Crested black macaques are social monkeys and, before their decline in the wild, were often seen in groups of up to 100 individuals (5). At present they are found in smaller groups. During the day they split into smaller units of 10 to 25 individuals, led by dominant males who police the group and prevent serious fights developing. They feed on figs, other fruit, vegetation, insects and small animals such as mice, crabs and lizards (6). Sometimes food is not eaten immediately but is stored in cheek pouches for a while. Individuals in the group maintain relationships by grooming each other and communicating vocally with grunts (4). Adult males 'yawn' to display their large canine teeth in order to assert dominance and avoid conflict (5). Breeding is non-seasonal and therefore occurs at any time of year. Females come into oestrous every 33 to 36 days and advertise their fertility with swollen pink bottoms. The females are monopolised by the group's dominant male, and after a gestation period of five and a half months a single infant is born (5). The offspring reach sexual maturity at four to six years and may live for up to 25 years (5).

Macaques are medium-sized to large-sized monkeys with stocky bodies. Some, like this species, possess only a very short rudimentary tail that has led in the past to the misidentification and naming of some macaques as apes, which completely lack tails (4). The crested black macaque is the most endangered of the seven macaque species found on the island of Sulawesi (formerly Celebes) in Indonesia (2) (5). This species is entirely black apart from its rump, which is a distinctive pink colour. In males the posterior is small and heart-shaped, and in females it is large, rounded and a darker pink (5). Its face is elongated with close-set brown eyes, and prominent ridges down the side of the nose (6). The body hair is quite sleek, though on the head it forms a distinctive punk-like crest, referred to by the common name, crested black macaque (4).

This species occurs on northeastern Sulawesi, Indonesia and the adjacent islands of Pulau Manadotua and Pulau Talise. It was historically found on Pulau Lembeh as well but has probably been extirpated from there. On Sulawesi it is found on the northern arm east of the Onggak Dumoga River and Mount Padang to the tip of the peninsula (Groves 2001). There is a sizeable introduced population on Pulau Bacan in the Maluku Islands (Indonesia).

Macaca nigra is one of seven species of macaques endemic to the island of Sulawesi in Indonesia (Fooden 1969). Celebes crested macaques are restricted to northeast Sulawesi and the adjacent islands of Pulau Manadotua and Pulau Talise. They are no longer found on Pulai Lembeh. An introduced population of at least 100,000 individuals resides on Pulau Bacan, in the Maluku Islands (Supriatna and Andayani 2008).

Biogeographic Regions: oriental (Introduced , Native )

Other Geographic Terms: island endemic

This species is endemic to Indonesia, where it occurs in north-eastern Sulawesi and on the adjacent islands of Pulau Lembeh, Pulau Manadotua and Pulau Talise; introduced to Pulau Bacan by humans (2).

Celebes crested macaques are large, diurnal primates with black faces and bodies. Their crania are more prognathic and baboon-like than those of other macaques (Sugardjito et al. 1989), and their crown hairs form a crest, pointing backward and up. Celebes crested macaques also have prominent kidney-shaped ischial callosities, which are bright pink, and a “nubbin-like” tail (Groves 2001). Because their tails are short (about 20 mm long), they are commonly misidentified as apes (Cawthorn Lang 2006).

The face of Celebes crested macaques is black and thinly covered with hairs, and their body hair is dense, coarse, and woolly (Fooden 1969). Juvenile Celebes crested macaques are paler than adults, appearing more brown than black. Juveniles have black hairs where the crest will develop (Fooden 1969). As adult males age, their hair grays and becomes grizzled, most noticeably in the upper body; this occasionally occurs on the arms of younger males as well (Hamada et al. 1988).

Celebes crested macaques are sexually dimorphic, with males typically measuring from 520 to 570 mm and weighing 9.9 kg and females measuring from 445 to 570 mm and weighing 5.5 kg (Thierry et al. 2004). Adult females are also paler in color than adult males (Fooden 1969).

Among the species of macaques present on the island of Sulawesi, Celebes crested macaques are most difficult to distinguish from Gorontalo macaques, Macaca nigrescens. Both have black skin, but the upper half of the body of Celebes crested macaques is darker and more dull (Hamada et al. 1988).

Range mass: 5 to 18 kg.

Average mass: Males 9.9 kg; Females 5.5 kg.

Range length: 445 to 570 mm.

Average length: Males 520 to 570 mm; Females 445 to 570 mm.

Sexual Dimorphism: male larger

Type for Macaca nigra
Catalog Number: USNM 217573
Collection: Smithsonian Institution, National Museum of Natural History, Department of Vertebrate Zoology, Division of Mammals
Sex/Stage: Male; Adult
Preparation: Skull
Collector(s): H. Raven
Year Collected: 1916
Locality: Pulo [= Pulau] Lembeh, Celebes, Sulawesi Utara, Indonesia, Asia

Habitat and Ecology

Systems

• Terrestrial

Celebes crested macaques and tarsiers are the only non-human primates to inhabit the forests of Sulawesi (Riley 2010). Celebes crested macaques live in the island’s forests at moderate elevations ranging from 700 to 1100 m above sea level, most likely determined by fruit abundance (Rosenbaum et al. 1998b). These areas experience relatively constant temperatures, although there are fluctuations in rainfall between dry and wet seasons (Supriatna and Andayani 2008). They prefer humid, tropical lowland and upland rainforests, but they are also known to frequent agricultural areas that have increasingly encroached on their habitat (Rosenbaum et al. 1998b; O’Brien and Kinnaird 1997).

Celebes crested macaques live in many protected areas including Gunung Lokon, Gunung Amban, Tangkoko Batuangus, Dua Saudara, and Batu Putih, and are also commonly found in captivity (Supriatna and Andayani 2008).

Range elevation: 700 to 1100 m.

Habitat Regions: tropical

Terrestrial Biomes: rainforest ; mountains

Other Habitat Features: agricultural

The crested black macaque inhabits primary and secondary tropical rainforest (2).

Celebes crested macaques are mostly frugivorous, and figs comprise the majority of their diet. However, they have a diverse diet, consuming more than 145 species of plants, mostly fruit (Dracontomelon spp. and Ficus spp.), as well as some invertebrates (O’Brien and Kinnaird 1997; Rosenbaum et al. 1998b; Riley 2007). When fruit is scarce, their diet is supplemented by insects, shoots, young leaves, and the stems of flowering plants (Riley 2007; Supriatna and Andayani 2008). Celebes crested macaques have been observed preying on fruit bats, forest geckos, frogs, and the eggs of the red junglefowl and black-naped monarch flycatcher (O’Brien and Kinnaird 1997). Members of this species also raid crops, including maize (Zea mays), yams (Dioscorea), cassava (Manihot esculenta), papayas (Carica papaya), bananas (Musa), and cacao (Theobroma cacao) (Bynum et al. 1999; Rosenbaum et al. 1998a; Riley, 2007).

Celebes crested macaques forage throughout the day, occasionally storing food reserves in their cheek pouches (Schulz 2009).

Animal Foods: mammals; amphibians; reptiles; eggs; insects; terrestrial non-insect arthropods; terrestrial worms

Plant Foods: leaves; wood, bark, or stems; seeds, grains, and nuts; fruit; flowers

Foraging Behavior: stores or caches food

Primary Diet: herbivore (Frugivore ); omnivore

Celebes crested macaques have no major predators or catarrhine competitors (Riley 2007). Many macaques on the island of Sulawesi are known to associate with birds, including hair-crested drongos and yellow-billed malkoha (Matsumura 2001; Whitten et al. 2002). These birds prey on the insects that leave the canopy when macaques pass through (Matsumura 2001).

As frugivores, Celebes crested macaques act as seed dispersers of rainforst trees, especially for species of Dracomelon and Ficus (Rosenbaum et al. 1998b).

Fecal tests of Celebes crested macaques revealed the presence of parasites including Balantidium sp., Entamoeba sp., Ascaris sp., and Trichostrongylus sp., all of which are found in both humans and nonhuman primates (Paulsen et al. 2006). Among captive Celebes crested macaques, 72.2% (13 out of a sample of 18 individuals) were parasitized, with a mean of 1.1 parasite taxon per individual (Jones-Engel et el. 2004). A study of Sulawesi macaques kept as pets, which included Celebes crested macaques, found 7 taxa of intestinal protozoa (Blastocystis hominis, Iodamoeba butschlii, Entamoeba coli, Entamoeba hartmanni, Chilomastrix mesnili, Endolimax nana, and Retortamonas intestinalis) and 3 taxa of nematodes (hookworm, Trichuris sp., and Ascaris sp.) (Jones-Engel et al. 2004).

Ecosystem Impact: disperses seeds

Mutualist Species:

• Dicrurus hottentotts
• Zanclostomus calyorhynchus

Commensal/Parasitic Species:

• Balantidium sp.
• Blastocystis hominis
• Iodamoeba butschlii
• Entamoeba coli
• Entamoeba hartmanni
• Chilomastrix mesnili
• Endolimax nana
• Retortamonas intestinalis
• hookworm Necator americanus
• Trichuris sp.
• Ascaris sp.

Celebes crested macaques have no major predators. They are, however, hunted by humans for their meat (Riley 2007; Durrell Wildlife Conservation Trust 2011).

Known Predators:

• Humans Homo sapiens

Celebes crested macaques utilize a variety of visual and physical behaviors to communicate. To indicate aggression, males stare with an open mouth, grin with the mouth closed, yawn, lunge, and chase (Nickelson and Lockard 1978; Reed et al. 1997). Submission and satisfaction are signaled with a grimace and lip smacking (Nickelson and Lockard 1978). As with other macaques, males are known to silently bare their teeth, postulated as a display of “peaceful intentions” (Thierry et al. 2000). Dominant males also bare their canines in a “yawn” to indicate supremacy (Assahad et al. 2008).

Celebes crested macaques also use vocalizations. They are known for a loud, bird-like call that may be a method of intervening in conflicts, signaling to keep away from other groups (Muroyama and Thierry 1998; Kinnaird and O’Brien 1999), or as a “signal of dominance to prevent contests between males for mates” (Neumann et al. 2010). Frequency and complexity of calls correlates with male rank, with high-ranking males emitting complex calls more frequently than subordinates (Assahad et al. 2008).

Communication Channels: visual ; tactile ; acoustic

Perception Channels: visual ; acoustic

The average life span of Celebes crested macaques is 18 years Cawthorn Lang 2006). The oldest individual in captivity survived 34 years (Schulz 2009).

Range lifespan

Status: captivity: 34 (high) years.

Average lifespan

Status: wild: 18 years.

Average lifespan

Status: captivity: 34 years.

Maximum longevity: 34 years (captivity)

Celebes crested macaques are promiscuous, or polygynandrous, as each individual (male and female) has multiple mates. Females exhibit sexual swelling in the anogenital area to indicate ovarian activity and sexual receptivity, sending visual and olfactory cues, which males sense by sniffing female genitalia (Engelhardt and Farajallah 2008). However, the degree to which males use visual, olfactory, and behavioral signs to determine female fertility remains unclear (Thierry et al. 2004).

Adult females approach high-ranking males for sex more often than they do low-ranking males, although all males copulate about the same amount (Reed et al. 1997). A female presents herself to a male by facing her swollen genitals, slightly raised, toward the male, and smacking her lips. The male holds the female’s legs and waist during copulation. The pair engages in reciprocal grooming following intercourse (Nickelson and Lockard 1978). There are two types of ejaculatory patterns among species of macaques. Male Celebes crested macaques mount the female multiple times but only ejaculate during the last mount (multiple-mount ejaculation) (Caldecott 1986).

Sexual swelling of the female anogenital area also occurs when females are not ovulating, which promotes non-procreative copulation (Thomson et al. 1992). Homosexual behavior such as displaying, mounting, and genetal contact has also been observed among males of this species (Soltis 2004).

Mating System: polygynandrous (promiscuous)

Little is known about the reproductive biology of Celebes crested macaques in the wild (Engelhardt and Farajallah 2008). They breed year-round, although periods of high ovarian activity among adult females occur from August to June, and birthing peaks between January and May (Engelhardt and Farajallah 2008).

Fertility of males and females overlaps (Bernstein et al. 1982). Females have a menstrual cycle of approximately 32 days, which is unusually long for macaques and occurs because of an extended follicular phase. Cycle length and maximal genital swelling period are extended when males are absent (Bernstein et al. 1982; Thomson et al. 1992).

Females give birth to a single offspring every 18 months. Gestation last 5.5 months. Offspring spend much less time with their mother at 4 months of age and are completely weaned by 1 year of age (Durrell Wildlife Conservation Trust 2011).

In most populations, the adult sex ratio is 3 females to 5 males (Thierry et al., 2004). Macaques are sexual bimaturists, with females reaching physical maturity faster than males, though the hormonal processes responsible for maturation are identical in both sexes (Thierry, 2007). Males produce sperm around 3 or 4 years of age, though they do not reach a body mass necessary for mating (physical maturity) for another 3 to 6 years (Thierry et al. 2004; Howard et al. 1999).

Breeding interval: Celebes crested macaques typically give birth once every 18 months.

Breeding season: Celebes crested macaques breed year-round, although breeding increases between August and June.

Average number of offspring: 1.

Range gestation period: 5 to 6 months.

Average weaning age: 12 months.

Range age at sexual or reproductive maturity (female): 4 to 6 years.

Range age at sexual or reproductive maturity (male): 4 to 6 years.

Key Reproductive Features: year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); viviparous ; post-partum estrous

Average birth mass: 461 g.

Average number of offspring: 1.

Little is known regarding parental care among Celebes crested macaques. Based on comparable patterns of behavior and social structure, parental investment in this species is likely similiar to that of other macaques: care is primarily maternal, though aunting behavior occurs, in which an infant is carried by juvenile females supervised by the mother (Cawthorn Lang 2006). Infants of this speices often cling to their mother's belly for protection (Durrell Wildlife Conservation Trust 2011).

Although sexual swelling in the anogenital area marks periods of ovulation, sexual swelling also occurs when females are not ovulating, promoting non-procreative copulation (Thomson et al. 1992). This may confuse paternity, increasing male parental care and decreasing the likelihood of infanticide (Soltis 2004).

Parental Investment: female parental care ; pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female)

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 0
Specimens with Barcodes: 2
Species With Barcodes: 1

Year Assessed

Assessor/s

Reviewer/s

Contributor/s

In 2008, the IUCN listed Celebes crested macaques as critically endangered (Supriatna and Andayani 2008). This species is also listed in CITES Appendix II. Population size has decreased by 80% in the last 40 years, primarily due to hunting and loss of habitat (Kyes et al. 2000; Laatung et al. 2006). There are estimated to be 4,000 to 6,000 individuals in the wild. On Bacan Island, there is an introduced population of 100,000 individuals, which is not included in the population estimate (Hamada et al. 1994), but could become the last surviving population of Celebes crested macaques in the near future (Riley 2010).

Main threats to the conservation of Celebes crested macaques include human population growth and land conversion (Bynum et al. 1999). Following recent spikes in human habitation, which in 2002 was 132 individuals per sq km, is large-scale deforestation by logging, the transmigration program, and cash crop farming, most of which is partially funded by the government (Whitten et al. 2002).

Humans cut down trees that are important to the diet and habitat of macaques, such as fig trees (Riley and Priston 2010). Human-macaque sharing of resources is occasionally possible, as with Arenga pinnata palm trees, from which macaques gather fruit and humans take palm fronds. However, deforestation for timber and firewood and clearing for agriculture permanently removes resources utilized by macaques (Riley, 2010).

Despite their critical conservation condition, the total area of protected land inhabited by Celebes crested macaques (16,848 ha) is less than any other macaques on the island of Sulawesi (Riley 2010).

CITES: appendix ii

IUCN Red List of Threatened Species: critically endangered

Classified as Critically Endangered (CR) on the IUCN Red List (1) and listed on Appendix II of CITES (3).

Population

Population Trend

Major Threats

This primate is threatened by over-hunting for food in Sulawesi where its meat is considered a delicacy. Its habitat is also threatened by human settlement, land clearing for agriculture and logging. While this is a problem in many areas worldwide, Sulawesi is particularly sensitive as it is an island and therefore has a limited amount of land for its wildlife and expanding human population (4).

Conservation Actions

International trade in this species is prohibited by its listing on Appendix II of the Convention on International Trade in Endangered Species (CITES) (3). Hunting on the island, however, is harder to regulate. This macaque is found in three areas on the island where hunting, logging and clearing is illegal. However, it is not clear how effective these regulations are (4). Captive breeding of this species has been successful at Jersey Zoo and a number of other zoos, and individuals may be reintroduced into the wild in the future. If this species is to survive, it is essential to address the problems of hunting and habitat loss on Sulawesi (5).

As humans reduce habitat, cut down fig trees, which are a key component to the diet of macaques, and as farms expand into forests, macaques have adapted to include crops in their diet, exploiting human resources (Richard et al. 1989; Riley and Priston 2010). Although local folklore describes humans and macaques as intertwined, crop raiding has earned them reputations as “pests” and “weed species” (Richard et al. 1989). Crop raiding is common among all species of macaques, including Celebes crested macaques (Riley and Priston 2010).

Certain diseases can also be transmitted among humans and macaques (Jones-Engel et al., 2005; Riley, 2010). There is evidence of macaque exposure to human respiratory viruses, including influenza A and parainfluenza 1, 2, and 3. This anthropozoonotic exchange may occur through individuals kept as pets, which come into contact with both wild macaques and humans (Jones-Engel et al. 2005).

Negative Impacts: injures humans (causes disease in humans , carries human disease); crop pest; causes or carries domestic animal disease

Celebes crested macaques play an important role in local folklore, and can serve as household pets (Hamada et al. 1988; Riley and Priston 2010). Considered a delicacy, they are hunted for bush meat and are served on holidays and special occasions (Schulz 2009; Durrell Wildlife Conservation Trust 2011). CITES prohibits live trade of this speices, though illegal trafficking may still occur (Supriatna and Andayani 2008).

Because Celebes crested macaques are the only non-human primate model for Type II diabetes mellitus, they are valuable in scientific research (Thomson et al. 1992).

This species also plays a considerable role in ecotourism (Schulz 2009).

Positive Impacts: pet trade ; food ; ecotourism ; research and education