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 in Tsaobis Leopard Park on the edge of the Namib desert in central Namibia. The Tsaobis baboon population is comprised of several troops of chacma baboons Papio ursinus. The first study of this baboon population began in 1990, and there has been continuous study of the population since 2000.

At present, we are working with two troops, each of about 60 individuals. Each baboon is individually recognisable, of known age (estimated through patterns of tooth eruption and wear) and of known relatedness to other individuals in the population (through microsatellite genotyping).

Tsaobis is a beautiful desert wilderness. The main feature of our study area is the dry riverbed and associated woodlands of the ephemeral Swakop River. It is in these woodlands that the baboons carry out most of their foraging over the winter months.

Immediately to the south of the riverbed, alluvial and gravel plains border rocky foothills and dramatic mountains, in which the sleeping cliffs of the baboons are found. The woodland along the riverbed is sustained by groundwater; beyond the river course there are only desert shrubs and small, sparse trees.

The climate is extremely arid: annual rainfall is only 123mm and shade temperatures can exceed 40oC in the austral summer. In addition to baboons, Tsaobis is home to a wide variety of wildlife including leopard, mountain zebra, kudu, springbok, and klipspringer.

Student Information:

PhD students play a key role in the research undertaken on the Tsaobis baboons. However, we regret that we will not be advertising any new PhD studentships over the coming months.

We are also always interested to receive proposals from PhD students who already have funding and are seeking an appropriate site for their study. Please contact the Project Director, Guy Cowlishaw.

Unfortunately, we are unable to accommodate students who wish to carry out undergraduate projects at Tsaobis.

Research Projects:

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 Alecia Carter, Julio Benavides, Elise Huchard, Andrew King, and Harry Marshall.

 

The Tsaobis Baboon Project is a long-term study of a desert baboon population in Namibia. The Project is based at the Institute of Zoology, the research arm of the Zoological Society of London, with collaborators at a variety of academic institutions around the country. The field site is located at Tsaobis Nature Park on the edge of the Namib Desert.

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

At present, we are working with two troops, each of about 60 animals. Each baboon is individually recognisable, of known age (estimated through patterns of tooth eruption and wear) and of known relatedness to other individuals in the population (established through microsatellite genotyping).
 

Volunteer Positions

Background

The first nine months of the 2014-2015 field season will run between late April 2014 and late January 2015. Our research will focus on three areas: (1) Sexual coercion and conflicts over reproduction, and (2) Social networks and information transfer, and (3) Climate, behaviour, and host-parasite dynamics. Fieldwork for these studies will involve data collection from two study troops. There will also be some vegetation and wider environmental monitoring activities, related to the baboons' use of the environment.

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. There are five start-and-end date options for these positions: two for 6-month volunteers (late April – late October; late July – late January) and three for 3-month volunteers (late April – late July; late July – late October; late October – late January). If you have a preference for which research area you would like to work on, please let us know. However, bear in mind that some of our studies will only be running in certain periods, as follows:

 

early 6 months late April to late October Sexual conflict, Social networks (3) & HP dynamics (3)
late 6 months late July to late January Sexual conflict, Host-parasite dynamics
early 3 months late April to late July Sexual conflict, Social networks
middle 3 months late July to late October Sexual conflict, Host-parasite dynamics
late 3 months late October to late January Sexual conflict, Host-parasite dynamics

 

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

Although we cannot afford to pay volunteers, 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 first six months of the field season. 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, early or late; 3-months, early, middle, or late), 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)
    - If you have a preference, which research area you would prefer to work in, i.e. Sexual coercion, Social networks, or Host-parasite dynamics; if you do not have a preference, please state no preference
  • 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 6th January 2014. We will notify successfully shortlisted candidates by the end of that day, and interviews will be held in London the following week, on Mon 13th and Tues 14th January 2014. 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 is a desert environment. The climate can be extremely hot, and there is very little rainfall. However, our fieldwork takes place primarily over the austral winter months, so the nights are cold (the temperature can fall as low as 0oC) and the days are usually fresh, sunny and warm. Rain falls during the summer months (between November and April, primarily January-February), but is light and erratic.

Landscape

Tsaobis is a beautiful desert wilderness. The main feature of our study area is the dry river bed and associated woodlands of the ephemeral Swakop River. It is in these woodlands that the baboons carry out most of their foraging over the winter months. Immediately to the south of the riverbed, alluvial and gravel plains encircle rocky foothills and mountains, in which the sleeping cliffs of the baboons are found. The woodland along the river bed is sustained by the watertable; beyond the river course there are only desert shrubs and small, sparse trees.

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).

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, often ascending and descending steep hills and slippery scree slopes as they follow the baboons. Towards the end of October, in the early summer months, the days also become hotter (40oC plus) and longer (14 hours). 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. On-site supervision and detailed guidelines will be provided at Tsaobis, describing how to work with the baboons in this desert landscape.

Vegetation mapping and surveying

At the beginning of the field season, and then at monthly intervals, vegetation mapping and phenological monitoring will also be undertaken. This work is an important complement to the baboon behavioural data, since it will describe the spatial and temporal availability of food patches to the baboons.

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

Accomodation 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 at outside 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 can be a little 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.

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

Due to the cold nights and hot days, field team members should bring light summer clothes plus sweatshirts and jumpers for the cooler weather. Woolly hat, gloves, and a warm fleece are essential for cold winter mornings and evenings (and for possible trips to the coast). 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 windcheater can also be useful for windy winter mornings. 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. 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 molded 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.
  • 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. 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.

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.

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 Sciences 271: 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.

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.

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.

2006

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.

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.

1994-1999

Cowlishaw G (1999b) 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.

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

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.