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Cercocebus Sanjei ) in the Udzungwa Mountains National Park, Tanzania

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Cercocebus Sanjei ) in the Udzungwa Mountains National Park, Tanzania 1 Social organisation, ecology and reproduction in the Sanje mangabey ( Cercocebus sanjei ) in the Udzungwa Mountains National Park, Tanzania A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science in Conservation Biology at Victoria University of Wellington By Kuruthumu Ally Mwamende School of Biological Sciences Victoria University of Wellington New Zealand 2009 1 2 Acknowledgments This study was funded by the New Zealand Aid for International Development (NZAID). Additional funding for field work was provided by School of biological sciences, Victoria University of Wellington (a research grant to Prof. Alan Dixson), and the Karori Wildlife Sanctuary (part of grant no. 24459/Award 1401 to Dr. Murray Williams). I would like to thank my supervisors Prof. Alan Dixson and Dr. Wayne Linklater, for their support in developing and shaping ideas for this project, before and during field and for their constructive editing during thesis writing. My supervisors also have encouraged and reassured me when I most needed it. I extend special thanks to Prof. Alan Dixson, my primary supervisor, for many useful advices such to ‘watch out-as things can get a little overwhelming’, for being a friend, and who, in his own way demanded my best. For all of this I extend my deepest thanks. For permission to work in the study area, I thank Tanzania National Parks Authority (TANAPA). The staffs of the Udzungwa Mountains National Park are thanked for their support, and help during fieldwork, especially Mr Matata Mushi, Chief Warden- in-charge, Joachim Tesha, Warden for protection, and Joram Mponjoli, Park Ecologist. Fieldwork would have not been possible without much logistical assistance of rangers from the Udzungwa Mountains National Park. Various people have contributed both knowingly and unknowingly to my study, and to all I owe a great debt. Thanks to my friends for comments on drafts of the thesis, including Alfan A. Rija and Lisa Bryant, Alfan also helped with data entry. Most data were collected by myself, between December 2007 and June 2008 with the help my assistants unless it stated otherwise. Throughout the entire 6.5 months of fieldwork, I was assisted by Ally Mihambo. Fridolin Mselewa, Sebastian Mvelege and Pius Mwenda provided valuable assistance in collecting ranging behaviour and feeding data. Mselewa, Mwenda and Mvelege also showed me the animal’s most used trails to make sure I don’t spend hours finding a way out in tangled vegetations; without their help the work of following the mangabeys would have been much harder and arduous task. Ruben Mwakisoma is thanked for help in tree identification in the field. In addition, Enock Peter and Eliatosha Mnguu assisted to monitor thermometer and rainfall gauge, arranged field supplies, and cook for us while working in the field. I am very thankful to all of these assistants. 2 3 I thank Mr Suleiman Haji of the National Herbarium of Tanzania (NHT) at the University of Dar es salaam for botanical expertise in plant identification. Several experts are thanked for statistical input, as indicated separately in the main chapters, including Nokuthaba Sibanda and Matthew Langen. Thanks also to Stephen Hartley for assistance with GIS and Cameron Jack for help in computer logistics. I also thank Patricia Stein for providing enough working space, and to Marry Murray for making sure we are ‘comfortable’ during writing phase. Alfan Rija, Fiona Hodge, Lisa Bryant, Mark Heath, Shruti Patel, Heather Murray and Chris Gibbons are thanked for their company, friendship and providing me with stimulating atmosphere and comfortable office. Special thanks to Prof. Dixson and family for their friendship, occasional visit invitations to their house and for making me feel at home in New Zealand. To Dr. Murray Williams, for his advices, and who pushed for funding and arranged travel for me to attend a conference to obtain the most needed experience, for reassurance and for being an interrogator at times. For all this, I extend my deepest thanks. I also thank the staff at Udzungwa Ecological Monitoring Centre (UEMC) for comfortable accommodation in Mang’ula, and permission to store equipment ‘for free’ while working in the field. The UEMC’s house attendants Shabani and Leonard are thanked for their help and hospitality, especially Leonard who kept the house clean, making it a comfortable place to stay after weekly trips from the forest. Thanks to researchers Drs. Andrew Marshall and Francesco Rovero, Trevor Jones, and Andrew Bowkett working in Udzungwa, for their support, and for sharing the forest work experience. Special thanks to Dr. Andrew Marshall, who introduced me, and for being an inspiration for me to study primates. Last but not least, I thank my husband, my family and friends, for their continued support in numerous ways, including my long time away from home, for their love when I was sick, and for friendship throughout my endeavour despite some of them not fully understanding ‘just what drove me to study ‘Nyani’. Finally, to Udzungwa ‘Ngolaga’ those endangered primates, for tolerating me to study their lives. Without their cooperation, this study would not have been feasible. 3 4 Abstract Understanding of the biology of threatened species is central to effective planning for sustainable management of wildlife. The Sanje mangabey, Cercocebus sanjei of Udzungwa Mountains National Park in Tanzania is under pressure from increasing human hunting, and habitat loss threatens its long-term survival. This thesis endeavoured to document the socio-biological and ecological aspects of this little- known species in a small habituated group of 62 individuals within the Mwanihana forest in Udzungwa Mountains National Park. I studied the social organisation, size and age-sex composition of one group by following and monitoring of the Sanje mangabeys over a six-month period. I also investigated the spatial-temporal interactions and associations between age and sex classes within a group. I assessed the Sanje mangabey’s ranging and movement patterns and monitored its reproductive behaviour by examining female sexual skin swellings and relationships between sexual swellings and sexual behaviour. Further, I investigated the species feeding habits and influences of seasonal parameters (rainfall and temperature) upon reproduction, feeding ecology and behaviour of this species. The Sanje mangabey social organisation is structured in a hierarchy characterised by dominance according to the sex and age of individuals. The study group was composed of individuals of all sexes and age classes; newborn babies, juveniles, sub adults and adults, males and females. Adult males were generally dominant over females and males of lower age. Males showed more agonistic behaviours to juveniles than did females and there were stable linear dominance hierarchies among individuals across the sexes (Landau’s dominance index, h’ =0.92, p =0.0281) and a high directional consistency index, DC =0.91) for both sexes. The linearity indices’ h’ were 0.84, P= 0.041, DC = 0.94 and 0.93, P = 0.0382, DC = 0.94) for males and females, respectively. Individuals spent about 8% on average of day grooming each other and the rate of grooming differed across and within sexes and age classes. There was a significant relationship between duration of the initiator’s first grooming episode and subsequent reciprocation by the receiver in all age classes ( P < 0.05). The independent effects of total grooming duration of the initiator also showed a significant difference and predicted total grooming duration of the recipient ( P = 4 5 0.0001). However, there was no evidence for grooming to be directed towards higher ranking Sanje mangabeys. The Sanje mangabey’s home range was 301 hectares and exhibited a mean daily range of 1760 metres within the Mwanihana forest. The range and movement patterns of this species differed significantly between seasons of the year. During the dry season (no rain) the group covered larger distances ( P = 0.001) between forest patches and moved significantly faster ( P = 0.001) than during wet (rain) season. The Sanje mangabey is omnivorous, feeding on plant material, invertebrates and vertebrates such as lizards. There were marked influences of temperature and rainfall on the feeding patterns. Mangabeys utilised the arboreal layer (>16m high) during cool and wet periods and were observed feeding mostly on the ground resources during hot and dry periods. The reproductive potential and social behaviour of the Sanje mangabey, however, did not vary with the different seasons of the year. Sanje mangabeys have a mean swelling cycle length of 32.1 ± 2.4 days. Individual females showed differences in swelling duration (F =12.43, P = 0.043) and no evidence of group variation in cycle length (H= 30.2, DF = 6, P < 0.05) was seen. Maximum swelling lasted for 4.4 ± 0.9 days on average and detumescence was 14.7 ± 3.9 days. The mean menses length was 3 ± 0.2 days (n = 3). Most cycling females, showed days with regular swellings of both sizes. The gestation length averaged 173 days and the interval between birth and resumption of the swelling cycle was observed to be 184 days. The sexual skin swellings appear to be sexually attractive signals of female reproductive condition. Although mating occurred in all stages of the female cycle, a peak was observed during maximum swelling size and breakdown. These findings illustrate the utility of sex swellings as signals used by males to determine reproductive condition.
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