Table Of ContentTHE REPRODUCTIVE TRAJECTORIES OF BACHELOR GELADAS
by
David J. Pappano
A dissertation submitted in partial fulfillment
of the requirements for the degree of
Doctor of Philosophy
(Anthropology)
in The University of Michigan
2013
Doctoral Committee:
Professor Jacinta C. Beehner, Chair
Professor Abigail W. Bigham
Professor John C. Mitani
Professor Elizabeth A. Tibbetts
© David J. Pappano
All rights reserved
2013
DEDICATION
For my parents, Teresa and Donald Pappano
For indulging a child’s curiosity in bats, snakes, and other animals
described in Golden Guides to Nature. Most parents would have just read
fairy tales to their children.
And my best friend and partner, Ashley
For your patience, kindness, and love in years past and those yet to come.
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ACKNOWLEDGEMENTS
Over the past six years, I have been fortunate to have the support of friends,
family, colleagues, and funding agencies that have made this dissertation possible. I am
grateful for their input at all levels of this endeavor—from conception to completion. My
time at the University of Michigan has been among the most formative and memorable
experiences of my life, and I hope that many of the friends and colleagues I have met
over these past years will continue to be a part of my life in the years to come.
This research would not have been possible without the generous support of
several government and non-profit funding agencies. A Graduate Research Fellowship
from the National Science Foundation and the James Spuhler Fellowship through the
Department of Anthropology largely funded my studies at Michigan. I am also grateful to
the National Science Foundation for support through a Doctoral Dissertation
Improvement Grant. The U.S. Fulbright Program and a Leakey Foundation General
Research Grant supported my fieldwork. I also thank the Rackham Graduate School for
additional support while writing the final parts of this dissertation.
My dissertation chair, Dr. Jacinta Beehner, deserves much praise for my
research trajectory. Jacinta has shown me how to develop a project and see it to
completion all while balancing her duties as director of the Core Assay Facility, co-
director of the University of Michigan Gelada Research Project, and mother to her two
small children. I met her and Dr. Thore Bergman the last week of December 2006 at
Princeton University. I had recently graduated college and was teaching high school
biology in Connecticut. They had just started their jobs at the University of Michigan and
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were preparing for a trip to Ethiopia in January to set up the gelada project. They needed
graduate students to study geladas in the formative years of the project, and I was
applying to fulfill that need. Against Thore’s warnings that “ice hockey goalies are crazy”
and probably her better judgment as well, she took a risk on me as one of their first
graduate students. For that, I am grateful.
I am also thankful for the guidance of my committee: Dr. John Mitani, Dr. Abigail
Bingham, and Dr. Elizabeth Tibbetts. I had the opportunity to teach two classes under
John while at the University of Michigan. In this lecture hall, John taught me how to
teach through splicing information and humor with thought-provoking questions. In my
early years of graduate school, I would attend the Behavioral Ecology Group organized
in part by Liz Tibbetts. This seminar provided me with weekly reminders that behavioral
ecology did not start and end with geladas (or primates for that matter). To Abby—the
only member of my committee that has fewer years at Michigan than I do—I am happy
to have had your input in these final steps of my dissertation. Finally, while he was not
on my committee, I am indebted to Thore Bergman for giving me the opportunity to
conduct research on the geladas of the Simien Mountains.
I owe a debt of gratitude to my friends at the University of Michigan and in
Ethiopia. My cohort in biological anthropology—Zachary Cofran, Dana Begun, and
Caroline VanSickle—have made this journey special. Also, to the newly minted Drs.
Noah Snyder-Mackler and Eila Roberts, your culinary skills in Ethiopia and help with
analyses at home have kept me sane. To the upstarts of the project—Marcela Benitez,
Elizabeth Johnson, Morgan Gustison, and Caitlin Barale—I wish the best of luck in their
future projects. Finally, to the camp managers and Ethiopian field workers of the gelada
project—Clay Wilton, Julie Jarvey, and Ambaye Fanta—your tireless efforts keeping the
campsite organized and functioning were greatly appreciated.
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Of the full-time project assistants, Esheti Jejaw deserves special recognition for
his role in this project. I met Esheti in 2008 during my first summer in Ethiopia. I hired
Esheti to be a full-time bachelor expert, which required him to find and name bachelors
that were associating with the project’s main study band. Because neither of us could
speak each other’s native language very well, we accomplished our goals that first
summer mostly through miming gelada behavior and speaking a pigeon “Amharenglish”.
By the time I returned my second summer, Esheti was using a Palm pilot to record data
and had begun to collect ad libitum data on dominance interactions and grooming within
bachelor groups. During my long haul of fieldwork in 2010-11, Esheti’s training came full
circle. He collected fecal samples for genetic and hormonal data, and was recording
focal animal samples everyday. Even though our communication has converged on a
mutual understanding of many Amharic-English phrases, I don’t know if I will ever be
able to convey just how grateful I am for his help in completing this work.
To my mother and father that fostered my love of science at an early age, this is
for your countless hours spent with me at the library or watching PBS Nature programs
at night. And to my brothers and sister, thank you for understanding in planning holidays
and events around my field schedule. Lastly, this thesis is for my best friend and partner
of the last seven years, Ashley. She agreed to marry me, now if I can only persuade her
to proofread this thesis.
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TABLE OF CONTENTS
Dedication .......................................................................................................................... ii
Acknowledgements ........................................................................................................... iii
List of Tables .................................................................................................................... vii
List of Figures ................................................................................................................. viii
Abstract ............................................................................................................................. ix
Chapter One: Introduction ................................................................................................. 1
Chapter Two: Testing extraction and storage parameters for a fecal hormone method . 14
Chapter Three: Demography and social behavior of bachelor geladas living in all-male
groups ............................................................................................................................. 32
Chapter Four: Social bonds of bachelor geladas living in all-male groups ..................... 59
Chapter Five: Social ‘predators’ within a multilevel primate society ............................... 88
Chapter Six: Testosterone and the reproductive trajectories of male geladas ............. 109
Chapter Seven: Conclusions ........................................................................................ 138
References .................................................................................................................... 143
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LIST OF TABLES
Table 2.1. A list of experiments conducted under controlled laboratory settings ............ 28
Table 3.1. A description of gelada male age classes ...................................................... 51
Table 3.2. Demography of all-male groups.. ................................................................... 52
Table 3.3. Stable periods of all-male groups. ................................................................. 53
Table 4.1. A summary of mixed models used in social bond analysis. ........................... 81
Table 5.1. Mixed model describing within-unit spacing (‘neighbor’ scan) ..................... 103
Table 5.2. Mixed model describing within-unit spacing (‘unit-spread’ scan) ................. 104
Table 5.3. Mixed model describing between-unit spacing (‘neighbor’ scan) ................. 105
Table 5.4. Mixed model describing between-unit spacing (‘unit-spread’ scan) ............. 106
Table 6.1. A summary of mixed models used in testosterone analysis. ....................... 132
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LIST OF FIGURES
Figure 2.1. Ambient temperature manipulation experiment results. ............................... 29
Figure 2.2. Freeze/thaw experiment results .................................................................... 30
Figure 2.3. Solution manipulation experiment results ..................................................... 31
Figure 3.1. Age class distribution for all bachelor geladas. ............................................. 54
Figure 3.2. Relatedness, age, and coresidence within an all-male group.. .................... 55
Figure 3.3. Elo-ratings of bachelors compared to adult females ..................................... 56
Figure 3.4. Elo-ratings for each bachelor male age class ............................................... 57
Figure 3.5. Comparison of grooming between bachelors and adult females .................. 58
Figure 4.1. Histograms of each social index and log-transformations ............................ 82
Figure 4.2. Scatterplot of composite sociality index (CSI) .............................................. 83
Figure 4.3. Relatedness, grooming, and proximity ......................................................... 84
Figure 4.4. Scatterplot of grooming index ....................................................................... 85
Figure 4.5. Scatterplot of proximity index ........................................................................ 86
Figure 4.6. Grooming equality index and bond strength ................................................. 87
Figure 5.1. Regression of bachelors and total number of geladas within a herd .......... 107
Figure 5.2. Effect of bachelor proximity on reproductive unit spread ............................ 108
Figure 6.1. Seasonal distribution of takeovers and aggression .................................... 133
Figure 6.2. Status and testosterone levels. ................................................................... 134
Figure 6.3. Testosterone levels during months of increased takeovers ........................ 135
Figure 6.4. Testosterone levels of bachelors ................................................................ 136
Figure 6.5. Testosterone levels and trajectories of “winners” and “losers” ................... 137
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ABSTRACT
Sexual selection strongly influences patterns of male behavior. Fertilizations
cannot be shared, therefore male-male relationships tend to be agonistic rather than
affiliative. Despite this fact, males in many species form all-male groups that vary in
composition and stability. All-male groups have been observed in many mammalian taxa
including cetaceans, ungulates, and primates. Because few studies have examined
these groups directly, we know little of how they form, the social relationships between
the males that comprise them, or if such relationships influence future reproductive
success. Geladas (Theropithecus gelada) are an Old World monkey species where
males disperse from their natal units and join all-male groups prior to acquiring
reproductive access to females. In this study, I examine the behavioral, hormonal, and
genetic factors that shape the reproductive trajectories of bachelor geladas living in all-
male groups.
This study has three major components. First, I address what factors, such as
age or kinship, influence the composition of all-male groups. I demonstrate that
relatedness influences all-male group formation, although only few pairs of males were
considered close kin. Second, I ask how bachelors interact within their groups. I show
that bachelors form stratified social bonds within their all-male groups. These bonds
were strongest between males that were closely related and similar in age. Third, I
attempt to answer how bachelors become dominant “leader” males. Males that
eventually became dominant had higher testosterone levels during the bachelor period. I
draw on extended field-based observations of wild geladas (2006-2011) as well as
hormonal and genetic data to answer my study questions. These findings represent the
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