Macaca hecki
Creators: name: Russell A. Mittermeier
name: Anthony B. Rylands
name: Don E. Wilson
Publisher: name: Lynx Edicions
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Primates
Cercopithecidae
Macaca
Macaca hecki

description: 5.
Heck’s Macaque
Macaca hecki
French: Macaque de Heck / German: Heck-Makak / Spanish: Macaco de Heck
Taxonomy. Papio (Inuus) hecki Matschie, 1901,
Buol, Tengah, Sulawesi, Indonesia.
M. heckiis a member of the silenus species-group of macaques, including M. siberu, M. pagensis, M. leonina, M. nemestrina, M. silenus, and the other Sulawesi species. It hybridizes with M. tonkeana in an area centered on the road from Tawaeli to Toboli that crosses the base of the northern peninsula, north-east of Palu, and possibly with M. nigrescens just east Gorontalo and in the basins of the Bolango and Bone rivers. In his 1980 review, C. P. Groves considered M. hecki to be a subspecies of M. tonkeana based on inferred intergradation. This classification was followed G. Corbet and J. Hill in their review The Mammals of the Indomalayan Region, published in 1992. The hecki-tonkeana hybrid zone was subsequently studied by E. Bynum and colleagues and found to be restricted (c.15 km by 7-5 km) and old, dating back to at least to 1900. This indicates that it is a zone of secondary contact and that there are barriers (prezygotic or postzygotic) to full introgression. In 2001, Groves listed heck: as a distinct species. Monotypic.
Distribution. NW Sulawesi from the base of the N peninsula (Isthmus of Palu) NE to just E of Gorontalo.
Descriptive notes. Head—body 58-68 cm (males) and 50-57 cm (females), tail 2-3 cm (males) and 2-3 cm (females); weight 8-10 kg (males) and 7-8 kg (females). Heck’s Macaque is black, with a brownish underside. Forearms are dark brown, and shanks are gray to light brown (i.e. paler than the trunk). Crown hair is lengthened but does not form a distinct crest, and cheeks are lighter colored and without elongated whiskers. Tail is very short and stubby, although not rudimentary. Ischial callosities are gray to yellow and kidney-shaped, with only a partial transverse furrow across them. Comparing the Gorontalo Macaque (M. nigrescens) with Heck’s Macaque, the Gorontalo Macaque has a baboon-like face, an elongated narrow crest, blackish limbs contrasting with a brown body, and gray ischial callosities. Heck’s Macaque has a short broad face and a short broad, somewhat upstanding crest, on the crown. It is black with strongly contrasting brown shanks (extending up the back of the thighs), weakly contrasting dark brown arms and underparts, and yellow, reniform, and only partially separated callosities. Comparing the Tonkean Macaque (M. tonkeana) with Heck’s Macaque, forearms and shanks of hindlimbs of the Tonkean Macaque are black like the body, underside is no paler than the upper side (it is in Heck’s Macaque), cheek whiskers have a pale grayish or brownish tint (not in Heck’s Macaque), rump patch is more clearly marked and bushy (not so in Heck’s Macaque), and ischial callosities are oval and orange-colored (similar to the Moor Macaque, M. maura, and the Booted Macaque, M. ochreata).
Habitat. Undisturbed or only lightly disturbed primary rainforest at elevations from close to sea level to 1600-1800 m. Seasons are not pronounced in these areas, with only one or two drier months each year and annual rainfall of 2500-3500 mm (higher in the mountainous regions). The extent to which Heck’s Macaque can thrive in degraded and secondary forest is not known.
Food and Feeding. The three macaque species of the northern peninsula, including Heck’s Macaque,are all principally frugivores/insectivores and are less folivorous than species to the south. When near human settlements, Sulawesi macaques raid subsistence and cash crops such as maize, yams, cassava, papaya, banana, and cacao, and are persecuted for doing so.
Breeding. Reproductive cycle of female Heck’s Macaquesis ¢.36 days. As in all Sulawesi macaques, females exhibit a pronounced, bright pink to red sexual swelling during the periovulatory period. A single offspring is born after a gestation of 174-196 days. Young are nursed for c¢.12 months and are sexually mature at c.4 years of age. Sexual patterns are broadly similar among Sulawesi macaques. The only behavioral patterns that appear to be specific to each taxon are their loud calls or chirp barks.
Activity patterns. Heck’s Macaques are diurnal and largely arboreal. The extent to which they become terrestrial, a trend for all macaques that are losing their forests,is unknown.
Movements, Home range and Social organization. Heck’s Macaque is one of the least studied of the Sulawesi macaques. It lives in multimale—multifemale groups averaging 20 individuals (range 7-30) and probably in large home ranges of more than a 100 ha. As in all Sulawesi macaques (and typical for the genus), males disperse and females are philopatric. Densities are 20-50 ind/km?.
Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Heck’s Macaque is fully protected by Indonesian Law. It is threatened mainly by loss of habitat, which is increasingly fragmented. The destruction of Sulawesi forests accelerated in the last 40 years, as it has for tropical forests worldwide; commercial logging, Indonesia’s transmigration program, the government-subsidized cash crop industry, and human population growth are all to blame. A report by C. Cannon and colleagues in 2007 reported that 80% of Sulawesi forests had been destroyed. Heck’s Macaque is hunted in some areas and persecuted as a crop pest. The fearis that intensive hunting for commercial trade within the ranges of the Crested Macaque (M. nigra) and the Gorontalo Macaque, which has resulted in their severely reduced numbers and endangered status, will move south in the northern peninsula into the range of Heck’s Macaque. In 1994, it was estimated that the total population was c.100,000, but it has certainly declined since then. Forest remains in inaccessible areas, but lowlands in the southern part of its distribution and coastal areas of the northern peninsula, in particular, have been largely converted to agriculture. The suitability of forests for Heck’s Macaques at higher elevations is unknown. In 1999, the lack of strictly protected areas in the range of Heck’s Macaque was seen as the biggest threat to its survival. Protection then was only afforded by a single nature reserve, Panua, of insufficient size (1500 ha), badly degraded and harboring low numbers of macaques. The Paguyaman River catchment was identified as of conservation value for Heck’s Macaque. Since 1999, Nantu Wildlife Reserve and a number of other reserves have been created that protect, or should protect, Heck’s Macaque: Dolangan, Mas Popaya Raja, Pangi Binanga, Pinjam/Tanjung Mantop, and Tangale reserves.
Bibliography. Abegg & Thierry (2002a), Bynum (1995, 1999), Bynum, Bynum, Froehlich & Supriatna (1997), Bynum, Bynum & Supriatna (1997), Bynum, Kohlhaas & Pramono (1999), Cannon et al. (2007), Clayton & Millner-Gulland (2000), Corbet & Hill (1992), Fooden (1969), Groves (1980, 2001), Juliandi et al. (2009), Lee, R.J. (1999, 2000), Lee, R.J. et al. (2005), Riley (2010a), Watanabe & Matsumura (1991), Thierry (2000b, 2007, 2011), Watanabe, Lapasere & Tantu (1991).
description: Published as part of Russell A. Mittermeier, Anthony B. Rylands & Don E. Wilson, 2013, Cercopithecidae, pp. 550-755 in Handbook of the Mammals of the World – Volume 3 Primates, Barcelona :Lynx Edicions on page 632, DOI: 10.5281/zenodo.6867065