Captive Environmental Influences on Behavior in Zoo Drills and Mandrills (Mandrillus), a Threatened Genus of Primate
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Portland State University PDXScholar Dissertations and Theses Dissertations and Theses 1996 Captive Environmental Influences on Behavior in Zoo Drills and Mandrills (Mandrillus), a Threatened Genus of Primate Erik Terdal Portland State University Follow this and additional works at: https://pdxscholar.library.pdx.edu/open_access_etds Part of the Biology Commons, Environmental Monitoring Commons, and the Zoology Commons Let us know how access to this document benefits ou.y Recommended Citation Terdal, Erik, "Captive Environmental Influences on Behavior in Zoo Drills and Mandrills (Mandrillus), a Threatened Genus of Primate" (1996). Dissertations and Theses. Paper 3014. https://doi.org/10.15760/etd.3009 This Dissertation is brought to you for free and open access. It has been accepted for inclusion in Dissertations and Theses by an authorized administrator of PDXScholar. Please contact us if we can make this document more accessible: [email protected]. CAPTIVE ENVIRONMENTAL INFLUENCES ON BEHAVIOR IN ZOO DRILLS AND MANDRILLS (MANDRILLUS ), A THREATENED GENUS OF PRIMATE by ERIKTERDAL A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in ENVIRONMENTAL SCIENCES AND RESOURCES: BIOLOGY Portland State University 1996 DISSERTATION APPROVAL The abstract and dissertation of Erik Terdal for the Doctor of Philosophy in Environmental Sciences and Resources: Biology were presented November 22,1995, and accepted by the dissertation committee and the doctoral program. COMMITTEE APPROVALS: orah Duffield Leonard Simpson n Dash , Richard Thortfs Marc Feldesman, Representative of the Office i ofof GraduateGraduate StudiesStudies /] DOCTORAL PROGRAM APPROVAL: Larry_arry Cra^wshaw, Director Environmental Sciences and Resources Ph.D. Program ACCEPTED FOR PORTLAND STATE UNIVERSITY BY THE LIBRARY ABSTRACT An abstract of the dissertation of Erik Terdal for the Doctor of Philosophy in Environmental Sciences and Resources: Biology presented November 22, 1995. Title: Captive Environmental Influences on Behavior in Zoo Drills and Mandrills ( Mandrillus), a Threatened Genus of Primate Drills (Mandrillus leucophaeus) are an endangered species of African monkey (Cercopithecidae), and their sole congener the mandrill (Mandrillus sphinx) is vulnerable to extinction. Both species are threatened in the wild by deforestation and hunting. Drills have a poor record of captive reproduction. Many individuals appear to have behavioral deficiencies which interfere with reproduction. Thus, the zoo population of drills does not serve as a “hedge” against the species’ total extinction: drills are endangered in captivity as well as in the wild. Mandrills, by contrast, reproduce well in captivity. Information on the behavior of mandrills in captivity may help zoo managers improve husbandry for both species. The intent of this research was to study the relationship between aspects of the captive environment and behaviors which lead to reproduction. A review of the literature on both drills and mandrills, in the wild and captivity, was used to suggest “essential characteristics” of the captive environment that may encourage animals of both species to engage in natural, active behaviors, to form cohesive dyads with opposite-sex adults, to develop affiliative bonds, and to engage in sexual behavior. Sixty-two drills and mandrills in 14 groups in the U.S. and Germany were studied with behavior sampling methods, using the Drill Species Survival Plan ethogram. Data were analyzed by multiple regression using transformed variables. No over-all species differences in behavior were found. Results suggested that two factors promoted natural activity: 1) environmental enrichment which provides positive reinforcement for active behaviors, and 2) an affiliative husbandry style by the animal’s keepers. Active animals were more likely to engage in social behaviors leading to copulation. Many of the non-reproducing drills appeared to be “passive and withdrawn,” and shared a constellation of signs that appeared to be analogous to human clinical depression. An etiological model for “passive and withdrawn” Mandrillus, based on biobehavioral theories of human depressive disorders, was developed to link early rearing conditions and environmental enrichment. This model was then used to develop a historic- demographic hypothesis for why mandrills have historically had greater reproductive success than drills in zoos. TABLE OF CONTENTS PAGE ACKNOWLEDGMENTS....................................................................................v LIST OF TABLES.............................................................................................viii LIST OF FIGURES............................................................................................ xi CHAPTER I INTRODUCTION..........................................................................1 Relating Captive Environment to Behavior and Reproduction ............................. 1 II LITERATURE REVIEW.............................................................. 4 History ............................................................................4 Taxonomy ....................................................................... 6 Distribution Status in the W ild ....................................................... 12 Habitat Destruction Hunting Status in Captivity ....................................................... 17 Why are Drills Reproducing Poorly in Captivity? Theory in Zoo Biology Mandrillus in the W ild ................................................ 24 Group Size and Composition Spatial Relationships Polyspecific Associations Habitat Structure Foraging and Feeding Humans Mandrillus in Captivity 34 Mandrills at CIRMF Development of Hypotheses Studies on Mandrillus in Zoos Exhibits and Husbandry Activity, Cohesion, Affiliative Bonding and Sexual Behavior Summary of Zoo Studies Relationg to Development of my Hypotheses III METHODS.........................................................................44 Mandrillus Groups: Composition, Exhibits and Husbandry ............................................44 Criteria used to Define Age Classes Exhibit and Husbandry Variables Behavioral Data .......................................................48 Ethogram Data Collection Methods ....................................... 57 Data Analysis .......................................................... 60 Proportion of Time Social Behavior Hourly Rates Screening Data Multiple Regression Analyses Statistical Procedures used for Visualization of Data IV RESULTS............................................................................69 Species Differences ................................................ 69 Hypotheses ............................................................. 71 Hypothesis One Hypothesis Two Hypothesis Three Hypothesis Four Overall Summary of Results V DISCUSSION OF RESULTS 92 Species Differences 92 Species Differences in Captive Environments Discussion of Results of Hypothesis Tests 96 Hypothesis One Hypotheses Two, Three and Four Proposed Experiments Two Examples of Reproductively Successful Drill Groups .. 107 Environments Activity Budgets Cohesion and Affiliative Bonding Sexual Behavior VI A PROPOSED MODEL TO EXPLAIN THE RESULTS . 120 A Constellation of Signs 120 Theory in Zoo Biology Revisited Towards a New Model The Human Model of Primate Psychopathology .124 Rationale for Using Humans as a Model Species for a Non-Human Primate Human Depressive Disorders .................................126 Effect of Early Rearing Environment on Behavior andNeurodevelopment in Laboratory Macaques ...............................131 A Proposed Experiment to Evaluate the Suggested Model ....................................134 A Historic-Demographic Hypothesis for the Species Difference in Captive Mandrillus Reproductive Success .............. 136 Conclusion.................................................................140 REFERENCES...........................................................................141 APPENDIX.................................................................................155 ACKNOWLEDGMENTS Scores of people in the U.S. and Germany helped me with this study. Even though I cannot thank them all by name, I appreciate all of their help. Jill Mellen (Metro Washington Park Zoo, Portland, OR, USA) has taught me since 1990 about mandrills and how to study them, and about all aspects of behavioral and zoo biology. Elizabeth Gadsby and Peter Jenkins (Drill Rehabilitation and Breeding Center, Calabar, Nigeria) were the people who led me to expand my interest in captive mandrills to drills, both in zoos and in Africa. At the Metro Washington Park Zoo, Dennis Pate, Rick Horton and Jan Baker helped me with planning the study and arranging the travel. Many people have helped me during my visits to several zoos in the U.S. since 1992. I especially wish to thank the dozens of keepers who have taught me so much about Mandrillus behavior and husbandry. Cathleen Cox (Los Angeles Zoo, Los Angeles, CA, USA) introduced me to drills and helped me to learn the Drill Species Survival Plan ethogram. Ed and Tammie Bettinger invited me to stay in their home when I visited Tulsa Zoo (Tulsa, OK, USA). In Atlanta, Gail and Nevin Lash invited me to stay in their home. I thank Debra Forthman (Zoo Atlanta, Atlanta, GA, USA) for making those arrangements. Zoo Atlanta’s curator, C. Dietrich Schaaf, shared with me his extensive knowledge about drills both in zoos and in Africa. Gail