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Social Organisation and Ecological Basis for Supergroup Formation In SOCIAL ORGANISATION AND ECOLOGICAL BASIS FOR SUPERGROUP FORMATION IN RWENZORI COLOBUS Alexandra Courtney Miller, BSc (Hons) This thesis is presented for the degree of Doctor of Philosophy of the University of Western Australia School of Human Sciences Anatomy and Human Biology 2019 THESIS DECLARATION I, Alexandra Courtney Miller, certify that: This thesis has been substantially accomplished during enrolment in the degree. This thesis does not contain material which has been accepted for the award of any other degree or diploma in my name, in any university or other tertiary institution. No part of this work will, in the future, be used in a submission in my name, for any other degree or diploma in any university or other tertiary institution without the prior approval of The University of Western Australia and where applicable, any partner institution responsible for the joint-award of this degree. This thesis does not contain any material previously published or written by another person, except where due reference has been made in the text. The work(s) are not in any way a violation or infringement of any copyright, trademark, patent, or other rights whatsoever of any person. The research involving animal data reported in this thesis was assessed and approved by The University of Western Australia Animal Ethics Committee. Approval #: RA/3/100/1468 The work described in this thesis was funded by ‘Research group: GRUETER,’ project grant number: 10301036, Primate Conservation Inc. and the Postgraduate Student Association of the University of Western Australia This thesis contains published work and/or work prepared for publication, some of which has been co-authored. Signature: Alexandra C Miller Date: 26/08/2019 II ABSTRACT Primates display a wide variety of social systems. A few primate species form “supergroups” which are shaped by a range of social and ecological selective pressures. Some supergroups of primates are socially organised into multilevel societies composed of multiple discrete social units nested within a larger social matrix. A characteristic of multilevel societies is that the higher levels can include hundreds of individuals. Among primates thus far, these societies have been observed in only a few taxa. In the high-altitude, mountainous forest of Nyungwe National Park, Rwanda, Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) form supergroups and were hypothesised to exhibit multilevel social organisation. This conjecture is based on similarities to Chinese snub-nosed monkeys in terms of large group size, montane habitat and seasonal consumption of lichen. Previous studies on the habituated supergroup have investigated the diet, activity and ranging patterns of the Rwenzori colobus. Here I present the first data on the ‘anatomy’ of a supergroup numbering 500+ individuals. This study aims to 1) determine the social organisation of the supergroup (multi-male, multi- female, vs multilevel), 2) examine the extent to which members of this group experience direct and indirect feeding competition, and 3) quantify the resource requirements of the supergroup, with a particular focus on the use of fallback foods during periods of preferred food scarcity. Rwenzori colobus were observed over a period of 13 months in 2016 and 2017. I extracted the social network structure from the time-stamped spatio-temporal distribution of passing individuals in a travelling progression identified to age-sex class. Core multi-male units (MMUs) with a mean of 1.73 adult males and 3.11 adult females, as well as one-male units, all-female units and bachelor units composed of adult and sub-adult males made up the substructure of the supergroup. In addition, proximity scans showed that adult males were in proximity to other adult males and sub-adult males more often than expected by chance, and III close-proximity resting clusters contained a mean of two adult-males. These results suggest that the Rwenzori colobus exhibit an internally sub-structured multilevel society with predominantly MMU core units as well as some one-male units (OMUs). This pattern differs from that observed in other nonhuman primates forming multilevel societies such as snub- nosed monkeys (Rhinopithecus spp.), geladas (Theropithecus gelada), Guinea baboons (Papio papio) and hamadryas baboons (Papio hamadryas). Due to the large group size of over 500 individuals, the Rwenzori colobus supergroup presents an opportunity to investigate the ecological preconditions underlying supergrouping. I used the patch depletion method to study feeding competition, comparing intake rate with movement rate in food patches. The colobus exhibited within-band scramble competition over young leaves, but not over mature leaves or fruit. Moreover, larger groups were able to occupy food patches for longer than smaller groups, indicating between-group contest for food patches. These findings suggest it may be the lack of competition for high-quality mature leaves, or the abundant lichen, that allows the formation of supergroups. Lastly, I investigated how the Rwenzori colobus supergroup responds to periods of resource scarcity. Specifically, I compared preferred food availability with consumption of potential fallback foods. Fruticose lichen (Usnea sp.) contributed >50% of the diet in months when preferred foods were less available, suggesting that these lichens constitute a fallback food for Rwenzori colobus supergroups. However, in a number of other montane forests in Burundi and Tanzania with similar tree species as well as fruticose lichen, Angolan colobus form small groups suggesting that lichen and high-quality leaves may create a resource base necessary to support colobus supergroups, but factors such as forest size, fragmentation, degradation and hunting by humans may impact group sizes. IV Table of Contents THESIS DECLARATION II ABSTRACT III CONTENTS V LIST OF FIGURES X LIST OF TABLES XII ACKNOWLEDGEMENTS XIV AUTHORSHIP DECLARATION XVIII CHAPTER ONE: General Introduction........................................................................ 1 Social organisation of colobines............................................................................................ 2 Angolan colobus................................................................................................................... 3 High elevation colobines....................................................................................................... 4 Multilevel societies: Hierarchical structuring........................................................................ 5 Case study: Hamadryas baboons.................................................................................... 6 Case study: Geladas....................................................................................................... 7 Case study: Snub-nosed monkeys.................................................................................. 7 Evolution of multilevel societies........................................................................................... 8 Bachelor threat............................................................................................................... 9 Fission-fusion.............................................................................................................. 10 Cultural transmission................................................................................................... 12 Ecological conditions needed for supergroup formation..................................................... 12 Thesis objectives and organisation...................................................................................... 15 Project significance............................................................................................................. 16 V CHAPTER TWO: Structure of a supergroup: A multilevel society containing multi-male units.............................................................................................................. 18 Abstract.............................................................................................................................. 19 Introduction....................................................................................................................... 19 Methods.............................................................................................................................. 24 Data collection............................................................................................................. 24 Progressions................................................................................................................. 25 Proximity scans............................................................................................................ 25 Data analysis........................................................................................................................ 26 Social network data...................................................................................................... 26 Selecting an optimal window size................................................................................ 27 Proximity scans: Nearest neighbour............................................................................. 30 Proximity scans: Close-proximity resting.................................................................... 30 Results................................................................................................................................ 31
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