Primate Society of Great BritainNo. 107JUNE 2012

THE PRIMATE SOCIETY OF GREAT BRITAIN� 5HJLVWHUHG�&KDULW\�1R�������� Officers: President: Prof. Kim Bard (10) (Centre for the Study of Emotion, Department of Psychology, University of Portsmouth, King Henry Building, King Henry I Street, Portsmouth, Hampshire, PO1 2DY) Hon. Secretary: Dr Sarah Elton (10) (Functional Morphology and Evolution Unit, Hull York Medical School, The University of York, Heslington, York, YO10 5DD) Hon. Treasurer Dr Clare Cunningham (Department of Psychology, School of Social and Health Sciences, University of Abertay, Kydd Building (Level 5), Bell Street, Dundee, DD1 1HG) Council Members: Mr G. Banes (Cambridge) – Student Representative (11) Dr S. Cheyne (Oxford University) (11) Dr D. Custance (Goldsmiths) (11) Dr G. Donati (Oxford Brookes) (12) Dr G. Forrester (Westminster) (12) Dr T. Humle (Kent) (12) Dr R. Kendal (Durham) – Research Working Party Convenor (10) Dr J. Lehmann (Roehampton) (10) Dr B. Majolo (Lincoln) (11) Dr S. O’Hara (Salford) (12) Dr C. Schwitzer (Bristol Zoo) (10) Dr S. Shultz (Oxford University) (11) Dr A. Smith Anglia Ruskin) – Meetings Officer (08) Dr S. Thorpe (Birmingham) (10) Convenors of Working Parties: Dr C. Harcourt (Dept of Veterinary Clinical Science, Leahurst, Chester High Road, Neston, Wirral, CH64 7TE, UK) - Convenor, Conservation Working Party Dr S. Hill (North of England Zoological Society, Chester Zoo, Caughall Road, Upton-by-Chester, Chester, CH2 1LH) - Convenor, Captive Care Working Party Assistant Members of Council: Dr M. Blanchard (Liverpool) - official Society archivist Dr S. Evans (Du Mond Conservancy, c/o Monkey Jungle Inc., PO Box 246, Miami, Florida 33170, USA) - for US membership Dr T.C. Rae (Roehampton) - Editor, 3ULPDWH�(\H�Mr C. Rosen MBE (IPPL) - Financial Advisor Honorary Members: Born Free Foundation Knowsley Safari Park Honorary Auditors: Messrs Morris & Co., Chester Subscription Rates: Annual subscription rates (send cheques and sterling drafts to the Treasurer): Full members and Associate members: Payment by Banker's Order £25.00 Payment by cheque, postal order, cash, credit card £27.50 Undergraduate and postgraduate student membership £15.00 (Membership of P.S.G.B. includes 3ULPDWH�(\H and supplements) Institutions wishing to receive 3ULPDWH�(\H�and supplements only: Annual subscription £30.00 Overseas subscriptions may be paid at longer intervals by arrangement with the Treasurer

EDITORIAL A recent, well-publicised permanent ban from keeping animals, imposed on a Midlands couple after they sold a young marmoset riddled with disease from neglect, has focused media attention again on the topic of keeping primates as pets. A bill has now been introduced to Parliament with the intent of making it illegal to own primates as pets in the UK (although the government has argued that existing regulation is sufficient). I was made aware of the issue rather forcefully during my term as PSGB Information Officer; one of the first pieces of correspondence I received in the role was from a couple looking for information on keeping callitrichids, as they were intending to set up a commercial monkey farm abroad. Having only really dealt with skeletal remains of primates, rather than actual living creatures, I was unprepared to speak for the Society on the issue; you can’t imagine my relief when I discovered that a pamphlet on the topic had been produced already. But pamphlets were already outdated as a means of efficient information dissemination even then. Now, in keeping with spirit of the age, a new website has been launched to provide up-to-date, scientific captive care advice (as reported in this issue). Sponsored by the University of Stirling, the National Centre for the Replacement, Refinement and Reduction of Animals in Research and our own PSGB Captive Care Working Party, marmosetcare.com is an accessible, well-illustrated and, perhaps most importantly, accurate source of information on all aspects of the care and well-being of these popular platyrrhines, as well as being a welcome addition to the discussion on keeping primates as pets. As primatologists, we understand more than anyone the appeal; after all, we like primates so much that we dedicated our professional lives to their study, care and/or conservation. But it is because we know so much about our primate relatives that we don’t keep them as pets ourselves. We know that they are wild animals, that most have evolved to live in groups, that they are, overwhelmingly, tropical organisms that can fare poorly in temperate climes. Although there are undoubtedly responsible owners, I would wager that many, if not most, are unaware of the basic biology of the animals in their care. It is hoped that this new resource will help people to understand the needs of the members of our order and perhaps persuade some to forgo ownership. And, finally, thanks to everyone who pointed out the error in the last editorial – to clarify, David Chivers was awarded the Occasional Medal for Conservation at the Winter 2011 meeting. Find a report of those proceedings, and much more, below.

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The articles and abstracts included in Primate Eye are not for citation or quotation without permission of the authors. The deadline for the next issue of Primate Eye is 15th September 2012. Items (manuscript or electronic in any standard format) for future issues should be sent to: Todd C. Rae Editor, Primate Eye Centre for Research in Evolutionary & Environmental Anthropology Department of Life Sciences Whitelands College The University of Roehampton Holybourne Avenue London SW15 4JD Email: editor@psgb.org Tel: UK (0)20 8392 3726 Fax: UK (0)20 8392 3610 PSGB correspondence unrelated to Primate Eye should be addressed to the Hon. Secretary. Notification of change of address should be sent to the Membership Secretary. The PSGB WebSite can be found at <www.psgb.org>

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REPORT: PSGB Winter Meeting 2011

Gardeners of the Forest: Primate Ecology and Forest Conservation 1st December 2011

Clifton Pavilion, Bristol Zoo Gardens The PSGB winter meeting of 2011 was held in conjunction with the Bristol Conservation and Science Foundation at Bristol Zoo Gardens. The theme of the meeting was ‘Gardeners of the Forest: Primate Ecology and Forest Conservation’, to tie in with the UN’s International Year of Forests. Successfully organised by Dr Christoph Schwitzer, Dr Sue Dow and Charlotte Bryant, the meeting consisted of 11 invited speakers and 17 posters. The day got off to an excellent start with the provision of coffee and pastries. There was time to mingle, collect a bag full of information about the symposium and get a first look at the posters. The symposium was introduced by Bryan Carroll, the director of Bristol Zoo. Ian Redmond then set the scene for the rest of the day’s talks, describing the various ways in which primates interact with their forest environment and their important role as gardeners. Next up was Jörg Ganzhorn who gave a controversial talk suggesting that lemurs may not play as important a role as seed dispersers as has previously been assumed. Helen Chatterjee then gave an interesting talk about the conservation of primates in China, using ecological niche modelling to investigate the distribution of, among others, the critically endangered Hainan gibbon. A very entertaining account of gibbon seed dispersal was given by David Chivers who was later duly awarded the Occasional Medal for Conservation. Joining David on the podium for the presentation was David Windmill who was awarded the Occasional Medal for Special Contributions to Primatology.

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David Chivers (r) receiving the Occasional Medal for Conservation from Hilary Box (c) and PSGB President Kim Bard (l).

David Windmill (c) receiving the Occasional Medal for Special Contributions to Primatology from Klaus Zuberbühler (r) and

PSGB President Kim Bard (l).

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The Napier Medal for an outstanding PhD thesis was awarded to Lauren Brent from the University of Roehampton. Unfortunately, Lauren was unable to attend to receive her award but Todd C. Rae was more than happy to step up and proudly collect the award on her behalf.

Napier Medal winner Lauren Brent A delicious buffet lunch was served, and people were given the chance to have a look around the zoo and enjoy the traditional English weather: pouring rain. Crowds gathered around the gorilla enclosure to see the tiny new infant’s foot – there must have been more of him somewhere! The first afternoon session predominantly focused on seed dispersal. Attendants were interested to hear about the importance of orangutans (Susan Cheyne), Amazonian frugivores (Caros Peres) and lemurs (Onja Razafindratsima) for seed dispersal, and the anthropogenic factors that threaten the survival of these species. On a different theme, Amanda Korstjens gave an engaging introduction to time budget models and their application. Tea and scones were provided to boost energy levels before the final session, and there was another opportunity to look at the posters and chat to the researchers who were displaying them. The final session focused on chimpanzees with a talk about their ability to remember the location of food in space and time by Karline Janmaat, and their role as seed dispersers in Rwanda by Nicole Gross-Camp. Next up was

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Vernon Reynolds who gave a sobering talk about the threats currently facing the chimpanzees of the Budongo Forest, ranging from the loss of the raffia palm trees to the possible construction of an oil pipeline. Despite the somewhat bleak subject of the talk, the subsequent discussions between Vernon Reynolds and members of the audience left us feeling that there may still be hope for the chimpanzees. A brief discussion by Bryan Carroll, Neil Maddison, Christoph Schwitzer and Ian Redmond and the customary raffle concluded what had been an extremely interesting and enjoyable symposium in a fantastic location. Charlotte Carne University of Roehampton

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PSGB Spring Meeting 2012

Organised by: Mark Kingston Jones, Aspinall Foundation

& Tatyana Humle, University of Kent

17th – 18th April 2012

Abstracts for oral presentations INVITED SPEAKER: Training and the Development of Survival -Critical Behaviors in Captive-Born Golden Lion Tamarins Beck, B. Smithsonian Conservation Biology Institute, Drake University and Iowa State University Email: benbeck21@gmail.com Between 1983 and 2003, 149 captive-born (CB) golden lion tamarins (Leontopithecus r. rosalia) (GLTs), were reintroduced to the wild, and 42 wild-born (WB) GLTs were translocated from imperiled forest patches to a protected area. Before reintroduction, some of the CB GLTs were trained intensively in cages, some were allowed to range freely in small wooded areas in zoos, and some received no pre-release behavioral preparation whatsoever. There was no significant difference in post-release survival under these three pre-release conditions. Post-release survival was enhanced by intensive post-release support, and was even higher among infants born in the wild to reintroduced parents. Translocated GLTs had the highest survival rates. Today, there are about 1,600 GLTs living in the wild, of which about 500 are descended from reintroduced CBs and 200 from translocated WBs. Both subpopulations are being monitored and managed as parts of a metapopulation. For this species, intensive post-release support is the most important factor in survival of reintroduced CBs, and translocation of WBs is more effective than reintroduction of CBs for establishing new subpopulations. More research is needed to understand why pre-release training is reported to enhance reintroduction survival in some species, but not in others. Growth and development in the genus Pan: a life history approach Brimacombe, C. University of Sheffield Email: prq09cb@sheffield.ac.uk Skeletal growth and development in chimpanzees and bonobos has been studied in a number of contexts that have focused on specific regions of development such as dental eruption, epiphyseal fusion at certain sites (e.g.

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the distal radius and ulna), and growth studies, typically of captive individuals. Virtually all of these studies have been conducted using separate samples and there exist few studies encompassing the full developmental spectrum of dental development, skeletal fusion, and long-bone dimensions in a single sample. Critically, no holistic studies to date have used a sample size significant enough to generate a statistically reliable general pattern of fusion in conjunction with dental development used to describe both these species. The present study charts the full spectrum of development using osteological collections from the Museum of Central Africa, the Powell-Cotton Museum, and the University of Zurich. Thirty-seven Pan paniscus and 177 Pan troglodytes individuals were examined. The Pan troglodytes sample encompassed two identified sub-species, P. t. troglodytes (n=72), and P. t. schweinfurthii (n=47). The majority of these individuals were wild shot. Epiphyseal fusion state was determined for a large number of post-cranial sites using a classification system based on McKern and Stewart (1957). Radiographs of dentition were taken and dentition was scored using the Demrijian (1979) method to produce dental scores. Long-bone lengths were measured for all long-bones. These data will be presented and comparisons between species and sub-species will be made with regards to the patterns determined. Pyrosequencing of Plant DNA in Chimpanzee Faeces Phillips, C.A., Pagani, L., Hunt, H., Knapp, L.A., Dickens, S., Lester, J., and Jones, M. University of Cambridge, Cambridge, UK, CB2 1TN Email: cap60@hermes.cam.ac.uk Directly observing feeding of apes reveals invaluable aspects of their diet, but such behavioural data can only be collected from habituated populations in fewer than one-in-five study sites. The identification of the whole range of food-items for apes ensures conservation efforts focus on habitat areas containing important food sources. Therefore, new approaches are needed to yield indirect data from remaining unhabituated populations. For diet, chimpanzee scats encountered on-site provide such an opportunity. Successful dietary analysis of faeces at molecular level has been achieved through pyrosequencing. This ‘high-throughput’ technique is well suited to dietary analysis as it can simultaneously sequence DNA from multiple plant and animal species present within a single sample. We extracted plant DNA from 73 faecal samples, collected from 10 adult chimpanzees in Kanyawara, Kibale National Park, Uganda using a primer set from the rbcL chloroplast gene region. Using this same primer set, we extracted DNA from 64 plant species known to be included within the diet of the Kanyawara chimpanzee community to use as a reference library to compare with findings from faecal samples. Faecal samples were pooled together and analysed as two ‘mega’ sample and pyrosequenced, using a Roche 454 Genome Sequencer

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where plants, of two fragment lengths (157bp and 550bp), were identified up to species level. We discuss the potential of this non-invasive method for dietary analysis of an ape population, especially for determining plant parts of partly- and fully-digested food-items in faeces, which are difficult or impossible to identify using current methods. Taking Personality Bias Seriously in Cognitive Research: A Case Example in Cebus apella Blake Morton, F., Lee, P.C. and Buchanan-Smith, H. Scottish Primate Research Group, Psychology, School of Natural Sciences, University of Stirling, Stirling UK FK9 4LA Email: morton.blake@gmail.com Personality influences decision-making and behaviour in animals, and regularly interacts with cognition. However, there is virtually no discussion among comparative psychologists on whether such interactions bias experimental outcomes, particularly in terms of how we distinguish what is “truly” cognitive as opposed to noise from personality. To illustrate the importance of these issues in the interpretation of cognitive tests, we provide a case example of a series of three problem-solving tasks individually administered (with free-choice participation) to eighteen capuchin monkeys (Cebus apella) at Living Links, RZSS. Using a combination of behavioural- and rater-based methods, the following personality types were identified: Dominance, Openness, Attentiveness, Shyness, and Sociability. Our results show that individual differences in Openness positively and significantly predicted the frequency at which monkeys participated in tasks. Among regularly participating monkeys (i.e. engaged in >75% trials), Dominance was negatively and significantly associated with monkeys’ average rates of learning and performance. Additionally, monkeys that progressed to the second phase of a two-phase experiment scored significantly lower on Dominance. Finally, Attentiveness was positively and significantly related to monkeys’ ability to remain focused on the experimenter during sessions. Studies involving animals should deal with an “unseen” bias from the personalities of their subjects which, in terms of cognitive research, may explain seemingly “cognitive” differences between sites, studies, or subjects.

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INVITED SPEAKER: The in situ and ex situ conservation of Javan primates in Indonesia Wedana, M. The Aspinall Foundation - Indonesia Email: madewedana@hotmail.com The Javan silvery gibbon (Hylobates moloch) and Javan leaf monkeys (the Javan ebony langur - Trachypithecus auratus - and Javan grizzled leaf monkey - Presbytis comata) are unarguably among some of the most threatened primate species in Indonesia. The Javan silvery gibbon was listed in 2010 as one of thirteen priority species for conservation action by the Ministry of Forestry, Republic of Indonesia (RoI). The official cooperation between The Aspinall Foundation and Ministry of Forestry established an MoU at the end of 2009 to work together to conserve these endemic primates of the Island of Java. The project is known as the Javan Primates Conservation Project. It started in 2010 and runs both in situ and ex situ conservation activities. The in situ programme involved a comprehensive survey of the Javan silvery gibbon to investigate and update information and data on their status in the wild. The survey established that no more than 2,000 individuals remain in the wild. Population and habitat threats were recorded including additional information as relevant to several specific sites. The Javan leaf monkeys face similar problems in their existing habitat; however, the study and survey of these species is still ongoing. The ex situ conservation programme involves the rescue and rehabilitation of these three primate species. The rehabilitation facility known as the Javan Primates Rehabilitation Center (JPRC) was established to rescue primates and rehabilitate them for release back to the wild. The JPRC currently cares for 7 rescued gibbon and 3 langurs. It has recently been expanding its facilities since an increasing number of gibbons and langurs need to be rescued. The JPRC plans to release currently rehabilitated individuals back to the wild later this year in 2012. Can ring-tailed lemurs, Lemur catta, show self-control under experimental conditions? Smith, K. and Genty, E. Royal Veterinary College/Institute of Zoology Email: katrionacsmith@hotmail.com Self-control can be defined as the ability to inhibit impulsive behaviour and can be considered adaptive in certain natural situations. A variety of species have been tested in a standard reverse-reward contingency test although the factors that affect the capacity to show self-control remain unknown. In this study, five ring-tailed lemurs, Lemur catta, were tested for the ability to inhibit a natural tendency to reach for the larger of two quantities of food. Subjects were offered a choice between four raisins and one raisin and

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under the reverse-reward contingency, subjects received the non-selected array. All subjects showed a strong, initial preference for the larger array so a large-or-none contingency was applied. In this training procedure, subjects were only rewarded following the selection of the smaller array. Three subjects learnt to reliably select the smaller of two arrays and were able to apply this rule in a return to the reverse-reward contingency and in subsequent trials where novel arrays were presented. A fourth subject learnt to consistently select the smaller array after a correction procedure where no reward was received until the smaller array was selected. These results demonstrate that ring-tailed lemurs are capable of showing a form of self-control and can add to the existing data in this area of behavioural science, particularly for prosimian species. The variable effect of rainfall on the condition of forest living baboons in Gashaka Gumti National Park, Nigeria Lodge, E., Ross, C. and MacLarnon, A. University of Roehampton, Whitelands College, Holybourne Avenue, London, SW15 4JD Email: e.lodge@roehampton.ac.uk As the main determinant of primary productivity in the tropics, rainfall is expected to increase food availability and therefore condition for herbivorous animals such as baboons. However, heavy rainfall is also associated with elevated disease risks. Our study used non-invasive methods to investigate the effect of rainfall on the energetics and physiology of adult female olive baboons. The study population is a geographical and climatic outlier, in comparison to the well studied baboon populations of southern and eastern African, experiencing high rainfall and large areas of closed forest. At Gashaka, higher levels of rainfall were associated with less time spent in energetically costly activities, lower energy expenditure, and higher urinary C-peptide levels (indicating higher energy balance), suggesting a positive influence of rainfall. However, rainfall was also positively correlated with glucocorticoid (stress hormone) levels and mortality, and, during the wet season, rainfall was negatively associated with energy intake. Thus, rainfall influences the baboons’ condition in two contrasting ways. At low to moderate levels, rainfall correlates positively with food availability and condition but, after a certain point, further increase in rainfall do not benefit the baboons, either because they already have sufficient food or because plant productivity stops increasing. Instead, further increases in rainfall are detrimental, increasing stress and mortality, perhaps due to an increased disease burden. This cost of heavy rainfall may be because the Gashaka population is at an ecological extreme for baboons, and may not be well adapted to cope with the high disease burden associated with heavy rainfall.

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INVITED SPEAKER: The PASA vet network – Helping PASA fulfil its welfare and conservation objectives Unwin, S. Veterinary Officer, Chester Zoo, UK and Veterinary Director of the Pan African Sanctuary Alliance (PASA) Email: Steve.unwin@chesterzoo.org

PASA (Pan African Sanctuary Alliance) is a network of primate conservation projects that aims to stem the flow of orphaned, injured and displaced primates in Africa. They do this by providing the highest standards of care to those already in sanctuaries, and to work closely with governments and local communities to ensure primates are valued, protected and secure in their natural habitat. To help fulfil this mission, PASA have created a successful environment for tackling wild primate health issues. This network is a unique, innovative and effective continuing professional development opportunity, based in Africa, which provides access to world class primate health teaching and materials. For example, the peer-reviewed PASA field veterinary manual is updated annually and made freely available via the PASA website. PASA’s animal health objectives are to 1) increase the capacity and training of African veterinarians and healthcare workers at PASA member sanctuaries and in regional wildlife offices, national parks and universities, 2) promote partnerships between wildlife veterinarians, zoologists and medics working in sanctuaries, in the field and in captivity, thus following the one world: one health ideal and 3) supply evidence-based data and research outcomes to assist PASA with measuring conservation impact from its activities. This presentation will highlight ways we are fulfilling these objectives. The effect of predation risk and resource distribution on the ranging behaviour of samango monkeys (Cercopithecus mitis erythrarchus) Coleman, B. Durham University Email: b.t.coleman@dur.ac.uk The risk of predation varies with environmental factors such as the degree of cover provided by vegetation, or the accessibility of an area to predators. These factors allows for the construction of a ‘landscape of fear’ indicating spatially where a prey species perceives higher predation risk. Visualising these landscapes allows great insights into the factors which drive an animal’s movements and spatial ecology. This study conducted on a group of medium sized arboreal monkeys (samango monkey; Cercopithecus mitis erythrarchus) in the Soutpansberg Mountains, South Africa, uses the ‘landscape of fear’ approach to demonstrate that ranging behaviour can largely be interpreted as an adaptive response to perceived eagle predation risk and the spatial distribution of a variety of resources. Results indicate

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that perceived predation risk can be an especially strong determinant of monkey space use, causing them to avoid otherwise beneficial areas. This study improves upon our understanding of predation risk as an extremely important driver of primate behaviour and evolution; a viewpoint historically overlooked by many primate studies. Predation risk and food availability influence fission-fusion dynamics of wild olive baboons (Papio hamadryas anubis) in Nigeria Alberts, N., Semple, S. and Lehmann, J. University of Roehampton, Whitelands College, Holybourne Avenue, London SW15 4JD Email: n.alberts@roehampton.ac.uk Increasingly, primate societies are described by their degree of fission-fusion dynamics, which indicates the extent to which groups vary in spatial cohesion and the temporal variation in the size and composition of parties. However, few studies have systematically quantified fission-fusion dynamics. Furthermore, while perceived predation risk and food availability are known to be key factors shaping primate societies, it is not clear if and how these factors influence fission-fusion processes. Here, we compare the degrees of fission-fusion dynamics of two troops (Gamgam & Kwano) of wild olive baboons in Nigeria, and relate findings to between-troop differences in levels of predation risk and food availability. We collected data over a 12-month period on party size, party composition, and spatial cohesion, along with measurements of predation risk and food abundance. We found that the degree of fission-fusion dynamics differed between troops, in correspondence with local differences in ecology. In the Gamgam home range, predation risk was higher and food more abundant. The Gamgam troop rarely fissioned and parties were frequently large. In the Kwano home range, by contrast, levels of predation risk and food availability were lower, and the troop was found to break up frequently into small parties. The Kwano troop showed more variation in the size of parties and in inter-individual distances than the Gamgam troop. However, the rate of party composition changes was similar between the two troops. Our results provide evidence that some aspects of primate fission-fusion dynamics are responses to local ecological conditions, but that other aspects may be less flexible.

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Duet pant hooting reflects both long-term and short-term social bonds in wild male chimpanzees Fedurek, P. and Slocombe, K.E. Department of Psychology, University of York, UK Email: pf517@york.ac.uk Male chimpanzees form strong and durable social bonds. Several indicators of social bonds have been recognised in this species, such as reciprocated grooming and coalitions. However, these well established bonding behaviours in primates are either time consuming (grooming) or require certain circumstances to occur (coalitions). In the proposed study we posit that male chimpanzee duet pant-hooting, a very common behaviour in these primates, is a more flexible but very reliable signal of affiliative relationships. The results of the study, which was conducted on the Kanyawara community in Kibale National Forest in Uganda, support this view showing that bonded males were more likely to be involved in duet pant-hooting than less bonded dyads (controlling for time spent in the same subgroup). We also compared frequencies of affiliative behaviours, such as feeding tolerance, mutual grooming, joint displays and support in agonistic interactions, between days in which dyads performed duet pant-hooting and days in which they did not. The results suggest that duet pant-hooting might also reflect affiliations between males on a short-term basis. Signalling temporary affiliations might be especially beneficial in primates such as chimpanzees living in highly flexible fission-fusion societies where valuable social partners are not always around. We suggest that pant-hoot chorusing in male chimpanzees might be an efficient and flexible signal of affiliative relationships that reflects both long-term and short-term social bonds. Bilateral asymmetry in humeral torsion: using the skeleton to assess patterns of arm preference in great apes and humans Barros, A. University College London Email: a.barros@ucl.ac.uk Approximately 90% of modern human groups display a preference for right-hand use. Because such a strong species-level right-hand preference has to date only been unequivocally identified in Homo sapiens, this trait is considered to be unique to this species. In apes, on the other hand, the evidence is controversial with contrasting findings regarding population-level handedness, raising questions about when this trait arose in hominins. Disparities in methods for quantifying and defining handedness, however, complicate cross-species comparative studies (e.g. behavioural observations in great apes, versus questionnaires in humans). Therefore, in order to compare human and non-human handedness patterns, we instead

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investigated the magnitude and directionality of skeletal asymmetries in humeral torsion in African apes (Gorilla gorilla and Pan troglodytes) and humans. Humeral torsion was chosen because it successfully predicts handedness in individual humans. Skeletal asymmetries were expressed as a score calculated through the unsigned (Absolute Asymmetry) and signed (Directional Asymmetry) differences between two measurements in a pair of skeletal elements for each individual. The aim was to identify whether human-like patterns of handedness are seen in African apes in order to investigate whether handedness, and in particular right-handedness, is a hominin-specific trait. Our results indicate the presence of a population-level right-handedness in humans but not in chimpanzees and gorillas, suggesting that right-handedness is a trait unique to humans. We do not reject that handedness exists in African apes, but suggest that unlike in humans, it is mediated by individual preferences. We thus support that handedness is apomorphic to hominins, probably as a result of a qualitative shift in behaviour, possibly linked to a shift in brain hemispheric specialization (related to either language or fine motor control), that has taken place since the Pan-hominin split. A comparative study across primate species on the evolution of cerebral lateralization: the validity of a quantitative Multidimensional Method Quaresminia, C.1, Forrester, G.S.2 and Vallortigara, G.1 1Center for Mind/Brain Sciences, University of Trento, Corso Bettini 31, 38068 Rovereto, Italy 2Department of Psychology, University of Westminster, 309 Regent Street, London, W1B 2UW E-mail: caterina.quaresmini@gmail.com Inconsistent methodological techniques have hindered comparisons of lateralized behaviours tied to dominant neural regions between human and non-human primates. A lack of valid inter-species comparisons has thus hindered our understanding of the evolution of species-specific cognitive capabilities. The present study investigated lateralized behaviour in both human and non-human primates by employing a quantitative Multidimensional Method. Specifically, we considered whether the context of interactions influenced the lateralization of behaviours underpinned by hemispheric dominance for specific neural processes. Two groups of zoo great apes, chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla gorilla), and a group of 3-4 year-old typical children were observed within their natural social environment. The handedness analysis revealed a significant right-hand/left hemisphere dominance during contact with inanimate targets (objects and environmental items) compared to animate targets (self and social partners) for both human and non-human species. These data suggest an evolutionary

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hemispheric specialization for object manipulation, shared by a common human/ape ancestor. Additionally, we reported a significant preference in keeping closest conspecifics to the left side of the body during manual activities, suggesting right hemisphere dominance in processing affective stimuli for apes, but not in children. These studies demonstrate the value of a quantitative method to investigate the evolutionary origins of hemispheric specialization underlying manipulative and social behaviours in human and non-human primates. Chimpanzee nest-building: anti-predation, thermoregulation or parasite-avoidance strategy? Koops, K. Leverhulme Centre for Human Evolutionary Studies, University of Cambridge Email: kk370@cam.ac.uk The construction of nests (or beds) for sleeping is a chimpanzee universal, yet little is known about the adaptive function of nest-building. I present an in-depth study of nest-building by chimpanzees at the Seringbara study site in the Nimba Mountains, Guinea, West Africa. I recorded 1520 chimpanzee nests over 28 months during three study periods between 2003-2008. I investigated where chimpanzees built their nests, both across the home range and in nest trees. I tested three, non-mutually exclusive, hypotheses for the function of arboreal nest-building. I assessed whether selectivity for nest tree characteristics reflected an anti-predator strategy, examined whether nesting patterns (both arboreal and terrestrial) and nesting height were influenced by variation in climatic conditions (temperature, humidity, wind), and measured mosquito densities at ground level and in trees at 10 m and related mosquito densities to nesting patterns. Chimpanzees preferred to nest above 1000 m and nested mainly in primary forest. They preferred relatively large trees with a low first branch, dense canopy and small leaves. No support was found for the anti-predation hypothesis, nor did mosquito densities explain arboreal nest-building. The thermoregulation hypothesis was supported, as both nesting patterns and nest-height variation across seasons reflected a humidity-avoidance strategy. Chimpanzees nested higher in trees and at higher altitudes in the wet season. In sum, chimpanzees were selective in their choice of nest sites, and tree-nesting patterns at Seringbara were best explained by a thermoregulation strategy of humidity avoidance.

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Abstracts for poster presentations Possible occurrence of diabetes in vervet monkeys (Cercopithecus aethiops) undergoing rehabilitation at Riverside Wildlife Rehabilitation Centre (RWRC), Limpopo Province - South Africa Albers, M. Oxford Brookes University Email: marlies.albers-2011@brookes.ac.uk Although the vervet monkey (Chlorocebus pygerythrus) is not endangered, there are centres in South Africa that specialize in their rehabilitation. Riverside Wildlife Rehabilitation Centre is such a centre. The centre was concerned about the occurrence of diabetes in domesticated vervet monkeys that were brought into the centre for rehabilitation. The occurrence of diabetes could reflect an inadequate diet prior to arrival at the centre. An improper diet may lead to a higher risk of developing diabetes. This research tested if monkeys recently brought to the Centre had a higher risk in developing diabetes than monkeys already at the centre based on their blood glucose values. The blood glucose levels were tested with Bayer bv. equipment. It was expected that monkeys that were recently brought into the centre would have higher blood glucose levels than the monkeys living in natural circumstances in the centre. However, the opposite was true. The domesticated vervet monkeys had significantly lower blood glucose levels than their semi-wild counterparts. I suggest that more rehabilitation centres start testing their (vervet) monkeys for the possible occurrence of diabetes when brought into rehabilitation centers. This research can prove very valuable when rehabilitating primates for future release. A comparison of the gastrointestinal parasites recovered from olive baboons (Papio anubis) and tantalus monkeys (Cercopithecus aethiops tantalus) in three habitats in Gashaka Gumti National Park, Nigeria Bailey, E. and Ross, C. University of Roehampton, Whitelands College, Holybourne Avenue, London, SW15 4JD Email: emmabailey4@googlemail.com We investigated the gastrointestinal parasite prevalence, abundance and seasonality in 3 troops of olive baboons (Papio anubis) and 2 troops of tantalus monkeys (Cercopithecus aethiops tantalus) in Gashaka Gumti National Park, Nigeria in 3 habitats. The groups included a group of each species in high altitude montane highlands and around a lowland village (in both cases these involved crop-raiding troops in contact with human settlements) and a group of baboons in a mixed forest habitat with minimal contact with humans. We collected faecal samples from each troop and these were analysed using the formol-ether concentration technique. Twelve

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parasite taxa were recovered, 3 protozoa and 9 helminths. Baboons had a higher parasite richness than C. a. tantalus monkeys which is best explained by their larger size and longer life span. We suggest that anthropogenic and altitudinal effects may have affected differences in the abundance of Schistosoma mansoni, Entamoeba sp. and Balantidium coli. A higher abundance of helminths in the non-crop-raiding baboon troop may be due to the nutritional benefits of crop-raiding enhancing the immune response to infection. We found seasonality affected the abundance of Balantidium coli, Trichuris sp., Entamoeba sp., Trichostrongylus sp., and Strongyloides sp. but not across all troops. Host age, sex and reproductive success are related to gastrointestinal parasite species richness in Papio anubis in Gashaka Gumti National park, Nigeria Boulton, R.A. and Ross, C. University of Roehampton, Whitelands College, Holybourne Avenue, London, SW15 4JD Email: rebeccaboulton87@googlemail.com Parasitism affects the viability of natural populations and parasites are an important factor driving the current human-induced global biodiversity crisis. Several studies have found that body mass, population density and social contact influence parasite species richness (PSR) within the same host clades and between populations of a species. Few studies have focused on the mechanisms underlying parasite aggregation at the level of the individual host. The aim of this study was to determine correlates of PSR at the level of the individual in a troop of olive baboons (Papio anubis) at the Gashaka Gumti National Park, Nigeria. A minimum of 2 faecal samples per individual were collected from 33 individuals in the Kwano troop and analysed for the presence of helminth and protozoan parasites using the modified Ridley formol-ether concentration technique. A total of 9 parasitic taxa were recovered. Results suggest that PSR is related to age, sex and birth rate in females. Parasite species richness (PSR) was higher in infants than all other age categories. Males displayed greater PSR than females across all age categories other than sub-adults, where the PSR is higher in females. Adult females with higher PSR scores reproduced less frequently. These patterns suggest that acquired immunity and the immunosuppressive role of reproductive hormones play a role in the distribution of gastrointestinal parasite fauna. Additionally it appears that PSR does influence reproductive success in females.

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Rehabilitation and release of rescued primates: A scientific approach Donaldson, A. Durham University, UK, and Colobus Trust, Kenya Email: andreadonaldson@yahoo.co.uk The Colobus Trust Primate Reintroduction Project, Diani Beach, Kenya meets the criteria of a welfare/reinforcement release and as such has combined welfare and conservation aims. Ecological and behavioural data collection on wild control groups began in September 2011, concentrating on identifying favoured and fallback feeding and refuge tree species. In addition a total count census of the primate populations in the Diani Forest has identified all troop locations within the wider release area. This information will allow a full habitat assessment to be conducted and the best possible release site to be located. Primate species currently being prepared for release are Zanzibar Sykes monkey (Cercopithecus mitis albogularis) and Hilgert’s vervet monkey (Chlorocebus pygerythrus hilgerti). Situated in a pre-release enclosure the individuals have been subjected to pre-release training, including predator awareness and wild food exposure. Using a soft release strategy the release troops will have a one-year extensive post-release monitoring period and comparative data collection will continue on the control troops. All methodology has been predominantly guided by the IUCN Guidelines for Non-human Primate Reintroductions (2002). The resulting scientific protocol developed during this two-year project, will be of direct interest to those working in the fields of reintroduction biology and any lessons learnt have real potential to make significant contributions to the slim body of knowledge that currently exists. Once completed, the protocol could be used to improve future reintroduction plans and model the performance and efficiency of different release strategies. Consequently, the programme offers a hugely exciting opportunity for setting standards for release programmes, whilst also furthering the welfare and conservation of African primates. Social Network Analysis to examine interaction patterns of immature chimpanzees (Pan troglodytes) in captivity Farooqi, S.H. and Koyama, N.F. Liverpool John Moores University, Liverpool, UK Email: sam_wild1211@yahoo.co.uk We used Social Network Analysis (SNA) to examine the pattern of affiliative interactions of immature chimpanzees in captivity. SNA provides quantifiable measures of an individual’s position in its social group, their number of interaction partners and a visual perspective of the spatial cohesiveness of individual networks. We collected data on five immature chimpanzees at Chester Zoo, UK from February 2008 to November 2009 using 15-minute focal animal sampling. Our analyses included social

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grooming and play interactions of immatures with other group members (excluding mothers as potential social partners). Our results showed that the node degree and tie strength for play networks was significantly higher than for grooming networks in each of the two years. Moreover, immatures were more central in their play networks and more peripheral in their grooming networks. Play networks expanded from 2008 to 2009 whereas grooming networks remained stable during the two years. Finally, there was no matrix correlation between the play network of immatures and the grooming network of mothers. These results suggest that immature chimpanzees used social play rather than grooming as a means to interact and integrate into adults’ social networks and supports the idea that play in infants and juveniles functions to strengthen an individual’s ties to its own social group and potentially improve social cohesion. Baboon rehabilitation: over 20 years of experience in rehabilitation and release of Chacma baboons Munro, S. and Griciute, J. C.A.R.E Email: joana.griciute@gmail.com C.A.R.E. (Centre for Animal Rehabilitation and Education) specialises in the rehabilitation of Chacma baboons (Papio cynocephalus ursinus) and has had incredible success in the release of their troops back into the wild. CARE was established in 1989 by Rita Miljo and is one of the first animal rehabilitation centres in South Africa. CARE has a unique method of rehabilitation of Chacma baboons, where individuals that are brought to the centre are placed in groupings with those similar in age and/or similar in behavioral markers. The process of rehabilitation can be broken down into age-based categories: infants (0-6 months), youngsters (6-12 months), juveniles (1-2 years), adolescences (2-5 years) and adults (5+ years). During 6-7 years of rehabilitation process, growing baboons have a possibility to observe and interact with other rehabilitant troops as well as the wild troop that is present on the facility’s property. These interactions aid to facilitate troops’ knowledge of proper foraging behavior, vocalizations and their proper contexts, as well as conflict resolution. By establishing these troops at young ages and allowing them to grow together, the cohesiveness of the troop is more strongly reinforced and these troops, that consists of natural sex/age ratio and social structure resembling that one of the wild baboons, are ready to be released back into the wild. From 1996, CARE has conducted 9 releases under various circumstances. CARE is conducting soft releases and they have been proven successful with average survival rates of 84% after the first year, and 70% after the second.

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Investigating handedness in free ranging captive primate populations Heaton, R. Email: ray.heaton@blueyonder.co.uk Four different colonies of captive baboons were observed in order to establish if there was a hand preference in the performance of manipulative tasks. Feeding behaviour in particular, and social interaction such as grooming and infant care were observed. Each colony consisted of more than fifty animals. Data were collected from individuals of known identity and from animals that were not individually identified. Differences found between the colonies such as species and hybridisation, colony/troop structure, and management will be discussed. Problems encountered in the study of these populations will be discussed as issues for research on free-ranging captive primate populations. Investigating the left-sided cradling preference in non-human primates Heaton, R. Email: ray.heaton@blueyonder.co.uk Left-sided cradling preference is well documented in humans; this is the tendency for mothers and other carers to cradle infants (independent from breast feeding/suckling) on the left side of the body. Observations from this study informed the first report of left-sided cradling behaviour in non-human primates, from a large multi species sample. Data will be presented from observations of chimpanzees, gorillas, orang-utans, gibbons, and monkey species which were carried out in captive situations. A strong left-sided cradling preference was found in chimpanzees and gorillas with less significant preference in orang-utans and gibbons. Many of the gorilla observations were undertaken at Howletts Wild Animal Park. These include ten mother-infant pairs with observations of one pair starting a few hours after the birth. The Howletts part of the study will be reported in some detail in order to give account of the group structure and management, and also the interesting parenting behaviour of some mothers. The possible evolutionary benefits of left-sided cradling will be discussed briefly. Female social relationships of wild Assamese macaques Macdonald, S., Schülke, O. and Ostner, J. Courant Research Centre, Evolution of Social Behaviour, Georg-August-University, Kellnerweg 6, 37077, Göttingen, Germany

Email: sally.macdonald@biologie.uni-goettingen.de Female primates invest a substantial amount of time forming and maintaining social relationships, much more so than males. To date much of

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the research aimed at improving our understanding of these relationships has focused primarily on female baboons. We aim to begin to broadening this work by providing the first detailed description of female social relationships in wild Assamese macaques (Macaca assemensis). We investigated how closely patterns in their social relationships match those described for female baboons, while also making some within-species comparisons with data collected from a provisioned temple site. Over 2300 hours of observational data were collected, during two one-year sampling periods (October 2007 - September 2008 and May 2010 - April 2011) from a single group in Phu Khieo Wildlife Sanctuary, Thailand. Subjects included all adult females present in the group during these two periods (N=12 and 15 respectively). Initial results revealed strong similarities in the overall strength and distribution of female social relationships between species but distinct differences in the directionality of grooming behaviour in particular. We also found strikingly similar dominance styles within-species, despite prominent differences in resource distribution between the groups investigated. These results provide both an important insight into female social structure in this species as well as useful comparative data for future research. Such broader cross-species comparisons not only improve our understanding of female primate social relationships in general but also have the potential to allow us to begin identifying possible universal properties of female social relationships. Primate research in the UK: a review Martindale, V. Oxford Brookes University Email: vicmartindale@aol.com Primates play a central role in pre-clinical and biomedical research efforts. While procedures on living animals in the UK are monitored and regulated through the Home Office under the terms of the Animals (Scientific Procedures) Act 1986, data on primate use are difficult to obtain. Review of data available on primate research in the UK was obtained from Home Office publications, published literature, Hansard (the Official Report of parliamentary debate) and Freedom of Information (FOI) requests. The UK is the largest user of non-human primates in the EU, using nearly half of all the primates used by member states. In 2010, the UK used 2,649 primates in 4,688 experiments. Although the Home Office does not collect data by species, a FOI request confirmed that currently there are only two species of macaque used in the UK: rhesus macaques (Macaca mulatta) and long-tailed macaques (Macaca fascicularis). The remaining 674 (25%) primates were marmosets and tamarins. However, numbers of each species and numbers used in specific areas of research were not collected. Neither is there a requirement to record the source of primates imported into the UK and whether they are F1 or F2+ generation. My findings highlight that

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despite the UK Freedom of Information Act 2000, which came into force in January 2005, there still remains the need for greater openness and accountability surrounding the use of primates as well as improved data collection by the Home Office. I discuss the potential conservation implications that this may have. Mitochondrial DNA diversity of a captive group of Hylobates moloch Mitchell, K.1, Harris, K.L.1, Kingston-Jones, M.2, Ford, M.2 and De Luna, C.1 1CEQAS, Writtle College, Chelmsford, Essex, CM1 3RR 2Howletts Wild Animal Park, Bekesbourne, Kent, CT4 5EL Email: katherine.mitchell@writtle.ac.uk Hylobates moloch is described as the most urgent conservation concern of all Asian primates. A future conservation initiative orchestrated by the Aspinall Foundation to minimise extinction risks for this species includes a captive breeding and reintroduction programme. Genetic variability has been described as the ‘clay of evolution’ (Amos and Harwood, 1998) which is a concept that is pertinent to a captively bred population who, after a (as yet unknown) number of generations may have undergone alterations at the genetic level as a cause of relaxed environmental pressures within the captive habitat. This study analysed genetic variability at the hypervariable-I region (HV-I) within the Control Region of mtDNA of 15 individuals housed at Howletts and Port Lympne zoological institutions. Although pertaining to uniparental genomic inheritance, it is the temporal scale of mutation of this genomic segment (purported to occur at a rate 5 to 10 times faster than in nuclear DNA) that this DNA marker was chosen. PCR product length totalled 511 nucleotide sites and a haplotype count of 5. Of the 511 loci 30 were found to be polymorphic, all with 2 alleles per locus. Nucleotide diversity (average over loci) was 0.011704 +/- 0.006630. Overall product length was longer than previously reported (e.g. Monda et al., 2007; Whittaker et al., 2007). However, although the HV-1 mtDNA region is described as highly variable and with the fastest mutation rate within the mtDNA molecule in primates, results within the study population indicate low levels of diversity. References: Amos, W., Harwood, J. (1998) Factors affecting levels of genetic diversity

in natural populations. Philosophical Transactions of the Royal Society of London B, 353; 177-186.

Monda, K., Simmons, R.E., Kressirer, P., Su, B., Woodruff, D.S. (2007) Mitochondrial DNA hypervariable region-1 sequence variation and phylogeny of the Concolor Gibbons, Nomascus. American Journal of Primatology, 69: 1285-1306.

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Crowd density effects on self-directed behaviours, affiliation, and agonism in zoo-housed callitrichids Scott, R.M. and Leavens, D.A. School of Psychology, University of Sussex Email: rebeccamscott@gmail.com The presence of visitors is a source of stress for zoo-housed primates, known as the ‘visitor effect’ (Hosey 2000). However, there is limited research on platyrrhines and a lack of consensus on behavioural indicators of welfare. Self-directed behaviours (SDBs) are an indicator of stress in primates (Maestripieri et al. 1992) and research has shown that the laterality of these SDBs could be indicative of asymmetries in emotional processing (Leavens et al., 2001, 2004). This study is the first to use SDBs as a measure of the visitor effect in callitrichids. Two golden lion tamarins (Leontopithecus rosalia) and three cotton-top tamarins (Saguinus oedipus) were observed for two weeks at London Zoo. The laterality and frequencies of SDBs, affiliative and agonistic behaviours and rates of withdrawal were recorded along with crowd density. The rates of SDBs significantly increased with crowd density (F(2,8)=20.47, p = .001). There was a significant left-hand bias for SDBs at medium (but not low or high) crowd densities (t(4)= 4.95, p = .008). In addition, agonistic behaviours and withdrawal increased with crowd density (both ps < .05), whereas, in contrast, affiliative behaviours decreased with increases in crowd density (F(2,6) = 7.2, p=.025). Overall, these findings confirmed that high crowd densities were associated with behavioural markers of mild stress in these tamarins. These results extend previous findings in catarrhines to platyrrhines. This was also the first study of lateral asymmetries in SDBs in callitrichids. These findings support the use of SDBs in assessments of primate well-being in captive environments. Saimiri scuireus’ resilience in the face of trade Smith, H. and Nijman, V. Oxford Brookes University Email: hels_smith@hotmail.co.uk Animal rescue centres provide refuge to primates who have been the victims of illegal trafficking and abuse. Ecuador is one of the most bio-diverse countries in the world and its centres are brimming with primates. Data were collected from 23 centres throughout the country over a five-year period which demonstrated that 46% of mammalian in-take was primates. Of these, 66% were less than two years old when they arrived at the centres but 75% would die in the first year. In this study data were collected from one Amazonian rescue centre over an 18-year period (1992 to 2010) with 765 primates brought into the centre. The majority were squirrel monkeys, Saimiri Sciureus [357 individuals]. This species appears to have greater

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resilience and survival rates than other primates. Similar to other species, 60% were under 2 years of age upon arrival but mortality rates were much lower; only 20% died in the first year. In fact, 61% went on to be released to live freely in the trees around the centre. There is little sexual bias to the squirrel monkeys coming into the centre with 150 males, 158 females and 49 which did not have their sex recorded. It could therefore be concluded that primates which have a lower presence in centres are not being illegally extracted at lower rates but are actually more negatively affected by the trauma of trade and simply do not survive long enough to get as far as the rescue centres. The effect of Non-Naturalistic Auditory Enrichment on the welfare of Moloch gibbons (Hylobates moloch) Wallace, E.K.1, Semple, S.1, Kingston-Jones, M.2 and MacLarnon, A.1 1 Roehampton University 2Howletts Wild Animal Park, Bekesbourne, Kent, CT4 5EL Email: emma.k.wallace@cantab.net The use of non-naturalistic auditory enrichment has never before been researched on Moloch gibbons (Hylobates moloch). Instrumental classical music with less than 90 beats per minute was played to eight individuals within two family groups at Howletts Zoo, Kent, UK. Playbacks were used for three of six weeks, alternating with control weeks involving no playbacks. Behavioural data were recorded for each individual as well as faecal samples being collected to analyse excreted cortisol metabolite concentrations. Data analysis showed that each individual reacted differently to the enrichment, for example, one adult male showed a significant increase in self-scratching, self-grooming and grooming others during music weeks whilst an adult female showed a significant decrease in self-scratching for the same weeks. Significant differences were seen for some of the eight individuals in brachiation, self-scratching, self-grooming, grooming others, being groomed by others and aggression received. No significant difference between the two conditions was found for the faecal cortisol concentrations and no consistent behavioural patterns could be seen across all of the individuals. It is unclear if the changes in behaviour showed a positive welfare change. The study was the first to use behavioural and physiological measures of stress to look at the impact of enrichment on groups of captive gibbons. It has shown that a combination of the two measures can be used when researching the effect of enrichment, as well as highlighting that individual differences are important when deciding upon and maintaining enrichment programmes.

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Examining regurgitation and reingestion in captive western lowland gorillas: stress, diet and rearing Winkler, S. and Lehmann, J. Roehampton University Email: sunshinediva4@hotmail.com Regurgitation and reingestion (R/R) is a behaviour that is only seen in captivity and very little is known about this behaviour and its causal factors. Many researchers have suggested several causal factors that influence R/R such as stress, boredom, diet, rearing history and inadequate environmental conditions. The aim of this study is to examine if stress, diet or rearing history affects the occurrence of R/R in captive western lowland gorillas. Studies have shown that self-directed behaviours (SDB) such as self-grooming, self-scratching and yawning can be used as an indicator of individual stress levels. Data were collected on the gorillas’ rates of both R/R and SDBs which were compared with their rearing history. Since the gorillas’ feeding schedule changed from day to day, the possible influence of diet on R/R rates was examined by looking at the differences in R/R rates at different times on different days. Results of this study indicated that hand-reared gorillas exhibited higher SDB rates than mother-reared. It was found that rates of R/R during the early afternoon (11:30am-12:30pm) were higher than R/R rates during any of the other times. Significant differences were found for R/R rates and day of the week as Sunday, Wednesday and Thursday all produced higher R/R rates than both Monday and Tuesday. These results suggest that diet may have an influence on the occurrence of R/R; however, this behaviour is very complex and more studies should be conducted to determine the exact causal factors of R/R. Correlating staff and behavioural ratings of chimpanzees’ novel task abilities Wood, L.1, Kendal, R.1, Hopper, L.2,3, Lambeth, S.3, Schapiro, S.J.3 and Flynn, E.1 1Durham University 2Georgia State University 3KCCMR, UTMDACC Email: l.a.n.wood@dur.ac.uk Chimpanzees (Pan troglodytes) exhibit individual differences which remain over time; hence, chimpanzees can be said to have ‘personalities’ (Freeman & Gosling, 2010). Husbandry employees play an important role in the in the welfare of captive chimpanzees and, with continued interaction, they can give valuable insight into such behavioural traits. The current study investigated whether husbandry employees could provide valid rating for the chimpanzees’ traits (consistent across raters) and whether these ratings reflected actual behaviour. Fifty-four chimpanzees group-housed

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chimpanzees, housed at the KCCMR facility in Bastrop, TX, USA were rated by three care technicians who had worked with the chimpanzees for between 18 and 40 months. Chimpanzees were rated on a seven-point Likert scale on the following traits: Inquisitiveness, Intelligence and Inventiveness (developed from a bottom-up approach with the same chimpanzees by Freeman et al., in prep). The validity of the ratings was tested with behavioural assessments, in which the chimpanzees, in their home group, were presented with four novel enrichment devices (food & non-food related). Results indicated (i) strong inter-rater reliability of staff behavioural ratings, (ii) each chimpanzee’s behaviour was highly correlated across enrichment devices, regardless of whether they contained food, and (iii) staff behavioural ratings predicted the chimpanzees’ behavioural responses to the novel enrichment devices. The current study suggests that captive facilities may be able to negate timely and costly behavioural assessments, instead relying on ratings from just a few experienced husbandry employees. Buton Macaques (Macaca ochreata brunnescens): crops, conflict, and behaviour on farms Wyper, W.M.1, Priston, N.E.C.2 and Lee, P.C.1 1 Behaviour and Evolution Research Group, School of Natural Sciences, University of Stirling, Stirling, UK 2 Department of Anthropology and Geography, Oxford Brookes University, Oxford, UK Email: rebecca.wyper@stir.ac.uk One consequence of anthropogenic habitat alteration is that many nonhuman primates are forced into conflict interactions with humans and their livelihood activities, especially through crop raiding. These problems are particularly acute for the endemic and threatened Buton Island macaque (Macaca ochreata brunnescens), in southeast Sulawesi, Indonesia. Our study investigated the crop-raiding behaviour of this species over time. Foods eaten and the behavioural repertoire exhibited by macaques during crop-raiding at and inside farm perimeters were observed over a period of 8 years (2002–2009). Storage organ crops (e.g. sweet potato) were abundant and most frequently raided by macaques. Individual macaques were most commonly observed to raid close (0–10 m) to farm perimeters. Activities such as feeding, resting, moving, and social interaction varied significantly as a function of penetration distance into the farm, but only marginally between age-sex classes. The annual average raid frequency per farm decreased over the latter years of the study period, raising questions about changes in macaque foraging and ranging behaviour over time and their response to farm management and mitigation strategies.

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Honest signals? Barbary macaque males cannot differentiate conception and post-conception female anogenital swellings Young, C.1, Majolo, B2, Heistermann, M.3, Schülke, O.1 and Ostner, J.1 1Courant Research Centre Evolution of Social Behaviour, Georg-August University Göttingen, D-37077, Germany 2 School of Psychology, University of Lincoln, Brayford Pool, Lincoln, LN6 7TS, UK 3Reproductive Biology Unit, German Primate Center, 37077 Göttingen, Germany Email: christopher.young@zentr.uni-goettingen.de Exaggerated sexual swellings have been shown to be a reliable indicator of ovulation in many cercopithecine primate females. Studies of sexual behaviour in Barbary macaques (Macaca sylvanus) concentrated on conception cycles and concluded that the maximal swelling period is indeed an honest signal of ovulation to males. However, a post-conception swelling may alter this perception if males cannot distinguish between the two swelling periods and thus do not alter their mating effort accordingly. Here we examine the behavioural consequences of conceptive and post-conceptive swellings in two wild groups of Barbary macaques of the Middle-Atlas Mountains, Morocco, using a combination of behavioural and hormone data and visual swelling scores. Relating the visually assessed period of maximum swellings (days -2 to -7 after deflation) to the hormonally determined 5-day fertile window (period of highest ovulation probability) within ovulatory cycles we found that ovulation most frequently took place within the maximum swelling period confirming previous results of the swelling being a reliable indicator in this species. Consequently, male sexual behaviour did not differ between the fertile period and the period of maximum swelling. However, females displayed, in 73% of cases, an anogential swelling 25-30 days following conception. Both male and female sexual behaviour did not differ between conceptive and post-conceptive swelling periods, indicating that Barbary macaque males are unable to differentiate between these two qualitatively different periods. Thus, the notion of Barbary macaques’ sexual swellings being an honest signal of timing ovulation needs to be re-considered.

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FUTURE MEETINGS

PSGB Winter Meeting 2012

Primate Biogeography

Wednesday 5th December 2012 ZSL, London

Organisers:

Sarah Elton, Durham University &

Helen Chatterjee, University College, London

We have an exciting range of topics from internationally-known speakers:

Alexander Harcourt, University of California, Davis. Human Biogeography. Is Man Merely a Monkey?

Jason Kamilar, Midwestern University

The Biogeography of Primate Communities: Ecological and Evolutionary Perspectives

Shawn Lehman, University of Toronto

Where the Wild Things Are: Lemur Conservation Biogeography in the 21st Century

Simon Bearder, Oxford Brookes University

(Osman Hill Memorial Lecturer) Primate Taxonomy in the Field: New Insights into Biodiversity,

Biogeography and Phylogeny.

Andrea Cardini, University of Modena and Reggio-Emilia Primate Biogeography in the Phenomic Era: Is There a Role for

Digital Taxonomy?

Plus titles to be confirmed from Terry Dawson (University of Dundee), Robert Marchant (University of York), Vincent Nijman (Oxford Brookes

University) and Sam Turvey (Zoological Society London)

The call for posters on all aspects of primate geography is also open (poster abstract deadline 31st October: the scientific programme committee will review

all abstracts and communicate decisions no later than 7th November).

You can get more details, submit an abstract and register via the website: <www.psgb.org/meeting_detail.php?ID=PSGB-Winter-Meeting-2012>

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PSGB Spring Meeting 2013

University of Lincoln

Organiser: Bonaventura Majolo, University of Lincoln

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ANNOUNCEMENT: <marmosetcare.com> There is a newly launched website that aims to promote the welfare of captive common marmosets (Callithrix jacchus) and illustrates their behavioural repertoire. Designed to be approachable and fun to use, as well as instructive, the site is interactive and is illustrated extensively with photos and videos. <marmosetcare.com> It represents an important new resource of information on common marmosets, the most-used New World primate in laboratory research and testing worldwide, and is therefore a worthy link item. Ana Isabel Moura Santos Departamento de Fisiologia Faculdade Ciências Médicas - UNL Campo Mártires da Pátria, 130 1169-056 Lisboa Portugal

ANNOUNCEMENT: NEW JOURNAL

Journal of Zoo and Aquarium Research (JZAR) The Research Committee of the European Association of Zoos and Aquaria (EAZA) is proud to announce that the Journal of Zoo and Aquarium Research (JZAR) is now open for submissions. JZAR will provide a forum for rapid publication of novel, peer-reviewed research papers, reviews, technical reports and evidence-based case studies. JZAR will be officially launched in summer 2012 as an open access online journal. Through their living collections, zoos and aquariums are uniquely placed to contribute to conservation-related research. Research categories covered by JZAR will include studies in pure and applied biological sciences, in situ conservation research and research aimed at developing other roles of zoos and aquariums. If you wish to submit a paper to JZAR, visit the holding page at www.jzar.org and find more information about the aims and scope of JZAR as well as the interim submission guidelines. The full website will be live by September 2012. On the holding page you will also find a list of example papers that are eligible for publication in JZAR. More information will follow soon. If you submit your paper now, it could be published as early as Autumn 2012. Christoph Schwitzer (on behalf of the EAZA Research Committee)

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CONSERVATION GRANT REPORT

Feeding ecology and conservation of the red-shanked doucs (Pygathrix nemaeus) at Son Tra Nature Reserve, Vietnam Larry Ulibarri <Larry.Ulibarri@yahoo.com> Awarded a PSGB grant in 2010 With the support of Primate Society of Great Britain, the ‘Son Tra Douc Research and Conservation Project’ was started. This project centers on the conservation of red-shanked doucs (Pygathrix nemaeus) on Son Tra Nature Reserve, Da Nang City, Vietnam. Red-shanked doucs are one of the most striking and beautiful primate species in the world, called the ‘monkey of 5 colors’ locally. They are classified as endangered and suffer from hunting and habitat loss throughout their range. Although some data are available on the species’ distribution, little is known on its ecology. Son Tra Nature Reserve is a reflection of the conservation situation in the country. This forest patch is home to one of the largest remaining populations of red-shanked doucs in Vietnam. It remains relatively intact, considering it is close to one of the largest cities in Vietnam. However, hunting, habitat loss and habitat fragmentation are threatening the doucs and biodiversity on Son Tra.

Part of a family from the habituation group showing (left to right) an adult female with newborn infant, alpha male of the family,

and a subadult male.

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This project gathers information about doucs’ social behavior, feeding ecology, ranging, and forest phenology. This study found that doucs live in multi-male multi-female families. Groups consist of multiple families, which have a daily and seasonal fission-fusion pattern. The sex ratio of adult females to adult males is typically 2:1. Care-taking of infants is done primarily by the mother, although adult males provide direct and indirect care-taking benefits. This project identified 62 different feeding plant species. Doucs feed on a variety of plants, mainly trees but also vines and climbers. Nutritional analysis of select food and non-food items is pending. However, doucs tend to select higher quality foods based on availability. This suggests their dental and gut morphology is an adaptation to fall-back foods.

This project also monitored the numerous threats to biodiversity on Son Tra. Illegal activities including hunting, fishing, timber and non-timber forest extraction are documented. Although this project does not have a direct law enforcement component, data on illegal activities are regularly provided to the protection authorities and the knowledge of these activities is beneficial for the protection of the doucs. Development of several tourist resorts and roads have serious negative impacts on biodiversity of Son Tra. Roads are fragmenting the forest into small blocks, and the doucs, as most other arboreal wildlife, cannot cross many of these roads due to the amount of forest removed and the steepness of the slopes created. This project assisted in receiving the government’s approval for a natural tree canopy bridge project which will connect forest patches. It also campaigned to halt road widening of existing roads on Son Tra but with limited success.

Finally, this project has an education component. Work with several governmental departments and Da Nang City consultant to raise awareness about the red-shanked doucs, Son Tra Nature Reserve, and the threats have resulted in two exhibitions. There are plans to continue a school based mobile exhibition, and expand the school curriculum to include Son Tra and the doucs. A VTV2 film has been produced on the Son Tra doucs and this project.

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Road construction on Son Tra is massive. This photo shows a small slope being cut into the forest. Doucs are not able to cross these large roads with exposed slopes. You can also see how vulnerable the doucs are, in some areas groups have been separated by road construction.

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PhD ABSTRACTS Human and Non-Human Primate Preferences for Faces and Facial Attractiveness For humans and non-human primates (NHPs) the face represents a particularly important source of social information providing a means of conspecific recognition and cues to personal details including sex, age, and emotional state. The human face may also be fundamental in the transmission to conspecifics of other forms of socially relevant information including the display of facial traits associated with sexual attraction and mate choice. A wealth of experimental literature indicates that humans display robust preferences for certain facial traits associated with facial attractiveness including preferences for bilateral facial symmetry, facial averageness and sexually dimorphic faces and facial features. It is thought that these preferences have evolved via sexual selection, and may be adaptive, due to the role that these specific facial features play in reliably signalling to others the possession of heritable genetic quality or ‘good genes’. Therefore, from an evolutionary perspective, it is possible that certain facial preferences may represent an evolutionary adaptation for the selection of potential mate quality. However, despite similarities between human and NHP face processing and recognition abilities, the shared evolutionary history and social importance of faces to primates in general, and the potential importance of these preferences in the mate choice decisions of NHPs, very little research has investigated the extent to which NHPs display comparable preferences to humans for these specific facial traits. Consequently, the aim of the following thesis was to comparatively assess the general and more specific preferences that humans and NHPs display for faces and for traits associated with facial attractiveness. Data were compiled from preference studies examining the visual preferences displayed by two species of NHP (brown capuchins (Cebus apella) and chimpanzees (Pan troglodytes)) for conspecific faces manipulated for those facial traits associated with attractiveness, and from a single study of brown capuchins examining their general visual preferences for various types of facial information. Comparative preference studies were also conducted upon human adults and infants examining the visual and declared preferences that they display for manipulations of facial attractiveness. Data showed that despite possessing general preferences for certain faces and facial information, generally NHPs displayed no significant preferences for those facial traits thought to influences judgements of attractiveness in humans. Possible reasons for this absence of preference for these particular

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facial traits and the evolutionary implications of these findings are discussed. Jack Alexander Fernall Griffey University of Stirling Nesting and Night-time Behaviours of Captive Chimpanzees (Pan troglodytes) Studies of nesting behaviours of free-ranging apes typically focus on ecological variables such as preferred tree species and areas within the home range, heights of nests, and nest group sizes. However, nesting in captive apes is rarely studied, despite the ubiquity of this sleep-related behaviour. The paucity of field data is often attributed to the inherent difficulty in observing what is essentially a night-time behaviour. Captive settings can provide researchers with an ideal opportunity to record nesting and sleep-related behaviours, yet such research on captive apes is also scant. Topics addressed include current practices in zoos regarding conditions for sleep in great apes, the potential effects of social and environmental factors on sleep site selection, the motor patterns involved in nest construction, preferred nesting structures and substrates, and nocturnal behaviours. This thesis documented and empirically tested hypotheses concerning nest-related activities in captive chimpanzees, with an aim to generate practical recommendations for enclosure design, sleeping areas, sleeping structures, and nesting substrates that have implications for the welfare of captive apes. As with the few reports that already exist, most chimpanzees in this research frequently constructed night nests. When building a nest, some techniques appeared to be universal across individuals and groups, where others were group-specific or occasionally characteristic of only certain individuals. An experiment showed that specific materials are preferred over others for nest building. Many chimpanzees appeared to express persistent preferences for particular sleeping sites, and for some this was to maintain proximity to kin or other closely bonded individuals. In one group, individual sleeping site preferences changed across seasons, although again this was subject to individual differences. Video analyses of night-time behaviours demonstrated that, although nests/sleep sites are primarily used for rest subsequent to retirement, a number of social and non-social activities were performed throughout the night. In conjunction with analysis of postural and orientation shifts, these data are unique in describing the nocturnal behaviours of chimpanzees out with a laboratory setting.

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Several aspects of nest-related behaviours showed a high degree of inter-and intra-group variation. Although this cautions against generalising findings across captive populations, research of this type has applied implications for the management of captive ape species, and can add to our as-yet meagre understanding of their nest and sleep-related behaviours. Louise Claire Lock University of Stirling Emerging Language: Cognition and Gestural Communication in Wild and Language-trained Chimpanzees (Pan troglodytes) An important element in understanding the evolutionary origin of human language is to explore homologous traits in cognition and communication between primates and humans. One proposed modality of language evolution is that of gestural communication, defined as communicative movements of hands without using or touching objects. While homologies between primate calls and language have been relatively well explored, we still have a limited understanding of how cognitive abilities may have shaped the characteristics of primate gestures. Chimpanzees (Pan troglodytes) are our closest living relatives and display some complex cognitive skills in various aspects of their gestural behaviour in captivity. However, it is not yet currently clear to what extent these abilities seen in captive apes are typical of chimpanzees in general and to what extent cognitive capacities observed in captive chimpanzees have been enhanced by the socio-cultural environment of captivity such as language training. In this Ph.D. research, I investigated the cognitive skills underlying gestural communication in both wild and language trained chimpanzees, with a special focus on the repertoire and the intentionality of production and comprehension. The study of cognitive skills underlying the production of the repertoire and the role of intentionality is important because these skills are cognitively demanding and are a prerequisite in human infants for their ability to acquire language. My research suggests that chimpanzee gestural communication is cognitively complex and may be homologous with the cognitive skills evident in pre-verbal infants on the cusp of language acquisition albeit only in imperative domain. Chimpanzees display a multifaceted and complex signal repertoire of manual gestures. These gestures are the prototypes, within which there is variation, and between which the boundaries are not clear-cut, but there is gradation apparent along several morphological components. Both wild and language trained chimpanzees communicate intentionally about their perceived desires and the actions that they want the recipients to undertake. They do not just express their emotions, but they communicate flexibly by adjusting their communicative tactics in response to the comprehension states of the

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recipient. Whilst chimpanzees communicate their intentions flexibly, the messages conveyed are specific. However, recipients comprehend gestures flexibly in light of the signaller’s overall intentions. Whilst wild and language trained chimpanzee gestural communication revealed similar cognitive characteristics, language trained chimpanzees outperformed wild apes in that they had ability to use signals which made distinctions that human deictic words can make. Whilst these differences between wild and language trained chimpanzees may be due to the different methodological approaches used, it is conceivable that language training may have influenced captive ape cognitive skills in the representational domain. These results from wild and language trained chimpanzees indicate that chimpanzees possess some form of cognitive skills necessary for language development and that cognitive skills underlying repertoire and use in chimpanzees are a shared capacity between humans, other apes and a common ancestor. These findings render theories of the gestural origins of language more plausible. Anna Roberts University of Stirling

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BOOK REVIEWS Please note: We actively encourage our members to volunteer reviews or proposals of relevant titles. If you would like to suggest a book or offer a review, please contact Book Reviews Editor Kara Moses on <mail@karamoses.com>. THE CHIMPANZEES OF BOSSOU AND NIMBA Tetsuro Matsuzawa, Tatyana Humle and Yukimaru Sugiyama (Eds.) (2011) Springer ISBN 978-4-431-53920-9 (Hardback) £135 Bossou has a special place in primatology and in the study of human evolution. It was not the first detailed investigation of wild chimpanzees, nor is it the most ‘wild’ of settings given the close co-existence of humans and apes in this part of Guinea. The significance of this study site lies in the habit of chimpanzees there to use stones as nut-cracking tools, and in the vision of the scientist who initiated the long-term project at Bossou, Yukimaru Sugiyama, to make that chimpanzee habit the focus of the project. This latest addition to the Primatology Monographs series provides an impressive overview of the varied approaches that have been used by a multitude of field workers to examine aspects of the stone-tool culture at Bossou and several other sites. From observational to experimental work both in the field and in captivity, to the recent application to archaeological survey techniques, the large number of chapters devoted to this theme (25% of all 40 chapters) is a reflection of the impact Bossou chimpanzees and their students have had on our understanding of the evolutionary origins of human technology. Despite the significance of the work on tool-use at Bossou, I suspect that the chapters on tools and culture would not be the main reason you decide to buy this expensive volume. Most of the chapters are rather concise and provide clear and well-organised synthesis of findings published over the years in greater detail in various journals. A compendium like this is great if you are just starting out in primatology and want to get your bearings in the field of chimpanzee tool-use and culture. It might be less useful if you are already familiar with this work and were looking for some fresh data on chimpanzees. If you belong to the latter category of readers you might find satisfaction elsewhere in the book. The Chimpanzees of Bossou and Nimba is structured into seven parts. Parts I and II provide a general introduction to the study site and population. The

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background on ecology and socio-ecology is well summarised but rather limited (chapter two). This is not surprising given that there has been less of an emphasis on such topics at Bossou (more information on ecology appears later in chapter 22 on crop-raiding). Sugiyama and Fujita’s chapter three on demography and reproduction, however, stands out as one of the few in this book that make the most of the longevity of the research project. They summarise previous findings and augment those with data on community composition through 2009, spanning 33 years in total. The picture which emerges is of a relatively small, isolated and highly gregarious community of about 20 individuals, that can easily be devastated by flu-like outbreaks, such as the one that reduced the community size to an all-time low of 12 individuals in 2003 (only up to 13 in 2010). Such insight can only be achieved through long-term efforts and the history of chimpanzee research and conservation at Bossou and in Guniea is the subject of several fascinating chapters (chapters one, four and five). Parts III and IV contain the material dealing with tool-use and culture, discussed earlier. Matsuzawa’s chapter 16 provides both an inspiring description of the logic and benefits of field experiments and an overall summary of how observations at the ‘Bureau’, the outdoor laboratory at Bossou played a key role for some of the most significant discoveries made by researchers there. This chapter also hints at the huge potential for longitudinal analyses on the ontogeny of tool-use offered by the vast video archive on nut-cracking spanning 20+ years. Part V combines five chapters under the heading of ‘Social Life and Social Intelligence’. They cover crop-raiding, road-crossing, object play, carrying mummified infants, as well as a study of the differences in subtle social behaviours between Bossou and Mahale chimpanzees. Without detracting from the quality of the individual chapters in this part, it would have been good to see more on the topic of ‘social life’. Given the small size of this community and the duration of research at Bossou, a long-term analysis of the dynamics of relationships and grouping would have provided a fascinating comparison to other chimpanzee communities. The final two sections of the book, part VI on adjacent communities and part VII on conservation, are the ones I found most engaging. Between them, fourteen chapters cover relatively recent work that is less well known than research presented earlier in the book. Importantly, these chapters set the Bossou chimpanzees in a broader regional perspective. Ongoing work in the Nimba Mountains, the Diécké Forest and Kpayee-Lepula show not just fascinating variation in the cultural behaviour of apes but also in the attitudes of local human populations towards them. While in some places chimpanzees are revered and protected, they are quite literally fair game in others. Connecting the isolated habitat of the Bossou chimpanzees to the forests of Nimba some 10 km away, via an ambitious reforestation project

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(chapter 37), is just one example of how pure research is often inextricably linked to conservation. It also is a sobering reminder that, once cleared, a forest can take a very long time and much human effort to recover. Last but not least, this book comes with a DVD. The effort taken in selecting clips for this compilation is evident. The videos are organised in seven comprehensive categories: Nut-cracking, Learning nut-cracking, Other tool-uses, The habitat and village, Play/Mother-offspring, Social relationships and Communication/other. While the book chapters and the videos could have been integrated better (by providing references to relevant videos throughout the text), the DVD is a much appreciated ‘bonus feature’, complete with informative subtitles and action replays. Having access to such material will allow researchers working at other field sites to compare notes on local behavioural idiosyncrasies. The videos also bring the material discussed in the book to life in a way that no amount of bar-charts, tables or photos ever will. The book/DVD combination will give you a very immediate sense of how lucky the researchers of Bossou and Nimba are to work with such fascinating animals and it is certainly worth having in one’s library. Alexander V. Georgiev Harvard University THE JAPANESE MACAQUES N. Nakagawa, M. Nakamichi, H. Sugiura (Eds.) (2010) Springer ISBN 978-4-431-53885-1 (Hardback) £126 This multi-authored volume was produced to commemorate the 23rd Congress of the International Primatological Society that was held in Kyoto, Japan in 2010. It represents a series of review chapters about major themes in primatology that go back to the earliest behavioural studies in the 1940s. The book starts with a series of spectacular colour photographs illustrating a stunning range of habitats and behaviours for this species including the now classic snow and hot-spring images, as well as food washing and stone handling. Perhaps the strangest picture is the group of macaques spelling out the Japanese word for monkey (��) by aggregating in a pattern of distributed food. Subsequently the book is divided into four broad categories: Overview of Field Research and Related Studies; Ecology and Conservation; Behaviours and Social relationships; and Recent Topics from Unique Approaches – although the grouping seems a little arbitrary in places. The title of the first section is somewhat misleading: although it does discuss the results of field research, it is actually rather less about specific field studies than material in subsequent chapters and in fact covers the

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history of Japanese macaque research, morphology and genetics. Chapter one is fascinating and documents the various ground-breaking observations that have been made about Japanese macaque behaviour over the years. Japanese macaques are unusual in that they live in a very densely populated, highly industrialised country and so human interaction is unavoidable for many populations. This has led to some very specific problems such as hybridisation with imported species that have been released into the wild following the closure of monkey parks. Chapter two is a very data-heavy chapter that includes extensive tables of measurements of Japanese macaques and shows the differences in these dimensions between provisioned and wild populations at different age classes. What is particularly interesting is the coverage of the changes associated with old age which are rarely documented for non-human primate species. Chapter three discusses the genetic differences between the various populations of Japanese macaque and their evolutionary origin from first colonisation which is presumed to have occurred approximately half a million years ago. Analyses use blood proteins, as well as mitochondrial and Y-chromosome genetics. Section two covers the ecological adaptations of different populations. Chapter four concentrates primarily on habitat and on regional variations in forest type, living in grassland, and the effects of logging. Chapter five, in contrast, concentrates specifically on feeding ecology and especially the seasonal variability that a temperate primate needs to cope with. This includes food choice, activity budgets, ranging behaviour, and how this varies by age and sex. Chapter six looks at the importance of Japanese macaques as seed dispersers with detailed species lists for seeds found in macaque faeces as well as evaluations of dispersion distances and germination effects. Chapter seven takes a different tack and reviews the conservation status of the Japanese macaque populations. Although this species is rated as ‘Least Concern’ by the IUCN Red List, it is noted that many populations within Japan are not well studied and that there have been considerable changes in population in the last 100 years. Crop-raiding is identified as a major problem that needs to be resolved. Section three starts with chapter eight which investigates vocal communication. This chapter analyses the variation of call structure and use between populations and discusses its possible role. Chapter nine introduces the idea of cultural inheritance in Japanese macaques. Cultural transmission of behaviour was first suggested in 1952 and since then numerous examples have been documented, primarily in provisioned Japanese macaque groups. Stone handling behaviour is one example of a solitary object play activity with no apparent adaptive value and has been the subject of detailed analysis since 1979. The bulk of this chapter is spent on a very detailed account of this behaviour and the various factors that influence its prevalence and transmission. Chapter ten concerns male and female mating

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strategies. Japanese macaques have multi-male multi-female groups which allows for complex strategy interactions between the sexes which have been extensively studied. Chapter 11 concerns the changes seen in female behaviour as they progress from infanthood through to old age, and discusses the information that can be obtained from long-term field studies. Chapter 12 continues on the theme of social structure but concentrates on the differences seen between different study populations. The final section is rather more varied than the others. Chapter 13 investigates behaviour-related candidate genes that have been identified in humans to see whether polymorphisms at these loci have demonstrable effects within Japanese macaques. Chapter 14 looks at the fatty acid content of the food eaten by Japanese macaques and contains a great deal of fatty acid composition data for a large variety of plants. Chapter 15 looks at the parasite loads of different populations of Japanese macaques and also whether there is any evidence of anti-parasite selection of plant foods. Again there is a very detailed table summarising the anti-parasitic activity of a large range of plants. Chapter 16 considers the recent history of Japanese macaques’ population numbers in more detail and whether there is evidence of a habitat bottleneck. Clearly these animals are relatively widespread within the country but can only cope with certain habitats which have been restricted in the past and may well become so again in the future. Chapter 17 discusses the effects of changes in management regimes on the impact of human-macaque conflicts. Of particular interest is the farmers' perceptions of agriculture in an area where it is not essential for primary food production but often considered as a habit or a hobby. Chapter 18 considers the role of play among juvenile Japanese macaques and considers the evidence for semantics among monkeys. The final chapter considers what can learnt from modelling social dynamics and the ecological value of complex social systems. All in all, this is a very dense book in terms of the amount of material covered in what is a relatively slim volume. The chapters are often fairly short and provide a rather terse introduction to each subject area followed by an enormous list of references, which quite often refer to papers in Japanese. It is certainly a very authoritative volume. The back cover claims that this is the most comprehensive book ever published in English on Japanese macaques and I see no reason to doubt this. There is a huge range of material here that will be of use to researchers in a great many fields of primatology. This is certainly a book that should be in everyone's institution library, and anyone interested in macaque biology should consider purchasing a copy (or at least accessing the relevant chapters online if your institution has the appropriate subscription). Bill Sellers University of Manchester

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MONKEYS ON THE EDGE: ECOLOGY AND MANAGEMENT OF LONG-TAILED MACAQUES AND THEIR INTERFACE WITH HUMANS M.G. Gumert, A. Fuentes & L. Jones-Engel (Eds) (2011) Cambridge University Press ISBN-13: 9780521764339 (Hardback) £60 Long-tailed macaques have one of the widest distributions of any non-human primate along with the dubious honour of being the world’s most invasive. They are well known for their ability to live commensally alongside humans, a situation which frequently culminates in negative consequences for one or both. This book sums up current knowledge of the species’ wild status and its apparent success as a colonising and commensal species. The employments of various strategies of conflict reduction at the human-macaque interface are described and further recommendations for research are made. Gumert et al. give a useful introduction to the species and the problems it currently faces in the wild, one of which is its popularity as a biomedical model which has led to an alarming decline in the species over large areas of its range. This decline is vividly described in the chapter on the species’ status in Laos and Cambodia. The authors state that the sheer scale of the capture of wild long-tailed macaques for the biomedical industry accompanied by a very high mortality rate will eventually mean the demise of whole populations in a region already under pressure from habitat destruction for large infrastructure projects as well as agriculture. The long-tailed macaque preference of living on the forest edge inevitably leads it into close proximity with humans and is the focus of the second part of the book. This interface is encouraged by public feeding of the monkeys for different reasons. Two chapters demonstrate the very different human reactions to this human-primate interface, negative on a university campus in Malaysia, and more tolerant in Bali where feeding the macaques is a tourist attraction, so although macaque numbers have increased so has the revenue the monkeys’ presence generates through tourism. The situation for the introduced long-tailed macaques of Mauritius is discussed by Sussman et al. The presence of the species on this island and the subsequent damage it is alleged to do to the endemic flora and fauna is a well-known conservation problem. To decrease the amount of human-macaque conflict through crop raiding etc, thousands of animals were captured for export and the population crashed from around 35,000 animals to around 10,000 in 2008. Another introduced population on Ngeaur Island is discussed within the context of the island’s colonial past, its other invasive species and future development. Here there is a complex situation where macaques are blamed for much crop loss and damage actually caused

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by other introduced species due to their association with the island’s colonial past. A second chapter on the introduced Mauritian macaques recounts the reasons why it has achieved pest status on the island and describes how farming and selling long-tailed macaques to the biomedical industry is supporting conservation efforts in Madagascar and Mauritius itself. This particular company’s efforts are laudable in wanting to “put something back” into species conservation but there are probably many conservationists, myself included, who would be uncomfortable with such an arrangement. There are many chapters in this book which lead me to wonder whether the long-tailed macaque itself is already in need of conservation funding in some parts of its range. On a positive note about species recovery at least, Southwick and Siddiq report on 50 years of research on rhesus macaques in northern India. Rhesus macaques have also been the subject of capture for medical export and in northern India, the population plummeted due to over-exploitation. However, such export is now completely banned and the rhesus macaque population has soared. This positive development has unfortunate consequences due to the huge increase in human-macaque conflict which has led to human mortalities in cities due to the fear the species can inspire amongst urban dwellers. This chapter touches on the complicated cultural and religious situation existing in India due to the behaviour and numbers of what is a religiously significant animal and the ethically demanding necessity of controlling its numbers without culling. Jones-Engel et al. examine possibilities for developing sustainable human-macaque communities by examining the many ways being used to mitigate human-macaque conflict. Case studies from Singapore and Hong Kong, where long-tailed and rhesus hybrids occur, are the subject of sterilisation programmes as well as the focus of public education campaigns. Singapore’s strict implementation of a non-feeding policy means they have some of the lowest levels of human-macaque conflicts in comparison to other areas of commensal macaque distribution. Summing up, the editors draw attention to the lack of information on sub-specific differences in the species and call for more research on the ecology and demographics of wild populations. This book is a timely reminder of what can happen to a “weed” macaque which has always seemingly occurred in large numbers over a huge geographical range. This book is in time to encourage more primatologists and conservationists to look more closely at what may be the world’s most invasive primate but which is also one of the most adaptable and one of the most exploited by the biomedical industry. Siân S. Waters Durham University

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PRIMATES OF GASHAKA: SOCIOECOLOGY AND CONSERVATION IN NIGERIA’S BIODIVERSITY HOTSPOT Volker Sommer and Caroline Ross (Eds) (2011) Springer ISBN 978-1-4419-7402-0 (Hardback) £117 This is an important publication about a little-known and only recently explored and studied area for African primates. A biodiversity hotspot still largely unexplored by field scientists, Gashaka Gumti National Park in eastern Nigeria holds important populations of West African chimpanzee (Pan troglodytes vellerosus, now ellioti), olive baboon (Papio anubis), forest Guenon and Colobus species. This area of Guinea savannah is described as one of the least known iconic primate habitats. Volker Sommer, an editor and a listed contributor to several of the chapters, has developed field study work with primates in Gashaka Gumti National Park over more than a decade and this has led to a permanent presence, the Gashaka Primate Project. Through their field station, established in 2000, and developing global network of workers and supporters, the project is able to address the conservation issues of the area, in particular the continued survival of the primate populations. The book reports on the primate populations at Gashaka, particularly chimpanzees and baboons, and places the populations of these species at Gashaka into a continental perspective. The sections covering the studies on chimpanzees include a detailed review of the conservation issues in eastern Nigeria, a review of cultural primatology mainly from insectivory and tool use from several field sites across Africa, and a comparative study with bonobos (Pan paniscus) from Salonga NP in Democratic Republic of Congo. The Nigerian-Cameroon chimpanzee P. t. vellerosus, the most genetically distinct subspecies of chimpanzee, survives in diminishing numbers of 3–5,000. From Vortex models of the numbers of animals arriving in sanctuaries, hunting pressure is calculated at an order of 2–14 times greater than sustainability. The Gashaka population (a contiguous population of approximately 1,000) is the only one that is not heavily hunted and there are suggestions to raise the conservation status of this subspecies from ‘endangered’ to ‘critically endangered’. Unlike at many other study sites such as Gombe, Tai and Budongo, there are no long-term detailed observational studies reported of the Gashaka chimpanzees, as they are rarely seen and not habituated to observers. Detailed field studies are reported which show varied and wide spread tool use in relation to the high degree of insectivory. Gashaka chimpanzees feed on army ants from nests but never eat termites. Termites and large-shelled

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nuts occur at Gashaka but in contrast to other sites, where these are consumed, Gashaka chimpanzees appear to show cultural difference in their feeding behaviour. A taxonomic review of P. t. vellerosus (Oates et al, 2008) was taking place as this book was in preparation and although mentioned in chapters one and 14, the revised subspecies name of P. t. ellioti is not used. The sections dealing with baboons include a detailed and current phylogeographical study of all species across Africa, crop-raiding, differences in life history of Gashaka baboons and flexibility in calling behaviour. Comparative studies, which are reported, have been neatly facilitated by detailed long-term studies of two troops in particular, one which regularly crop-raids and another which utilises only wild food. Crop-raiding is reported to bring life history benefits but other dietary influences i.e. the consumption of the African black plum is shown to have a contraceptive effect on these baboons. Other primate species are reported in the detailed habitat survey but a field study of only one other, the Putty-nosed monkey (Cercopithecus nictitans martinii), a species of Guenon, is reported. In this study, alarm-calling behaviour is shown to have interesting differences compared to other Guenon species. Other parts of the book give an account of the Ethno-Primatology of the region, the Bush Pharmacy, the Fulnai pastoralist people, and the future prospects for the Nigerian-Cameroon chimpanzee subspecies survival. In Nigeria’s Taraba region chimpanzees are protected in some societies by food taboos, whilst in other situations they are preferentially hunted because of their similarity to humans, which makes their body parts a more powerful ingredient in folk medicine. The fascinating section on the use of plants by human and non-human primates presents a database of over 300 plant species used. Apparent self-medication by leaf swallowing in chimpanzees is reported. Fulnai pastoralist people have occupied the grassland sections of the park for over 20 years and now their diversification into enclaves of subsistence farming is increasing the areas of human wildlife conflict in the National Park. Although an edited volume, this book is more than just an account of studies of primate species in the area because it addresses and considers in detail the issues for biodiversity conservation. It achieves the stated aim of the editors for the contributions to be of an interdisciplinary dimension from across the African continent. It celebrates the development and the current success of the work at Gashaka, by careful reference to all involved over time and those currently involved at all levels of responsibility. All sections

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of the book are well illustrated by figures, maps, and photographs. Each chapter is well supported by references and several chapters have useful appendices. This volume is published at the tenth anniversary of Gashaka Primate Project which is now the largest research base of this kind in West Africa. It is to be hoped that Volker Sommer and his collaborators can find funding and ways to continue the project and conserve these primate populations in perpetuity. In common with other similar scientific publications this book has a high price which may unfortunately make it unobtainable to some except as a library loan. Together with the recently published Primates of West Africa, a field guide by John Oates (Conservation International 2011), Primates of Gashaka makes excellent source material and much more, for those interested in nature conservation and primates in West Africa. Ray Heaton Ape Alliance UNDERSTANDING ANIMAL WELFARE: THE SCIENCE IN ITS CULTURAL CONTEXT David Fraser (2008) Wiley ISBN: 978-1-4051-3695-2 (Softback) £37.99 Animals are integral to all human societies and their presence is ubiquitous. The synthesis of how human values and animal-welfare science have developed together is a great achievement and provides essential reading for all interested in animal welfare, but also anthropologists and the growing number of people captivated by human-animal interactions. Despite the study of animal welfare science being relatively new, the methods and emphasis employed by those in this field have managed to diversify remarkably. In his Understanding of Animal Welfare, Fraser provides three books in one; the first section is a highly engaging review of the development of animal-welfare science, which outlines how socio-economic factors and subjective opinions have shaped the field; the second section provides complete and easily accessible reviews of the parameters measured to assess animal welfare, including examples from key studies in the area; and finally, the third section attempts the impossible and provides the reader with thoughts on how to interpret data in light of the social context in which they are collected. The first section of this book provides an engrossing historical review of the evolution and development of animal-welfare science; which was a personal highlight for me. I was particularly intrigued by the concept that welfare

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science was principally determined by subjective notions of what was felt to be most important to animal welfare, which in essence is driven by societal values or needs, including economics and religious views. And, that it was these precepts which then influenced what objective empirical data were collected, analysed and evaluated to measure animal welfare. Thus our highly objective science was founded on principles derived from subjective conjecture. Fraser provides background to the development of the divergent subject which we now recognise as the study of animals’ minds, bodies and natures. By contrast, the second section of the book provides reviews of subjects which most of us working in animal welfare will be familiar with; providing overviews of the different topics which are investigated and the different parameters used in the study of animal welfare, such as the study of abnormal behaviour and stress, and the use of indicators of endocrine function. However, Fraser’s authority on this subject navigates the reader through a maze of contradictory information, and there is a lot to be gained from his interpretation and representation of this information. The broad scope of this section in particular has been deceptively and expertly condensed and is a must-read for anyone interested in collecting data for animal-welfare studies, as he provides succinct discussions of the merits and pitfalls of approaches and indices used in animal-welfare science. Finally, Fraser pulls everything together in a synthesis and pragmatic approach about how to proceed in the study of animal-welfare science. By considering the relative merits of the different approaches used to assess animal welfare, the interplay of objectivity and subjectivity in science and the final section ‘coda’, for those very pushed for time and want a brilliant summation of Fraser’s views and thoughts on the subject; though I would recommend that this is a good refresher and shouldn’t really replace reading this great book in its entirety. On another level, Fraser writes in a companionable and effortless style, which makes this complex subject easy to digest. Never patronising, always interested and interesting, Fraser communicates a wealth of experience and knowledge in the subject of animal-welfare science. I have already recommended this book to my students, and all I come into contact with; I do the same in this review! Vicky Melfi Paignton Zoo Environmental Park

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THE PRIMATE SOCIETY OF GREAT BRITAIN founded on May 22 1967 by the Primate Group of the Zoological Society of London

Founding Council: J.R. NAPIER (President), R.P. MICHAEL (Hon.Sec.), R.J. ANDREW (Hon.Treasurer), E.H. ASHTON, L.D. BROOKES, C.R. COID, P. COTES, J.H. CROOK, J. DAVIES, R.N. T-W-FIENNES, R.A. HINDE, G.H. MANLEY, I. ROWLANDS, A.C. WARREN, L. WEISKRANTZ Past Presidents: 1967 - 1970 J.R. NAPIER 1986 - 1969 B. WOOD 1970 - 1973 R.P. MICHAEL 1989 - 1993 R.I.M. DUNBAR 1973 - 1976 R.N. T-W-FIENNES 1993 - 1998 H.O. BOX 1976 - 1979 M.H. DAY 1998 - 2001 P.C. LEE 1979 - 1982 R.D. MARTIN 2001 - 2005 R. BARTON 1982 - 1986 D.J. CHIVERS 2005 - 2010 A. MACLARNON Past Secretaries: 1967 - 1970 R.P. MICHAEL 1987 - 1990 R.C. HUBRECHT 1970 - 1974 K.R. HOBBS 1990 - 1993 P.C. LEE 1974 - 1975 V. REYNOLDS 1993 - 1996 C. ROSS 1975 - 1978 R.D. MARTIN 1996 - 1999 H. BUCHANAN-SMITH 1978 - 1981 A.F. DIXSON 1999 - 2005 J. LYCETT 1981 - 1985 S.K. BEARDER 2005 - 2010 C. SCHAFFNER 1985 - 1987 H.O. BOX Past Treasurers: 1967 - 1970 R.J. ANDREW 1987 - 1990 S. KINGSLEY 1970 - 1974 A. JOLLY 1990 - 1994 R. CROMPTON 1974 - 1977 D.J. CHIVERS 1994 - 1998 G. R. HOSEY 1977 - 1980 E.B. KEVERNE 1998 - 2002 C. EVANS 1980 - 1984 L. AIELLO 2002 - 2009 R. HILL 1984 - 1987 A. MACLARNON 2009 - 2012 G. BROWN Past Editors: 1974 - 1977 N.R. CHALMERS 1993 - 1996 D. BRANDON-JONES 1977 - 1993 J.C. INGRAM 1996 - 2006 W. SELLERS Osman Hill Memorial Lecturers (established 1977): 1978 M.H. DAY 1996 T. ROWELL 1980 R.A. HINDE 1998 C.B. STRINGER 1982 F. BOURLIERE 2000 A.F. DIXSON 1984 P.M. BUTLER 2002 I. TATTERSALL 1986 J.P. HEARN 2004 C. VAN SCHAIK 1988 H. KUMMER 2006 A. JOLLY 1990 R.D. MARTIN 2008 W. MCGREW 1992 J.H. CROOK 2010 A. WHITEN 1994 R.I.M. DUNBAR Napier Memorial Medal Winners (established 1991): 1991 CHRISTOPHER PRYCE 2003 SUSANNE SHULTZ 1993 MARTA LAHR 2005 CORRI WAITT 1995 CARLOS DREWS 2007 ANNIKA PAUKNER 1997 NICOLA KOYAMA 2009 ERIK WILLEMS 1999 MARK COLLARD 2011 LAUREN BRENT 2001 RUSSELL HILL Occasional Medal Winners (established 1996): 1997 JANE GOODALL CBE: Conservation 2007 CYRIL ROSEN MBE: Conservation 2008 STEPHEN NASH: Special Contributions to Primatology 2011 DAVID CHIVERS: Conservation 2011 DAVID WINDMILL: Special Contributions to Primatology Charles A. Lockwood Medal Winners (established 2009): 2009 CLAIRE SANTORELLI 2010 STEPHEN MONTGOMERY 2011 NIENKE ALBERTS

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Editorial … … … … … … … 1 Report: PSGB Winter Meeting 2011 – Gardeners of the Forest: Primate Ecology and Forest Conservation … … 3 PSGB Spring Meeting 2012 – Port Lympne, Kent Paper Abstracts … … … … … 7 Poster Abstracts … … … … … 17 Future Meetings: Winter 2012 – Primate Biogeography … … … 30 Spring 2013 – TBC, Lincoln … … … … 31 Announcement: marmosetcare.com … … … 32 Announcement: Journal of Zoo and Aquarium Research (JZAR) 32 PhD Abstracts: Human and Non-Human Primate Preferences for Faces and Facial Attractiveness – Jack Griffey … … … 36 Nesting and Night-time Behaviours of Captive Chimpanzees – Louise Lock … … … … … 37 Emerging Language: Cognition and Gestural Communication in Wild and Language-trained Chimpanzees (Pan troglodytes) – Anna Roberts … … … 38 Book Reviews: The Chimpanzees of Bossou and Nimba … … 40 The Japanese Macaques … … … … 42 Monkeys on the Edge: Ecology and Management of Long- tailed Macaques and Their Interface with Humans … 45 Primates of Gashaka: Socioecology and Conservation in Nigeria’s Biodiversity Hotspot … … … 47 Understanding Animal Welfare: The Science in its Cultural Context … … … … … 49 ,661�����������