between 24 and 34 million in 
pristine North America. By 1800, the pop- 
ulation had declined by an estimated 50- 
65%. Deer rebounded slightly during the first 
half of the 19th Century, owing to the dis- 
placement of many native Americans from 
the East, but a resurgence of market hunt- 
ing, this time by Americans of European 
descent, forced the number of white-tailed 
deer to a low of between 300,000 and 
500,000 by 1900 (McCabe and McCabe, 
1984). 

Market hunting. —Market hunting flour- 
ished after the Civil War. Firearms 1m- 
proved, first with breech-loaders and then 
with repeating rifles and shotguns. During 
the same period, railroad transportation 
greatly expanded wild game markets to bur- 
geoning eastern populations. 

The white-tailed deer, of course, was not 
the only species to decline in the face of 
more efficient market hunting. American 
bison were slaughtered first for subsistence 
and later for the market value of their 
tongues and hides. Naturalist and anthro- 
pologist George Bird Grinnell, hunting bi- 
son along the Republican River in 1872, 
found the species even then in such serious 
decline that he thought extinction likely 


(Reiger, 1972). In 1874, Congress passed 
legislation to prohibit the killing of female 
bison by Americans of European descent, 
but President Grant gave the bill a pocket 
veto (McHugh, 1972). Further interest in 
protecting bison dissipated two years later 
with news that Custer and five companies 
of the 7th Cavalry had died at the Little 
Bighorn. Thereafter, European Americans 
accepted Phil Sheridan’s praise for bison 
hunters who were busily destroying the “‘In- 
dians’ commissary” (McHugh, 1972). 

The early conservation movement in North 
America. — The near extinction of the bison 
provided a rallying point for America’s first 
movement for wildlife preservation. This 
movement, beginning in the 1880s, resulted 
from pressure by sportsmen’s groups that 
flourished during that period, and from na- 
ture enthusiasts, who took much of their 
sentiment from 19th Century romanticism 
(Dunlap, 1988). Prompted by the American 
Ornithologists’ Union, Congress estab- 
lished the Office of Economic Ornithology 
and Mammalogy within the U.S. Depart- 
ment of Agriculture in 1885. Forerunner of 
the Bureau of Biological Survey and U.S. 
Fish and Wildlife Service, this new Office 
had Clinton Hart Merriam as its first chief. 

The early preservation movement gath- 
ered momentum in the 1890s with devel- 
opment of “realistic” nature stories, by Er- 
nest Thompson Seton and others. These 
stories attempted to use the science of that 
time (including the now discredited “‘sci- 
ence”’ of animal psychology) as a vehicle to 
deliver a moral message, and gained wide 
readership through popular magazines 
(Dunlap, 1988). 

Despite growing sentiment in favor of 
wildlife preservation, market hunting con- 
tinued. By 1900, most states had laws reg- 
ulating hunting, but inconsistencies be- 
tween neighboring states, together with ease 
of transporting wild animal products from 
one state to another, allowed de facto mar- 
ket hunting to continue. Growing sentiment 
in favor of wildlife protection led Congress 
to pass the Lacey Act in 1900. The Lacey 


CONSERVATION 423 


Act, drawing on Congressional authority to 
regulate interstate commerce, made inter- 
state shipment of game taken in violation 
of state laws a federal offense. In addition, 
the Lacey Act imposed federal restrictions 
on importation of exotic wildlife. 

Sentiment toward predators was an en- 
tirely different matter. Neither hunters nor 
nature lovers of the early 20th Century ap- 
preciated the value of carnivores. The same 
sentimental view that advocated protection 
for “noble”’ species like the elk (Cervus ela- 
phus) depicted predators such as the gray 
wolf (Canis lupus) as cruel, cunning, de- 
structive, and even dangerous. Lacking a 
lobby, predators of the time did not lack 
opponents; stockmen looked to the federal 
government for support in their war on 
predators. 

Responding to the stockmen’s wishes, 
Congress authorized the expenditure of the 
first federal funds for predator control in 
1914. The following year, the Bureau of Bi- 
ological Survey hired professional trappers 
and began implementing its Congressional 
mandate. 

Direct legal protection. —Through the ear- 
ly years of the 20th Century, efforts to aid 
wild mammals focused almost entirely upon 
direct legal protection. Motives stemmed 
from the desire of sportsmen to increase 
their hunting opportunities and from nature 
enthusiasts whose interest in wildlife was 
sentimental and aesthetic. Zoologists (the 
term ““mammalogist’”’ was not then in gen- 
eral use) had little direct involvement with 
efforts to improve the status of wildlife. 
Those who specialized in mammals studied 
taxonomy and made inferences concerning 
phylogeny. Moreover, many early mammal 
specialists lacked formal academic prepa- 
ration, having learned mammalogy through 
apprenticeships. 

In the absence of science, wildlife con- 
servationists developed measures based on 
cultural tradition, sentiment, and dogma, 
and used the law as the main vehicle for 
implementation. Given the rudimentary 
state of ecology at the time, such an ap- 


proach may have been unavoidable. The 
drawback of such a non-scientific basis was 
that its effectiveness and progress could not 
be objectively measured and evaluated. A 
program’s success, aside from a few obvious 
cases in which wild populations greatly ex- 
panded or declined, simply could not be 
determined. Ineffective or misguided pro- 
grams, such as the “buck laws” that pro- 
tected female cervids, were sustained for de- 
cades. 


After 1919 


By the time that the ASM was founded 
in 1919, the term “‘conservation”’ had come 
into general use. Gifford Pinchot first used 
the word in its modern context, feeling the 
need for a term that included the taking of 
a sustainable yield from a managed resource 
(Trefethen, 1975). To sport hunters, of 
course, Pinchot’s goal of sustainable yield, 
developed initially for commercial timber, 
applied equally well to game. 

Application of Pinchot’s principles to wild 
mammals required information obtainable 
only through field studies. Given the limited 
development of ecological principles at the 
time, almost nonexistent funding for re- 
search, and the shortage of qualified field 
workers, field data would be long in coming. 
Wildlife conservation as applied to game 
would continue to be based on tradition and 
implemented through arbitrary seasons and 
bag limits that may have had little to do 
with biological reality. 

Controversy over policy on mammalian 
predators. — Popular sentiment in the years 
between the World Wars still favored the 
destruction of medium-to-large carnivores, 
both to protect livestock and to protect pop- 
ular game animals. Gradually, however, 
many of the naturalists and biologists with 
the Bureau of Biological Survey became 
concerned over the decline of large mam- 
malian predators and the accidental killings 
of other wild animals. Others accepted more 
traditional views of predators and em- 


424 SHAW AND SCHMIDLY 


barked enthusiastically on their agency’s 
mission to eradicate them. Neither side 
could seek answers in science, as not even 
the most basic field studies of food habits, 
behavior, and population ecology of wild 
predatory mammals existed. Given that 
many of the ASM’s founders, including its 
first president, C. Hart Merriam, were past 
or present employees of the Bureau, that 
controversy was bound to divide the new 
society as well. 

Open opposition to government predator 
control flared at the society’s 1924 meeting, 
where two Survey biologists, Edward A. 
Goldman and W. B. Bell, were called upon 
to defend their agency’s policy (Dunlap, 
1988). Thus began a protracted and often 
bitter controversy that would erupt from 
time to time for nearly half a century. The 
controversy was propelled not only by a lack 
of field data, but also by a fundamental 
question concerning the mission of the Sur- 
vey and of its successor, the U.S. Fish and 
Wildlife Service. Critics of predator control 
contended that the agency should work on 
behalf of publicly-owned wildlife, as it did 
in most other programs. Predator control 
was another matter. With cooperative fund- 
ing from states and livestock growers, it was 
becoming a service for the benefit of the 
livestock industry. 

As the predator control controversy con- 
tinued, gradual progress was made on the 
conservation and management of game spe- 
cies. Game recovery turned out to require 
more than mere legal protection. Changes 
in the land, brought about through agricul- 
ture, grazing, mining, and the clearing of 
forests took place at about the same time as 
excessive commercial hunting. Thus, with- 
out some type of habitat restoration, game 
protection often could not succeed. 

Science-based conservation programs in 
universities. —Early in the 20th Century, 
Frederick Clements (1916) gave the world 
his theory of plant succession and Victor 
Shelford (1913) described the concept of 
natural animal communities. These pio- 
neering treatises laid the theoretical foun- 


dations for the study of natural communi- 
ties by describing the process of plant 
succession and by presenting criteria for de- 
fining the original biomes or major habitat 
associations of North America. Wildlife 
conservation could now take advantage of 
these discoveries and did so, albeit slowly 
at first. What was needed was a formal text- 
book and academic programs in wildlife 
conservation and management. 

The unifying textbook (Leopold, 1933) 
appeared and, shortly thereafter, its author 
accepted a professorship in game manage- 
ment at the University of Wisconsin, the 
first of its kind in the United States. Leopold 
and his students provided some of the first 
ecological studies on wild animals that could 
be applied directly to conservation and 
management. 

Academic programs in wildlife manage- 
ment received another important boost 
through one of the many ideas of J. N. 
“Ding” Darling. Darling helped set up a 
special research unit at Iowa State Univer- 
sity, paying some of the initial costs himself. 
The U.S. Fish and Wildlife Service expand- 
ed Darling’s prototype into a series of Wild- 
life Cooperative Research Units at major 
universities to bolster graduate programs in 
wildlife conservation and management. 

Public funding for conservation. —Through 
the mid-1930s, state wildlife conservation 
agencies received virtually all of their funds 
from the sale of hunting and fishing licenses. 
These funds were generally insufficient for 
wildlife research and, more importantly, the 
money from license sales was controlled by 
state legislatures, who often transferred 
funds to state projects unrelated to wildlife. 

The solution to the problems of inade- 
quate funding, and the allocation of fish and 
game monies to other state projects, came 
in the form of the Federal Aid to Wildlife 
Restoration Act in 1937. Often called sim- 
ply the Pittman-Robertson (P-R) Act, it was 
arguably the most important federal legis- 
lation affecting American wildlife. The Act 
placed a federal excise tax on the manufac- 
ture of sporting arms and ammunition, and 


CONSERVATION 425 


redistributed the revenue, via federal au- 
thorities, to state wildlife conservation 
agencies on a matching basis. 

To qualify for this federal aid, each state 
had to pass enabling legislation ensuring that 
all funds collected through license sales 
would be used only for fish and wildlife pur- 
poses. State wildlife agencies now had a 
broader, more sustainable source of fund- 
ing, and one that was virtually immune to 
policial manipulation. Within a year, 43 of 
the then 48 states complied, and the other 
five followed soon thereafter (Williamson, 
1987). In 1939, P-R apportioned $890,000 
to the states. By 1986, that figure had grown 
to over $107 million (Kallman, 1987). 

Federal aid funds were earmarked for 
wildlife restoration, not for law enforce- 
ment. These monies made possible much 
of the desperately needed research on wild- 
life habitat problems and on the implemen- 
tation of solutions. Finally, legal regulations 
of harvests were being supplemented by 
habitat improvement. 

Progress after World War IT.—The pros- 
perity after World War IJ prompted many 
changes in wildlife conservation. Returning 
servicemen exchanged uniforms for hunting 
garb and state license sales boomed. Cor- 
respondingly, P-R reapportionment soared 
from $817,500 in 1945 to nearly $11 mil- 
lion in 1949 (Kallman, 1987). Increased 
revenue led to more wildlife research and 
management. 

The postwar years brought about increas- 
es in international cooperation and trade. 
As international concerns in general grew, 
so did interest in wildlife management and 
conservation on a global scale. The Inter- 
national Union for Conservation of Nature 
and Natural Resources (IUCN) was formed 
in 1948 as an independent international or- 
ganization to promote wise and sustainable 
use of the world’s natural resources. Mem- 
bership in the IUCN consisted of national 
government, governmental agencies con- 
cerned with conservation, and private or 
non-governmental organizations (NGOs). 
Leadership from the IUCN has helped de- 


velop international treaties on behalf of 
wildlife. 

In 1961, another important NGO, the 
World Wildlife Fund, came into being. The 
World Wildlife Fund’s primary mission was 
to raise money on behalf of vanishing spe- 
cies throughout the Earth. Both the IUCN 
and the World Wildlife Fund were based in 
Switzerland. 

The first postwar international conven- 
tion affecting wild mammals was the Inter- 
national Convention for the Regulation of 
Whaling, which met in Washington, D.C., 
late in 1946. Superceding the earlier 1931 
Convention, this one established the Inter- 
national Whaling Commission (IWC), 
charged with reviewing harvests and estab- 
lishing quotas. The Commission issued few 
restrictions until the early 1960s when, faced 
with clear evidence of depleted stocks and 
an international lobby opposed to whaling, 
it gradually shifted toward more protection. 
In 1982, the IWC agreed to set commercial 
whaling quotas at zero by 1986 and to re- 
view the effects of this protection on whale 
stocks by 1990 (Lyster, 1985). 

Sustainable harvests.—Detailed under- 
standing of the effects of harvest on wild 
mammals has been slow in coming because 
the species most likely to be affected by har- 
vest are large, have low rates of increase, 
and long generation times. These traits make 
conclusive field investigations lengthy and 
expensive. Furthermore, large mammals fall 
under the jurisdiction of established wildlife 
agencies, subject to their own priorities and 
pressures exerted by various interest groups. 
Such agencies are often reluctant to approve 
the sort of long-term, high-visibility field 
investigations that would be required to im- 
prove the predictability of the effects of game 
harvests. 

Game harvests have remained imprecise 
and unrefined since the turn of the century. 
About the best that can be said about tra- 
ditional seasons and bag limits is that, with 
rare exception, they avoid overharvests. 
Even into the 1980s, a leading specialist in 
the harvest of large mammals concluded that 


426 SHAW AND SCHMIDLY 


the principle change in hunting regulations 
in the United States over the past several 
decades was a relaxation of the ban against 
hunting on Sundays (Caughley, 1985). 

Broader public interest.— Although regu- 
lation of hunting changed little in postwar 
years, public interest in non-game species 
has increased substantially. Concern over 
rare and endangered species led to passage 
of the first Endangered Species Act in 1966. 
More symbolic than substantive, the Act 
did little more than authorize the Secretary 
of the Interior to develop and maintain a 
list of vanishing wildlife threatened with ex- 
tinction. 

The environmental movement in the late 
1960s led to passage of the Endangered Spe- 
cies Act of 1969, curbing imports on wild 
animals (and parts thereof) threatened in 
their native lands. Four years later another 
Endangered Species Act retained refined el- 
ements from its two predecessors and ex- 
tended federal protection to native wildlife 
threatened with extinction. Section 6 of this 
Act provided for federal funds for use by 
state wildlife agencies on behalf of endan- 
gered species. Since the Act’s Section 7 pro- 
tected critical habitat of endangered species 
from any development using federal funds, 
it provided for interagency consultation to 
resolve conflicts and suggest alternatives 
(Yaffee, 1988). 

In 1972, Congress passed the Marine 
Mammal Protection Act (MMPA). This Act 
applied to all marine mammals and placed 
a moratorium on their harvest or harass- 
ment. It also established regulatory author- 
ity over commercial use of marine mam- 
mals and products made from them. Finally, 
recognizing that marine mammals play 1m- 
portant roles in marine ecosystems, the Act 
prohibited reduction of marine mammal 
populations to the point that they cease to 
perform their ecological functions (Dunlap, 
1988; Trefethen, 1975). 

Exploitation vs. protection.—One of the 
most persistent controversies in wild mam- 
mal conservation is the conflict over con- 
trolled exploitation versus preservation. 


With its long and generally successful tra- 
dition in game management, wildlife con- 
servation in the United States and Canada 
generally leans toward controlled exploita- 
tion, principally through sport hunting. Not 
only can sport hunting help populations re- 
cover, it can provide landowners with in- 
centives to maintain natural habitat and can 
generate important revenue. Nonetheless, 
the preservationist view—that the best way 
to ensure survival of wild animals is through 
complete protection from exploitation—has 
gained favor during the past 2 decades. 
Management of endangered species in most 
cases precludes exploitation. Populations of 
many furbearing and, especially, marine 
mammals have recovered well when afford- 
ed complete protection. Each approach can 
work under some conditions, but decisions 
often are clouded by ideological divisions 
between the two camps. This division pre- 
vents some private conservation organiza- 
tions from working together more effective- 
ly and presenting a united front on broader 
conservation issues. 

Given proper habitat, most North Amer- 
ican game mammals fare quite well, wheth- 
er subjected to regulated hunting or afforded 
complete protection. Wild species found in 
increasingly crowded developing nations, 
however, may not be so fortunate. While 
tourism attracted by the large mammals of 
East Africa offers justification for protec- 
tion of wildlife in national parks, un- 
checked human population growth in 
nations like Kenya may soon overcome that 
advantage (Myers, 1979, 1985). Rather than 
have parks steadily converted to subsistence 
farms, a better strategy may be to employ 
would-be farmers in a sustainable harvest 
of wild mammals and in processing them 
for sale. Unfortunately, there is no clear an- 
swer. Just as either controlled harvest or 
complete protection can ensure the survival 
of most species of wild mammals in North 
America, either strategy could result in ex- 
tinction in the poorer, more crowded de- 
veloping nations. 

Even in North America, the debate over 


CONSERVATION 427 


exploitation continues among professional 
mammalogists and wildlife managers. One 
important example is game ranching, used 
in various forms in Europe, New Zealand, 
South Africa, the United States, and Can- 
ada. Game ranching 1s practiced on private 
land and involves to some degree the “‘pri- 
vatization” of what is usually regarded as 
public property. Proponents argue that game 
ranching offers important economic incen- 
tives to private landowners who would oth- 
erwise convert wildlife habitat to more prof- 
itable uses. While the practice may require 
intensive management and acceptance of 
some rather artificial conditions, it may of- 
fer the only real hope for retaining large wild 
mammals on private lands. 

Legislation aimed at encouraging private 
game ranching in Alberta, Canada, recently 
generated sharp controversy. Geist (1988) 
argued that privatization would undermine 
what has generally been successful wildlife 
conservation. Further, any shift from public 
to private ownership would leave popula- 
tions of large wild mammals at the whims 
of market forces. When market demand was 
high, incentives to overharvest would be 
powerful. Conversely, when market de- 
mand slacked off, neglect would ensue. 

If Geist’s (1988) arguments are valid, and 
if they apply to wild mammals outside of 
Alberta, then they challenge a basic premise 
of the IUCN’s World Conservation Strat- 
egy. Can wild mammals be exploited on a 
sustainable basis by market forces? Put an- 
other way, can markets themselves become 
sufficiently stabilized to ensure the long-term 
survival of wild mammals? And, if privately 
owned wild mammals are successfully es- 
tablished, will their wild counterparts be re- 
garded as competitors to be destroyed? 

The international wildlife trade. —Just as 
unregulated market hunters in the United 
States depleted wild mammals in the 19th 
Century, unregulated international com- 
merce in wild mammals and parts thereof 
began to threaten numerous species by the 
mid-20th Century. After a decade of 
prompting by the IUCN, a Convention on 


International Trade in Endangered Species 
(CITES) convened in Washington, D.C., in 
March, 1973. The Convention decided to 
list the more imperiled species in its Ap- 
pendix I and to require both an export per- 
mit from the country of origin and an im- 
port permit from the country of destination. 
Species less critically threatened, but none- 
theless rare, are listed in its Appendix IJ and 
require an export permit from the country 
of origin. In both cases, permits are issued 
by a “scientific authority,” typically a wild- 
life or natural resource management agency. 

Practically speaking, legal trade of Ap- 
pendix I is negligible between signatories. 
Appendix II listings, however, allow trade 
at the discretion of the originating country 
but require record keeping and regular re- 
porting. These public records prove useful 
in monitoring trade and population trends 
for periodic status review. 

At the 1976 review meeting of the Con- 
vention in Berne, Switzerland, members 
voted to adopt strict criteria for listing and 
delisting species. Under these ‘“‘Berne cri- 
teria,” the information required for listing 
a species need not be as detailed or conclu- 
sive as that for delisting. This arrangement 
reversed the traditional burden of proof, 
placing it on those who advocate exploita- 
tion rather than on those who urge protec- 
tion. Predictably, controversy ensued, but 
the rationale of erring on the side of pro- 
tection prevailed. 

Projections of global declines in wild 
mammals. — Despite the considerable prog- 
ress in conservation during the 1960s and 
70s, the 1980s opened with extraordinarily 
pessimistic projections for the Earth’s wild 
species. Deforestation, particularly of the 
little-known but species-rich tropical moist 
forests, was accelerating. Field studies 
showed that the recovery potential or resil- 
iency of tropical moist forests was far lower 
than that of temperate forests. International 
trade in wildlife and products from wildlife 
increased, spurred by rising demand in con- 
sumer nations and by increasing effort to 
use natural resources, such as wildlife and 


428 


forest products, to balance trade and to off- 
set growing indebtedness incurred by pro- 
ducer nations. 

Besides local habitat losses and heavier 
commercial exploitation, wild species be- 
gan facing threats from large-scale impacts 
to their environments. Ocean dumping and 
its resulting pollution increased in both scope 
and intensity. Atmospheric threats, first 
from acid precipitation and later from de- 
pletion of atmospheric ozone and increases 
in “greenhouse” gases, caused unprece- 
dented effects upon entire biomes. Thus, 7he 
Global 2000 Report to the President of the 
United States in 1980 projected that from 
15 to 20% of the world’s wild species, if 
current trends continued, would be extinct 
by the year 2000 (Barney, 1980). 

While the task force labored over The 
Global 2000 Report ... the IUCN devel- 
oped a comprehensive plan to offset some 
of the report’s more dire projections. The 
UCN’s World Conservation Strategy rec- 
ognized that humanity would continue to 
exploit the Earth’s seas and soils, but sought 
to thwart exploitation’s impact by shifting 
it toward sustainable development. This ba- 
sic change is analogous to the difference be- 
tween mining a nonrenewable resource and 
cropping a renewable one. 

Insofar as wild mammals were con- 
cerned, the World Conservation Strategy of- 
fered several recommendations, aimed 
principally at large mammals. First, a series 
of large nature reserves (of sufficient size to 
sustain wild populations of large mammals) 
should be established. Second, controlled 
exploitation, ranging from traditional sport 
hunting to less conventional game cropping, 
should be permitted in or around such ar- 
eas. Properly done, such harvest would al- 
low sustainable exploitation of meat and 
trophies, as well as providing employment 
and revenue. This is especially important in 
developing nations. Finally, the World Con- 
servation Strategy recommended preserving 
wild species of mammals because of the ge- 
netic diversity their populations contain, 
potentially useful for the improvement of 


SHAW AND SCHMIDLY 


existing livestock and for the creation of 
“new” domesticated animals in the future 
(IUCN, 1980). In short, the IUCN’s plan 
presented conservation as an integral part 
of economic development, rather than as 
the antithesis to it. 

Many conventional types of development 
clearly are not sustainable. One of these is 
the large-scale clearing and conversion of 
tropical moist forests, either for commercial 
logging or for conversion of lowland forest 
to farms and pastures. Once the primary 
forests are cleared, recovery of the ecosys- 
tems to anything resembling their original 
state becomes unlikely. Tropical forests hold 
their nutrients not in soils, but in decaying 
plant and animal matter near the soil sur- 
face. Clearing and burning deprives the al- 
tered ecosystem of nutrients needed for re- 
covery, and land surfaces become exposed 
for the first time to the direct effects of sun 
and wind. Insect and pest outbreaks follow, 
and remaining patches of tropical moist for- 
est succumb to isolation and the combined 
physical and biological changes along their 
edges (Lovejoy et al., 1986). 


New Approaches to the 
Conservation of Mammals 


Threats to the long-term survival of free- 
living wild mammals are larger and more 
complex than ever before. Participants in a 
recent conference in Washington, D.C., ex- 
amined the effects of atmospheric changes, 
largely the “greenhouse effect,’ on biodi- 
versity (Peters, 1988). The climatic changes 
brought about by increasing levels of at- 
mospheric carbon dioxide could trigger sig- 
nificant geographic shifts in plant and ani- 
mal communities. Thus national parks and 
other reserves, already suspected as being 
of insufficient size, may prove even less ef- 
fective at sustaining wild mammals as cli- 
mates shift. One possible solution would be 
to leave or develop north-south corridors 
of natural habitat between protected areas 


CONSERVATION 429 


in an effort to accommodate climatically- 
induced shifts in geographic ranges. 

As threats increase in scale, so must ef- 
forts in ecological research. Ecological stud- 
ies of wild mammals, particularly large spe- 
cies, increasingly are being carried out with 
the replication and controls needed in good 
experimental designs. In addition, the larger 
the scope of ecological investigations, the 
greater the cost. Thus, large-scale studies 
can become prohibitively expensive. One 
elegant and straightforward solution to these 
problems is the systematic use of wildlife 
management as scientific research (Mac- 
Nab, 1983). Rather than apply one general 
management practice to a region the size of 
a state, wildlife agencies could deliberately 
vary practices, be they harvest levels, hab- 
itat improvements, or other options, in ways 
that would allow direct comparisons and 
evaluations. Some areas could be left alone 
to serve as “‘controls.’’ This systematic ap- 
proach to management would require more 
careful planning and more detailed moni- 
toring, but their potential benefits would 
certainly be worth the extra effort. 

A similar framework with which to in- 
tegrate wildlife management with research 
is called comprehensive planning (Crowe, 
1983). Adopted to varying degrees by some 
state wildlife agencies, comprehensive plan- 
ning provides for periodic review of man- 
agement practices using pre-established cri- 
teria. A particular program in wildlife 
management is planned, implemented, re- 
viewed, and then reassessed routinely, thus 
allowing for improvement or, if necessary, 
replacement. Done properly, comprehen- 
sive planning not only provides important 
new research, but also reduces the political 
machinations that occur within agencies. 
Rather than deciding on a program’s fate 
purely through competing political forces, 
agencies can evaluate it through analysis of 
field data. Even when a management pro- 
gram completely fails to meet its objective, 
useful information can be obtained and the 
effort justified. 

A potentially far-reaching technique for 


large-scale field studies is a collection of 
computer software packages known as Geo- 
graphical Information Systems (GIS). GIS 
links attribute data (e.g., biogeographical 
province, biome type, species occurrence, 
topographic features) with positions on the 
earth (McLaren and Briggs, 1993). Two 
principal approaches are inventory, consist- 
ing of descriptive data, mapping, and da- 
tabase management, and analysis, com- 
prised of modeling and statistical treatments 
(Berry, 1993). Commonly used in natural 
resource management since the 1980s, GIS 
applications also are indispensible in de- 
tailed spatial studies of mammalian ecology 
(August, 1993). 

Gap analysis is a particular application of 
GIS designed to target spatial ““gaps”’ in state- 
wide habitat protection systems. Once 
identified, such “‘gaps’’ often can be filled 
to ensure adequate protection of threatened 
species and rare natural communities. Gap 
analysis offers the advantages of identifying 
needs of several species at once as well as 
presenting a more proactive approach in 
which conservation measures may be taken 
before situations become desperate (Scott et 
al., 1991). 

Wild mammals and the maintenance of 
biodiversity. — Wildlife conservation began 
as game management, with the aim of pro- 
ducing a “surplus” for sport hunting. Game 
management could be improved by field 
studies of the ecology of a game species in 
general and its responses to harvest and 
changes in land use in particular. Thus, game 
management succeeded by meeting the 
needs of game species one at a time. It 
seemed only reasonable in the early days of 
endangered species conservation to contin- 
ue this tradition from game management, 
except that the objective was restoration 
rather than harvest. 

Effective as it was for mostly temperate 
game mammals, this single-species man- 
agement proved inadequate in the face of 
such serious and widespread threats as de- 
forestation and increased international traf- 
ficking of wildlife. Of the roughly 4,100 spe- 


430 SHAW AND SCHMIDLY 


cies of mammals on Earth, only a small 
fraction has been studied sufficiently to per- 
mit development of detailed conservation 
plans. Conservationists began to realize that 
there was neither enough time nor resources 
to rely exclusively on single-species man- 
agement. New challenges required new ap- 
proaches. Professional wildlife conserva- 
tion began to shift from efforts to save 
“species A”’ (typically a large mammal with 
popular appeal) to preserving biodiversity 
on an ecosystem level. 

This biodiversity approach offers two dis- 
tinct advantages over single-species man- 
agement. First, it allows more efficient al- 
location of time and resources. Instead of 
4,100 management plans for wild mam- 
mals, it can rely on protecting reserves lo- 
cated in the roughly 193 biogeographical 
provinces or principal habitat types on which 
those 4,100 mammals depend for survival 
in the wild. Second, it recognizes the im- 
perative of saving self-sustaining ecosys- 
tems, a goal consistent with the IUCN’s view 
of sustainable uses of natural resources. 

Concern for preserving biodiversity be- 
gan attracting biologists from outside the 
traditional ranks of wildlife management. 
Population geneticists and evolutionary 
ecologists started to supplement their basic 
research with investigations into sustaining 
biodiversity. A new field, conservation bi- 
ology, appeared along with an edited book 
of the same name in 1980 (Soulé and Wil- 
cox, 1980). In 1987, the Society for Con- 
servation Biology was established with its 
journal, Conservation Biology. 

This new discipline is more broadly based 
than conventional wildlife management. 
Although conservation biology is interdis- 
ciplinary and includes many specialties, two 
of the more longstanding ones featured here 
are conservation genetics and insular ecol- 
ogy. 

Conservation genetics. —The importance 
of conservation genetics escaped the notice 
of most wildlife managers, who knew that 
genetic depletion posed a problem for do- 


mesticated mammals but saw little evidence 
of its practical significance to wild ones. 
Thriving populations of white-tailed deer, 
for example, founded from only a few in- 
dividuals, suggested that wild species had a 
greater resistence to genetic problems im- 
posed by small, isolated populations. 

The first clues that wild species might suf- 
fer from inbreeding appeared in studies of 
captive-bred zoo mammals in which more 
inbred populations consistently produced 
fewer surviving offspring than did less in- 
bred ones (Ralls et al., 1979). Why was there 
such a marked difference between the zoo 
populations and their free-living, trans- 
planted counterparts? Part of the answer 
stems from the fact that small populations 
of wild animals lose genetic variation in two 
stages (Franklin, 1980). The first is the so- 
called “founder effect” (Mayr, 1963), which 
occurs when a population undergoes sudden 
and severe numerical reduction, leaving only 
a small number of surviving “founders.” 
Fewer founders mean that rarer genes are 
likely to be lost for future generations, re- 
ducing genetic variation. 

The founder effect may be followed by 
additional genetic loss through inbreeding, 
genetic drift, or both. At low numbers, close 
relatives are likely to breed with one anoth- 
er. Also, small populations suffer from ge- 
netic drift, the loss of rarer genes by chance. 
These processes deplete genetic variability 
for each generation that a population is kept 
at low numbers. 

Zoo populations have been subjected to 
both stages of genetic losses. Reintroduced 
game populations typically experience only 
the founder effect, quickly increasing their 
numbers and reducing the effects of in- 
breeding and genetic drift. 

Insular ecology.—Insular ecology is an 
applied version of the classic theories of is- 
land biogeography (MacArthur and Wilson, 
1967). These theories predict that islands 
will be colonized by wild species at a rate 
inversely proportional to distance from the 
mainland. Moreover, species on islands be- 


CONSERVATION 431 


come extinct at rates inversely proportional 
to island size, the rationale being that the 
smaller the island, the smaller the popula- 
tions, and the smaller the populations, the 
greater the threat of extinction. 

These basic theories seem simple and 
plausible enough, and they are supported by 
studies of land-bridge islands separated from 
mainlands since sea levels rose at the end 
of the Pleistocene (Wilcox, 1980). When the 
islands were peninsulas, they presumably 
contained the same levels of species diver- 
sity that occurred on the mainland to which 
they were connected. Since the time of 1so- 
lation by rising sea water can be reliably 
estimated, comparisons of historical levels 
of species on those islands can be compared 
with diversity on mainlands. The resulting 
differences compare closely with those pre- 
dicted by island biogeography (Wilson and 
Willis, 1975). 

Protected areas of natural habitat are in- 
creasingly fragmented and isolated from one 
another in human-dominated landscapes. 
The species preserved on such habitat is- 
lands may be subject to the same general 
patterns of extinction incurred by species 
on land-bridge islands. To the extent that 
the analogy holds (a matter of some debate 
at the time of this writing), isolated habitat 
preserves may lose many species. Mammals 
seem to be the class of vertebrates most 
vulnerable to this effect of insular ecology 
(Wilcox, 1980). Except for bats, mammals 
are more limited than birds in their dis- 
persal, so their prospects for recolonizing 
isolated habitat preserves are limited. 
Mammals have higher metabolic demands 
than do reptiles and amphibians, and thus 
require larger areas over which to forage. 
Large mammalian carnivores, empirically 
recognized as ‘“‘extinction prone”’ (Ter- 
borgh, 1974), require the largest areas of all 
and it may be that this area requirement is 
at least as important in their demise as con- 
flict with human activity. Indeed, one recent 
model predicted that no national park or 
other habitat reserve anywhere in the world 


was large enough to sustain a population of 
large carnivores indefinitely (Belovsky, 
1987). 

An increasing role for mammalogists. — 
Perhaps the broadest question of all con- 
cerns the role that the discipline of mam- 
malogy will and should play in conserva- 
tion. Mammalogists concerned with 
conservation like to think that their research 
will lead to more effective conservation. Al- 
though distinguished authorities (Caughley, 
1985; Geist, 1988) have argued that early 
wildlife conservation in North America suc- 
ceeded despite the science of the day and 
not because of it, the situations in which we 
find ourselves today differ markedly from 
those that confronted Hornaday and Grin- 
nell. The forces contributing to the loss of 
wild mammals are not just local market 
hunters, but large-scale habitat disruption, 
unprecedented atmospheric and oceanic 
pollution, and international trafficking. 
Many of these new threats are highly tech- 
nical and require the technological assis- 
tance and conceptual innovation that can 
best be provided by science. Mammalogists 
will play an increasingly important role in 
ensuring the survival of wild mammals into 
the 21st Century and beyond. 


Summary and Conclusions 


The conservation of wild mammals be- 
gan largely with direct legal protection aimed 
at curbing excessive hunting and trapping. 
Such measures were based little on science, 
but the fact that many depleted populations 
recovered, especially those of North Amer- 
ican ungulates, attested to the utility of legal 
protection in dealing with overhunting. 

In recent decades, habitat alterations, 
habitat fragmentation, and genetic deple- 
tion have joined overharvesting as threats 
to the survival of wild species of mammals. 
These new threats have stimulated devel- 
opment of new scientific subdisciplines, such 
as conservation biology, and emerging tech- 


432 SHAW AND SCHMIDLY 


nologies, including geographical informa- 
tion systems. In the future, the conservation 
and management of wild mammals will de- 
pend even more on mammalogy and related 
sciences. 


Literature Cited 


Aucust, P. V. 1993. GIS and mammalogy: building 
a database. Pp. 11-26, in GIS applications in mam- 
malogy (S. B. McLaren and J. K. Braun, eds.). Okla- 
homa Museum of Natural History, Norman, 41 pp. 

Barney, G. O. 1980. The global 2000 report to the 
President of the United States. Vol 1: the summary 
report. Pergamon Press, New York, 360 pp. 

BeLovsky, G.E. 1987. Extinction models and mam- 
malian persistence. Pp. 35-57, in Viable populations 
for conservation (M. E. Soulé, ed.). Cambridge Uni- 
versity Press, New York, 189 pp. 

Berry, J. K. 1993. The application of GIS to mam- 
malogy: basic concepts. Pp. 4-10, in GIS applica- 
tions in mammalogy (S. B. McLaren and J. K. Braun, 
eds.). Oklahoma Museum of Natural History, Nor- 
man, 41 pp. 

CAUGHLEY, G. 1985. Harvesting of wildlife: past, 
present, and future. Pp. 3-14, in Game harvest man- 
agement (S. L. Beasom and S. F. Roberson, eds.). 
Caesar Kleberg Wildlife Research Institute, Kings- 
ville, Texas, 374 pp. 

CLEMENTS, F. E. 1916. Plant succession. Carnegie 
Institute Publication, New York, 242:1-512. 

Crowe, D. M. 1983. Comprehensive planning for 
wildlife resources. Wyoming Fish and Game De- 
partment, Laramie, 143 pp. 

Duntap, T. R. 1988. Saving America’s wildlife. 
Princeton University Press, Princeton, New Jersey, 
222 pp. 

FRANKLIN, I. R. 1980. Evolutionary change in small 
populations. Pp. 135-149, in Conservation biology 
(M. E. Soulé and B. A. Wilcox, eds.). Sinauer As- 
sociates, Sunderland, Massachusetts, 395 pp. 

Geist, V. 1988. How markets in wildlife meat and 
parts, and the sale of hunting privileges, jeopardize 
wildlife conservation. Conservation Biology, 2:15- 
26. 

IUCN. 1980. World conservation strategy. Interna- 
tional Union for Conservation of Nature and Nat- 
ural Resources, Gland, Switzerland, 40 pp. 

KALLMAN, H. (ED.). 1987. Restoring America’s wild- 
life, 1937-1987: the first 50 years of the Federal Aid 
in Wildlife Restoration (Pittman-Robertson) Act. 
U.S. Department of the Interior Fish and Wildlife 
Service, Washington, D.C., 394 pp. 

LeopoLtp, A. 1933. Game management. Charles 
Scribner’s Sons, New York, 418 pp. 

Lovejoy, T. E., ET AL. 1986. Edge and other effects 
of isolation on Amazon forest fragments. Pp. 257- 
285, in Conservation biology: the science of scarcity 
and diversity (M. E. Soulé, ed.). Sinauer Associates, 
Sunderland, Massachusetts, 584 pp. 

LysTer, S. 1985. International wildlife law: an anal- 


ysis of international treaties concerned with the con- 
servation of wildlife. Grotius Publications Limited, 
Cambridge, England, 470 pp. 

MacArtTHor, R. H., AND E. O. WILSON. 1967. The 
theory of island biogeography. Princeton University 
Press, Princeton, New Jersey, 203 pp. 

MacNas, J. 1983. Wildlife management as scientific 
experimentation. Wildlife Society Bulletin, 11:397- 
401. 

MATTHIESSEN, P. 1987. Wildlife in America. (revised 
edition) Viking/Penguin Press, New York, 332 pp. 

Mayr, E. 1963. Animal species and evolution. Har- 
vard University Press, Cambridge, Massachusetts, 
797 pp. 

McLaren, S. B., AND J. M. Briccs. 1993. Introduc- 
tion. Pp. 1-3, in GIS applications in mammalogy (S. 
B. McLaren and J. K. Braun, eds.). Oklahoma Mu- 
seum of Natural History, Norman, 41 pp. 

McCasg, R. E., AND T. R. McCabe. 1984. Of slings 
and arrows: an historical retrospective. Pp. 10-72, 
in White-tailed deer: ecology and management (L. 
K. Halls, ed.). Wildlife Management Institute and 
Stackpole Books, Harrisburg, Pennsylvania, 870 pp. 

McHuau, T. 1972. The time of the buffalo. Alfred 
A. Knopf, New York, 339 pp. 

Myers, N. 1979. The sinking ark. Pergamon Press, 
New York, 307 pp. 

1985. Conservation of Africa’s cats: prob- 
lems and opportunities. Pp. 437-446, in Cats of the 
world: biology, conservation, and management (S. 
D. Miller and D. D. Everett, eds.). National Wildlife 
Federation, Washington, D.C., 501 pp. 

Peters, R. L. 1988. Effects of global warming on 
species and habitats: an overview. Endangered Spe- 
cies Update, 5:1-8. 

RALLS, K., K. BRUGGER, AND J. BALLou. 1979. In- 
breeding and juvenile mortality in small populations 
of ungulates. Science, 206:1101-1103. 

Reicer, J. F. 1972. The passing of the Great West: 
selected papers of George Bird Grinnell. Winchester 
Press, New York, 182 pp. 

Scott, M. S., B. Csuti, K. Smitu, J. E. Estes, AND S. 
Caicco. 1991. Gap analysis of species richness and 
vegetative cover: an integrated biodiversity conser- 
vation strategy. Pp. 282-297, in Balancing on the 
brink of extinction: the Endangered Species Act and 
lessons for the future (K. A. Kohm, ed.). Island Press, 
Washington, D.C., 315 pp. 

SHELFORD, V.E. 1913. Animal communities in tem- 
perate America. University of Chicago Press, Chi- 
cago, 362 pp. 

Sou.é, M. E., AND B. A. WiLcox (EDs.). 1980. Con- 
servation biology. Sinauer Associates, Sunderland, 
Massachusetts, 395 pp. 

TERBORGH, J. 1974. Preservation of natural diversity: 
the problem of extinction-prone species. BioScience, 
24:715-722. 

TREFETHEN, J. B. 1975. An American crusade for 
wildlife. Winchester Press and the Boone and Crock- 
ett Club, New York, 409 pp. 

Witcox, B. 1980. Insular ecology and conservation. 
Pp. 95-117, in Conservation biology (M. E. Soulé 
and B. A. Wilcox, eds.). Sinauer Associates, Sun- 
derland, Massachusetts, 395 pp. 

WILLiAMson, L. L. 1987. Evolution of a landmark 


CONSERVATION 433 


law. Pp. 1-17, in Restoring America’s wildlife, 1937- of communities (M. L. Cody and J. M. Diamond, 

1987: the first 50 years of the Federal Aid in Wildlife eds.). Harvard University Press, Cambridge, Mas- 

Restoration (Pittman-Robertson) Act (H. Kallman, sachusetts, 545 pp. 

ed.). U.S. Department of the Interior Fish and Wild- YAFFEE, S. L. 1988. Endangered species protection 

life Service, Washington, D.C., 394 pp. through interagency consultation. Endangered Spe- 
WILson, E. O., AND E. O. Wituis. 1975. Applied bio- cies Update, 5:14-19. 


geography. Pp. 522-534, in Ecology and evolution 


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