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Bai Use in Forest Elephants (Loxodonta Africana Cyclotis): Ecology, Sociality & Risk

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Bai Use in Forest Elephants (Loxodonta Africana Cyclotis): Ecology, Sociality & Risk Bai Use in Forest Elephants (Loxodonta africana cyclotis): Ecology, Sociality & Risk Victoria L. Fishlock A thesis submitted for the degree of Doctor of Philosophy Psychology School of Natural Sciences University of Stirling August 2010 i Declaration I declare that the work undertaken and reported within this thesis is my own and has not been submitted in consideration of any other degree or award. Victoria Louise Fishlock i Abstract Forest elephant (Loxodonta africana cyclotis) sociality is relatively little-studied due to the difficulties of making direct observations in rainforests. In Central Africa elephants aggregate at large natural forest clearings known as bais, which have been postulated to offer social benefits in addition to nutritional resources. This thesis explores the role of these clearings as social arenas by examining bai use within three main themes; ecology, sociality and risk factors. Seasonal changes in elephant use of the Maya Nord bai (Republic of Congo) are described, along with the demography of the visiting population. Elephant visit rate was highly variable; the number of elephants using Maya Nord in an observation day ranged from 0 to 117 animals. This variability was unrelated to local resource availability and productivity suggesting that bai use occurs year round. Elephants in Odzala-Kokoua do not show high fidelity to a single clearing; 454 elephants were individually identified and re-sighted an average of 1.76 times (range 1-10) during the twelve month study period. Previous bai studies have yet to quantify how elephants associate with one another within the bai area. This study examines socio-spatial organisation and associate choice using two 2 measures of association within the 0.23 km bai area; aggregations (all elephants present in the clearing) and parties (elephants spatially co-ordinated in activity and movement) and distinguishes these from parties that range together (i.e. arrive and leave together). Social network analyses (SocProg) were used to describe inter- and intra-sexual multi-level organisation in the bai environment, and to illustrate the non-random nature of elephant ii aggregations and parties. Bais were shown to function as social arenas; female elephants showed active choice of certain associates and active avoidance of others when creating parties, whereas males were less discriminatory. Parties formed in the clearing (mean size= 3.93, SE= 0.186) were larger than ranging parties (mean size= 2.71, SE= 0.084) and elephants stayed for 50% longer in the clearing when they associated with individuals from outside their ranging party. Inter- and intra-sexual relationships were maintained within the clearing, and these are suggested to offer elephants essential opportunities for social learning. The patterning and nature of the relationships observed at the Maya Nord clearing indicates that forest elephants use a fission-fusion social structure similar to that of savannah elephants (Loxodonta africana africana); relationships are significantly structured by age- and sex- and underpinned by individual identity. Old experienced females hold key roles for forest elephants, and male relationships are superimposed on the network of female associations. Odzala-Kokoua elephants use bais to maintain their social relationships despite being highly sensitive to the anthropogenic risks involved in using these open areas. The results of this study suggest that forest and savannah elephants lie on the same social continuum, balancing social “pulls” to aggregate against the ecological “pushes” that force groups to fission. Previous models of savannah elephant sociality construct levels of association and social complexity upwards from the basic mother-calf unit (e.g. Wittemyer & Getz 2007). My results suggest that it may be more appropriate to consider elephant sociality and associations as in dynamic equilibrium between social and ecological iii influences acting at all levels of grouping, and to explicitly test how these underlie the opportunity costs that elephants are willing to pay in order to maintain social groupings. iv Acknowledgements No PhD student is an island, and I owe a great many people thanks for their support. My first call of gratitude is to my field team, who waded through swamps, carried in our equipment and food, and made it possible for me to spend a year at Maya Nord. They are; Ngounda “Esomali” Eric, Ngouma “la Joie” Fréderic, Mimiessi “Sitak” Rock and, most crucially, Andembo “Tranquillise-toi” Remy. Remy’s unfailing friendship, support and experience ensured the success of the project and my sanity, especially during the long hard hours when elephants didn’t show up, or appeared only to run away again. He taught me a great deal about Congo and Odzala, and I am deeply indebted. My original opportunity to work in the rainforests of Central Africa was brought about by Liz Rogers who, in doing so, quite literally changed my life. Thanks are due to her and my colleagues at Nouabalé-Ndoki, especially Thomas Breuer and Mireille Ndoundou-Hockemba for that first chance. Steve Blake was responsible for my choice of field site and his support, all the way from the Galapagos, got me through some tough times in the field. Mr Djoni-Djimbi Bourges always backed this project and it is through his permission and MEF-Congo that I was allowed to work in Odzala, despite challenging times for the Park. The PNOK, ECOFAC and WCS-Congo staff helped me get into, and stay in the field. Financial support for this study came from generous awards from the Leverhulme Trust, the Panton Trust, the Richard Hughes Scholarship and the University of Stirling. Various people have offered support, advice and friendship, both in Congo and elsewhere. Bryan Curran gave all of these generously and there isn’t enough gin in the world to repay the debt to the “full service organisation”, which is probably a good thing for his liver. Also in Central Africa, Phil Henschel helped me immensely in the early days and loaned a camera trap which the hyaenas avoided and then ate (sorry again about that), and Andrea Turkalo let me come and stay at elephant heaven and shared a wealth of knowledge. In Stirling, Peter Hancock, Kate Howie, Andy Hoyle, Clare Leadbetter and Giles Willoughby helped with computing and modelling traumas, and David Lusseau put me on the right track with SocProg. My family and friends have been incredibly supportive and understanding of my long absences and have helped me cope when it all seemed too much, mainly by making me think about things other than elephants! My parents and brother taught me to believe in myself and follow my passion, even when it meant disappearing to the depths of Congo. Although in fact my mother may well be partly to blame, since her last-ditch response to the endless “but why….?” I uttered as a child was, invariably, “elephants”. I like to think she was right. My longsuffering Dad heroically handled UK v logistics for me. Special thanks to Andy Hoyle (for maths help and Lego Indy therapy), Elizabeth Mackenzie, Louise Taylor, Kim Hockings, Michelle Klail, Sarah-Jane Vick, Betsy Herrelko (for being my “PA” when I disappeared to Kenya!), Ida-Maria Skavhaug (for living with me), Cat Macleod and all the other Stirling PhDs. Helen & Colin McMillan provided a welcome refuge from the office (and gin), and Mary-Ann Craig was simply always there. My supervisors here at Stirling have been, to put it bluntly, amazing. Christine Caldwell has let me talk “at” her endlessly and let me bounce my ideas into some sort of clarity, with great patience and good humour. Phyllis Lee has been simply extraordinary and her guidance and experience is written throughout these pages. Her joy in elephants and her wealth of knowledge have constantly comforted and inspired me. For the dubious honour of dedicating this thesis, I follow Rich Parnell’s example: like his gorillas, my study subjects have little need of such a thing, though they have given me much pleasure (and occasionally scared me witless) and I am incredibly privileged to have spent time with them. Instead, this thesis is for all those who know Africa’s “heart of darkness” is a place of vivid life, light and beauty, and who work so hard to keep it that way in a harsh timber- and ivory-hungry world. vi Table of Contents Declaration ......................................................................................................................... i Abstract ............................................................................................................................. ii Acknowledgements .......................................................................................................... v Table of Contents ............................................................................................................ vii Chapter 1 Introduction ................................................................................................ 2 1.1 Sociality: Ecological and Evolutionary Considerations ....................................... 2 1.2 The Challenge of Fission-Fusion Sociality .......................................................... 5 1.3 Elephants as a Study Species ............................................................................. 8 1.4 Studying Social Systems ................................................................................... 18 1.5 Thesis Aims and Objectives .............................................................................. 22 1.6 Thesis Outline ..................................................................................................
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