2009 Vol 99.pdf (2.45mb) - Primate Society of Great Britain

Primate Society of Great Britain
No. 99
OCTOBER 2009

THE PRIMATE SOCIETY OF GREAT BRITAIN Registered Charity No. 290185
Officers:
President: Prof. Ann Maclarnon (06)
(Centre for Research in Evolutionary Anthropology,
School of Human & Life Sciences, Roehampton
University, Holybourne Avenue, London SW15 4JD)
Hon. Secretary: Dr Colleen Schaffner (06)
(Psychology Department, University of Chester,
Chester CH1 4BJ)
Hon. Treasurer and Dr Gillian Brown (09)
Membership Secretary:
School of Psychology, University of St Andrews,
St Mary’s College, South Street,
St Andrews, Fife, Scotland, KY16 9JP, UK
e-mail: membership@psgb.org
Council Members:
Ms K. Balolia (University College London – Student Representative (09)
Prof. H. Buchanan-Smith (Stirling) (08)
Dr H. Chatterjee (University College, London) (09)
Dr C. Cunningham (University of Abertay Dundee) – Book Review Editor (09)
Prof. R. Dunbar (Oxford University) (09)
Dr S. Elton (Hull York Medical School) – Marketing Committee Convenor (08)
Dr A. Fletcher (University of Chester) (09)
Dr A. Korstjens (Bournemouth) (07)
Dr N. Newton-Fisher (Kent) (08)
Dr V. Nijman (Oxford Brookes University) (09)
G. Norton (Anglia Ruskin) (07)
Dr S. Pika (Manchester) (08)
Dr K. Pullen (Paignton Zoo) (09)
Dr J. Setchell (Durham) – Information Officer (07)
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 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, Primate Eye
Mr C. Rosen MBE (IPPL) - Financial Advisor
Dr A. Smith (Anglia Ruskin) – Meetings Officer
Honorary Auditors: Messrs Morris & Co., Chester
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The content of Primate Eye is printed on recycled paper by Top Copy, Bristol BS16 6JE

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EDITORIAL
I have a confession to make – I’m addicted to popular science books. I find
it hard to pass a Waterstone’s without whipping out my credit card (and
loyalty card!) for yet another paperback on primates, evolution,
palaeontology, evolutionary psychology, etc. After all, the book that
effectively started primatology, On the Origin of Species by Means of
Natural Selection (C. Darwin, 1859, John Murray, London), was a popular
science tome. Sadly, not all of the recent titles live up to this illustrious
precursor.
I recently had the bad luck to shell out proper folding money for the
paperback edition of Terence Kealey’s Sex, Science & Profits (2009,
Vintage, London). It promised so much; the subtitle is ‘How People
Evolved to Make Money’. I had always thought that evolutionary
economics had great potential – there could be loads of recent research on
biological markets and complex systems such as networks that could inform
such a work. Instead, it turned out to be an exemplar of a worrying trend –
the tendency for authors to take a fairly transparent polemic and ‘sex it up’
(in this case, literally) with a bit of evolution. The central theme of the book
gradually emerges: public funding of science is bad. It is effectively a profree-market
tirade claiming that we slow down the ‘progress’ of civilization
(or his favourite surrogate, technological innovation) by using tax revenues
to pay for research, with some cherry-picked bits of the (mostly older and
often popular) literature sprinkled in to provide a scientific gloss. It doesn’t
wash, for a multitude of reasons, but even the ‘science-y’ bits fall down; his
quoting of S. J. Gould conflates punctuated equilibria with spandrels, and
the attached footnote cites a 1980s book chapter on eighteenth century US
technology! ∗
On the other hand, I also picked up the paperback of Ben Goldacre’s
wonderful Bad Science (2009, Harper Perennial, London). Here is an
engaging, entertaining work full of really good analyses of really bad
science and science reporting. Rather than try to use science as a
justification for a particular point of view, a position that Kealey shares with
the eugenics advocates he censures, Goldacre gives the reader the benefit of
the doubt in terms of being able to understand the concepts involved in the
original research (including statistics), and provides some careful, readable
explanations of the scientific enterprise. His examples show how it can all
go wrong, and be massively misunderstood by a public (and journalists) not
familiar with the process, but he cites enough truly worthy works to
convince the reader that it doesn’t have to be so always. And it’s
∗ To be scrupulously honest, the correct source is listed, as well, but all of
the endnotes for this chapter are listed in seemingly random order.

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occasionally funny, too. Reading this work restored my faith in popular
science books generally and ensured that I’d be racking up Waterstone’s
loyalty points for some time to come.
As for the present issue of Primate Eye, we’ve got plenty to keep you
occupied – meeting abstracts, meeting announcements, ‘proper’ book
reviews, and all of the other goodies you’ve come to expect. Don’t forget, if
you’ve got your own announcement, query, tirade, or point of view that
may of interest to the greater primatological community, don’t hesitate to
contact us.
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 January 2010. 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 Anthropology
School of Human and Life Sciences
Whitelands College
Roehampton University
Holybourne Avenue
London SW15 4JD
Email:
Tel: Intl. +44 (0)20 8392 3726
Fax: Intl. +44 (0)20 8392 3527
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 .

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REPORT: PSGB SPRING MEETING 2009
Form & Function
16/17 April 2009, Bournemouth University
The 2009 Spring Meeting of the Primate Society of Great Britain, held at
the Landesdown Campus of Bournemouth University on the 16th and 17th
of April, was on the theme of ‘Form and Function’. The organisers Mandy
Korstjens, Jo Setchell and Bridget Waller successfully put together a
stimulating array of speakers and events, a feat greatly appreciated by those
in attendance. What would have once, perhaps, been anatomical overload
was instead a vibrant meeting integrating behaviour and ecology into the
study of morphology. In its entirety, the meeting included extended talks
from five plenary speakers, an invited public lecture by Prof. Jan A.R.A.M.
van Hooff, twelve oral presentations (many by students), two workshops, a
poster session, the first Charles A. Lockwood Student Prize for best student
oral presentation, and an excursion to Marwell Zoo.
Five plenary speakers set the tone of the meeting: Gabrielle Macho offered
unique insights into extant great ape and extinct hominid morphology and
behaviour, Sarah Elton highlighted the contribution of a multitude of factors
to guenon morphology and variation in both the crania and postcrania, Todd
Rae used cercopithecine postcrania to outline problems (and solutions?)
pertaining to phylogeny and function in morphological studies, Robin
Crompton reported on recent developments in understanding the
biomechanics of the hominids responsible for the Laetoli footprints, and
Robin Dunbar outlined the need for, and use of, time budget models to
better understand primate evolution.
Organisers, volunteers and award winners from the PSGB Spring Meeting 2009:
Form & Function. L-R: Claire Santorelli (Charles A. Lockwood Student Prize
recipient), Catherine Martine and Emmanuelle Amiral (volunteers),
Mandy Korstjens and Jo Setchell (organisers),
Emma Nelson (Student Poster Prize recipient), and Bridget Waller (organiser).

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The shorter talks, comprising the majority of the meeting, were on a variety
of topics, not restricted to form and function. Jacklyn Ellis presented results
on stress and anxiety responses in the olive baboons of Gashaka, Lauren
Brent demonstrated the application of social network analysis to rhesus
macaques, and, despite technological problems, Denise Bailey from the
Jane Goodall Institute gave an enthusiastic presentation on their current
work and need for continued financial support. Katharine Balolia examined
the relationships between extended cranial growth and social behaviour in
the great apes, Lisa Riley advocated the necessity of cognitive enrichment
in captive chimpanzees, Nick Davis looked at the effect of various stressors
on captive spider monkeys, and Erica Kempf summarised the nature of
primate interactions with aquatic environments (in no way associated with
the widely derided aquatic ape hypothesis).
Marina Kenyon argued that variation in communication between whitecheeked
gibbons was linked to numerous factors, Emma Nelson inferred an
association between 2D:4D digit ratios and dominance rank for rhesus
macaques, Bill McGrew tested competing hypotheses on factors influencing
allo-parental caretaking in captive cotton-top tamarins, and Mark Harrison
provided evidence that energetic pressures are a driving factor in orangutan
body size evolution. In recognition of the dedication and inspiration that
paleoanthropologist and former Council member Charlie Lockwood
provided to students, this year saw the first Charles A. Lockwood Student
Prize. The deserved winner of the specially designed medal was Claire
Santorelli from the University of Chester, who gave a deeply interesting talk
on vocal traditions and acoustic variation in wild spider monkeys from two
geographically disparate communities in Mexico and Costa Rica.
Over the two days of the conference, several posters, mostly by students,
were exhibited. These covered a range of topics, some of which addressed
the theme of form and function, with others examining behavioural ecology,
conservation and potential research opportunities. Of particular interest was
the mix of work at differing stages: that is, completed, ongoing, exploratory
and future research. The Student Poster Prize was awarded to Emma Nelson
for her study on predicting the social systems of extinct hominid taxa using
digit ratios.
A highlight of the meeting was the public lecture, given by Professor Jan
A.R.A.M. van Hoff from Utrecht University, exploring the shared history of
Darwinian understanding of the natural world and behavioural investigation
of primate facial expressions. Throughout the talk, Professor van Hoff
brought his talk to life with a range of animated facial expressions that
served to illustrate his underlying scientific arguments. As one of the
‘characters’ in primatology, Jan A.R.A.M. van Hoff delighted in amusing
and entertaining his audience with his animation.

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The meeting also included a new venture: student orientated workshops, the
first on presentation skills and the second on geometric morphometrics.
Colleen Schaffner and Todd Rae gave an extremely entertaining
performance on how (and how not!) to give a good presentation. The key
points were: know your audience, avoid wordy slides, keep audience
attention on you rather than your slides, and end with a strong take-home
message. For the morphometrics workshop, Andrea Cardini, with
interjections and additions from Sarah Elton, outlined some of the uses and
potential pitfalls of using increasingly popular geometric morphometrics
techniques for quantifying size and shape. This functioned as both a solid
introduction for novices, whilst giving clarification on trickier aspects of the
associated methodology for those with more experience of the technique.
The meeting ended with a Saturday trip to Marwell Zoo, with the admission
fee kindly waived by the zoo. The visit included a guided tour of, and
behind the scenes access to, the siamang enclosure. Other primates visited
at the zoo included lemurs, tamarins, colobines and macaques. Other
interesting animals included capybaras, okapis, meerkats and giant
anteaters. Towards the end of the excursion a stretched golf cart with
multiple VIPs passed the group, with one youngster shouting “life is good”.
After a very pleasant and intellectually satisfying trip to Bournemouth, who
were we to disagree?
Alex Bjarnason & Katharine Balolia
University College London

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PSGB WINTER MEETING 2009
Primate Stress: Causes, Responses and Consequences
1 st and 2 nd December 2009
Zoological Society of London, Meeting Rooms. ZSL, Regent’s Park,
London
Meeting Outline
This stimulating meeting will focus on state-of-the-art research pertaining to
stress in primates covering behavioural, physiological and neurological
responses to stressors in the physical and social environment both in
captivity and the wild. The consequences of these responses for the
individual’s biological fitness will be assessed. The meeting will consist of
a number of invited presentations from eminent, international scientists in
the field as well as proffered papers spread, in themed sessions, over two
days. There will also be one or more poster sessions. We are delighted to
announce the conference will feature two plenary presentations and several
other invited papers from all from internationally respected researchers.
Call for Early Registration
We are requesting early registrations for this conference. Due to the high
interest in this exciting two-day meeting and the limited capacity at the
venue, we encourage pre-registration to guarantee attendance. Early
registrations will be accepted until 17:00 on 29 th November 2009 as subject
to availability. As the deadline for abstract submission has now past we
regret that we are unable to accept abstracts for oral presentations.
Payment Details
The programme will commence 09:00 on 1 st December and continue until
17:45 on 2 nd December. Registration costs include attendance at the two-day
meeting and all teas and coffees. Additional details will be posted on the
PSGB webpage closer to the date of the conference and e-mailed directly to
registrants. We are able to accept payment either online by credit or debit
card, or by personal cheque.
Online payment: You can pay securely online
with a credit or debit card. You will not be charged any fee for using this
secure, online payment service.
Cheque: Please make a cheque payable to Primate Society of Great Britain
and send it, together with a hard copy of the completed registration form
(below), to:

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Tessa Smith
Department of Biological Sciences
University of Chester
Parkgate Road
Chester CH1 4BJ
Alternatively you could send the form online by pressing the ‘Submit form
only’ option.
You will be sent a receipt when payment has been received.
Student
Member
1-Day 2-
Days
Registration Fees
Student Nonmember
Ordinary /
Associate
Member
1-Day 2-Days 1-Day 2-
Days
Non-member
1-Day
2-Days
Cost £20 £35 £35 £60 £35 £60 £50 £90
Organising Committee
Paul Honess (Oxford University), Stuart Semple (Roehampton
University), Tessa Smith (University of Chester)

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Provisional Programme
The organisers are pleased to present the provisional programme for the
PSGB Winter Meeting 2009. NB. Please note this programme is subject to
change.
1 st December 2009
09:00 Registration
09:45 Opening remarks
10:00 PLENARY
Risk, resilience, and gene x environment
Suomi, Stephen
interactions in primates
11:00 Refreshments & Posters
11:30 INVITED
Enrichment, chronic stress and its Garner, Joseph
impact on research data
12:00 Glucocorticoid responses to environmental Davis, Nick
and social stressors in zoo-housed spider
monkeys
12:30 The effect of weaning stress on the social Eckardt, Winnie
behaviour of mountain gorillas
13:00 Napier Medal Presentation, PSGB AGM & Lunch
14:30 INVITED
Stress revisited: a critical evaluation of Koolhaas, Jaap
the stress concept
15:00 Ratings of zoo chimpanzees (Pan troglodytes)
and orangutans (Pongo pygmaeus and Pongo
abelii) are not anthropomorphic projections
15:30 The influence of visitor density on the Cooper, Tara
behaviour of zoo-housed gorillas (Gorilla
gorilla gorilla) and chimpanzees (Pan
troglodytes)
16:00 Refreshments & Posters
16:30 INVITED
Social subordination stress effects Shively, Carol
behavior, pathophysiology, and disease
risk in adult cynomolgus macaques
17:00 Stress and attentional bias in rhesus Bethell, Emily
macaques
17:30 Subjective well-being is genetically Adams, Mark
correlated with personality in orangutans
18:00 Posters & Close

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2 nd December 2009
08:30 Registration
09:20 Opening remarks
09:30 PLENARY
Primate conservation: synergy among Chapman, Colin
stress, nutrition, climate change and
disease
10:30 Refreshments & Posters
11:00 INVITED
The common marmoset: species-typical Pryce,
stress and stress experiments to
Christopher
understand human disorders
11:30 Sex specific life history and climate- Dirks, Wendy
related stresses recorded in Papio
hamadryas dental tissues
12:00 Impact of human disturbance on stress, Zommers, Zinta
disease and conservation of chimpanzees,
Pan troglodytes, in Budongo Forest,
Uganda
12:30 Lunch & Posters
14:00 INVITED
Distress alleviation in monkeys and apes Aureli, Filipo
14:30 Individual differences in anxiety level Majolo,
affect reconciliation in wild Japanese Bonaventura
macaques (Macaca fuscata yakui)
15:00 Social capital and physiological stress in Brent, Lauren
female rhesus macaques
15:30 Refreshments & Posters
16:00 The use of visual barriers to alleviate Honess, Paul
stress-related hair-pulling in rhesus
macaques (Macaca mulatta)
16:30 Is training for cognitive testing stressful? Herrelko, Elizabeth
A comparison of self-directed behaviours
of chimpanzees (Pan troglodytes)
17:00 The effects of an enhanced socialisation Tasker, Louisa
programme on behaviour, welfare and
cardiac responses of newly acquired
cynomolgus macaques (Macaca
fascicularis) during six-week
acclimatisation period
17:30 Close

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Posters
Was the primate stress system a selective force
for increased hominin intelligence?
Linear enamel hypoplasias in baboons
Integrating measures of behaviour, physical
health and physiology to produce an overall
assessment of welfare in the cynomolgus
macaque (Macaca fascicularis)
Hair loss in captive primates: A long-term four
species comparison
Evidence for differential susceptibility to
putative childhood stress effects: The role of
relationship status
Investigating the role of social interactions on
adrenal activity in female Barbary macaques
(Macaca sylvanus)
Softly, softly, catchy monkeys
Victim behaviour following aggression in
captive Sulawesi crested macaques
Cummins, John
Williams, Chris
Tasker, Louisa
Hopper, Lydia
Boothroyd, Lynda
Edwards, Katie
Owen, Yvonne
Watkin, Nia
ABSTRACTS – ORAL PRESENTATIONS
1st December
PLENARY LECTURE
RISK, RESILIENCE, AND GENE X ENVIRONMENT
INTERACTIONS IN PRIMATES
Stephen J. Suomi
Laboratory of Comparative Ethology, Eunice Kennedy Shriver National
Institute of Child Health and Human Development, National Institutes of
Health, DHHS, Bethesda, MD 20892-7971, USA
Presenter email: ss148k@nih.gov
Recent research with both humans and rhesus monkeys has provided
compelling evidence of gene-environment (G x E) interactions throughout
development. For example, a specific polymorphism of the serotonin
transporter (5-HTT) gene is associated with deficits in infant
neurobehavioral functioning, poor control of aggression and low serotonin
metabolism during juvenile and adolescent development, and excessive
alcohol consumption in early adulthood in monkeys reared with peers but

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not in monkeys reared by their mother. One interpretation of these findings
is that secure attachment relationships somehow confer resiliency to
individuals who carry alleles that may otherwise increase their risk for
adverse developmental outcomes (“maternal buffering”). Similar patterns of
apparent “buffering” have been demonstrated for G x E interactions
involving several other genes with functionally equivalent polymorphisms
in both humans and rhesus monkeys. Recent research has suggested that
much of this “buffering” may be taking place in the context of early face-toface
interactions between rhesus monkey mothers and their infants.
Moreover, the allelic variation seen in these genes in rhesus monkeys and
humans but apparently not in other primate species may actually contribute
to their remarkable adaptability and resilience at the species level.
INVITED LECTURE
ENRICHMENT, CHRONIC STRESS AND ITS IMPACT ON
RESEARCH DATA
Joseph Garner
Animal Sciences, Purdue University, West Lafayette, IN, USA
Presenter email: jgarner@purdue.edu
Behaviour is what makes an animal an animal, and almost every aspect of
animals has evolved as a consequence of behaviour: the digestive system is
dependent on feeding behaviour; the reproductive system upon mating
behaviour; the skeleto-muscular system exists to allow locomotive
behaviour; even the immune system is intimately interwoven with
behaviour. The major role of behaviour is to allow animals to control their
environment, and to survive in environments that are physiologically
inhospitable, or even lethal. Consequently a lack of control over even
innocuous stimuli can induce widespread and devastating stress responses.
Biologically relevant enrichments provide animals the mean to control
stressors in the environment with suitable species-typical behaviours. Thus
properly designed, and experimentally ratified enrichment, should render
animals more normal and less variable. Conversely, barren environments,
which lack any meaningful control for the animal are likely to induce
widespread stress-mediated changes in animals, and such animals will be
abnormal and of little relevance as research models. For example,
experimental vaccines may be ineffective in barren housing, but extremely
effective under enriched conditions, as a product of stress-induced immune
suppression in barren housing. As such, the call for enriched housing to
show that it is no different from barren housing is nonsensical – instead the
burden of proof should be on barren, unenriched housing to show that
animals housed therein are not abnormal. Using mice as an example, this
talk will review these concepts, and illustrate the central role of the animal’s
natural history, sensory world, and behaviour (the animal’s point of view) in

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designing and validating enrichments. In particular, real world examples
will be discussed, where failing to take the animal’s point of view has lead
to the adoption of ‘enrichments’ or ‘refinements’ that actually impair
wellbeing; where taking the animal’s point of view provides superior
solutions to wellbeing issues; and where suitably enriched animals are
demonstrably more ‘normal’ and yield superior research data.
GLUCOCORTICOID RESPONSES TO ENVIRONMENTAL AND
SOCIAL STRESSORS IN ZOO-HOUSED SPIDER MONKEYS
Nick Davis 1,2,3 , Colleen M. Schaffner 1 , Tessa E. Smith 2
1 Department of Psychology, University of Chester, Parkgate Road, Chester,
CH1 4BJ
2 Department of Biology, University of Chester, Parkgate Road, Chester,
CH1 4BJ
3 North of England Zoological Society, Upton-by-Chester, Chester,
CH2 1LH
Presenter email: n.davis@chesterzoo.org
There is minimal quantitative data on the physiological effects of stressors
experienced by zoo-housed primates. Over a 7-year period we monitored
the physiological responses to a variety of environmental and social
stressors in a group of 8-13 spider monkeys (Ateles geoffroyi rufiventris)
housed at Chester Zoo. We assessed the impact of visitors, aggression,
reproductive events, separation and the introduction of a new male on the
spider monkeys’ urinary cortisol. We found that cortisol was positively
associated with increased visitor numbers. For aggression, reproductive
events and separation we investigated the physiological stress responses the
week prior, the day of and the week following an event. Aggressive events
were associated with the largest increase in urinary cortisol with targets
experiencing a seven-fold increase in cortisol on the day of severe
aggression and bystanders having a 20-fold increase on the day of lethal
aggression. In addition, cortisol levels in bystanders were still significantly
elevated the week following lethal aggression. For reproductive events,
cortisol levels were significantly elevated in mothers the week prior to and
the day of birth, and elevated in bystander females on the day of birth. In
terms of animal separations, cortisol levels were significantly elevated in
separated individuals only when separations were greater than 24 hours.
Surprisingly, the introduction of a new male produced little change in
urinary cortisol among the resident females. The findings of our study
contribute to the understanding of the physiological responses to stressors in
a zoo environment and have implications for animal management.
Keywords: zoos, Ateles, stress response

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THE EFFECT OF WEANING STRESS ON THE SOCIAL
BEHAVIOUR OF MOUNTAIN GORILLAS
W. Eckardt 1,2 , A. W. Fletcher 1 , K. Fawcett 2
1 University of Chester, UK, 2 Karisoke Research Center, Dian Fossey
Gorilla Fund International
Presenter email: w.eckardt@chester.ac.uk
The weaning process in primates can be extremely stressful, in particular for
offspring who are weaned at an early age compared to their peers. Infants
often respond to weaning with temporary depression and regression to
infantile behaviour, such as seeking more contact with the mother and
extending the length of suckling bouts. Many studies have investigated
behavioural changes in mother-offspring interactions during the course of
the weaning process. However, changes in social interactions involving
other group members, such as frequency of playing and agonistic behaviour,
and the potential long-term impact of these on the offspring’s social
development have been, in the most part, neglected. In this study, the effect
of weaning on the frequency of playful and agonistic behaviour was
investigated in 16 mountain gorillas (Gorilla beringei beringei) (30-54
months old) living in the Virunga massif (Rwanda). Pre- and post-weaning
behaviour were compared within individuals as well as between individuals
within the same age range. There was found to be a strong decline in play
and an increase in agonistic interactions after weaning, with early-weaned
offspring being more strongly affected. In primates, including humans, the
combination of a decrease in play behaviour and an increase in agonistic
behaviour can be indicative of stress. The potential impact of such
behavioural changes on the offspring’s social development will be
discussed taking the weaning age and sex of offspring into account.
Keywords: weaning stress, wild mountain gorillas, play, agonistic
behaviour
INVITED LECTURE
STRESS REVISITED: A CRITICAL EVALUATION OF THE
STRESS CONCEPT
Jaap M. Koolhaas
Behavioral Physiology, University of Groningen, P.O.Box 14, 9750 AA
Haren, The Netherlands
Presenter email: J.M.Koolhaas@rug.nl
The stress concept has been subject of much scientific debate. This paper
will highlight a biologically oriented framework of interpretation to
evaluate both the adaptive and maladaptive nature of the stress response in
relation to the existing environmental demands. Recent evidence shows that
the physiological response to clearly negative, uncontrollable situations may

14
be as large as the response to positive conditions such as food intake or
sexual behavior. It will be argued that it is not the magnitude of the
physiological response that dissociates a negative from a positive stimulus
but the speed of recovery of the response. Stress-related terminology should
be restricted to uncontrollable and unpredictable conditions. These
conditions may lead to a variety of behavioral and neurobiological changes
in the organism. To facilitate the interpretation of these stress-induced
changes in terms of their adaptive and/or maladaptive nature, the concepts
of regulatory range and regulatory capacity of allostatic mechanisms will be
introduced. These concepts are also helpful in understanding individual
differences in coping style and animal welfare in relation to environmental
demands. In stress research, much can be gained by more carefully
exploiting the biological basis of animals using ecologically relevant
models. This allows a fundamental analysis of factors modulating the
individual adaptive capacity and hence the individual vulnerability to
disease.
Key words: stressor, stress response, adaptation, coping style, allostasis
RATINGS OF ZOO CHIMPANZEES (PAN TROGLODYTES) AND
ORANGUTANS (PONGO PYGMAEUS AND PONGO ABELII) ARE
NOT ANTHROPOMORPHIC PROJECTIONS
Alexander Weiss 1,2 , Mark J. Adams 1
1 Psychology, School of Philosophy, Psychology and Language Sciences,
The University of Edinburgh, 2 Scottish Primate Research Group
Email: alex.weiss@ed.ac.uk
If researchers wish to assess subjective states such as stress in animals, they
will have to go beyond behavioural measures. These measures might
involve hormonal assays whereas others might rely on the judgement or
impressions of researchers and personnel who are intimately familiar with
the individual animals. Despite findings to the contrary, a major barrier to
using the latter is the continuing charge that these types of measures, like
so-called subjective personality ratings, are partly or entirely the product of
anthropomorphic projection. We will demonstrate two new forms of
principal components analysis using existing personality data and test the
hypothesis that personality dimensions in nonhuman primates are the
products of anthropomorphic projections. Subjects were chimpanzees
housed in U.S. and Australian zoos (N=202); chimpanzees housed in
Japanese zoos, research institutes and sanctuaries (N=151); and orangutans
housed in U.S., Canadian, Australian and Singaporean zoos (N=174). We
show that a) the personality dimensions emerging from the covariance
matrix of animal effects do not markedly differ from the dimensions that
were described in previous studies and b) the personality dimensions
emerging from covariance matrix of rater effects differ somewhat from the

15
animal dimensions. We also show that c) the dimensions emerging from
covariance matrix of rater effects differ between the three samples. These
findings are contrary to predictions from the hypothesis that ratings are
anthropomorphic projections, highlight the ways in which raters perceive
personality, and suggest a new method of assessing ratings of personality,
stress, and related constructs.
Keywords: personality, chimpanzees, orangutans, anthropomorphism
THE INFLUENCE OF VISITOR DENSITY ON THE BEHAVIOUR
OF ZOO-HOUSED GORILLAS (GORILLA GORILLA GORILLA)
AND CHIMPANZEES (PAN TROGLODYTES)
Tara Cooper, Deborah L. Wells, Peter G. Hepper
School of Psychology, Queen’s University Belfast, Belfast, BT7 1NN, UK
Presenter email: tjenkins01@qub.ac.uk
Visitors to zoos can influence the behaviour of captive primates, although it
is still unclear whether the size of the human audience has an impact upon
the welfare of such animals. This study explored the effect of fluctuations in
visitor numbers on the behaviour of zoo-housed gorillas and chimpanzees.
Chimpanzees (n=6) and gorillas (n=6) housed in Belfast Zoological
Gardens were studied during three, 4-day long, periods of visitor density.
Period 1 represented a relatively stable phase of visitor density (3000 per day). Visitor
numbers reverted back to relative normality during Period 3 (

16
INVITED LECTURE
SOCIAL SUBORDINATION STRESS EFFECTS BEHAVIOR,
PATHOPHYSIOLOGY, AND DISEASE RISK IN ADULT
CYNOMOLGUS MACAQUES
Carol A. Shively, S.L. Willard, T.B. Clarkson, C.K. Henkel, D.P. Friedman,
T.C. Register
Wake Forest University School of Medicine, Winston-Salem, NC, USA
Presenter email: cshively@wfubmc.edu
While there are benefits to social group-living, there are also costs, one of
which is the stress that can result from the increased social density and
proximity inherent in a gregarious social organization. Social stress may
increase the risk of disease, and promote pathophysiological changes that
increase the risk of disease. We have studied disease susceptibility in adult
female cynomolgus monkeys (Macaca fascicularis) housed in small social
groups of 4 females each. Linear social status hierarchies form and are
stable over long time periods. In order to approximate as closely as possible
the human condition, the monkeys are fed a Western diet containing
moderate amounts of fat and cholesterol. Socially subordinate females
appear behaviourally and physiologically to be stressed relative to
dominants. They receive more aggression, are groomed less, spend more
time vigilant, less time being groomed, and more time alone than
dominants. Subordinates are hypercortisolemic, have higher heart rates in
response to a standardized stressor, suppressed reproductive function, and
decreased dopamine receptor 2 binding potential in the striatum. They are
more likely than dominants to deposit fat in the viscera, develop more
coronary artery atherosclerosis, and exhibit depressive behaviour.
Depressed monkeys are distinct from nondepressed subordinates
behaviourally and physiologically. For example, depressed monkeys, like
depressed human beings, have alterations in serotonin receptor 1a binding
throughout many areas of the brain which mediate emotional responses to
environmental events, and smaller anterior hippocampi. Thus, the stress of
social subordination in these small captive groups has extensive effects on
the behaviour, physiology and health of cynomolgus macaques.
Keywords: Stress, cortisol, serotonin, hippocampus, atherosclerosis,
visceral obesity, depression

17
STRESS AND ATTENTIONAL BIAS IN RHESUS MACAQUES
Emily J. Bethell 1,2 , Stuart Semple 1 , Mandy Holmes 1,2 , Ann MacLarnon 1
1 Centre for Research in Evolutionary Anthropology, Whitelands College,
Roehampton University, Holybourne Avenue, Roehampton, London , SW15
4JD, 2 School of Psychology, Birkbeck College, University of London, Malet
Street, London, WC1E 7HX
Presenter email: E.Bethell@roehampton.ac.uk
Cognitive biases provide a measure of psychological wellbeing in humans.
People high in anxiety, for example, demonstrate a bias to attend to
threatening information. This is known as attentional bias and is implicated
in the onset and maintenance of psychological disorders in humans.
Measuring psychological wellbeing in other species has remained
problematic for welfare researchers. However, it is possible that the
diagnostic and therapeutic cognitive methods used with humans may be
adapted for use with other species. We present data from a novel method for
testing attentional bias in rhesus macaques. Monkeys were shown pairs of
pictures of threatening versus non-threatening conspecific faces. Latency
and duration of gaze towards the faces was coded. Following a stressor
monkeys showed a different pattern of gaze towards face pairs than when
no stressor had been given. We discuss our findings in terms of existing
vigilance and avoidance theories of attentional bias for threatening
information in humans, and suggest the method presented provides a
valuable means of measuring psychological wellbeing in non-human
primates.
Keywords: attentional bias, stress, macaque, psychological wellbeing
SUBJECTIVE WELL-BEING IS GENETICALLY CORRELATED
WITH PERSONALITY IN ORANGUTANS
Mark J. Adams 1 , Alexander Weiss 1,2
1 Psychology, School of Philosophy, Psychology and Language Sciences,
The University of Edinburgh, 2 Scottish Primate Research Group
Presenter email: m.j.adams-2@sms.ed.ac.uk
Subjective well-being, or happiness, captures the balance of positive and
negative moods in an individual, and is related to personality, health, and
stress responses in humans and other primates. Differences in personality
and subjective well-being are described by dimensions that appear to be
conserved across primate lineages. To understand the potential for these
traits to co-evolve, it is essential to estimate their heritabilities and genetic
correlations. Orangutans have been shown to have one well-being
dimension in addition to five personality dimensions: Extraversion,
Dominance, Neuroticism, Agreeableness, and Intellect. We assessed
personality and subjective well-being in 174 zoo-housed orangutans. To

18
reveal the evolvability of personality and subjective well-being in
orangutans, we estimated trait heritability simultaneously with a
multivariate animal model using Markov chain Monte Carlo. Heritability
point estimates for the five personality traits ranged from 0.20 for
Extraversion to .43 for Intellect. The heritability of subjective well-being
was 0.56. Consistent with findings on humans and chimpanzees, orangutan
happiness was genetically correlated with Extraversion (r A =0.54),
Dominance (r A =0.52), and Neuroticism (r A =-0.70). Thus, it is possible for
genetic differences in well-being to be maintained by correlated responses
to selection for personality. These findings suggest the importance of
accounting for individual differences in primate personality when assessing
the fitness consequences of stress response.
Keywords: Pongo pygmaeus, Pongo abelii, heritability, quantitative
genetics
2 nd December
PLENARY LECTURE
PRIMATE CONSERVATION: SYNERGY AMONG STRESS,
NUTRITION, CLIMATE CHANGE, AND DISEASE
Colin Chapman
Department of Anthropology and McGill School of Environment, McGill
University, Montreal, Quebec, Canada
Presenter email: colin.chapman@mcgill.ca
Tropical countries, many of which harbour primate populations, are losing
approximately 12.5 million ha of forest annually; these forests are also
being degraded by logging and forest fires, which are not considered
“deforestation”. In addition to loss of forest habitat primate populations
continue to suffer high levels of commercial and subsistence hunting. Thus,
the future for primates looks grim without even considering the largely
unknown impacts of global climate change and newly emerging infectious
diseases, like Ebola. In this talk, we synthesize 26 to 36 years of population
and habitat data to determine the potential causes of group density changes
for five species of primates in Kibale National Park, Uganda. We focus both
on areas within the park that were disturbed in the late 1960s and on forest
fragments that lie outside the protected area. Within the national park,
mangabey and black-and-white colobus group density increased, blue
monkeys declined, while redtails and red colobus were stable in all forest
areas. For blue monkeys and mangabeys, there were no significant changes
in food availability over time that could account for changes in their group
density. For redtails, neither group density nor food availability changed
over time. For black-and-white colobus, a decrease in food availability over
time in the unlogged forest surprisingly coincided with an increase in group

19
density. Finally, while red colobus food availability and quality increased
over time in the heavily logged area, their group density was stable in all
areas. In general, all forest fragment populations declined, and this seemed
to be a result of a synergy among stress, nutritional, and disease. Analyses
of data on disease and cortisol levels suggests that disease levels are
increasing as would be predicted based on climate change in the region, and
increasing contact with humans that fosters disease transmission. We
suggest that these populations are generally in non-equilibrium states. Thus,
large protected areas will be required to ensure that declines in some areas
are compensated by increases in areas with different histories.
INVITED LECTURE
THE COMMON MARMOSET: SPECIES-TYPICAL STRESS AND
STRESS EXPERIMENTS TO UNDERSTAND HUMAN DISORDERS
Christopher Pryce
Preclinical Laboratory, University Psychiatric Clinic, University of Zurich,
Switzerland
Presenter email: christopher.pryce@bli.uzh.ch
Based on over 50 years of study by field and laboratory scientists, the
common marmoset (Callithrix jacchus) is a primate species for which there
is sufficient scientific knowledge to maintain captive groups in speciesappropriate,
i.e., stress-minimizing environments. Only because of this is it
possible to have the control conditions against which to study the stress
biology of the common marmoset, both in its own right and to provide
insights into human stress-related disorders, their mechanisms and
treatment. This talk will review studies by my colleagues and me on this
species and these subjects. As neonates, common marmosets exhibit high
blood basal levels of stress hormones (ACTH, cortisol), and these decline
gradually across maturation. Marmoset infants, like human infants, are
already able to exhibit stress hormone increases in response to social and
physical stressors. Marmoset urinary cortisol titres are positively correlated
with blood and CSF cortisol titres, so that urinary cortisol provides a
meaningful biomarker for bioactive cortisol availability, both under basal
and stress conditions. In stable, intact family groups of marmosets, social
stress is minimal. In adult twin sister pairs separated from their parents,
agonistic behaviour is rare in the absence of an unrelated male but typical in
the presence of an unrelated male. Subordinate sisters either suppress
ovulation and avoid aggression or are attacked and ovulation is suppressed:
the reduced oestrogen levels of such anovulatory females are associated
with reduced cortisol, which therefore does not serve as a stress marker
under these social/reproductive conditions. Common marmosets exposed
prenatally to high corticoid levels, to model the treatment used in at-risk
human premature births, exhibit increased feeding, reduced social play and

20
impaired motor behaviour as they develop. Marmoset infants that are
temporarily separated from their parents (early deprivation, ED) but
otherwise receive typical parental care, exhibit short-term increases in
cortisol and long-term, allostatic increases in blood pressure and
noradrenaline. Neither basal nor stress cortisol levels are altered long-term
by ED; nonetheless, brain expression of the cortisol receptors is reduced
long-term and in the hippocampus specifically. ED marmosets exhibit
increased anxiety and depression-like reduced motivation for reward.
Human early life stress is also associated with reduced hippocampal cortisol
receptor expression as well as increased risk of depression/suicide. Welldesigned
studies in the common marmoset, that utilise our knowledge of the
species to minimise uncontrolled stress, can provide valuable insights into
stress-related human disorders, their underlying causes and potential
treatments.
Keywords: common marmoset, cortisol, receptors, development, control
group
SEX SPECIFIC LIFE HISTORY AND CLIMATE RELATED
STRESSES RECORDED IN PAPIO HAMADRYAS DENTAL
TISSUES
Wendy Dirks 1 , Donald J. Reid 1 , Louise T. Humphrey 2 , M. Christopher
Dean 3 , Teresa E. Jeffries 4
1 Oral Biology, School of Dental Sciences, Newcastle University, Newcastle
upon Tyne, 2 Departments of Palaeontology and 4 Mineralogy, Natural
History Museum, London, 3 Department of Anatomy and Cell Biology,
University College London, London
Presenter email: Wendy.Dirks@ncl.ac.uk
Teeth develop incrementally, recording the history of their own formation
and the chronology of stress episodes in the lives of the individuals in
whose mouths they form. In histological tooth sections, daily and longer
period incremental lines are visible in dental tissues. Within this regular
pattern, stress episodes appear as accentuated lines that can be cross
matched between teeth. Our work focuses on using this recorded pattern to
compare the ages at which stresses occur in male and female baboons from
populations living under different ecological conditions. Our sample
consists of pure Papio hamadryas anubis and P. h. hamadryas-anubis
hybrids, with three males and two females from Ethiopia and one male and
one female from Uganda. In the Ethiopian female baboons, we compared
the ages at stress to rainfall records from their lifetimes and in the two
Ugandan baboons, we compared the ages at stress to the weaning process
revealed by spatial changes in strontium/calcium ratio in the enamel. All
seven baboons show stresses at the age when their mothers would
experience postpartum oestrus, but this occurs at an earlier age in Uganda

21
than in Ethiopia. Females exhibit stress at menarche, while males show
stress at the age at which they would disperse from their natal groups. One
hybrid male showed more frequent stress than the anubis males, perhaps
reflecting less ability to buffer environmental stress with increasing body
size during growth. Ethiopian males showed stresses at yearly intervals that
seem to correlate with the bimodal pattern of rainfall.
Keywords: baboon, histology, accentuated increments
IMPACT OF HUMAN DISTURBANCE ON STRESS, DISEASE AND
CONSERVATION OF CHIMPANZEES, PAN TROGLODYTES, IN
BUDONGO FOREST, UGANDA
Zinta Zommers 1 , Tobias Deschner 2 , Fabian Leendertz 3
1 Wildlife Conservation Research Unit, University of Oxford, 2 Max Planck
Institute for Evolutionary Anthropology, 3 Robert Koch Institute
Presenter email: zinta.zommers@zoo.ox.ac.uk
Across Africa, chimpanzee populations are endangered by habitat loss and
hunting. For conservation to succeed, it is critical to determine how
chimpanzees adapt to human and environmental stressors. Yet, little is
known about the mechanisms by which chimpanzees respond to
disturbance. While chimpanzees may adapt to changes through the stress
response, chronic elevation of stress hormones may result in reproductive
failure and disease. This research project examines the impact of human
disturbance on chimpanzees in Budongo Forest, Uganda. It attempts to
identify whether human disturbance results in increased stress, and whether
or not increase stress influences disease burdens. Between 2006 and 2008,
four communities of Budongo chimpanzees, each with different disturbance
regimes, were studied. Feces and urine samples were collected
opportunistically during focal follows. Behavioural data were also recorded.
Over 800 urine samples have been analyzed for cortisol and other stress
hormones using Liquid Chromatography-Mass Spectroscopy. Density
Gradient Gel Electrophoresis is being used to characterize bacteria
assemblages in feces samples to determine if there are differences between
individuals in different sites, and between individuals with different levels
of stress. By comparing data from the four sites, we will begin to identify
the impact of human disturbance on stress and on disease levels in
chimpanzees. We expect chimpanzees in disturbed sites to have different
bacterial assemblages than chimpanzees in undisturbed sites. Preliminary
results indicate that social factors may play a greater role in stress than
human disturbance. However analysis is ongoing. Research will increase
understanding of wildlife endocrinology and disease ecology.
Keywords: stress, chimpanzee, disease

22
INVITED LECTURE
DISTRESS ALLEVIATION IN MONKEYS AND APES
Filippo Aureli
Research Centre in Evolutionary Anthropology and Palaeoecology,
Liverpool John Moores University, Liverpool, UK
Presenter email: F.Aureli@ljmu.ac.uk
Various measures have been used to quantify stress and related phenomena,
such as distress and anxiety, in monkeys and apes. There are physiological
measures such as the increase in cortisol and heart rate. Self-directed
activities, such as self-scratching and self-grooming, have been successfully
used as behavioral indicators. Whereas stress can be an adaptive response
preparing an organism for appropriate actions, the prolonged activation of
such a response can be deleterious. Thus, mechanisms for alleviating
distress are expected to be selected for. The individual perceiving distress
may actively seek ways to reduce it. In other cases, third parties may
provide distress alleviation. Less is known about these latter mechanisms,
but they are particularly interesting for their potential altruistic nature and
cognitive underpinnings. For both types of mechanisms it is important not
only to describe their potential in distress alleviation, but to fully evaluate
their effectiveness.
Keywords: anxiety, self-scratching, grooming, consolation
INDIVIDUAL DIFFERENCES IN ANXIETY LEVEL AFFECT
RECONCILIATION IN WILD JAPANESE MACAQUES (MACACA
FUSCATA YAKUI)
Bonaventura Majolo
Department of Psychology, University of Lincoln, Brayford Pool, Lincoln,
LN6 7TS, UK
Presenter email: bmajolo@lincoln.ac.uk
Studies conducted on reconciliation (i.e., the post-conflict exchange of
friendly behaviour between former opponents) have mainly investigated
intra-individual sources of variation in post-conflict behaviour, showing that
animals have a stronger increase in anxiety and are more likely to reconcile
after conflicts with valuable partners, such as kin. Much less attention has
been given to how differences between individuals in emotional profiles
affect post-conflict behaviour. The aim of this study was to analyse whether
inter-individual differences in baseline anxiety levels predicted the
magnitude of the increase in anxiety following a conflict and the occurrence
of reconciliation. Data were collected on two groups of wild Japanese
macaques (Macaca fuscata yakui). Animals having a higher baseline level
of anxiety had a more dramatic anxious response following a conflict while

23
controlling for a series of factors (e.g. relationship quality between
opponents). These more anxious animals were also less likely to reconcile
than more relaxed individuals. Therefore, more anxious animals face some
social costs by being less able to cope with the post-conflict condition.
These findings would suggest that inter-individual differences in anxiety
levels are somehow maladaptive. However, the results of this study may be
interpreted and discussed, following the concept of behavioural syndrome,
as tradeoffs between benefits and costs across conditions.
Keywords: competition, emotion, scratching, stress
SOCIAL CAPITAL AND PHYSIOLOGICAL STRESS IN FEMALE
RHESUS MACAQUES
Lauren J.N. Brent 1 , Stuart Semple 1 , Michael Heistermann 2 , Ann
MacLarnon 1
1 Centre for Research in Evolutionary Anthropology, Roehampton
University, London, UK, 2 Department of Reproductive Biology, German
Primate Centre, Göttingen, Germany
Presenter email: L.Brent@roehampton.ac.uk
An emerging body of evidence suggests that social factors influence the
stress responses of group-living species. Among those factors, affiliative
interactions between individuals and members of their social environment
have been linked to stress reduction. Social capital, or the degree to which
an individual is integrated into society, represents the constraints and
opportunities faced by individuals in their social lives and has been
associated with lower stress levels in humans. Social network analysis is a
relatively new technique to primatology, but is one that provides a range of
measures of social capital which account for both direct and indirect
connections between individuals. Here I use social network techniques to
explore the relationship between social capital and physiological stress in
female rhesus macaques. Behavioural data were collected over a 9-month
period for 21 adult females from a free-ranging group on Cayo Santiago
(Puerto Rico). Faecal samples were analysed for glucocorticoid metabolite
concentrations using enzyme immunoassays and network measures of social
capital were generated using UCINET and SOCPROG software. A
significant negative relationship was found between one measure of social
capital and physiological stress levels, indicating that social capital may
help individuals cope with the stressors of group-living. A connection
between social capital and stress reduction in female rhesus macaques
complements research in human and non-human primates which has shown
that greater social capital is associated with improved health, reduced
mortality and increased reproductive success.
Keywords: social capital, stress, social network analysis, rhesus macaques

24
THE USE OF VISUAL BARRIERS TO ALLEVIATE STRESS-
RELATED HAIR-PULLING IN RHESUS MACAQUES (MACACA
MULATTA)
Paul Honess 1 , Yingying Jiang 2 , Julia McDonnell 2
1 Department of Veterinary Services, University of Oxford, Parks Road,
Oxford. OX1 3PT, UK 2 Dept of Zoology, University of Oxford, South Parks
Road, Oxford, OX1 3PS, UK
Presenter email: paul.honess@vet.ox.ac.uk
Captive conditions place a number of stresses on captive primates. One of
the manifestations of this, among macaques held for biomedical research,
can be diffuse or localised alopecia resulting from abnormal hair-pulling
behaviour. While alopecia can result from a number of non-behavioural
causes, it is the association with stress-related behaviour which is attracting
the attention of regulators, veterinary surgeons and primate welfare
biologists, as it may provide a practical index of stress. A number of
foraging and grooming-related enrichment studies have shown a degree of
success in reducing hair-pulling behaviour. This presentation describes a
study examining the effect of an increase in the provision of visual barriers
on hair-pulling among twenty-five group-housed rhesus macaques (Macaca
mulatta) in a breeding facility in the UK. The introduction of extra visual
barriers produced a significant reduction in hair-pulling behaviour, but also
had other beneficial effects: significantly decreasing aggression and
significantly increasing both affiliative behaviour and the amount of time
spent foraging. This study demonstrates that this simple and relatively
cheap environmental enrichment option not only has a dramatic effect on
hair-pulling behaviour but also has significant benefits for other aspects of
animal welfare.
Keywords: hair-pulling, alopecia, stress, visual barriers
IS TRAINING FOR COGNITIVE TESTING STRESSFUL? A
COMPARISON OF SELF-DIRECTED BEHAVIOURS OF
CHIMPANZEES (PAN TROGLODYTES)
Elizabeth S. Herrelko, Sarah-Jane Vick, & Hannah M. Buchanan-Smith
Department of Psychology, University of Stirling, UK
Presenter email: bherrelko@gmail.com
An important aspect of research with animals is ensuring the study is as
stress-free as possible for both the welfare of the animals and the quality of
the research. As with many cognitive studies, the ChimpCam Project, an
examination of chimpanzee (Pan troglodytes) cognition with the use of
cameras and computer tests, needed to start out with training. The goal of
this study was to identify whether or not training for cognitive testing
elicited higher rates of self-directed behaviours (SDBs) when compared to

25
other activities, as expected if the chimpanzees were anxious, uncertain or
frustrated by the training. SDBs of 11 chimpanzees (6 males, 5 females)
collected during a five-month period were compared between conditions:
(1) baseline (non-training situations) and two training conditions (2) an
ongoing, two-year programme of husbandry training (off-exhibit, i.e.,
station/stay while being closed into an area) and (3) training for cognitive
testing (i.e., introducing new on-exhibit areas and targeting to test
windows). Preliminary findings indicate that the mean rate per minute of
SDBs during training for cognitive tasks did not differ from husbandry
training, nor did SDBs during overall training differ from those exhibited
during activities outside training. There was considerable individual
variation in rates of SBDs that are not easily explained by level of training
nor willingness to participate. We conclude that training chimpanzees for
husbandry and for cognitive tests does not compromise welfare, but rather
that their repeated interest in the challenges offered, and their willingness to
participate, implies that the training is enriching and mentally stimulating.
Keywords: cognition, training, self-directed behaviours, welfare
THE EFFECTS OF AN ENHANCED SOCIALISATION
PROGRAMME ON BEHAVIOUR, WELFARE AND CARDIAC
RESPONSES OF NEWLY ACQUIRED CYNOMOLGUS
MACAQUES (MACACA FASCICULARIS) DURING SIX-WEEK
ACCLIMATISATION PERIOD
Lou Tasker 1 , Janet Kelly 2 , Simon Sutcliffe 2 , Hannah M. Buchanan-Smith 1
1 Department of Psychology, University of Stirling, UK,
2 Covance
Laboratories Ltd, Otley Road, Harrogate, North Yorkshire, UK
Presenter email: Louisa.Tasker@stir.ac.uk
Non-human primates may instinctively perceive humans as a threat,
reacting fearfully or aggressively when in close proximity. Humans are,
however, a necessary feature of the lives of all captive animals, and close
contact (including visual, auditory, olfactory stimuli) and handling is often
regularly required in laboratories. Animal care guidelines recommend
positive staff-animal relationships to enhance primate welfare. Habituating
and socialising primates to the behaviour, sight, sound and smell of humans,
will help to avoid or reduce fear responses and facilitate handling for
routine husbandry such as weighing and physical examination. Our aim was
to determine the effects of a six-week enhanced socialisation programme
with animal care staff on newly acquired cynomolgus macaques (Macaca
fascicularis) in a laboratory. We compare a range of welfare measures
recorded from control (N=40) and a matched group (N=40) of male and
female juvenile macaques subject to a socialisation programme based upon
pairing care staff contact and routine events with food rewards e.g. small
pieces of dried fruit. We compare data between the two groups on the

26
behavioural responses to care staff during handling and husbandry, changes
in body weight, body condition and alopecia scores, and physical health
over six weeks from arrival at a new facility. We also investigate the effects
of socialisation on cardiac parameters (heart rate, electrocardiogram
waveforms and blood pressure) recorded at baseline by non-invasive digital
electrocardiogram (ECG) and in-direct high definition oscillometry blood
pressure. Our results provide support for the importance of positive
relationships for both primate welfare and quality of science.
Keywords: cynomolgus macaque, welfare, socialisation, refinement
ABSTRACTS – POSTER PRESENTATIONS
WAS THE PRIMATE STRESS SYSTEM A SELECTIVE FORCE
FOR INCREASED HOMININ INTELLIGENCE?
John Cummins
Presenter email: deva_research@lineone.net
The evolution of primate intelligence likely led to increased episodes
of chronic psychological stress, as individuals lived less ‘in the moment’
and were able to prospect about future outcomes. The increase in
psychological stress impacts adversely on physical health, longevity and
reproductive fitness, i.e., allostatic load, potentially leading to an
evolutionary bottleneck, limiting the evolution of intelligence, unless
resolved through a selective mechanism. A cognitive framework involving
an increased interrelationship between the primate stress system and
hominin intelligence is proposed. Non-human primates may have a highlevel,
late-stage comparator, receiving input from many specialised parallel
modules. Operating serially, it accurately compares actual and goal states to
assess the animal’s effectiveness. It sends a record of effectiveness to longterm
memory, and corrective feedback to the specialist modules. It also
sends excitatory and inhibitory signals to the stress system. I propose a
chance variation in hominins, raising the threshold of excitation of the stress
system along these pathways, with feedback causing the comparator to
assess that the animal is more in control of the situation than is accurate.
This potentially dangerous event was not ‘washed out’ through learning, but
caused far-reaching systemic effects. It introduced flexibility to the
comparator and enabled it to model a variety of relationships between goal
and actual states, representing real or illusory control, creating domain –
general intelligence. Flexibility was also ‘sucked down’ differentially by the
specialist modules according to their ecological requirements, leading to
increased domain-specific intelligence. Flexible syntax in the language
module is offered as an example.
Keywords: primate, Hominin, stress, intelligence

27
LINEAR ENAMEL HYPOPLASIAS IN BABOONS
Chris Williams
Department of Archaeology, University of Sheffield, S1 4ET, UK
Presenter email: prp06cjw@sheffield.ac.uk
Linear enamel hypoplasias (LEH) are enamel defects that provide a record
of periods of physiological stress on tooth formation. This study uses
macroscopic and microscopic methods to analyze the expression of LEH in
baboons (subspecies of Papio hamadryas). The study sample consists of 85
wild baboon specimens from the collection at the Royal College of
Surgeons, London. Only permanent teeth with an estimated 80% of crown
height present with perikymata visible over much of the enamel surface
were included. The frequencies of LEH were analysed for both individuals
and tooth class by sex and jaw. No significant difference was found
between males and females in the frequency and distribution of LEH
defects. The mandibular dentition (20% of teeth affected) exhibited more
defects than the maxillary dentition (11.5%). I1s and I2s were found to have
the highest frequencies of LEH (48.8% and 25.4%, respectively, across both
jaws). A subsample of 16 mandibular incisors exhibiting LEH was selected
for microscopic analysis. Tooth impressions were sputter-coated and
viewed under reflected-light microscopy. Defect width, the distance of the
defect from the cementum-enamel junction (CEJ) and the number of
perikymata within the defect were recorded. The majority of the LEH were
located in the cervical third of the tooth crown. Defect widths ranged from
140 to 990µm (mean=470µm), containing between 2-8 perikymata
(mean=4.8). The results of the analyses were compared with available
published data for other non-human primates in order to assess the effect of
dental development patterns on the expression of LEH.
Keywords: enamel hypoplasia, dental defect, developmental stress, baboons
INTEGRATING MEASURES OF BEHAVIOUR, PHYSICAL
HEALTH AND PHYSIOLOGY TO PRODUCE AN OVERALL
ASSESSMENT OF WELFARE IN THE CYNOMOLGUS MACAQUE
(MACACA FASCICULARIS)
Lou Tasker 1* , Janet Kelly 2 , Hannah M. Buchanan-Smith 1
1 Department of Psychology, University of Stirling, UK,
2 Covance
Laboratories Ltd, Otley Road, Harrogate, North Yorkshire, UK
Presenter email: Louisa.Tasker@stir.ac.uk
Care staff need to assess primate wellbeing accurately in order to promote
refinements and to ensure any changes made to the environment, husbandry
or procedures are appropriate in promoting positive welfare, for example
through enrichment, and by preventing or minimising stress. This is no easy
task as there are species and individual differences in response to

28
environments. Furthermore welfare is a complex state and no single
measure of it is universally accepted. In this presentation we outline the
development and use of a multi-dimensional welfare assessment for
cynomolgus macaques (Macaca fascicularis); the most frequently used nonhuman
primate in laboratories. Our aim was to construct an assessment
framework that could be used to evaluate the effectiveness of planned
changes to husbandry events. We built upon current laboratory assessments
for animal well-being, using a stepwise rationale for inclusion of multiple
species-specific measures based upon their feasibility, validity, reliability
and sensitivity to detect changes in welfare status. We review behavioural,
physiological and physical health data from 10 cohorts of group-housed
male and female juvenile macaques (N=350) subject to normal housing and
husbandry procedures in the laboratory. We advocate using a range of
welfare measures to provide care staff with a surveillance framework for
monitoring the well-being of cynomolgus macaques. This framework is
being used successfully to assess targeted changes to routine handling and
husbandry, ensuring they effectively enhance macaque quality of life.
Keywords: cynomolgus macaque, welfare, integrating measures
HAIR LOSS IN CAPTIVE PRIMATES: A LONG-TERM FOUR
SPECIES COMPARISON
Lydia M. Hopper 1 , Susan P. Lambeth 2 , Mary Owens 2 , Ty McAdams 2 ,
Ricky Merino 2 , Leigh Ann Long 2 , Lawrence E. Williams 2 , Steven J.
Schapiro 2
1 Department of Psychology, South Road, Durham University, Durham,
DH1 3LE, UK, 2 Department of Veterinary Sciences, The Michale E. Keeling
Center for Comparative Medicine and Research, The University of Texas
M.D. Anderson Cancer Center, Bastrop, Texas, USA
Presenter email: lydia.hopper@durham.ac.uk
We report the current status of a long-term assessment of hair-pulling
behaviours by chimpanzees (Pan troglodytes), rhesus macaques (Macaca
mulatta), squirrel monkeys (Saimiri boliviensis boliviensis, S. b. peruviensis
and S. sciureus) and owl monkeys (Aotus nancymaae, A. azarae, A.
vociferans) housed at The Michale E. Keeling Center for Comparative
Medicine and Research (KCCMR), The University of Texas M.D.
Anderson Cancer Center, USA. There are many hypotheses surrounding
hair loss in primates, with potential causes related to hormone changes,
stress, seasonal changes, housing, breeding, aggression, sex and age. A
recent review concluded that it is a maladaptive behaviour, most likely
related to chronic stress-related husbandry deficiencies; the Institute for
Laboratory Animal Research associated hair-pulling with chronic exposure
to environmental stressors. Our on-going study aims to rate primate hair
loss at KCCMR quarterly for a minimum of 2 years. This regular data

29
collection will (i) allow for the analysis of baseline hair loss, (ii) identify
possible risk factors and, in the future, (iii) assess the ameliorating effects of
novel environmental enrichment. Hair loss is scored using a random sample
of no less than 15% of each species within our primate population
(chimpanzees N=29, rhesus moneys N=151, squirrel monkeys N=61, and
owl monkeys N=36). A total of 277 primates were randomly sampled to
guarantee the inclusion of adult males, adult females, adolescent males and
adolescent females. For each species we consider ‘season’
(spring/summer/fall/winter), ‘sex’ (male/female), ‘age’ (adult/subadult) and
‘group size’ as correlates for hair-pulling levels.
Keywords: hair loss, chimpanzees, rhesus macaques, squirrel monkeys, owl
monkeys
EVIDENCE FOR DIFFERENTIAL SUSCEPTIBILITY TO
PUTATIVE CHILDHOOD STRESS EFFECTS: THE ROLE OF
RELATIONSHIP STATUS
Lynda G. Boothroyd 1 , David I. Perrett
1 Department of Psychology, Durham University, UK, 2 School of
Psychology, University of St Andrews, UK
Presenter email: l.g.boothroyd@dur.ac.uk
Father absence and family stress have long been associated with age of
menarche and it has previously been reported (Boothroyd & Perrett 2008:
study 1) that there is also an association between family background and
facial attraction. In the current study, however, childhood stress in the form
of father absence or a poor father-daughter relationship were associated
with a difference in facial preferences and age of menarche only amongst
those who were single or in unhappy/uncommitted relationships. Those who
reported that they were in happy and committed relationships showed no
difference between father absent and present individuals in terms of either
current facial preferences or past pubertal development. Thus tendency to
be in a positive relationship in adulthood seems to indicate some critical
difference between subgroups of father absent/high stress women. This
further supports the notion (proposed by, e.g., Belsky 1997) that some
individuals may be more susceptible to the “effects” of rearing environment
than others.
Keywords: menarche, Homo sapiens, father absence, mate choice

30
INVESTIGATING THE ROLE OF SOCIAL INTERACTIONS ON
ADRENAL ACTIVITY IN FEMALE BARBARY MACAQUES
(MACACA SYLVANUS)
Katie L. Edwards 1,2 , Susan L. Walker 2 , Rebecca Bodenham 1 , Susanne
Shultz 1,3
1 School of Biological Sciences, University of Liverpool, Liverpool, UK,
2 North of England Zoological Society, Chester Zoo, Chester, UK, 3 Institute
of Cognitive and Evolutionary Anthropology, Oxford University, Oxford,
UK
Presenter email: K.L.Edwards@liverpool.ac.uk
Activation of the Hypothalamic-Pituitary-Adrenal (HPA) axis and the
subsequent release of glucocorticoids from the adrenal gland allows
individuals to adapt to changes in their environment. Although this process
is part of the body’s normal response to a stressor, prolonged elevation of
glucocorticoids can have deleterious effects on a wide range of processes
including health and reproduction. In primates, social stressors may be one
way in which adrenal activity is increased, but social systems are complex
and the relationship between dominance rank and glucocorticoid
concentration across species is not consistent. Previous studies have used
average glucocorticoid concentrations to draw links between either
dominance rank or particular social behaviours and adrenal activity.
However, hormone concentrations are transient, and this approach could be
misleading. We hypothesise that adrenal activity is dependant on the
individual, and related to the specific behaviours in which that individual is
involved. In the current study, eight female semi-free ranging Barbary
macaques (Macaca sylvanus) at Trentham Monkey Forest (Staffordshire,
UK) were followed for consecutive days, so that observed rates of social
behaviours could be analysed in respect to the glucocorticoid concentration
present in faeces the following day. Data were analysed in two ways, 1)
assessing the relationship between average glucocorticoid concentrations
and average rates of particular behaviours, and 2) analysing daily
glucocorticoid concentrations within individuals in relation to specific
social behaviours. By comparing the outcome of these analyses, it is clear
that the influence of behaviour on adrenal activity is complex, and
determining what factors predict adrenal activity can vary depending on the
analysis employed.
Keywords: glucocorticoids, behaviour, primates, Macaca

31
SOFTLY, SOFTLY, CATCHY MONKEYS
Yvonne Owen
Writtle College, UK
Presenter email: Yvonne.owen@ntlworld.com / 98272056@writtle.ac.uk
Traditional techniques used to capture New World monkeys, such as net
captures, can induce high levels of acute stress resulting in physiological
and behavioural responses detrimental to welfare. Alternatively, training
animals to voluntarily participate in husbandry and/or veterinary practices
using operant conditioning via positive reinforcement training is accepted as
a humane process which can reduce stress and improve welfare. A practical
application for training red-bellied tamarins (S. labiatus) had been identified
within Paradise Wildlife Park (Hertfordshire, UK). This case study aimed to
establish whether it was possible to use operant conditioning to train a
family of five red-bellied tamarins to voluntarily enter a transportation box
and remain calm for one minute, and whether this technique could be used
to reduce their experience of stress during capture and containment. A study
of two separate net capture processes was conducted and measures of
locomotion and vocalisations were recorded and compared to the same
measures recorded during the capture of the trained tamarins. Although
analysing stress-induced vocalisations as a quantitative measure of stress
has been studied in rodents and farm animals this measure remains
relatively novel. Net-captured monkeys exhibited rapid and erratic
locomotion and emitted long, sustained and high frequency vocalisations
during capture, whereas the trained tamarins exhibited minimal locomotion
and emitted only four brief vocalisations at RMS -35 dB during capture.
This study indicates that the use of operant conditioning via positive
reinforcement training did reduce potential for stress and improve welfare
during the capture and containment of the study group.
Keywords: stress, welfare, transportation, operant conditioning
VICTIM BEHAVIOUR FOLLOWING AGGRESSION IN CAPTIVE
SULAWESI CRESTED MACAQUES
Nia Watkin, Sean J. O’Hara
School of Biological Sciences, University of Liverpool, UK
Presenter email: n.watkin@me.com; sean.ohara@liv.ac.uk
Primates have complex social groups compared with most other non-human
mammals. There are many benefits associated with group living but there
are also associated costs. Most of these costs relate to conflicts of interest or
compromises, for example, over direction of travel or time scheduling.
Disputes, erupting as a result of within-group competition, are the most
serious form of internal conflict as they can lead to aggression between
group-mates causing injury, stress and damage to social relationships. A key
question in understanding social evolution is how do victims of aggression

32
deal with the negative consequences of group living? We evaluated this by
measuring post-conflict behaviour in a group of 31 captive Sulawesi
macaques (Macaca nigra) at Chester Zoo. Our results showed victims, but
not their aggressor, increased their level of group monitoring, selfscratching,
lick-smacking, teeth-baring and vocalisation compared to
matched-control periods. They played and foraged less. In addition, victims
received increased lip-smacking, vocalisation and aggression from groupmates
who also groomed and played with victims less than during matchedcontrol
periods. However, compared to their aggressor, victims engaged in
more post-conflict affiliative interactions (e.g., embraces, allogrooming).
Given the suite of behaviours suggesting foraging and other beneficial
activities are tempered while the victim anticipates further attack we
interpret these findings as evidence that aggression is costly and stressful
for victims. Post-conflict affiliative interactions, however, appear to
mitigate this negative aspect of group living for victims.
Keywords: affiliation, post-conflict, stress, victim

33
REGISTRATION FORM FOR THE WINTER 2009 MEETING
OF THE
PRIMATE SOCIETY OF GREAT BRITAIN
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Parkgate Road, Chester CH1 4BJ
You will be sent a receipt when payment has been received.

34
FUTURE MEETINGS
PSGB Spring Meeting 2010
Social Cognition and Evolution
7th and 8th April 2010
University of Abertay Dundee
The 2010 PSGB Spring Meeting will be hosted by the Department of
Psychology, University of Abertay Dundee. We are delighted to announce
the following keynote speakers:
Susanne Shultz, Institute for Cognitive and Evolutionary Anthropology,
University of Oxford
Thomas Bugnyar, Department for Neurobiology and Biology of Cognition,
University of Vienna and the Konrad Lorenz Research Center
James Anderson, Behaviour and Evolution Research Group, Department
of Psychology, University of Stirling
Carel van Schaik, Anthropological Institute and Museum, University of
Zürich
Plenary presentations will be on the theme of ‘Social Cognition and
Evolution’; however, we welcome proffered papers on any area of
interest to members of PSGB. We particularly encourage submissions
from postgraduate students and early career researchers, with prizes being
awarded for the best student presentations, including the Charles Lockwood
Medal for best oral
presentation and a further award for best poster presentation. Bursaries are
available to student members of PSGB to help with travel costs for those
presenting at the conference; visit student pages for
further information.
The conference will take place on Wednesday 7th April (full day) and
Thursday 8th April (half day). As well as oral and poster presentation
sessions, we will run a number of short skills workshops aimed at early
career researchers (see details below).
Confirmed workshop:
How to Get Published
Dr Sarah Elton, Functional Morphology and Evolution Research Unit, Hull
York Medical School
Dr Jo Setchell, Department of Anthroplogy, University of Durham
Proposals for further workshops are welcomed. Please contact the organiser,
Dr Clare Cunningham.

35
Primate Society of Great Britain Céilidh
All delegates are invited to the PSGB Céilidh that will take place on the
evening of Wednesday 7th April at the APEX Hotel in Dundee. This venue
is walking distance from the university and town centre. Tickets will be
available soon from the organisers so book early to avoid disappointment.
The estimated ticket prices are £10.00 for students/unwaged and £15.00 for
those that earn. This will include a buffet and dancing till midnight. Kilt
hire is also available locally for those wishing to get back to their Scottish
roots (details from Dr Clare Cunningham).
For further information visit
Or contact the organiser:
Dr Clare Cunningham
Department of Psychology
University of Abertay
Kydd Building
Bell Street
Dundee
DD1 1HG
Tel: 01382 308583
Email:
PSGB Winter Meeting 2010
The year 2010 will mark the 50th anniversary of the start of Dr Jane
Goodall’s ground-breaking chimpanzee research at Gombe Stream,
Tanganyika (now Tanzania). We are delighted to announce that the Primate
Society of Great Britain will hold its two-day Winter Meeting at the
Zoological Society of London’s Meeting Rooms (ZSL, Regent’s Park,
London) on 8th-9th December 2010, in celebration of the different types of
research done at, or inspired by, Gombe over the past 50 years, and looking
forward to the future. We are still in the early stages of planning this event,
but invited speakers will include Dr Jane Goodall, DBE (Founder, the Jane
Goodall Institute , Tara Golshan (Executive
Director, Education, the Jane Goodall Institute-UK
, as well as other key primatologists who have
been involved in the work at Gombe Stream National Park. So, please
‘watch this space’ and the PSGB’s website, for future announcements about
what should be a truly exciting event!

36
ANNOUNCEMENT
Nominations for PSGB Council Members 2010
In order to expedite the Annual General Meeting (AGM) and to adhere
more closely to the processes outlined in our constitution, Council is posting
the nominations for new Council members in advance of the AGM. The
current list of sitting Council members is found on the inside front cover;
individuals with an (07) after their name will step down at the AGM in
December. The nominated individuals, if deemed elected, will serve no
longer than three years, commencing immediately following the AGM in
December. We try to nominate Council members with a range of expertise,
experience and geographic location in order to have balanced views on the
matters considered by Council. The nominated individuals have confirmed
their willingness to serve on Council and are full members in good standing.
Please be advised that any member may put forward the names of any other
eligible members in place of those on the list prepared by the Council; such
nominations shall be seconded by two other members and forwarded in
writing to the Secretary at least one week before the date of the meeting. If
no fresh nominations are received, the Council’s nominees will thereby be
deemed elected. Otherwise election shall be by ballot at the meeting.
Proposed Council members:
Dr Rachel Kendal -Durham University
Dr Sonja E. Koski - Cambridge University
Dr Julia Lehman - Roehampton University
Dr Christoph Schwitzer -Bristol Zoo Gardens
Dr Susannah Thorpe - University of Birmingham
Thank you,
Colleen Schaffner, Secretary

37
BOOK REVIEWS
SPIDER MONKEYS: BEHAVIOUR, ECOLOGY AND EVOLUTION
Christina Campbell (Ed.) (2008)
Cambridge University Press
ISBN 978-0-521-86750-4 (Hardback) £65.00
A book concentrating on spider monkeys is long overdue and this one
succeeds in consolidating and reviewing current information and, in a few
chapters, presents new information available from long-term field studies.
The book begins with the indispensable section on taxonomy, morphology
and evolution. The latter two topics are dealt with by Alfred Rosenberger
and others who comment on how spider monkey anatomy more resembles
that of gibbons than other ateline species. Using a variety of methods,
Andrew Collins then presents the most up-to-date case regarding the issue
of spider monkey taxonomy at least to the species level. The author stresses
the need for further work on the identification of subspecies particularly in
Central America which will provide a “consensus of use” for
conservationists and researchers alike.
The second section of the book deals with ecology with a comprehensive
review of the sites of wild spider monkeys across 13 field sites by Anthony
Di Fiore, Andres Link and Lawrence Dew. The chapter includes an
extensive table listing the plants eaten by spider monkeys at eight field sites
across Central and South America. Rob Wallace reviews habitat use and
ranging patterns and the final chapter in this section by Lawrence Dew
reviews the spider monkey’s role as a seed disperser in comparison with
woolly monkeys. He concludes that spider monkeys are unique in their
dispersal of large fleshy-fruited seeds and, as such, are a key indicator
species for the health of a Neotropical rain forest.
Behaviour and reproduction are dealt with in section three. Dionisius
Youlatos describes locomotion and positional behaviour using a series of
very useful line drawings to illustrate locomotor modes and postures.
Gabriel Ramos-Fernandez reviews communication studies and this is
followed by Filippo Aurelli and Colleen Schaffner who review spider
monkey social interactions and social relationships and include a very clear
explanation of their fission fusion social system. An excellent and very
interesting review of current data but, as the authors make clear, it reveals
more questions about spider monkey social behaviour than answers.
Christina Campbell and Nicole Gibson combine data from captivity and
from two study sites to present information on spider monkey reproductive
and sexual behaviour. They also lament the lack of data available. Laura
Vick presents new, albeit preliminary, data from her long-term study on the
development of gender differentiated social roles in the Yucatan spider

38
monkey. Her chapter is followed by Yukiko Shimooka and others
describing the current knowledge on demography and group composition.
The final section deals with spider monkeys and their interactions with
humans. The spider monkey conservation chapter concludes that if existing
protected areas are managed and enforced, spider monkeys could continue
to exist, albeit with the loss of a few subspecies. However, this may be a bit
optimistic in my experience of working in and around protected areas in
Central America. I was pleased to see a chapter on spider monkey
ethnoprimatology by Laura Cormier and Bernardo Urbani, who clearly
demonstrate the importance of spider monkeys to indigenous peoples as
both a food and cultural resource. These authors clearly establish that the
species is hunted throughout its range and a chapter might have been
included with data collated from the various studies carried out on hunting
in the Neotropics. This information might then be used to assess the
sustainability of spider monkeys into the future if hunting continues at the
current rate.
It’s a great pity this book is so expensive because it is a very useful resource
for those of us interested in spider monkeys. However, it does reveal large
gaps in our current knowledge. Hopefully this will inspire more people to
study spider monkeys particularly in the many areas where no research
presently occurs.
SIÂN S. WATERS
Durham University
SEXUAL SELECTION AND THE ORIGINS OF HUMAN MATING
Alan F. Dixson (2009).
Oxford University Press
ISBN: 9780199559428 (Hardback) £70.00
Evolutionary psychology has resulted in an explosion of research that, while
providing some fascinating insights into human mating behaviour, has been
heavily criticized for adhering to a number of unfounded assumptions. Alan
Dixson’s Sexual Selection and the Origins of Human Mating Systems,
scheduled to coincide with the 200th anniversary of Darwin’s birth,
provides an invaluable analysis of cross-species research in order to place
human sexuality in the context of a wealth of empirical evidence relating to
our evolutionary history. He begins, for example, with a discussion of
diverse approaches to the analysis of human mating systems (including
advances in the fossil record, genetics and our understanding of postcopulatory
sexual selection and primate social lives), each of which is
employed to address arguments surrounding issues such as sperm
competition, cryptic female choice, copulatory patterns, oestrus and sexual

39
dimorphisms in primates. The framework for evaluation is the basic
classification of primate mating systems (monogamy, polygyny, polyandry,
multi-male/multi-female and dispersed), and diverse topics are drawn
together to develop a model of the evolution of human mating behaviour
that best fits the cross-species data.
The book falls broadly into two halves, with the first addressing the issue of
post-copulatory selection in humans. An excellent review of the history of
research into testes-to-body-weight ratio (which has, on occasion, been
cited as support for an evolutionary history of polygyny - or selection via
sperm competition - in humans) leads to the conclusion that human data are
consistent with that of polygynous and monogamous primate species, with
little support for post-copulatory selection pressures. The critique continues
with comparative analyses of male reproductive biology and physiology,
sperm morphology, ejaculate concentration and female orgasm to further
allay claims that there is evidence for sperm competition or that the function
of female orgasm is to exert selection for the sperm of favoured males. The
second half addresses a variety of additional issues, dispelling further myths
surrounding human behaviour. Analysis of occurrence of face-to-face
copulation across primate species, for example, leads to the conclusion that
it is unlikely to have evolved in humans to promote female orgasm and that
this, along with prolonged intromissions, are likely to have a occurred as the
result of alternative pressures such as reduced risk of predation. Likewise,
the author argues that humans are not an anomalous species that have “lost”
oestrus, rather that the evidence is more consistent with a general pattern of
cross-cycle receptivity across primate species. Sexual dimorphisms are
compared, incorporating those often discussed in the human literature (e.g.,
body size and composition, hair distribution and facial structure) as well as
some that are less well understood (e.g., age at puberty). This
comprehensive review of a substantial volume of evidence relating to a
diverse series of topics culminates in the conclusion that human data are
most consistent with evolution under a polygynous to monogamous mating
system with little post-copulatory selection. This has implications for a
number of the assumptions that have sometimes been inherent in
evolutionary psychology research.
The quantity and variety of empirical data discussed, sourced from a wide
range of disciplines, make this book an ideal resource for anyone interested
in evolutionary psychology, human sexuality or primate mating systems.
The graphical representations of comparative data help to dispel some of the
myths regarding human mating systems that have developed in the last
decades. While the negative evaluation of evolutionary psychology
(especially in the conclusion) may be perceived by many as unfounded (and
could perhaps have been countered by a selection of work the author
considers to be “good science”), this is the most thorough discussion of

40
primate mating systems to date and will provide an excellent source of
reference for discussion of human sexuality in the future.
FHIONNA MOORE
University of Abertay Dundee
CODING AND REDUNDANCY: MAN-MADE AND ANIMAL-
EVOLVED SIGNALS
Jack P. Hailman (2008)
Cambridge, Harvard University Press
ISBN: 9780674027954 (Hardback) £25.95
This book is not specifically related to primates. It brings examples from a
wide range of animals, from insects to humans, which makes the book very
attractive, innovative and suitable to a broad audience, from ordinary
readers to experts in animal behaviour and communication. Jack Hailman
starts with a clear and concise review on ethology and animal
communication, connecting concepts of information theory (e.g., entropy)
to the way that animal’s signals encode information. Hailman then classifies
communication signals used by human (e.g., pedestrian crosswalk
markings, railroad semaphore, vehicle turn indicators, Braille) and by nonhuman
animals (e.g., contact calls, body colour patterns and features,
mating behaviours), showing how alike both groups encode information. He
discusses comparatively about three main strategies of communication
codes (i.e., binary, multi-valued and multivariate coding) and how the
coding principles can be used to pass the information more effectively and
economically. Furthermore, he considers redundancy in signalling and
shows how sometimes it can actually improve information transmission,
instead of being associated with inefficiency and waste of energy. Hailman
managed to explain very nicely about the quantitative measures related to
information theory and coding strategies. However, the math can be skipped
and the book is still understandable and very enjoyable! Here we have a
nice volume whose text is complemented with excellent illustrative figures,
that certainly helps the reader to clarify about the various coding strategies
explored throughout the book. Concise tables at the end of some chapters
summarize well all the examples discussed and are very useful to later
refresh the reader’s mind. Although the examples on non-human primates
focus mainly on acoustic signalling, the book can still be used
comparatively to explain and understand a great deal on primate
communication signals in general. After reading this book, people will
certainly look at man-made communication signals in a different way, often
making the comparative links to the animal-evolved signals.
BRUNA BEZERRA
University of Bristol

41
BOOKS OF INTEREST
BLOOD WOOD
Penny Morgan (2009)
Authorhouse
ISBN: 9781438905754 (Paperback) £17.09
This is the second novel from author Penny Morgan who wrote Prime
Witness, previously reviewed in Primate Eye. These books are worthy of
mention here as they attempt to bring many of the issues facing primatology
into mainstream fiction; an endeavour that can only help in promoting
public awareness of many of the difficulties facing primate species
worldwide. This book shifts focus from cognitive studies on captive apes
that threaten to blur the line between humans and chimpanzees to issues
surrounding habitat loss in their native forests. As before, the main crux of
the story is a ‘whodunit’ with the murder of an employee from Forest
Watch, sent to Cameroon to assess levels of illegal logging occurring within
the first few chapters. What ensues is an investigation that crosses two
continents that reveals power and corruption at the highest levels of
government that threatens to promote the extinction of many ape species.
Characters return from the previous tale including Caro the languagetrained
bonobo who aided the police in apprehending the culprit in the last
story; however, this time she is living with her own kind in a managed
population of chimpanzees in Africa.
In the last book, from the very beginning, I found myself wanting to know
who had committed the crime and why. The way the story was structured
captured the reader in just a few pages. With this book, there was not the
same immediacy and I think this was due to a greater complexity in the
relationships between the individual characters and associations that
contribute to the story. I did find the many acronyms for government
departments, centres and administrative bodies that proliferate through the
story somewhat confusing. This made the book difficult to read at times and
I didn’t find this as compelling as Prime Witness until the later chapters
when all the elements began to come together. Then I couldn’t put it down.
So overall, this was a good book that again tackles issues relating to
primatology and brings them to the attention of a potentially new audience.
CLARE CUNNINGHAM
University of Abertay Dundee

42
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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 - 1996 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
Past Secretaries:
1967 - 1970 R.P. MICHAEL 1985 - 1987 H.O. BOX
1970 - 1974 K.R. HOBBS 1987 - 1990 R.C. HUBRECHT
1974 - 1975 V. REYNOLDS 1990 - 1993 P.C. LEE
1975 - 1978 R.D. MARTIN 1993 - 1996 C. ROSS
1978 - 1981 A.F. DIXSON 1996 - 1999 H. BUCHANAN-SMITH
1981 - 1985 S.K. BEARDER 1999 - 2005 J. LYCETT
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
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 1994 R.I.M. DUNBAR
1980 R.A. HINDE 1996 T. ROWELL
1982 F. BOURLIERE 1998 C.B. STRINGER
1984 P.M. BUTLER 2000 A.F. DIXSON
1986 J.P. HEARN 2002 I. TATTERSALL
1988 H. KUMMER 2004 C. VAN SCHAIK
1990 R.D. MARTIN 2006 A. JOLLY
1992 J.H. CROOK 2008 W. MCGREW
Napier Memorial Medal Winners (established 1991):
1991 CHRISTOPHER PRYCE: Endocrine and social correlates of maternal behaviour in
callitrichids
1993 MARTA LAHR: The origins of modern humans: A test of the multiregional
hypothesis
1995 CARLOS DREWS: Psychological warfare and the management of relationships
between male baboons
1996 NICOLA KOYAMA: Reconciliation behaviour in wild Japanese macaques
1999 MARK COLLARD: Morphological evolution of the hominoids and papionins:
implications for palaeoanthropological cladistics
2001 RUSSELL HILL: Ecological and demographic determinants of time budgets in
baboons: implications for cross-population models of baboon socioecology
2003 SUSANNE SHULTZ: Of monkeys and eagles: predator-prey interactions in the Taï
National Park, Côte d'Ivoire
2005 CORRI WAITT: Facial attractiveness among rhesus macaques (Macaca mulatta):
manipulating and measuring preferences for conspecifics' facial characteristics.
2007 ANNIKA PAUKNER: Secondary representational abilities in nonhuman primates
(Macaca nemestrina, Cebus apella)
Occasional Medal Winners (established 1996):
1997 JANE GOODALL CBE: Conservation
2007 CYRIL ROSEN MBE: Conservation
2008 STEPHEN NASH: Special Contributions to Primatology

Primate Society of Great Britain
PRIMATE EYE No 99 OCTOBER 2009
• CONTENTS •
Page
Editorial … … … … … … … 1
Report: Spring Meeting 2009 – Form and Function … … 3
PSGB Winter Meeting 2009 – Primate Stress: Causes, Responses
and Consequences
Draft Schedule … … … … … 6
Paper & Poster Abstracts … … … … 10
Pre-registration Form … … … … 33
Future Meetings:
Spring 2010
Winter 2010
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34
35
Announcement: Nominations for PSGB Council Members 2010 36
Announcement: Charles A. Lockwood Student Prize 2009 … 36
Book Reviews:
Spider Monkeys: Behaviour, Ecology and Evolution … 37
Sexual Selection and the Origins of Human Mating … 38
Coding and Redundancy: Man-Made and Animal-Evolved
Signals … … … … … … 40
Books of Interest:
Blood Wood … … … … … 41
ISSN 0305-8417