Macaca sylvanus
Barbary macaque

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)

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)

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)

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)

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)

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)

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)

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)

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)

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)

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)

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

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)