Macaca sylvanusBarbary macaque

Geographic Range

Macaca sylvanus inhabits Morocco, Algeria, and Gibraltar. The majority of M. sylvanus are found in the Middle and High Atlas mountains and in the Rif mountain regions of Morocco. A smaller population is located in the Tellian Atlas mountains of Algeria. Within the Atlas mountains, M. sylvanus is restricted to the Grand Kabylie mountains, the Petit Kabylie mountains, and the Chiffa Gorges. In Gibraltar, a population of about 200 are maintained by constant reintroduction of new animals. They are the only non-human primates in Europe. (Deag, 1977; Mittermeier and Rylands, 2008)


Barbary macaques prefer habitats consisting of high altitude mountains, cliffs, and gorges. Although they prefer high altitude habitats, up to 2600 m, they can also be found at sea level. Their primary habitat is cedar forests, but they are also found in mixed forests of cedar and holm-cork oak, pure oak forests, shrubby rock outcrops along coasts, and occasionally in grasslands at low elevations. During the winter they are highly arboreal, but become more terrestrial during summer. (Deag, 1977; Fa, 1982; Mittermeier and Rylands, 2008; Ménard and Vallet, 1997; Rowe, 1996; Taub, 1977)

  • Range elevation
    0 to 2600 m
    0.00 to 8530.18 ft

Physical Description

Barbary macaques are covered in thick fur that presumably helps protect them from cold temperatures. Body color ranges from yellowish-gray to darker grayish-brown. Their chests and stomachs are much lighter than the rest of their bodies while their faces are often dark pink. Barbary macaques have noticeably short tails, ranging from 1 to 2 cm long. Females average 450 mm from head to tail, weigh about 11 kg, and display estrus with large anogenital swellings. Males are larger, weighing 16 kg and range in length from 550 to 600 mm. Like all cercopithecids, Barbary macaques have large cheek pouches for carrying food. The dental formula of a Barbary macaque is I2/2, C1/1, P2/2, M3/3. (Deag, 1977; Mittermeier and Rylands, 2008; Richardson, 2009; Rowe, 1996; Taub, 1980)

  • Sexual Dimorphism
  • male larger
  • sexes colored or patterned differently
  • Range mass
    11 to 16 kg
    24.23 to 35.24 lb
  • Range length
    450 to 600 mm
    17.72 to 23.62 in


Barbary macaques are polygynandrous, as males and females have multiple mates. Females display estrus with large anogenital swellings. Females initiate and terminate matings and compete with each other for mates by interrupting copulation. Male rank has little effect which individuals females choose to mate with. By the time estrus is complete, each female has mated with all or nearly all males in her troop. Often, females continue to copulate even when conception is impossible. At the end of breeding season, the combined number of copulations by all the females in a troop can number in the hundreds. Female promiscuity is thought to mask the true identity of an infant's father resulting in paternal support from more than one male. (Brauch, et al., 2007; Kuester and Paul, 1996; Paul, et al., 1993a; Paul, et al., 1993b; Pfefferle, et al., 2007; Rowe, 1996; Small, 1990; Soltis, 2004; Taub, 1977; Taub, 1980)

Although male Barbary macaques have little influence over who they mate with, dominant individuals tend to mate more often than subordinates and tend to mate more with dominant females. Despite competition for females, male Barbary macaques show high tolerance for each other. They compensate for intense sperm competition by having large testical-size to body-size ratios. Older males have more breeding success than younger individuals. Males use three mating strategies when attracting females. Individuals using "proximity-possession" remain in close proximity to a female, which usually ensures one's opportunity to mate. Others use the "pertinacious strategy", where they closely follow a female until they are noticed and allowed to mate. Lastly, lowest ranking males use a "peripheralize and attract strategy", where they stay far away from the female, but shake branches or carry other infants in full view of the female in order to attract her attention. Successful males then mate with a female, only mounting her once for a short period of time before both part ways. (Brauch, et al., 2007; Kuester and Paul, 1996; Paul, et al., 1993a; Paul, et al., 1993b; Pfefferle, et al., 2007; Rowe, 1996; Small, 1990; Soltis, 2004; Taub, 1977; Taub, 1980)

Barbary macaques are very social. Once infants are born, the entire troop takes part in caring for infants (i.e., cooperative breeding). (Brauch, et al., 2007; Kuester and Paul, 1996; Paul, et al., 1993a; Paul, et al., 1993b; Pfefferle, et al., 2007; Small, 1990; Soltis, 2004; Taub, 1977; Taub, 1980)

Mating season of Barbary macaques begins in November and ends in December. Gestation last for 164.2 days, on average, and one offspring is usually born between April and June. Barbary macaques typically reach sexual maturity around 46 months of age, regardless of gender. Females remain in estrus for about 1 month and have their first offspring around 5 years of age. Interbirth interval ranges from 13 to 36 months, and the first offspring is generally smaller than subsequent offspring. Females reach menopause during the last 5 years of their lives, though estrus may continue for a few more years. Females often continue to copulate, even when conception is no longer possible. (Bowman and Lee, 2005; Ménard, et al., 2001; Paul and Kuester, 1996; Rowe, 1996; Taub, 1980; Thierry, 2004)

Although no formal research has been conducted on Barbary macaque infants, in general, newborn primates are altricial and require intense care. Female rhesus macaque, a close relative to Barbary macaques, stay in close contact with their newborns for the first 3 months after birth. In young macaques, weight is often used as a substitute for age. Rhesus macaques are weaned between 1,300 to 1,400 grams. Depending on environmental conditions and resource abundance, infants can be weaned between 200 to 362 days. Average birth weight for Barbary macaques is 450 g. (Bowman and Lee, 2005; Ménard, et al., 2001; Paul and Kuester, 1996; Rowe, 1996; Taub, 1980; Thierry, 2004)

  • Breeding interval
    Barbary macaques breed once a year.
  • Breeding season
    November to December
  • Range number of offspring
    1 to 2
  • Average number of offspring
  • Average number of offspring
  • Range gestation period
    158 to 170 days
  • Average gestation period
    164.2 days
  • Average age at sexual or reproductive maturity (female)
    46 months
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    1399 days
  • Average age at sexual or reproductive maturity (male)
    46 months
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    2007 days

Barbary macaques live in matrilineal societies with youngest daughter ascendancy, thus newborn females outrank everyone else in the immediate family except their mothers. Alloparenting is common among Barbary macaques. The entire troop shows interest in newborns and displays friendly teeth-chattering or lip-smacking towards infants. Unlike many primates, Barbary macaques actively take care of unweaned infants more than weaned infants. Females often help "baby-sit" infants of other females. Infants of closely related females and of the highest ranking female usually receive the most attention. Nulliparous females (i.e., females without offspring) engage in infant-carrying more than other females, but this has not been linked to an increase in the survival of their own infants. Females who have recently lost infants are also more likely to carry others' infants. Research suggests that extensive infant handling may allow females to rise in rank. Infants receiving extra care do not experience higher survival rates than those who do not receive extra care. (Chapais, 2004; Kuester and Paul, 1986; Paul and Kuester, 1996; Rowe, 1996; Small, 1990; Thierry, 2004)

Due to the promiscuous mating system of Barbary macaques, males have no way of knowing which infant is theirs. As a result, they provide paternal care to infants in their troop. For example, adult males group around infants to protect them from approaching predators. In general, males appear to show preference to male infants. Young-adult males tend to develop strong bonds with male infants, and older adult males prefer to take care of any infants from high-ranking females. Females appear to show mating preference to males that provide the most paternal care to their offspring. (Chapais, 2004; Kuester and Paul, 1986; Paul and Kuester, 1996; Rowe, 1996; Small, 1990; Thierry, 2004)

  • Parental Investment
  • altricial
  • male parental care
  • female parental care
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • male
      • female
  • pre-independence
    • protecting
      • male
  • post-independence association with parents
  • extended period of juvenile learning
  • maternal position in the dominance hierarchy affects status of young


Barbary macaques typically live 22 years in the wild. Males rarely live more than 25 years, and females appear to live slightly longer than males. Infants have a 10% mortality rate in the wild. No data has been reported for captive Barbary macaques. (Kuester and Paul, 1999; Mehlman, 1989)


Barbary macaques are diurnal and do most of their foraging and traveling during the day. At night, 2 to 3 individuals may sleep together in same-sex clusters. An average troop has 24 individuals but can contain up to 59 individuals and consist of a 1:1 males-female ratio. Like many social primates, Barbary macaques allogroom one another, which is an important daily ritual that builds strong bonds between individuals, removes parasites, and lowers stress. Males and females form their own hierarchies. Female hierarchies are strictly matrilinear, and each female holds a specific rank with the troop. Newborn daughters inherit the next highest position under her mother, above their older sisters and other in the troop. Females protect social status via aggression, and rank is relatively stable and rarely changes. Male have looser dominance hierarchies and show less aggression towards one another. Compared to most primates, Barbary macaques they are a relatively peaceful species, and nearly half of all intraspecific interactions are aggressive. Most aggressive interactions are restricted to chases and retreats, and 20% of aggressive acts are followed by reconciliatory acts between individuals. (Deag, 1973; Kuester and Paul, 1999; Mehlman, 1989; Ménard and Vallet, 1993; Preuschoft, et al., 1998; Rowe, 1996; Shutt, et al., 2007; Thierry, 2004)

  • Range territory size
    3.07 to 9.01 km^2
  • Average territory size
    7.25 km^2

Home Range

Home ranges of Barbary macaque troops are between 3.07 and 9.01 km^2, with an average 7.2 km^2. Home ranges may overlap with those of other troops by 39% to 100%. (Deag, 1973; Kuester and Paul, 1999; Mehlman, 1989; Ménard and Vallet, 1993; Preuschoft, et al., 1998; Rowe, 1996; Shutt, et al., 2007; Thierry, 2004)

Communication and Perception

Barbary macaques display a variety of facial expressions to show emotions. Females show rounded-mouth threats towards other females as a sign of aggression and dominance. Bared teeth show submission. Lip-smacking and teeth-chattering are signs of appeasement which are often directed at a dominant individual or towards infants. Barbary macaques also display relaxed, open-mouthed, play faces which are thought express happiness. (Fischer, 1998; Fischer, et al., 1998; Hammerschmidt and Fischer, 1998; Hammerschmidt and Todt, 1995; Hammerschmidt, et al., 1994; Mehlman, 1982; Preuschoft, 1992; Preuschoft, et al., 1998; Shutt, et al., 2007; Small, 1989)

Sounds is a crucial part of communication in primates. Barbary macaques scream and grunt at trespassing troops. They also use a loud, high-pitched "ah-ah" call to warn troop members of potential danger. Mating calls from females during copulation have been shown to increase the likelihood of ejaculation in males. Barbary macaques are capable of recognizing individuals by calls, and mothers can recognize their infants by their cries. Young cry a string of high-pitched calls at dusk, presumably to find their mother in sleeping clusters. Troops are familiar with the vocalizations of neighboring troops as well. Studies have shown calls are learned through experience and different social groups may use different dialects. (Fischer, 1998; Fischer, et al., 1998; Hammerschmidt and Fischer, 1998; Hammerschmidt and Todt, 1995; Hammerschmidt, et al., 1994; Mehlman, 1982; Preuschoft, 1992; Preuschoft, et al., 1998; Shutt, et al., 2007; Small, 1989)

Food Habits

Due to dramatic changes in climate in the mountains throughout the year, the diet of Barbary macaques changes seasonally. During spring, they eat various vegetation and feast on caterpillars that live in oak tress. By summer, fruits are plentiful along with other small seeds, roots, and fungi. Barbary macaques become terrestrial foragers during spring and summer to acquire these foods. Occasionally, they also eat small vertebrates such as frogs and tadpoles. Oaks produce acorns during fall, which Barbary macaques feed on during this time. During periods of particularly high mast production, macaques may subsist on acorns for more than half the year. During winter, ground forage becomes limited and Barbary macaques become arboreal again. Arboreal forage during winter consists of the leaves, seeds, and bark of evergreens. (Melhman, 1988; Ménard and Vallet, 1997; Rowe, 1996)

  • Animal Foods
  • amphibians
  • insects
  • Plant Foods
  • leaves
  • roots and tubers
  • wood, bark, or stems
  • seeds, grains, and nuts
  • fruit
  • flowers
  • Other Foods
  • fungus


The primary predator of Barbary macaques are large eagles that patrol the mountains for prey. At least one troop member is constantly vigilant for danger. Barbary macaques give a special high-frequency "ah-ah" alarm call when eagles are spotted. Upon hearing this call, they quickly escape to the lower canopy to hide. Less common predators consist of golden jackals and red foxes. (Deag, 1973; Mehlman, 1982)

Ecosystem Roles

Macaca sylvanus is omnivorous and consumes insects, fruit, and other plant materials. This species is an important seed disperser in the mountains where they reside. They are also an important prey item for eagles, golden jackals, and red foxes. Macaca sylvanus is host to a number of ecto- and endoparasites including flatworms, roundworms, sucking lice, parasitic Protozoa (Giardia), and the viruses that cause canine distemper (Paramyxoviridae) and encephalomyocarditis (Picornaviridae), which can sometimes be fatal in M. sylvanus. (Canelli, et al., 2010; Cohn, et al., 2007; Martin, 1950; Mehlman, 1982; San Román and Horsley, 2009)

  • Ecosystem Impact
  • disperses seeds
Commensal/Parasitic Species
  • flatworms (Trematoda)
  • roundworms (Nematoda)
  • sucking lice (Anoplura)
  • parasitic protozoa (Giardia)
  • Paramyxoviridae
  • Picornaviridae

Economic Importance for Humans: Positive

Due to their low numbers in the wild, Barbary macaques are not commonly used in labs, but some labs still utilize them for biomedical research. There is also a small illegal pet trade for them. In Gibraltar they attract a large number of tourists. (Deag, 1977)

Economic Importance for Humans: Negative

Barbary macaques Macaca sylvanus occasionally raid gardens or farms, which has led to trapping and illegal poaching. (Deag, 1977)

  • Negative Impacts
  • crop pest

Conservation Status

Barbary macaques are listed as endangered on teh IUCN's Red List of Threatened Species. In Morocco and Algeria they are a federally protected species, but are still in danger of local extinction. Their greatest threat is habitat loss, predominantly from logging. As a result, macaques are pushed farther up mountains into nutrient poor areas where survival is more difficult. Resource competition with domestic goats has become an increasing problem as well. Recently, Barbary macaques have changed their feeding patterns to incorporate more bark and flowers so they can survive. Minor threats include trapping, illegal poaching, and death by herding dogs. About 300 infants are taken out of Morocco annually for pet trade. Barbary macaques are listed under Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). (Ciani, et al., 2005; Deag, 1977; Mehlman, 1989; Mittermeier and Rylands, 2008; San Román and Horsley, 2009; van Lavieren and Wich, 2010)


Judy Jinn (author), University of Michigan-Ann Arbor, Phil Myers (editor), University of Michigan-Ann Arbor, John Berini (editor), Animal Diversity Web Staff.



living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map


uses sound to communicate


young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.


Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.


uses smells or other chemicals to communicate

cooperative breeder

helpers provide assistance in raising young that are not their own

  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates


humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.


animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females


union of egg and spermatozoan


forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.


offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males


having the capacity to move from one place to another.


This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.


an animal that mainly eats all kinds of things, including plants and animals

pet trade

the business of buying and selling animals for people to keep in their homes as pets.


the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

scrub forest

scrub forests develop in areas that experience dry seasons.

seasonal breeding

breeding is confined to a particular season


reproduction that includes combining the genetic contribution of two individuals, a male and a female


associates with others of its species; forms social groups.


uses touch to communicate


that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).


Living on the ground.


defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.


A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.


uses sight to communicate


reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.


Bowman, J., P. Lee. 2005. Growth and threshold weaning weights among captive Rhesus macaques. American Journal of Physical Anthropology, 96/2: 159-175. Accessed March 25, 2011 at

Brauch, K., D. Pfeffrle, K. Hodges, U. Möhle, J. Fischer, M. Heistermann. 2007. Female sexual behavior and sexual swelling size as potential cues for males to discern the female fertile phase in free-ranging Barbary macaques (Macaca sylvanus) of Gibraltar. Hormones and Behavior, 52/3: 375-383. Accessed March 12, 2011 at

Canelli, E., A. Luppi, A. Lavazza, D. Lelli, E. Sozzi, A. Moreno-Martin, D. Gelmetti, E. Pascotto, C. Sandri, W. Magnone, P. Cordioli. 2010. Encephalomyocarditis virus infection in an Italian zoo. Virology Journal, 7/64: 1-7.

Chapais, B. 2004. How kinship generates dominance structures: a comparative perspective. Pp. 186-187 in B Thierry, M Singh, W Kaumanns, eds. Macaque societies: a model for the study of social organization. United Kingdom: Cambridge University Press.

Ciani, A., L. Palentini, M. Arahou, L. Martinoli, C. Capiluppi, M. Mouna. 2005. Population decline of Macaca sylvanus in the middle atlas of Morocco. Biological Conservation, 121/4: 635-641. Accessed March 12, 2011 at

Cohn, D., V. Smith, M. Pizzaro, L. Jones-Engel, G. Engel, A. Fuentes, E. Shaw, J. Cortes. 2007. Pediculosis in Macaca sylvanus of Gibraltar. Veterinary Parasitology, 145/1-2: 116-119.

Deag, J. 1973. Intergroup encounters in the wild Barbary macaque (Macaca sylvanus l.). Pp. 316-370 in R Michael, J Crook, eds. Comparative ecology and behaviour of primates. New York: Academic Press.

Deag, J. 1977. The status of the Barbary Macaque Macaca sylvanus in captivity and factors influencing its distribution in the wild. Pp. 268-285 in P of Monaco, G Bourne, eds. Primate Conservation. New York, York: Academic Press.

Fa, J. 1982. A survey of population and habitat of the barbary macaque (macaca sylavanus) l. in north morocco. Biological Conservation, 24/1: 45-66. Accessed March 11, 2011 at

Fischer, J. 1998. Barbary macaques categorize shrill barks into two call types. Animal Behaviour, 55/4: 799-807. Accessed March 12, 2011 at

Fischer, J., K. Hammerschmidt, D. Todt. 1998. Local variation in Barbary macaque shrill barks. Animal Behaviour, 56/3: 623-629. Accessed March 12, 2011 at

Gachot-Neveu, H., N. Menard. 2004. Gene flow, dispersal patterns, and social organization. Pp. 117-130 in B Thierry, M Singh, W Kaumanns, eds. Macaque societies: a model for the study of social organization. United Kingdom: Cambridge University Press.

Hammerschmidt, K., V. Ansorge, J. Fischer, D. Todt. 1994. Dusk calling in barbary macaques (Macaca sylvanus): Demand for social shelter. American Journal of Primatology, 32/4: 277-289. Accessed March 12, 2011 at

Hammerschmidt, K., J. Fischer. 1998. Maternal discrimination of offspring vocalizations in Barbary macaques (Macaca sylvanus). Primates, 39/2: 231-236. Accessed March 12, 2011 at

Hammerschmidt, K., D. Todt. 1995. Individual differences in vocalisations of young Barbary macaques (Macaca sylvanus): a multi-parametric analysis to identify critical cues in acoustic signalling. Behaviour, 132/5: 381-399. Accessed March 12, 2011 at

Kuester, J., A. Paul. 1996. Female-female competition and male mate choice in Barbary macaques (Macaca sylvanus). Behaviour, 133/9: 763-790. Accessed March 12, 2011 at

Kuester, J., A. Paul. 1999. Male migration in Barbary macaques (Macaca sylvanus) at Affenberg Salem. International Journal of Primatology, 20/1: 85-106. Accessed March 12, 2011 at

Kuester, J., A. Paul. 1986. Male-infant relationships in semifree-ranging Barbary macaques (Macaca sylvanus) of affenberg salem/FRG: Testing the “male care” hypothesis. American Journal of Primatology, 10/4: 315-327. Accessed March 21, 2011 at

Martin, L. 1950. Susceptibility of monkeys (Macaca sylvanus) to distemper. Bull. -Acad. vet. Fr., 23: 457-462.

Mehlman, P. 1989. Comparative Density, Demography, and Ranging Behavior of Barbary Macaques (Macaca sylvanus) in Marginal and Prime Conifer Habitats. International Journal of Primatology, 10/4: 269-292.

Mehlman, P. 1982. Aspects of the ecology and conservation of the Barbary macaque in the fir forest habitat of the Moroccan Rif mountains. Pp. 165-199 in J Fa, ed. The Barbary macaque. New York: Plenum Publishing Corporation.

Mehlman, P., R. Parkhill. 1988. Intergroup interactions in wild Barbary macaques (Macaca sylvanus), Ghomaran Rif mountains, Morocco. American Journal of Primatology, 15/1: 31-44. Accessed March 12, 2011 at

Melhman, P. 1988. Food resources of the wild Barbary macaque (Macaca sylvanus) in high-altitude fir forest, Ghomaran Rif, Morocco. Journal of Zoology, 214/3: 469-490. Accessed March 22, 2011 at

Mittermeier, R., A. Rylands. 2008. "Macaca sylvanus" (On-line). Accessed March 21, 2011 at

Ménard, N., D. Vallet. 1997. Behavioral responses of Barbary macaques (Macaca sylvanus) to variations in environmental conditions in Algeri. American Journal of Primatology, 43/4: 285-304. Accessed March 12, 2011 at;2-T/abstract.

Ménard, N., D. Vallet. 1993. Population dynamics of Macaca sylvanus in Algeria: An 8-year study. American Journal of Primatology, 30/2: 101-118. Accessed March 12, 2011 at

Ménard, N., F. von Segesser, W. Scheffrahn, J. Pastorini, D. Vallet, B. Gaci, R. Martin, A. Gautier-Hion. 2001. Is male–infant caretaking related to paternity and/or mating activities in wild Barbary macaques (Macaca sylvanus)?. Comptes Rendus de l'Académie des Sciences - Series III - Sciences de la Vie, 324/7: 601-610. Accessed March 12, 2011 at

Paul, A., J. Kuester. 1996. Infant handling by female Barbary macaques ( Macaca sylvanus ) at Affenberg Salem: testing functional and evolutionary hypotheses. Behavioral Ecology and Sociobiology, 39/2: 133-145. Accessed March 12, 2011 at

Paul, A., J. Kuester, D. Podzuweit. 1993. Reproductive senescence and terminal investment in female Barbary macaques (Macaca sylvanus) at Salem. Fologia Primatologica, 14/1: 105-124. Accessed March 12, 2011 at

Paul, A., J. Kuester, A. Timme, J. Arnemann. 1993. The association between rank, mating effort, and reproductive success in male Barbary macaques (Macaca sylvanus). Primates, 34/9: 491-502. Accessed March 12, 2011 at

Paul, A., D. Tommen. 1984. Timing of birth, female reproductive success and infant sex ration in semi-free ranging barbary macaque (Macaca sylvanus). Folia Primatologica, 42/1: 2-16. Accessed March 12, 2011 at

Pfefferle, D., K. Brauch, M. Heistermann, J. Hodges, J. Fischer. 2007. Female Barbary macaque (Macaca sylvanus) copulation calls do not reveal the fertile phase but influence mating outcome. Proceedings of the Royal Society, 275: 571-578. Accessed March 12, 2011 at

Preuschoft, S. 1992. “Laughter” and “smile” in Barbary macaques (Macaca sylvanus). Ethology, 91/3: 220-236. Accessed March 22, 2011 at

Preuschoft, S., A. Paul, J. Keuster. 1998. Dominance styles of female and male Barbary macaques (Macaca sylvanus). Behaviour, 135/6: 731-735. Accessed March 12, 2011 at

Prud'homme, J., B. Chapais. 1993. Rank relations among sisters in semi-free-ranging Barbary macaques (Macaca sylvanus). International Journal of Primatology, 14/3: 405-430. Accessed March 12, 2011 at

Richardson, M. 2009. "Barbary macaque" (On-line). Accessed March 21, 2011 at

Rowe, N. 1996. The pictorial guide to the living primates. East Hampton, NY: Pogonias Press.

San Román, S., S. Horsley. 2009. "Deforestation spells doom for Barbary Macaque" (On-line). Accessed March 21, 2011 at

Shutt, K., A. MacLarnon, M. Hiestermann, S. Semple. 2007. Grooming in Barbary macaques: better to give than to receive?. Biology Letters, 3: 231-233. Accessed March 12, 2011 at

Small, M. 1989. Alloparental behaviour in Barbary macaques, Macaca sylvanus. Animal Behaviour, 39/2: 297-306. Accessed March 12, 2011 at

Small, M. 1990. Promsicuity in barbary macaques (Macaca sylvanus). American Journal of Primatology, 20/4: 267-282. Accessed March 12, 2011 at

Soltis, J. 2004. Mating systems. Pp. 137-150 in B Thierry, M Singh, W Kaumanns, eds. Macaque societies: a model for the study of social organization. United Kingdom: Cambridge University Press.

Taub, D. 1980. Female choice and mating strategies among wild Barbary macaques (Macaca sylvanus). Pp. 287-367 in D Lindburg, ed. The macaques: studies in ecology, behavior, and evolution. New York, NY: Van Nostrand Reinhold Company.

Taub, D. 1977. Geographic Distribution and Habitat Diversity of the Barbary Macaque Macaca sylvanus L.. Folio Primatologica, 27/2: 108-133. Accessed March 12, 2011 at

Theirry, B., M. Singh, W. Kaumanns. 2004. Macaque societies: a model for the study of social organization. United Kingdom: Cambridge University Press.

Thierry, B. 2004. Social epigenesis. Pp. 267-288 in B Thierry, M Singh, W Kaumanns, eds. Macaque societies: a model for the study of social organization. United Kingdom: Cambridge University Press.

van Lavieren, E., S. Wich. 2010. Decline of the Endangered Barbary macaque Macaca sylvanus in the cedar forest of the Middle Atlas Mountains, Morocco. Oryx, 44: 133-138. Accessed March 12, 2011 at