Tsaobis Baboon Project

Baboon on rock outcrop

The Tsaobis Baboon Project is a long-term study of a desert baboon population in Namibia. Our work is carried out in collaboration with Tsaobis Nature Park and in affiliation with the Gobabeb Research and Training Centre.

Research permission in Namibia is kindly provided by the Ministry of Lands and Resettlement and the Ministry of Environment and Tourism. In the UK, the Tsaobis Baboon Project is based at the Institute of Zoology, the research arm of ZSL.

Our research is undertaken with collaborators in various institutions, including the departments of Zoology and Biological Anthropology at the University of Cambridge, the department of Zoology at the University of Oxford, the Institute for Integrative Biology at the University of Liverpool, the Institute for Evolutionary Science at the University of Montpellier, the Centre for Research in Evolutionary Anthropology at Roehampton University London, and the Fenner School of Environment and Society at the Australian National University.

 

Project Background:

The aim of the Tsaobis Baboon Project is to carry out fundamental research in behavioural ecology and population ecology using desert baboons as a model system. Our work also has a strong conservation science theme with its interest in the effects of complex social structure on population dynamics and extinction.

The Tsaobis Baboon Project is based at Tsaobis Nature Park on the edge of the Namib Desert in central Namibia. At present, we work with two troops of chacma baboons (Papio ursinus), each of about 60 individuals. Each baboon is individually recognisable, of known age, and of known relatedness to other individuals in the population.

Research on the Tsaobis baboons began in 1990, and has been ongoing on an annual basis since 2000. To date, over 50 scientific publications have been produced from our research, including several articles in the journals Nature, Current Biology, and Proceedings B.  For a full bibliography, please see Publications.

 

Volunteer Information

Each year, we recruit volunteers to assist with fieldwork on the Tsaobis Baboon Project. There are six positions for volunteers currently available for the last nine months of the 2014-2015 field season, from late October 2014 to early August 2015. Positions are available for three-month and six-month periods. The travel and subsistence costs of volunteers in the field will be covered by the project, and a contribution will also be made towards the costs of the return flight.

If you would like more information, or to apply for one of these positions, further details are provided under Information for Volunteers.

The deadline for applications is 9.00am Monday 16th June 2014

Future volunteer opportunities: We are currently awaiting news on grant funding to decide if there will be a second recruitment round for an extended 2015 field season (most likely from August to October inclusive). If so, this round will be  in six months time (December 2014).  If not, the next recruitment round will be in 18 months time (December 2015) for the 2016 field season.

Student Information:

Research

Since its inception, a variety of research studies have been carried out on the behaviour, ecology, genetics, and health of the Tsaobis baboons. These projects have included studies on personality, social foraging, friendships and paternal care, leadership and group coordination, sexual signalling and mate choice, and predation risk.

Recent completed PhD projects include those by Julio Benavides, Alecia Carter, Elise Huchard, Andrew King, and Harry Marshall.

Volunteer Positions

 

Background

We are currently recruiting volunteer field assistants in the last nine months of our 2014-2015 field season, between late October 2014 and early August 2015. Fieldwork will primarily involve daily follows of baboon troops on foot, collecting data on the behaviour of individually recognisable animals, together with monthly insect/vegetation surveys and wider ecological monitoring related to the baboons' environment. The primary research focus will be on baboon host-parasite dynamics.

 

Positions Available

We are currently looking to fill six volunteer places. These six places will be filled with a combination of 6-month and 3-month positions, as follows:

6-month positions:  late October – late April (desert summer)
  early February – early August (summer-winter transition)
3-month positions: early February – early May (summer-winter transition)
  early May – early August (desert winter)

 

Qualifications

Volunteers are generally expected to be graduates with a good degree in the biological sciences. The Tsaobis Baboon Project is an excellent opportunity for graduate students to gain experience of field research before commencing a PhD. Applicants should be very physically fit, enthusiastic and hard working. Previous experience of fieldwork and a full driving licence are essential.

Payment & Conditions

We will cover all work-related travel and subsistence costs in Namibia (including food and accommodation at Tsaobis). The volunteer will be expected to cover the cost of their visa (about £100), but we will cover the costs of their insurance. In addition, we will contribute a minimum of £75 towards the cost of the air ticket following the successful completion of the volunteer’s fieldwork. All data collected during the course of the fieldwork will be the property of the Tsaobis Baboon Project, although volunteers will be fully acknowledged in all relevant papers arising from the Project.

Application

If you would like to apply for a volunteer position, you will need to submit a CV and covering letter.

  • The CV should be up to date. It should list at least two referees including their e-mail contact details. Please state whether or not you have a driving licence (for manual gear shift), and if so for how long you have held it.
  • The covering letter should explain why you would like to work on the Project. In the letter you must also specify the volunteer period for which you are applying (6-months, ‘desert summer’ or ‘summer-winter transition’; 3-months, ‘summer-winter transition’ or ‘desert winter’), together with the earliest and latest dates (day and month) that you would be available to take-up these positions (this will assist us in finalising the exact start and end dates for all of the team)
  • Before submission, please check that your application covers all of these preceding points. Incomplete applications will be at a disadvantage.
  • Both CV and covering letter should be submitted as a single Word document (with the covering letter on a separate page preceding the CV) and sent by e-mail.
  • Your e-mail application should have the subject header "volunteer application:" followed by your name. The e-mail should be sent to the Project leader, Dr Guy Cowlishaw, at the following address: "guy.cowlishaw@ioz.ac.uk".

Applications must be received by 9am Monday 16th June 2014. We will notify successfully shortlisted candidates the same week, and interviews will be held in London two weeks later on Mon 30th June, Tues 1st July, and Weds 2nd July. Applicants should keep these dates free for interview, since no other dates will be available. Telephone/skype interviews will be possible for overseas applicants.

 

Further Information

Additional details on the Tsaobis Baboon Project and the volunteer work are given below, under six headings: The Study Site, Working Conditions, Living Conditions, Personal Details, What To Bring, and Previous Baboon Research at Tsaobis. Additional information about Namibia can be found in the standard travel guides, such as "Footprint" and "Bradt".

 

The Study Site

Weather

Tsaobis experiences the characteristic extremes of desert environments. In winter time (mid-April to mid-October), the nights are cold (occasionally as low as 0oC) while the days are dry and hot. In summer time (mid-October to mid-April), warm nights are followed by scorching days when the temperatures can soar to more than 40oC in the shade. Some days in summer can also be quite humid, when rare clouds build for spectacular thunderstorms with torrential rain.

Landscape

Tsaobis is a beautiful desert wilderness. The landscape is a mixture of alluvial and gravel plains, rocky hills, and nearby mountain peaks. Most vegetation comprises herbs, small shrubs, and dwarf trees, although the dry sandy bed of the ephemeral Swakop River that cuts through this landscape supports several woodland groves. Following summer rains, the desert springs into life and the baboons forage across the hills and plains. In the winter time, as the summer growth dies back, the baboons forage primarily in the Swakop woodlands. Throughout the year, the baboons sleep on cliffs in the rocky hills.

Wildlife

Tsaobis is home to a variety of wildlife including mountain zebra, kudu, springbok, klipspringer and, of course, baboons! Predators include jackals and, occasionally, leopards. Birds and reptiles also abound.

Location

Tsaobis is centrally located in Namibia. It is only a 3.5 hour drive from Windhoek (the capital). The nearest town, Karibib, is a one-hour drive away and comprises a petrol station, bank, post office, and basic shops.

 

Working Conditions

Daily Schedule

Data collection requires full-day follows of the study troops. This ensures that the location of the sleeping cliff used by the baboons each night is known, which in turn facilitates their pick-up by observers early the next morning. If the day begins late, or ends early, the baboons can be lost and may take several days to relocate. Full-day follows require that the field team have already had breakfast, prepared their pack lunches, and are ready to leave camp in good time before dawn; it also means that they will not get back before nightfall. The most distant baboon sleeping cliffs are more than an hour away from camp, so a prompt start in the morning is essential (e.g. 4:30am). The baboons are then accompanied until they reach their sleeping cliff at dusk (e.g. 6pm, returning back to camp by 7pm). Although these are long days, the weekly schedule ensures that there are regular breaks for all the team (see below). In the peak summer months, when the days are at their longest and hottest, we will switch to a half-day system where observers will either do the morning or afternoon shift.

Working with baboons in the Tsaobis landscape

Observers spend the day on foot in the company of the baboons as they traverse the rocky, mountainous terrain around the Swakop riverbed. This landscape is beautiful but also physically demanding. Field team members travel (with backpacks) up to 10km a day, ascending and descending steep hills and slippery scree slopes as they follow the baboons. The baboons are habituated to the presence of human observers, allowing data to be collected from close proximity without causing disturbance, but observers must always act carefully and responsibly when in the company of these wild animals. Data collection is largely conducted with handheld computers. Faecal and urine samples are also collected on a routine basis. On-site supervision and detailed guidelines will be provided at Tsaobis, describing how to work with the baboons in this desert landscape.

Environmental surveys and monitoring

At monthly intervals throughout the field season, insect and plant phenology surveys will be conducted. These are an important complement to the baboon behavioural data, since they describe the spatial and temporal availability of food items/patches to the baboons. Additional environmental surveys and monitoring may also be conducted, depending on current projects and research priorities.

Weekly work schedule

Since data collection from the baboons requires consecutive full-day follows, these "field days" usually take place over two-day stints with an intervening one-day break. The latter are primarily "rest" days, but there will also be some office and general housekeeping duties (e.g. updating databases, cooking supper). Trips will be made to Windhoek about once a month for supplies, and volunteers will also be expected to assist occasionally with these trips. Although the field-office/rest day cycle of 2-1 days is the normal routine, this schedule necessarily retains flexibility throughout the field season.

 

Living Conditions

Accommodation

Accommodation is based around a small two-roomed bungalow. The bungalow has a kitchen, bathroom and office. Members of the field team sleep in their own tents which are pitched under trees close to the bungalow, but use the living area of the bungalow for rest and relaxation.

Telephone and e-mail

The Project has access to the rest camp telephone and wi-fi (although the latter is relatively slow and often erratic). Both phone and wi-fi are available to volunteers, although the use of the former will be charged to volunteers at the local rates.

Visitors

Members of the field team may receive visitors at Tsaobis, but volunteers should consult the Project prior to making arrangements as permission from relevant landowners needs to be sought. Care must be taken that visitors do not disrupt the field routine and all plans and arrangements for visitors must be confirmed with the Project Director prior to the visit. Unfortunately, accommodation for visitors cannot be guaranteed at the field site and it will not be possible for visitors to accompany the volunteers when working with the baboons.

 

Personal Details

Preparation

In addition to organising equipment (see below), there will be various tasks and pieces of paperwork that volunteers must complete before they can travel to Tsaobis. Work on two key tasks should be initiated as soon as the volunteer has been firmly accepted onto the Project. These tasks are:

  • put a deposit down on a flight, and
  • arrange vaccinations (see below)

We will organise the visa paperwork and travel insurance for the volunteer. In addition, where possible, we encourage volunteers to do a little background reading on baboons before coming to Tsaobis. In particular, Louise Barrett's "Baboons: Survivors of the African Continent" (BBC, 2000) is an excellent introduction to the world of baboons, while Robert Sapolsky's "A Primate Memoir" (Vintage, 2002) is a wonderful account of fieldwork with baboons.

Health and safety, general

All volunteers must be aware of their blood group and will need to ensure that they are up-to-date with their vaccinations. For the most recent information and treatment, volunteers are recommended to visit their local travel clinic or GP. This should be done as soon as the volunteer has been accepted onto the Project, since some vaccination courses can require several months between the first and last injection. Because the volunteers will be working with wild primates, vaccinations for hepatitis B and rabies must be obtained in addition to the standard vaccinations for Namibia. There is no malaria in this region of the country, but if volunteers plan to travel further north before or after the field season, e.g. to Etosha National Park, anti-malarials will be required.

Health and safety at Tsaobis

Because fieldwork at Tsaobis is conducted on foot and takes place from dawn to dusk, often over difficult terrain, volunteers should be physically fit. Due to the dry desert climate, Tsaobis is a healthy place to work, but volunteers will need to take appropriate steps to avoid dehydration, sunstroke and excessive sun exposure. This includes carrying plenty of water, wearing wide-brimmed hats with appropriate clothing and sunglasses, and the regular application of sunblock. In peak summer months, volunteers may find that long-sleeved shirts and long trousers are more appropriate than t-shirt and shorts.

Health and safety outside Tsaobis

Although the tar and gravel roads in Namibia are generally excellent, all volunteers should drive carefully and at reasonable speed, and never drive at night. Although Namibia is generally a safe country to visit, when in Windhoek (and other urban areas) it is a sensible precaution to stay alert and avoid walking the streets at night, especially alone. For further details on health and safety in Namibia, please refer to an up-to-date travel guide.

 

What to Bring

Clothes

In winter time, due to the cold nights and hot days, field team members should bring sweatshirts and jumpers as well as light summer clothes. Woolly hat, gloves, and a warm fleece are essential for cold winter mornings and evenings (and for possible trips to the coast). Such clothes are not needed in summer time. At all times of year, field clothes should include long-sleeved shirts and long trousers, in addition to t-shirts and shorts. Avoid wearing dark colours if possible; lighter colours are much cooler. A wide-brimmed hat and sunglasses are essential. A small lightweight waterproof windcheater is also indispensable for windy winter mornings and wet summer days. In addition to field clothes, some smart clothes will also be useful (e.g. for visits to Windhoek). Footwear should include a robust pair of walking boots and a sturdy pair of sandals.

Equipment

Personal equipment should include:

  • A tent. Ideally your tent should be three person, since it will also double as your private retreat. If you are working at Tsaobis in the summertime, it must also be waterproof. A mattress will be available, but your tent will need to be large enough to accommodate it. The mattress dimensions are 190cm x 88cm x 15cm.
  • A 3-season sleeping bag
  • A comfortable backpack for daily use in the field. Ideally bring a backpack with a waist strap as well as shoulder straps, so that the weight of the bag can be carried on the hips rather than the back. Backpacks that are moulded to allow air to pass between the bag and your back (e.g. the Berghaus "freeflow" design) are cool to wear. Backpacks should be a minimum of 25-30 litres volume. If you are working at Tsaobis in summertime, your backpack should also have a raincover.
  • A comfortable pair of hiking boots. Ideally these should be lightweight but robust (with ankle support).
  • A good set of binoculars, ideally 8x40 or 10x40. Avoid poor binoculars: they will make your work difficult and frustrating.
  • A spare pair of spectacles (if used)
  • A torch (ideally a head torch)
  • Ankle-length gaiters
  • A simple compass and whistle
  • A sewing kit
  • A water bottle can also be helpful, although most of the time we simply re-use the bottles that come with bottled water and soft drinks.
  • Many previous volunteers have found it useful to bring a laptop, camera, MP3 player, e-reader, and/or books to read. A weather-proof bag, or just a ziplock bag, can be useful to keep valuable electronic equipment safe from dust and sand (and, in summer time, water). Compact/travel games are always welcome, as are DVDs.
  • Field guides can usually be purchased in Windhoek (mammals, birds, trees, herps, etc), and some are also available for reference at Tsaobis.

For personal expenses, volunteers can bring cash or traveller’s cheques, and should also be able to make withdrawals from ATMs using a standard debit card. Credit cards are widely accepted and are recommended for emergencies.

2014

Carter AJ,Marshall HH, Heinsohn R, Cowlishaw G. (2014) Personality predicts the propensity for social learning in a wild social primate. PeerJ 2:e283; DOI 10.7717/peerj.283.

Williams D, Pettorelli N, Henschel J, Cowlishaw G, Douglas C. Alien tree invasion and mammal distribution along an ephemeral river in Namibia. Journal of African Ecology [early online]

2013

Carter AJ, Feeney WE, Marshall HH, Cowlishaw G, Heinsohn R. (2013) Animal personality: what are behavioural ecologists measuring? Biological Reviews 88: 465-475.

Carter AJ, Marshall HH, Heinsohn R & Cowlishaw G. (2013) Personality only predicts decision-making when information is unreliable. Animal Behaviour 86: 633-639.

Huchard E, Charpentier MJ, Marshall H, King AJ, Knapp LA, Cowlishaw G. (2013) Paternal effects on access to resources in a promiscuous primate society. Behavioural Ecology 24: 229-236.

Marshall HH, Carter AJ, Ashford A, Rowcliffe JMR, Cowlishaw G. (2013) How do foragers decide when to leave a patch? A test of alternative models under natural and experimental conditions. Journal of Animal Ecology 82: 894-902.

Cowlishaw G (2013) Papio ursinus Chacma baboon. In: Mammals of Africa. Volume II: Primates (eds T. Butynski, J. Kingdon, J. Kalina), pp. 225-228. Bloomsbury: London.

2012

Benavides J, Huchard E, Pettorelli N, King AJ, Brown ME, Archer CE, Appleton CC, Raymond M, Cowlishaw G (2012) From parasite encounter to infection: multiple-scale drivers of parasite richness in a wild social primate. American Journal of Physical Anthropology 147: 52-63.

Carter AJ, Marshall HH, Heinsohn R, Cowlishaw G (2012a) Evaluating animal personalities: do observer assessments and experimental tests measure the same thing? Behavioural Ecology and Sociobiology 66: 153-60.

Carter AJ, Marshall HH, Heinsohn R, Cowlishaw G (2012b) How not to measure boldness: demonstration of a jingle fallacy in a wild social primate. Animal Behaviour 84 : 603-609

Marshall HH, Carter AJ, Coulson T, Rowcliffe JM, Cowlishaw G (2012). Exploring foraging decisions in a social primate using discrete-choice models.American Naturalist 180: 481-495.

Marshall HH, Carter AJ, Rowcliffe JM, Cowlishaw G. (2012) Linking social foraging behaviour with individual time budgets and emergent group-level phenomena. Animal Behaviour 84: 1295-1305.

Rands SA, Pettifor RA, Rowcliffe JM & Cowlishaw G (2012) State-dependent foraging rules for social animals in selfish herds. In Anil Seth, Tony Prescott & Joanna Bryson (eds) Modelling Natural Action Selection. Cambridge University Press: Cambridge. [reprinted from Proceedings B: Biological Sciences271: 2613-2620.]

2011

Huchard E, Cowlishaw G (2011) Female-female aggression around mating: an extra cost of sociality in a multimale primate society. Behavioural Ecology 22: 1003-1011.

King AJ, Clark FE, Cowlishaw G (2011) The dining etiquette of desert baboons: the roles of social bonds, kinship, and dominance in co-feeding networks. American Journal of Primatology 73: 768-774.

King AJ, Sueur C, Huchard E, Cowlishaw G (2011) A rule-of-thumb based on social affiliation explains collective movements in desert baboons. Animal Behaviour 82: 1337-1345.

Meise K, Keller C, Cowlishaw G, Fischer J (2011) Sources of acoustic variation: implications for production specificity and call categorization in chacma baboon (Papio ursinus) grunts. Journal of the Acoustical Society of America 129: 1631-1641.

2010

Célérier A, Huchard E, Alvergne A, Féjan D, Plard F, Cowlishaw G, Raymond M, Knapp LA & Bonadonna F (2010) Detective mice assess relatedness in baboons using olfactory cues. Journal of Experimental Biology 213: 1399-1405. [with commentary]

Huchard E, Alvergne A, Féjan D, Knapp LA, Cowlishaw G & Raymond M (2010) More than friends? Behavioural and genetic aspects of heterosexual associations in wild chacma baboons. Behavioural Ecology and Sociobiology 64: 769-781.

Huchard E, Knapp LA, Wang J, Raymond M & Cowlishaw G (2010) MHC, mate choice and heterozygote advantage in a wild social primate. Molecular Ecology 19: 2545-2561.

Huchard E, Raymond M, Benavides J, Marshall H, Knapp LA & Cowlishaw G (2010) A female signal reflects MHC genotype in a social primate. BMC Evolutionary Biology 10: 96.

Wang J, Brekke P, Huchard E, Knapp L, & Cowlishaw G (2010) Estimation of parameters of inbreeding and genetic drift in populations with overlapping generations. Evolution 64: 1704-1718.

2009

Alvergne A, Huchard E, Caillaud D, Charpentier MJE, Setchell JM, Ruppli C, Fejan D, Martinez L, Cowlishaw G & Raymond M (2009) Human ability to visually recognize kin within primates. International Journal of Primatology 30: 199-210.

Huchard E, Benavides J, Setchell J, Charpentier MJE, Alvergne A, King A, Knapp LA, Cowlishaw G & Raymond M (2009). The importance of shape in sexual signals: evidence from primate sexual swellings. Behavioural Ecology and Sociobiology 63: 1231-1242.

Huchard E, Courtiol A, Benavides J, Knapp LA, Raymond M & Cowlishaw G (2009) Can fertility signals lead to quality signals? Insights from the evolution of primate sexual swellings. Proceedings B: Biological Sciences 276: 1889-1897.

King AJ & Cowlishaw G (2009a) Feeding benefits drive interspecific associations between rock kestrels and desert baboons. Journal of Zoology 277: 111-118.

King AJ & Cowlishaw G (2009b) Leaders, followers, and group decision-making. Communicative and Integrative Biology 2: 2.

King AJ & Cowlishaw G (2009c) All together now: behavioural synchrony in baboons. Animal Behaviour 78: 1381-1387.

King AJ, Isaac NJB & Cowlishaw G (2009) Ecological, social, and reproductive factors shape producer-scrounger dynamics in baboons. Behavioral Ecology 20: 1039-1049.

Zinner d, Groeneveld LF, Keller C, Roos C (2009) Mitochondrial phylogeography of baboons (Papio spp.) – Indication for introgressive hybridization? BMC Evolutionary Biology 9:83.

2008

Huchard E, Weill M, Raymond M, Cowlishaw G & Knapp LA (2008) Polymorphism, haplotype composition, and selection in the Mhc-DRB of wild baboons.Immunogenetics 60: 585-598.

King AJ, Douglas C, Huchard E, Isaac N & Cowlishaw G (2008) Dominance and affiliation mediate despotism in a social primate. Current Biology 18: 1833-1838. [with commentary]

Rands SA, Cowlishaw G, Pettifor RA, Rowcliffe JM & Johnstone RA. (2008) The emergence of leaders and followers in foraging pairs when the qualities of individuals differ. BMC Evolutionary Biology 8: 51.

2006-2007

King A & Cowlishaw G (2007) When to use social information: the advantage of large group size in individual decision making. Biology Letters 3: 137-139.

Huchard E, Cowlishaw G, Raymond M, Weill M & Knapp LAP (2006) Molecular study of Mhc-DRB in wild chacma baboons reveals high variability and evidence for transpecies inheritance. Immunogenetics 58: 805-816.

Rands SA, Pettifor RA, Rowcliffe JM & Cowlishaw G (2006) Social foraging and dominance relationships: the effects of socially-mediated interference.Behavioural Ecology and Sociobiology 60: 572-581.

2000-2005

Cowlishaw G, Lawes MJ, Lightbody M, Martin A, Pettifor R & Rowcliffe JM (2004) A simple rule for the costs of vigilance: empirical evidence from a social forager. Proceedings B: Biological Sciences 271: 27-33. [related research on samango monkeys, Cercopithecus mitis erythrarcus]

Rands SA, Pettifor RA, Rowcliffe JM & Cowlishaw G (2004) State-dependent foraging rules for social animals in selfish herds. Proceedings B: Biological Sciences 271: 2613-2620.

Rands SA, Cowlishaw G, Pettifor RA, Rowcliffe JM & Johnstone RA (2003) The spontaneous emergence of leaders and followers in foraging groups.Nature 423: 432-434.

Hill RA & Cowlishaw G (2002) Foraging female baboons exhibit similar patterns of antipredator vigilance across two populations. In: Eat or be eaten: predation sensitive foraging in primates (ed LE Miller), pp.187-204. Cambridge University Press: Cambridge.

1997-1999

Cowlishaw G (1999) Ecological and social determinants of spacing behaviour in desert baboon groups. Behavioural Ecology and Sociobiology 45: 67-77.

Cowlishaw G (1998) The role of vigilance in the survival and reproductive strategies of desert baboons. Behaviour 135: 431-452.

Cowlishaw G (1997a) Trade-offs between foraging and predation risk determine habitat use in a desert baboon population. Animal Behaviour 53: 667-686.

Cowlishaw G (1997b) Refuge use and predation risk in a desert baboon population. Animal Behaviour 54: 241-253.

Cowlishaw G (1997c) Alarm calling and implications for risk perception in a desert baboon population. Ethology 103: 384-394.

Cowlishaw G & Davies JG (1997) Flora of the Pro-Namib Desert Swakop River Catchment: community classification and implications for desert vegetation sampling. Journal of Arid Environments 36: 271-290.

1994-1996

Cowlishaw G & O'Connell SM (1996) Male-male competition, paternity certainty and copulation calls in female baboons. Animal Behaviour 51: 235-238.

Davies JG & Cowlishaw G (1996) Interspecific competition between baboons and black kites following baboon-ungulate predation in the Namib desert. Journal of Arid Environments 34: 247-249.

Cowlishaw G (1995) Behavioural patterns during baboon group encounters: the role of resource competition and male reproductive strategies. Behaviour 132: 75-86.

O'Connell SM & Cowlishaw G (1995) The post-copulation withdrawal response in female baboons: a functional analysis. Primates 36: 441-446.

Cowlishaw G (1994) Vulnerability to predation in baboon populations. Behaviour 131: 293-304.

O'Connell SM & Cowlishaw G (1994) Infanticide avoidance, sperm competition and mate choice: the function of copulation calls in female baboons.Animal Behaviour 48: 687-694.