Barbary macaques are yellowish gray to grayish brown with paler underparts and a dark pinkish face (MacDonald 1985).The Barbary macaque has an extremely short vestigial tail. This tail reduction is a uniquely derived feature of the Barbary macaque and resulted in the popular misnomer of Barbary Ape.
Morphological evidence suggests an early divergence of M. sylvanus from other extant macaques. Barbary macaques lack some morphological specialisations of other macaques (Groves 1989). Adult males have ischial callosities that fuse across the midline, unlike other macaques that have lost this condition (Groves 1989). High-resolution restriction site mapping of the mitochondrial ribosomal genes confirm that Macaca sylvanus is the sister clade to all Asian macaques (Morales and Melnick 1998).The ancestral macaque stock originated in Africa ca. 7-6 million years ago (Ma) and dispersed into to Eurasia ca. 6–5 Ma (Fooden 2007). Species that are closely related to M. sylvanus have been recorded in the European fossil record dating from the Middle Pliocene to the Middle Pleistocene (Gentili et al 1998). Macaques became extinct in Europe during the Pleistocene. The Gibraltar colony of Barbary macaques was reintroduced, probably in historic times.The divergence of the Moroccan population and the two main Algerian subpopulations can be dated to approximately 1.6 million years ago, assuming an origin of 5.5 million years ago for the genus Macaca. The distinction between Moroccan and Algerian haplotypes provides a basis for determining of the origin of the Gibraltar macaques. The Gibraltar sample includes both Algerian and Moroccan haplotypes, implying a dual origin of the founding females (Modolo et al 2005).
Barbary macaques show moderate sexual dimorphism in body size and the larger males reach attain sexual maturity and adult body weight later than females.Average body weight in wild-collected adults is 9.9kg in females and 14.5kg in males (Fooden, 2007).
Lifespans of up to is 30 years have been recorded a semi free-ranging population (Fooden, 2007).
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.
Biogeographic Regions: palearctic (Native )
In Morocco, M. sylvanus can still be found in the Rif mountains (northern Morocco) and the Middle and High Atlas mountains (central and southern Morocco). In Algeria, it is found in the Tellian Atlas (Petite Kabylie and Grande Kabylie mountains, and an isolated population in the Chréa National Park) (northern Algeria). More specifically, in Morocco the High Atlas populations (two anciently separated populations) are found in the Bou Tferda valley to Demnat region and around the Ourika Valley, respectively; the Rif populations are primarily on Mounts Lakraa, Tissouka, Tazoute, Bouhacham, and Djebel Moussa. In Algeria, from west to east, Barbary Macaque populations are distributed as follows: (a) Chiffa gorges (Chréa National Park); (b) Djurdjura forests and rocky cliffs (Djurdjura National Park, Grande-Kabylie); (c) Akfadou forests, including a small ‘subpopulation’ recently settled in degraded forests and maquis, near El-Kseur (Grande-Kabylie and Petite-Kabylie); (d) Cap Carbon, Aiguades and Pic des Singes (Gouraya National Park; Béjaïa, Petite-Kabylie); (e) Chaabet-el-Akhra Gorges (Kherrata; Béjaïa, Petite-Kabylie); (f) ‘Massif des Babors’ forests (Sétif and Béjaïa, Petite-Kabylie); (g) Guerrouch forest (Taza National Park; Jijel, Petite-Kabylie). Additionally, a Barbary Macaque population might still occur in Djebel Bouzegza (Boumerdes, Grande-Kabylie) (F. Belbachir pers. comm. 2007).
The species occurs from sea level to 2,600 m (Cuzin 2003), but in Morocco at least it is most common at altitudes above 1,000 m (F. Cuzin pers. comm. 2007).
Barbary macaques (Macaca sylvanus) are now restricted to isolated forest fragments in the Atlas and Rif Mountain ranges in Morocco and Algeria. The Moroccan and Algerian populations are separated by a gap of about 700km (Fooden 2007).The species was once more widely distributed in North Africa. A large and long-established semi-wild colony is present on Gibraltar.An 18th century document preserved in the British Museum notes that “a great quantity” of Barbary macaques was introduced into the British garrison in 1740 (Fooden 2007).The Barbary macaque is now the only species of the genus Macaca found outside Asia.
Barbary macaques inhabit mid to high altitude mixed cedar and oak forest, and scrub and cliffs (MacDonald 1985).In Algeria monkey troops were observed living in non-deciduous cedar-green oak forest, deciduous oak forest and on rocky mountainous ridges lacking arboreal vegetation. Monkey densities were higher in cedar forest than in deciduous forest, and the ceder forest troop had a more balanced sex ratio and better survival of young animals. The mountain troop had an unstable aging population with low population density (Menart et al 1985).
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.
Range mass: 11 to 16 kg.
Range length: 450 to 600 mm.
Other Physical Features: endothermic ; homoiothermic; bilateral symmetry
Sexual Dimorphism: male larger; sexes colored or patterned differently
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.
Range elevation: 0 to 2600 m.
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: savanna or grassland ; forest ; scrub forest ; mountains
Habitat and Ecology
In Algeria, occupied habitats include mixed cedar (Cedrus atlantica) and oak forests; humid zen oak (Quercus canariensis) and cork oak (Quercus suber) mixes; Algerian fir (Abies numidica), cedar and zen oak mixed forests; and gorges dominated by scrub vegetation (Taub 1977; F. Belbachir pers. comm. 2007). Moroccan habitats include high cedar forests (Cedrus atlantica), cedar/holm oak (Quercus ilex) mixtures, pure holm oak forests and cliffs and gorges dominated by scrub vegetation. The cedar/oak forests of the Middle Atlas region contain the largest remaining population of M. sylvanus, and are considered to be optimal habitat for the species (Camperio Ciani et al. 2001). In habitats with cedar, macaques can reach densities of 25-40 individuals per km2 or greater, whereas in habitats without cedar much lower densities of 5-7 individuals per km2 are reported (Fa 1984; Mehlman 1989). In all occupied habitats, there is a species of oak available (F. Cuzin pers. comm., 2007).
Its diet is primarily composed of cedar (Cedrus atlantica) and the oak (Quercus sp.), which make up over 50% of its total intake. It eats fruits (33% of its intake), tree leaves (16%), and other plant parts (24%).
M. sylvanus lives in social groups of up to 80 individuals of both sexes with a modal size of 40 individuals (Ménard 2002). Life span in the wild is known to be up to 22 years (Lindenfors 2002). Females reach sexual maturity between 3.5 and 4 years of age and males between 4.5 and 7 years of age. In the wild, average age at first birth is 5.3 years for females (Ménard and Vallet 1993, 1996; Lindenfors 2002), and the birth interval is 1.3 years (Taub 1974 in Fa 1984; Ménard and Vallet 1993, 1996).
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.
Animal Foods: amphibians; insects
Plant Foods: leaves; roots and tubers; wood, bark, or stems; seeds, grains, and nuts; fruit; flowers
Other Foods: fungus
Primary Diet: omnivore
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.
Ecosystem Impact: disperses seeds
- flatworms (Trematoda)
- roundworms (Nematoda)
- sucking lice (Anoplura)
- parasitic protozoa (Giardia)
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.
- eagles (Accipitridae)
- golden jackal (Canis aureus)
- red fox (Vulpes vulpes)
This list may not be complete but is based on published studies.
Life History and Behavior
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.
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.
Communication Channels: visual ; tactile ; acoustic
Perception Channels: visual ; tactile ; acoustic ; chemical
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.
Status: wild: 22 years.
Status: wild: 22 years.
Status: wild: 22.0 years.
Status: wild: 20.0 years.
Status: captivity: 17.0 years.
Lifespan, longevity, and ageing
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.
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.
Barbary macaques are very social. Once infants are born, the entire troop takes part in caring for infants (i.e., cooperative breeding).
Mating System: polygynandrous (promiscuous) ; cooperative breeder
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.
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.
Breeding interval: Barbary macaques breed once a year.
Breeding season: November to December
Range number of offspring: 1 to 2.
Average number of offspring: 1.
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 (male): 46 months.
Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization ; viviparous
Average birth mass: 450 g.
Average number of offspring: 1.5.
Average age at sexual or reproductive maturity (male)
Sex: male: 2007 days.
Average age at sexual or reproductive maturity (female)
Sex: female: 1399 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.
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.
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
Molecular Biology and Genetics
Barcode data: Macaca sylvanus
Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.
See the BOLD taxonomy browser for more complete information about this specimen and other sequences.
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Download FASTA File
Statistics of barcoding coverage: Macaca sylvanus
Public Records: 2
Specimens with Barcodes: 4
Species With Barcodes: 1
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).
CITES: appendix ii
IUCN Red List of Threatened Species: endangered
IUCN Red List Assessment
Red List Category
Red List Criteria
- 1994Vulnerable(Groombridge 1994)
- 1990Vulnerable(IUCN 1990)
- 1988Vulnerable(IUCN Conservation Monitoring Centre 1988)
- 1986Vulnerable(IUCN Conservation Monitoring Centre 1986)
According to Moroccan authorities the number of macaques has increased over the last decade (van Lavieren 2006). However, a number of detailed surveys have documented marked declines in population density, along with losses of a number of sites (Von Segesser et al. 1999; Camperio Ciani et al. 2005; van Lavieren 2006). The remaining subpopulations in the Rif mountains and High Atlas mountains of Morocco are fragmented and sometimes small. Fourteen isolated populations were identified in the High Atlas (Cuzin 2003). Algerian subpopulations are also fragmented (Mehlman 1989; Fa 1984; Von Segesser et al. 1999). In 1999, just seven widely separated isolated populations existed, whereas 35 years ago they were found in at least six additional localities in Algeria (Von Segesser et al. 1999). The remaining isolated populations are now completely separated by distances of 50-100 km, with no corridors. The only area where M. sylvanus is thought to occur in relative abundance is the cedar forest area of the central Middle Atlas, which represents the largest refuge of the North African forest ecosystem. In the 1980s it was estimated that 65-75% of the world's remaining population of Macaca sylvanus lived in this area (Camperio Ciani 1986). The populations in the Middle Atlas outside the central region were found to be much lower in density (Taub 1975). The central region of the Middle Atlas thus has a crucial role in the survival of the species. In the mid-1970s, population densities in the central Middle Atlas cedar forest were variously estimated at 60-70 individuals per km2 (Deag 1974) or 43 per km2 (Taub 1975). Subsequent studies showed that the population density in this region declined from 44 to 25 individuals per km2 over the last two decades of the 20th century (Camperio Ciani et al. 1999). In 2005, surveys in the Middle Atlas showed an average density of 15 to 20 individuals per km2 (van Lavieren 2006), in some areas dropping to an average density of 7-10 individuals per km2 (Camperio Ciani et al. 2005). Surveys conducted in 2006-2007 show that some habitat fragments contained only one or a few small groups. The population density in these fragments was little more than 0 individuals per km2 and local extinction appears imminent (N. Ménard pers. comm. 2008).
These data indicate that, in the central Middle Atlas (the global stronghold of the species), average population density has declined by c.50-80% over the last 30 years. Because of political unrest, many other macaque populations have not been censused since 1990 (K. de Smet pers. comm. 2007), so the population trend in these areas cannot be quantified but a declining trend is suspected.
The destruction and degradation of the macaque’s forest habitat is the most serious threat to the species. Severe habitat loss, fragmentation and degradation have been caused by domestic and industrial consumption of wood, use of fire, clearing for cultivation and overgrazing by sheep and goat herds (Taub 1977, Fa 1984, Camperio Ciani 1986, Menard and Vallet, 1993). This is exacerbated by poor forestry management (often Cedrus atlanticus is favoured and efforts are made to reduce Quercus ilex, but the latter is a crucial resource for macaques: Ménard and Vallet, 1986, 1997).). Additionally, in recent years shepherds have increasingly settled near water sources. As shepherd tribes move into the forest, they often enclose open water sources with cement wells so they can extract water for their herds. As a result, macaques and other wildlife have been excluded from water sources in areas where it was previously accessible to them (Camperio Ciani et al. 2003).
Live trade is also a significant threat to the wild M. sylvanus population. Most of the specimens taken from the wild are for the international pet trade. In comparison with the international trade, offtake for local purposes is relatively low, although M. sylvanus are kept fairly frequently as pets in Morocco (van Lavieren 2004) and local commercial use (remunerated photography in tourist areas and restaurants) has been reported in Algeria (F. Belbachir pers. comm. 2007). M. sylvanus seems to be rarely used for food, except for some old reports in Algeria (Deag 1977). Reports of capture for the international pet trade date back to 1977 (Deag 1977), and since then the trade has increased markedly (van Lavieren 2004). Sanctuaries and zoos in Europe have become overstocked with Barbary Macaque infants offered to them by authorities and ex-owners, most infants coming straight from the wild (van Lavieren 2004). Infant M. sylvanus are offered openly and covertly for sale on markets all over Morocco, and prices of up to 200 Euro per animal have been recorded (van Lavieren 2004). Prices of about 100 Euros per animal have been recorded in Algeria, with Chaabet-el-Akhra gorges (Béjaïa) and Yakouren (Akfadou) being critical locations where illegal trade occur (F. Belbachir pers. comm. 2007). It is estimated that up to 300 infants are taken annually from the Moroccan habitats (van Lavieren 2004). The maximum sustainable offtake of macaques in the Central Middle Atlas region has been estimated at between 200 and 250 individuals per year (van Lavieren 2004). If the estimate of 300 macaques taken annually from the wild is correct, then the offtake exceeds sustainability by up to 50% per year in this population.
Over the last few decades there have been repeated surveys of the species in the wild, mainly carried out by scientists from outside the range states. The national authorities are aware of these surveys but do not always support or agree with the outcome (E. van Lavieren pers. comm. 2006). The Moroccan authorities initiated studies on the ecology, demography and genetics of the Moroccan Middle Atlas population to run from 2006-2008 (M. Mouna pers. comm. 2006, N. Menard pers. comm. 2008). These studies are conducted by N. Menard’s team and will provide new information on population demographics and density.
In both Morocco and Algeria, the national forestry departments are responsible for the management and the protection of flora and fauna. There has been much debate about appropriate management of M. sylvanus. The national forestry department in Morocco contends that there are too many macaques in the region (due to the disappearance of predators such as the leopard Panthera pardus), and that they are responsible for the degradation of cedar forests through their bark stripping behaviour. Consequently culls and translocations have been carried out in some areas (F. Cuzin pers. comm. 2007). However, field studies and surveys indicate that the population is declining and that bark stripping behaviour is potentially induced by water shortage (Camperio Ciani et al. 2001, 2003), although the latter assertion is controversial as bark-stripping has been observed to occur when drinking water is freely available and water content in bark is lower than that found in other available food resources (Menard and Qarro 1999). An alternative hypothesis is that Barbary Macaques strip cedar bark when some nutrients they search for are unavailable in other food resources (N. Ménard unpubl. data).
A number of activities have been undertaken to increase public awareness and reduce illegal trade in this species. AAP (a Netherlands-based sanctuary for exotic animals) has initiated a project to combat illegal trade of Barbary Macaques into Europe, involving inter alia awareness raising among potential buyers, cooperation with authorities in the consuming countries and training of customs officers in Spain (E. van Lavieren pers. comm. 2006). In Algeria, the ecological association Amazer-N’-Kefrida carried out public education and awareness raising campaigns in 2006 and 2007, against illegal trade, commercial uses, and improper artificial feeding of Barbary Macaques, closely collaborating with the Gendarmerie Nationale (National Gendarmery), the Algerian Customs, the Laboratory of Ecology and Environment of the Université de Béjaïa, the General Forests Directorate, and the National Parks (F. Belbachir pers. comm. 2007).
Most habitats of M. sylvanus in Algeria have national park status. This is not the case for all the habitats in Morocco. A part of the Rif mountains has national park status, and there are plans to make the cedar/mixed forest in the central Middle Atlas a national park, but a large part of the species' habitat in Morocco lies outside protected areas. Protected areas containing populations of this species include the Toubkal National Park, Eastern High Atlas N.P., Ifran N.P., Talassemtane N.P., Bou Hachem Reserve and Djebel Moussa N.P. in Morocco, and the Djurdjura, Taza, Chréa, and Gouraya National Parks in Algeria. However, the parks in Algeria and Morocco suffer from significant human impact, and all these areas required much stricter protection than is currently in place. Barbary Macaques breed well in captivity. The possibility of reintroducing animals to northern Tunisia, where they went extinct in the 1900s, should also be studied.
Relevance to Humans and Ecosystems
Barbary macaques Macaca sylvanus occasionally raid gardens or farms, which has led to trapping and illegal poaching.
Negative Impacts: crop pest
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.
Positive Impacts: pet trade ; ecotourism ; research and education
The Barbary macaque (Macaca sylvanus), Barbary ape, or magot is a species of macaque unique for its distribution outside Asia and for its vestigial tail. Found in the Atlas Mountains of Algeria and Morocco and El-Kouf National Park in Libya along with a small population of unknown origin in Gibraltar, the Barbary macaque is one of the best-known Old World monkey species.
The Barbary macaque is of particular interest because males play an atypical role in rearing young. Despite uncertain paternity, males are integral to raising infants. Generally, Barbary macaques of all ages and sexes contribute in alloparental care of young.
Macaque diets consist primarily of plants and insects and they are found in a variety of habitats. Males live to a maximum of 25 years while females may live up to 30 years. Besides humans, they are the only free-living primates in Europe. Although the species is commonly referred to as the "Barbary ape", the Barbary macaque is actually a true monkey. Its name refers to the Barbary Coast of Northern Africa.
The Barbary macaque population of Gibraltar is the only such population outside of Northern Africa and the only population of wild monkeys in Europe. The Rock of Gibraltar is populated by approximately 230 macaques.
The monkey is yellowish-brown to grey with a lighter underside. The Barbary macaque has a body length of around 556.8mm in females and 634.3mm in males and mean body weight is reported to be 9.9±1.03 kg in females and around 14.5 kg in males. Its face is dark pink and its tail is vestigial, measuring anywhere from 4 to 22mm. Males often have a more prominent tail. The front limbs of this monkey are longer than its hind limbs. Females are smaller than males.
The Barbary macaque is mainly found in the Atlas and Rif Mountain ranges of Morocco and Algeria and Jebel Akhdar in Libya. It is the only species of macaque that is distributed outside of Asia. These animals can occupy a variety of habitats, such as cedar, fir, and oak forests, or grasslands, scrub, rocky ridges full of vegetation. Most Barbary macaques inhabit cedar forests currently, however, this could reflect the present habitat availability rather than a specific preference for this habitat.
The diet of a Barbary macaque consists of a mixture of plants and insect prey. M. sylvanus consume a large variety of gymnosperms and angiosperms. Almost every part of the plant is eaten, including flowers, fruits, seeds, seedlings, leaves, buds, bark, gum, stems, roots, bulbs, and corns. Common prey caught and consumed by Barbary macaques are snails, earthworms, scorpions, spiders, centipedes, millipedes, grasshoppers, termites, water striders, scale insects, beetles, butterflies, moths, ants, and even tadpoles.
The Barbary macaque is gregarious, forming mixed groups of several females and males. Troops can have 10 to 100 individuals and are matriarchal, with their hierarchy determined by lineage to the lead female. Unlike other macaques, the males participate in rearing the young. Males may spend a considerable amount of time playing with and grooming infants. In this way, a strong social bond is formed between males and juveniles, both the male's own offspring and those of others in the troop. This may be a result of selectivity on the part of the females, who may prefer highly parental males.
The mating season runs from November through March. The gestation period is 147 to 192 days, and females usually have only one offspring per pregnancy. Females rear twins in rare instances. Offspring reach maturity at three to four years of age, and may live for 20 years or more.
Grooming other Barbary macaques leads to lower stress levels for the individuals that do the grooming. While stress levels do not appear to be reduced in animals that are groomed, grooming more individuals leads to even lower stress levels; this is a benefit that might outweigh the costs to the groomer, which include less time to participate in other activities such as foraging. The mechanism for reducing stress may be explained by the social relationships (and support) that are formed by grooming.
Male Barbary macaques interfere in conflicts and form coalitions with other males, usually with related males rather than with unrelated males. These relationships suggest that males do so in order to indirectly increase their own fitness. Furthermore, males form coalitions with closely related kin more often than they do with distantly related kin. These coalitions are not permanent and may change frequently as male ranking within the group changes. Although males are more likely to form coalitions with males who have helped them in the past, this is not as important as relatedness in determining coalitions. Males avoid conflicting with higher ranking males and will more frequently form coalitions with the higher ranking male in a conflict. Close grouping of males occur when infant Barbary macaques are present. Interactions between males are commonly initiated when a male presents an infant macaque to an adult male who is not caring for an infant, or when an unattached male approaches males who are caring for infants. This behavior leads to a type of social buffering which reduces the number of antagonistic interactions among males in a group.
An open mouth display by the Barbary macaque is used most commonly by juvenile macaques as a sign of playfulness.
The main purpose of calls in Barbary macaques is to alert other group members to possible dangers such as predators. Barbary macaques can discriminate calls by individuals in their own group from those by individuals in other groups of conspecific macaques. Neither genetic variation nor habitat differences are likely causes of acoustic variation in the calls of different social groups. Instead, minor variations in acoustic structure among groups similar to the vocal accommodation seen in humans are the likely cause. However, acoustic characteristics such as pitch and loudness are varied based on the vocalizations of individuals they associate with, and social situations play a role in the acoustic structure of calls.
Barbary macaque females have the ability to recognize their own offspring’s calls through a variety of acoustic parameters. Mothers demonstrate different behaviors upon hearing the calls of other infant macaques as opposed to the calls of their own offspring. Because female macaques can distinguish among a variety of parameters, infant calls do not have to differ dramatically for mothers to be able to recognize their own infant’s call. More parameters for vocalizations lead to more reliable identification of calls in both infants and in adult macaques so it is not surprising that the same acoustic characteristics that are heard in infant calls are also heard in adult calls.
Although Barbary macaques are sexually active at all points during a female’s reproductive cycle, male Barbary macaques determine a female’s most fertile period by sexual swellings on the female. Mating is most common during a female’s most fertile period. The swelling size of the female reaches a maximum around the time of ovulation, suggesting that size helps a male predict when he should mate. This is further supported by the fact that male ejaculation peaks at the same time that female sexual swelling peaks. There is not a sufficient change in female sexual behavior around the time of ovulation in order to demonstrate to the male that the female is fertile. The swellings, therefore, appear necessary for predicting fertility.
Barbary macaque females differ from other non-human primates in that they often mate with a majority of the males in their social group. While females are active in choosing sexual associations, the mating behavior of macaque social groups is not entirely determined by female choice. These multiple matings by females decreases the certainty of paternity of male Barbary macaques and may lead them to care for all infants within the group. In order for a male to ensure his reproductive success, he must maximize his time spent around the females in the group during their fertile periods. Injuries to male macaques peaks during the fertile period which points to male-male competition as an important determinant of male reproductive success. Not allowing a female to mate with other males, however, would be costly to the male since doing so would not allow him to mate with more females.
Unlike other macaques where most parental care comes from the mother, Barbary macaques from all age and sex groups participate in alloparental care of infants. Male care of infants has been of particular interest to research because high levels of care from males is uncommon in groups where paternity is highly uncertain. Males even act as true alloparents of infant macaques by carrying them and caring for them for hours at a time as opposed to just demonstrating more casual interactions with the infants. Female social status plays a role in female alloparental interactions with infants. Higher ranking females have more interactions whereas younger, lower ranking females have less access to infants.
Interaction with the environment
Barbary macaques can cause major damage to the trees in their prime habitat, the Atlas Cedar forests in Morocco. Since deforestation in Morocco has become a major environmental problem in recent years, research has been conducted to determine the cause of the bark stripping behavior demonstrated by these macaques. Cedar trees are also vital to this population of Barbary macaques as an area with cedars can support a much higher density of macaques than one without them. A lack of a water source and exclusion of monkeys from water sources are major causes of cedar bark stripping behavior in Barbary macaques. Density of macaques, however, is less correlated with the behavior than the other causes considered.
The Barbary macaque was named by Carl Linnaeus in 1758, along with numerous other species named in that same year. The scientific name is Macaca sylvanus. Phylogenetic and molecular analysis with other primates has been done. Studies concerning interspecific DNA variation within the genus Macaca show that the Barbary macaque is most related to the Asian macaques. Scientists have also studied the Y-chromosome, however, this has been unfruitful. One study indicates that the Barbary macaque has origins in Morocco and Algeria. The results of a phylogenetic analysis show that M. sylvanus chromosomes resemble M. mnilatta with the exception of chromosomes 1, 4, 9, and 16. Relative to humans, M. sylvanus has a homolog of chromosome 13.
Relationship with humans
Wild populations of Barbary macaques have suffered a major decline in recent years to the point of being declared in 2009 as an endangered species by the IUCN. Three-quarters of the world population are located in the Middle Atlas Mountains.
This species is also poached for live specimens as pets in the illegal pet trade, and for clandestine collectors. Spain is the main entry point in Europe. Today, no accurate data exist on the location and number of individuals out of their habitat. An unknown number of individuals are included in zoological collections, at other institutions, in private hands, in storage, or waiting to be relocated to appropriate destinations.
The habitat of the Barbary macaque is under threat from increased logging activity. As such, they are listed as endangered by the IUCN Red List. Local farmers view the monkeys as pests, and engage in extermination of the species. Once common throughout northern Africa and southern Europe, only an estimated 12,000 to 21,000 Barbary macaques are left in Morocco and Algeria. Once, their distribution was much more extensive, reaching Tunisia and Libya. Their range is no longer continuous, with only isolated areas of range remaining. During the Pleistocene, this species inhabited the Mediterranean coasts and Europe, reaching Italy, Hungary, Spain, Portugal and France, and as far north as Germany and the British Isles. The species decreased with the arrival of the Ice Age, becoming extinct in the Iberian Peninsula 30,000 years ago.
The skull of a Barbary macaque was discovered during excavation in the 1970s at the pre-Christian Navan Fort in County Armagh, Northern Ireland. Carbon dating tests suggest it died there in the third century BC.
Tourists interact with wild monkeys in many places, tourists may be encouraged to feed, photograph and touch them. Tourism may aid conservation but may also stress the animals. Aggressive behaviour by tourists stressed the animals physiologically. Stress related hormones were found in their droppings. Other tourist actions made the monkeys uneasy without apparent physiological stress. Stress behaviour in Barbary macaques increased when humans got too close to them, touched them or even photographed them. When tourists got too close macaques were more aggressive and less friendly to each other. High tourist numbers are not a problem for the macaques provided tourists stay at a distance. Macaques that interacted with tourists had more parasites, possibly due to crowding at feeding sites. Macaques in contact with tourists further were fatter with less healthy fur than their counterparts that did not receive unhealthy food from humans. Guidelines are needed to avoid tourists stressing the animals unnecessarily. 
Many of the mistaken ideas about human anatomy contained in the writings of Galen are apparently due to his use of these animals, the only anthropoid available to him, in dissections. Strong cultural taboos of his time prevented his performing any actual dissections of human cadavers, even in his role as physician and teacher of physicians.
The last wild population in Europe is that of Gibraltar, which unlike that of North Africa, is thriving. Currently, there are around 230 individuals living on the Rock of Gibraltar, and they form groups of up to 75 and will occasionally enter towns.
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