Edited by
Mary E. Glenn and Marina Cords
Kluwer Academic / Plenum Publishers, 2002
TABLE OF CONTENTS
Preface
Mary E. Glenn and Marina Cords
Part I Evolutionary Biology and Biogeography
1 The Guenons: An Overview of Diversity and Taxonomy 3
Thomas M. Butyoski
2 Y-chromosomal Window onto the History of Terrestrial Adaptation in the Cercopithecini 15
Anthony J. Tosi, Paul J. Buzzard,
Juan Carlos Morales, and Don J. Melnick
3 Molecular Timescale and Gene Tree Incongruence in the Guenons 27
Todd R. Disotell and Ryan L. Raaum
4 Phylogeny of the Cercopithecus Ihoesti Group Revisited: Combining Multiple Character Sets 37
Jean-Pierre Gautier, Regine Vercauteren Drubbel, and Pierre Deleporte
5 Terrestriality and the Maintenance of the Disjunct Geographical Distribution in the Ihoesti Group 49
Beth A. Kaplin
6 Biogeographic Analysis of Central African Forest Guenons 61
Marc Colyn and Pierre Deleporte
7 Hybridization between Red-tailed Monkeys (Cercopithecus ascanius)
and Blue Monkeys (C. mitts) in East African Forests 79
Kate M. Detwiler
8 A Genetic Study of a Translocated Guenon: Cercopithecus mona on Grenada 99
K. Ann Horsburgh, Elizabeth Matisoo-Smith, Mary E. Glenn, and Keith J. Bensen
Part II Behavior
9 Diversity of Guenon Positional Behavior 113
W. Scott McGraw
10 Unique Behavior of the Mona Monkey (Cercopithecus mona): All-Male
Groups and Copulation Calls 133
Mary E. Glenn, Reiko Matsuda, and Keith J. Bensen
11 Group Fission in Red-tailed Monkeys (Cercopithecus ascanius) in Kibale National Park, Uganda 147
Tammy L. Windfelder and Jeremiah S. Lwanga
12 Interindividual Proximity and Surveillance of Associates in Comparative Perspective 161
Adrian Treves and Pascal Baguma
13 Why Vervet Monkeys (Cercopithecus aethiops) Live in Multimale Groups 173
Lynne A. Isbell, Dorothy L. Cheney, and Robert M. Seyfarth
14 When are there Influxes in Blue Monkey Groups? 189
Marina Cords
15 Costs and Benefits of Alternative Mating Strategies in Samango Monkey Males 203
Mairi C. Macleod, Caroline Ross, and Michael J. Lawes
16 Female Reproductive Endocrinology in Wild Blue Monkeys:
A Preliminary Assessment and Discussion of Potential Adaptive Functions 217
Karen Pazol, Ann A. Carlson, and Toni E. Ziegler
17` Grooming and Social Cohesion in Patas Monkeys and Other Guenons 233
Janice Chism and William Rogers
18 Development of Mother-Infant Relationships and Infant Behavior in
Wild Blue Monkeys (Cercopithecus mitis stuhlmanni) 245
Steffen Forster and Marina Cords
19 Influence of Foraging Adaptations on Play Activity in Red-tailed and
Blue Monkeys with Comparisons to Red Colobus Monkeys 273
Eric A. Worch
20 Effects of Natural and Sexual Selection on the Evolution of Guenon Loud Calls 289
Klaus Zuberbuhler
Part III Ecology
21 Resource Switching and Species Coexistence in Guenons: A Community Analysis of Dietary Flexibility 309
Joanna E. Lambert
22 Variation in the Diets of Cercopithecus Species: Differences within
Forests, among Forests, and across Species 325
Colin A. Chapman, Lauren J. Chapman, Marina Cords, Joel
Mwangi Gathua, Annie Gautier-Hion, Joanna E. Lambert,
Karyn Rode, Caroline E. G. Tutin, and Lee J. T. White
23 Diet of the Roloway Monkey, Cercopithecus diana roloway, in Bia National Park, Ghana 351
Sheila H. Curtin
Part IV Conservation
24 Conservation of Fragmented Populations of Cercopithecus mitis in
South Africa: the Role of Reintroduction, Corridors and Metapopulation Ecology 375
Michael J. Lawes
25 Assessing Extinction Risk in Cercopithecus Monkeys 393
Tharcisse Ukizintambara and Christophe Thebaud
26 Conservation of the Guenons: An Overview of Status, Threats, and Recommendations 411
Thomas M. Butynski
Epilogue 425
Marina Cords and Mary E. Glenn
Index 431
PREFACE
The African monkeys commonly referred to as the guenons make up the largest and most diverse primate group
on the African continent. Their pelage variations are arguably the greatest of any group of mammals in the world,
ranging from drab browns to multicolored experiments in body design. Some species live mainly in savanna-woodlands
while others live almost entirely in the upper canopy of rain forests. Social organization varies from monogamy to
unimale/multifemale groups to multimale/multifemale groups. The fascinating diversity of these monkeys has not been
lost on scientists interested in evolutionary biology. Field and laboratory researchers have begun to examine many
aspects of the biology of guenons, from molecular variation within and among species, to their roles in the ecosystems
of which they are a part.
Interestingly, and most likely because many of the species live in dense forests, the vast majority of research on
guenons has taken place during the past few decades. For example, the first and until recently last major scientific
symposium devoted to the study of wild guenons occurred at the XIX International Ethological Conference in 1985.
It was appropriately entitled the "Biology, Phylogeny, and Speciation of Forest Cercopithecines.
" Out of this symposium came the groundbreaking publication, A Primate Radiation: Evolutionary Biology of the African
Guenons. It became, like its symposium predecessor, the first and last book dedicated to guenon field research.
Almost all the major conceptual frameworks for our understanding of guenon biology were outlined within it.
Since that publication, however, new guenon species and subspecies have been discovered, previously unstudied species
have become the subject of long-term research projects, knowledge of the better-known species has greatly increased,
and original hypotheses have been challenged. It was for these reasons that we (Mary Glenn and Marina Cords) and
Annie Gautier-Hion decided to organize another scientific symposium focusing on guenon field research at the XVIII Congress
of the International Primatological Society in Adelaide, Australia in January 2001. We decided to invite as many
guenon biologists as possible, including those who had been studying guenons for decades, and most importantly,
those who were just starting out. So many of our colleagues agreed to present their work that we extended the symposium
to two days. The excitement generated by all the new research inevitably led to the proposition of producing a new guenon
book. Another call for contributions went out and we received an even greater response than before, a total of forty-nine
authors representing ten different countries. The results of all the authors' tremendous efforts are presented in this
volume.
The intensive research on a number of the member taxa has provided substantial grist for debate on the process of speciation
as well as the definition of species. This debate continues and is evidenced by the disagreement about how the taxonomic
organization of these monkeys should be arranged. Depending on which version you accept, the guenons include between
twenty-three and thirty-six species. There is an even greater variety of controversial subspecies within Cercopithecus,
Miopithecus, Allenopithecus, Erythrocebus, and Chlorocebus, the viability of the last four genera themselves being open
to debate. Inevitably the question of taxonomy arose while we were organizing this book. After much debate and consultation
with taxonomic experts (some of whom are also authors in this volume), we decided to use the consensus taxonomy developed
for guenons by Peter Grubb, Tom Butyoski, John Oates, Simon Bearder, Todd Disotell, Colin Groves, and Tom Struhsaker
at the IUCN/SSC Primate Specialist Group meeting held in Orlando, Florida in February 2000. Any authors who have deviated
from this taxonomy have explained their reasons in their respective chapters and generation.
We decided to follow the precedent of A Primate Radiation, as well as more recent field guides, to include within
the guenon group all monkeys in the genera Cercopithecus, Allenopithecus, Chlorocebus, Miopithecus, and Erythrocebus.
Over the years, the term guenon has been used to describe both this wider set of primate taxa as well as only those within
Cercopithecus. There is no clear precedent or distinct historical root for the usage of "guenon." Our usage of the term
is meant to provide a simple manner in which to refer to this group of closely related species and genera.
We organized the chapters into four general categories. Befitting any treatise on a diverse radiation of organisms,
the first section deals with evolutionary biology and biogeography. The chapters in this section readily demonstrate
the continuing challenge associated with understanding the complicated evolutionary relationships of guenons.
The second section is devoted to behavioral studies, and, as in A Primate Radiation, it is the largest. Our ever-increasing
appreciation of guenon behavioral diversity has led researchers to question many original ideas about their behavior,
including those related to male relationships and reproductive strategies, group structure and social organization,
and vocalizations. Investigations of ecological relationships make up the third section. As with the previous section,
many original concepts have been reassessed with the increase in single species studies conducted over many years in
multiple locations, as well as information about previously unstudied guenons. For example, dietary flexibility in some
species appears to be more common than previously thought. Finally, and most notably different from A Primate Radiation
is the fourth section devoted to conservation. The rate at which African primates, including the guenons, are becoming
threatened with extinction has increased dramatically over the past decade and a half. Contributions reflect both an
increasingly sophisticated understanding of extinction processes as well as the extreme urgency for immediate conservation
action with regard to many of the guenon taxa.
The juxtaposition of many new discoveries with the realization that some species or subspecies of guenons may not exist
by the time another book dedicated to guenon biology is published is sobering. A perusal of the chapters, however, is
cause for optimism. Some of the contributors to A Primate Radiation are included here. Their unflagging devotion to the
taxa is inspiring. In addition, some authors began their careers after the original guenon book was published.
Some of them are so new to the field that this is their first guenon publication. Most encouraging are the chapters by
African authors. The continued interest of so many people representing the past, present, and future of guenon biology
can only be a cause for hope and reaffirms the importance of studying these fascinating beings.
There are a number of people who deserve our gratitude. We wish to thank the contributing authors for their hard work,
diligence and excellent manuscripts. Many of the authors also provided comments and reviews for other chapters in the book,
for which we are grateful. We are indebted to the series editor, Russell Tuttle, for his meticulous copyediting and reviews,
hard work, support and patience. We also thank Thelma Rowell, Keith Bensen and several anonymous reviewers for their helpful
criticism and insightful comments on the manuscripts; our volume profited greatly from their suggestions, and we appreciate
the amount of time it took to review so many manuscripts. Special thanks must go to Alisha Clompus for allowing us to
publish her beautiful sketches, and to Tammy Smart for her assistance. Finally, this publication would not have been
possible without help and encouragement from Andrea Macaluso, Life Sciences Editor, and the many other staff members at
Kluwer Academic Plenum Publishers.
We close with a dedication of this volume to three biologists who have contributed greatly to our knowledge of guenon
biology. Two are senior researchers who did not prepare chapters for this volume. Still, their academic and research
progeny are well represented in our list of authors. Thelma Rowell was one of the first scientists to study guenons in the
wild and in captivity more than 30 years ago: her work inspired curiosity and questions, and her enthusiasm inspired a
next-generation of students and their research. Tom Strubsaker was similarly a pioneer in field studies of wild guenons,
working in multiple sites across the African continent. He has been a dedicated advocate for conservation of guenon habitat,
indeed for African forests in general. This book owes much, both directly and indirectly, to the inspiration of both Thelma
and Tom.
We also dedicate this volume to Joel Mwangi Gathua, who was just starting his research career. He received his first field
training as an undergraduate from Thelma Rowell, and went on to complete his Ph.D. studying the ecology of red-tailed monkeys
in a forest in his native Kenya. As a biologist from a guenon-habitat country, with a deep interest in understanding and
conserving forest organisms and ecosystems, he stands as an example of where future research must go. Mwangi died tragically y
oung, but we hope his example of dedication, care and energy will inspire other young African scholars especially for many
years to come.
Mary E. Glenn
Marina Cords
Arcata, California and New York, 2002
CONTRIBUTORS
Pascal Baguma
Makerere University Biological Field
Station, P.O. Box 409, Fort Portal,
Uganda
Keith J. Bensen
Windward Islands Research and
Education Foundation, 11 East Main
Street, Bayshore, NY 11706, USA
Thomas M. Butynski
Zoo Atlanta's Africa Biodiversity
Conservation Program, National
Museums of Kenya, P.O. Box 24434
Nairobi, Kenya
Paul J. Buzzard
Department of Anthropology, Columbia
University, New York, NY 10027, USA
Ann A. Carlson
Large Animal Research Group
Department of Zoology, University of
Cambridge, Downing Street, Cambridge
CB2 3EJ, UK
Colin A. Chapman
Department of Zoology, University of
Florida, Gainesville, FL 32611, USA
Lauren J. Chapman
Department of Zoology, University of
Florida, Gainesville, FL 32611, USA
Dorothy L. Cheney
Department of Biology, University of
Pennsylvania, Philadelphia, PA 19104,
USA
Janice Chism
Department of Biology, Winthrop
University, Rock Hill, SC 29733, USA
Marc Colyn
UMR 6552, CNRS - Universite de
Rennes 1, Station Biologique, 35380
Raimpont, France
Marina Cords
Department of Ecology, Evolution
and Environmental Biology,
Columbia University, New York, NY
10027, USA
Sheila H. Curtin
942 Shevlin Drive, El Cerrito, CA 94530,
USA
Pierre Deleporte
UMR 6552, CNRS - Universite de
Rennes 1, Station Biologique, 35380
Paimpont, France
Kate M. Detwiler
Department of Anthropology, New York
University, 25 Waverly Place, New York,
NY 10003, USA
Todd R. Disotell
Department of Anthropology,
New York University, 25 Waverly Place, New York,
NY 10003, USA
Steffen Forster
Department of Zoology, Technische
Universitat Braunschweig,
Braunschweig, Germany
Joel Mwangi Gathua
Formerly, Mammalogy Department,
National Museums of Kenya, Nairobi,
Kenya
Jean-Pierre Gautier
UMR 6552, CNRS - Universite de
Rennes 1, Station Biologique, 35380
Paimpont, France
Annie Gautier-Hion
UMR 6552, CNRS - Universite de
Rennes I, Station Biologique, 35380
Paimpont, France
Mary E. Glenn
Department of Anthropology,
Humboldt State University, Arcata, CA
95521, USA
K. Ann Horsburgh
Department of Anthropology, 701 East
Kirkwood Avenue, Indiana University,
Bloomington, IN 47405, USA
Lynne A. Isbell
Department of Anthropology, University
of California, Davis, CA 95616, USA
Beth A. Kaplin
Department of Environmental Studies,
Antioch New England Graduate
School, 40 Avon Street, Keene,
NH 03431, USA
Joanna E. Lambert
Department of Anthropology, University
of Oregon, Eugene, OR 97403, USA
Michael J. Lawes
School of Botany and Zoology, Forest
Biodiversity Programme, University of
Natal, Pietermaritzburg, South Africa
Jeremiah S. Lwanga
Makerere University Biological Field
Station, RO. Box 409, Fort Portal,
Uganda
Mairi C. Macleod
School of Life Sciences, University of
Surrey Roehampton, West Hill, London
SW15 3SN, UK
Elizabeth Matisoo-Smith
Department of Anthropology, The
University of Auckland, Private Bag
92019, Auckland, New Zealand
Reiko Matsuda
Department of Anthropology, Graduate
School and University Center of the City
University of New York, New York, NY
10016, USA
W. Scott McGraw
Department of Anthropology, The Ohio
State University, 1680 University Drive,
Mansfield, OH 44906, USA
Don J. Melnich
Center for Environmental Research and
Conservation, Columbia University, New
York, NY 10027, USA
Juan Carlos Morales
Center for Environmental Research and
Conservation, Columbia University, New
York, NY 10027, USA
Karen Pazol
Yerkes Regional Primate Research
Center, Emory University,
Lawrenceville, GA 30043, USA
Ryan L. Raaum
Department of Anthropology, New York
University, 25 Waverly Place, New York,
NY 10003, USA
Karyn Rode
Department of Zoology, University of
Florida, Gainesville, FL 32611, USA
William Rogers
Department of Biology, Winthrop
University, Rock Hill, SC 29733, USA
Caroline Ross
School of Life Sciences, University of
Surrey Roehampton, West Hill, London
SW15 3SN, UK
Robert M. Seyfarth
Department of Psychology, University of
Pennsylvania, Philadelphia, PA 19104,
USA
Christophe Thebaud
UMR 5552 CNRS/UPS, 13 Avenue du
Colonel Roche, BP 4072, F-31029,
Toulouse Cedex 4, France
Anthony J. Tosi
Department of Anthropology,
Columbia University, New York, NY
10027, USA
Adrian Treves
Center for Applied Biodiversity Science,
Conservation International, 65
Greenway Terrace, Princeton, NJ 08540,
USA
Caroline E.G. Tuhn
Centre International de Recherches
Medicales de Franceville, Gabon
Tharcisse Ukizintambara
School of Biological Sciences,
University of East Anglia,
Norwich NR4 7TJ, UK
Regine Vercauteren Drubbel
Universite Libre de Bruxelles -
Anthropologie, CP 192 - B 1050
Bruxelles, Belgium
Lee J.T White
Wildlife Conservation Society, 185th
Street and Southern Boulevard, Bronx,
NY 10460, USA
Tammy L. Windielder
Department of Anthropology, University
of Notre Dame, Notre Dame, IN 46556,
USA
Eric A. Worch
Education Department, University of
Michigan, Flint, M1 48502, USA
Toni E. Ziegler
Department of Psychology and
Wisconsin Regional Primate Research
Center, University of Wisconsin, 1223
Capitol Court, Madison, W1 53715, USA
Klaus Zuberbuhler
School of Psychology, University of St.
Andrews, St. Andrews, Fife KY16 9JU,
UK
WHERE TO ORDER
Andrea Macaluso, Editor
Kluwer Academic / Plenum Publishers
233 Spring Street
New York, NY 10013
Telephone: (212) 620-8007
Fax: (212) 463-0742
Email: amacaluso@wkap.com
Price: $139.00 (Hardbound), ISBN 0-306-47346-1
Posted Date: 03-25-03
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