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<document id="B5828260F180EB0C2305C5E37665E873" ID-CLB-Dataset="100956" ID-DOI="10.5281/zenodo.6867065" ID-GBIF-Dataset="3d520847-5163-4b5c-87bf-2cdceb781098" ID-ISBN="978-84-96553-89-7" ID-Zenodo-Dep="6867065" IM.illustrations_requiresApprovalFor="plazi" IM.metadata_requiresApprovalFor="plazi" IM.taxonomicNames_requiresApprovalFor="plazi" checkinTime="1658256599114" checkinUser="jonas" docAuthor="Russell A. Mittermeier, Anthony B. Rylands &amp; Don E. Wilson" docDate="2013" docId="CE199B17FFC4FFC1FFFE6227F8C1F671" docLanguage="en" docName="hbmw_3_Cercopithecidae_0550.pdf.imf" docOrigin="Handbook of the Mammals of the World Volume 3 Primates, Barcelona: Lynx Edicions" docTitle="Macaca hecki" docType="treatment" docVersion="7" lastPageNumber="632" masterDocId="3220E36FFFC1FFC4FF596510FFB7FFFC" masterDocTitle="Cercopithecidae" masterLastPageNumber="755" masterPageNumber="550" pageNumber="632" updateTime="1699359875441" updateUser="ExternalLinkService">
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<figureCitation id="DE8B3684FFC4FFC1FFFE6227FF70F899" box="[167,199,1847,1893]" captionStart="Plate 36: Cercopithecidae" captionStartId="2.[108,138,3367,3392]" captionTargetBox="[14,2754,17,3635]" captionTargetPageId="1" captionText="1. Barbary Macaque (Macaca sylvanus), 2. Lion-tailed Macaque (Macaca silenus), 3. Crested Macaque (Macaca migra), 4. Gorontalo Macaque (Macaca nigrescens), 5. Hecks Macaque (Macaca hecki), 6. Tonkean Macaque (Macaca tonkeana), 7. Moor Macaque (Macaca maura), 8. Booted Macaque (Macaca ochreata), 9. Pagai Macaque (Macaca pagensis), 10. Siberut Macaque (Macaca siberu), 11. Sunda Pig-tailed Macaque (Macaca nemestrina), 12. Northern Pig-tailed Macaque (Macaca leonina), 13. Toque Macaque (Macaca sinica), 14. Bonnet Macaque (Macaca radiata)" figureDoi="http://doi.org/10.5281/zenodo.6867431" httpUri="https://zenodo.org/record/6867431/files/figure.png" pageId="5" pageNumber="632">5.</figureCitation>
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<heading id="1D479D6DFFC4FFC1FF816227FD9CF899" box="[216,555,1847,1893]" pageId="5" pageNumber="632">
<vernacularName id="C8B35A2FFFC4FFC1FF816227FD9CF899" box="[216,555,1847,1893]" pageId="5" pageNumber="632">Hecks Macaque</vernacularName>
</heading>
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<paragraph id="460F2A01FFC4FFC1FD396227FCE8F899" blockId="5.[165,966,1847,1934]" box="[608,863,1847,1893]" pageId="5" pageNumber="632">
<heading id="1D479D6DFFC4FFC1FD396227FCE8F899" box="[608,863,1847,1893]" pageId="5" pageNumber="632">
<taxonomicName id="81B05182FFC4FFC1FD396227FCE8F899" ID-CoL="3WWNG" baseAuthorityName="Matschie" baseAuthorityYear="1901" box="[608,863,1847,1893]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="hecki">
<emphasis id="74C4F613FFC4FFC1FD396227FCE8F899" box="[608,863,1847,1893]" italics="true" pageId="5" pageNumber="632">Macaca hecki</emphasis>
</taxonomicName>
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</paragraph>
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<subSubSection id="0EAA798AFFC4FFC1FFFF6267FC72F870" box="[166,965,1911,1932]" pageId="5" pageNumber="632" type="vernacular_names">
<paragraph id="460F2A01FFC4FFC1FFFF6267FC72F870" blockId="5.[165,966,1847,1934]" box="[166,965,1911,1932]" pageId="5" pageNumber="632">
<heading id="1D479D6DFFC4FFC1FFFF6267FC72F870" box="[166,965,1911,1932]" pageId="5" pageNumber="632">
<emphasis id="74C4F613FFC4FFC1FFFF6267FF44F870" bold="true" box="[166,243,1911,1932]" pageId="5" pageNumber="632">French:</emphasis>
<vernacularName id="C8B35A2FFFC4FFC1FFA46267FE06F870" box="[253,433,1911,1932]" pageId="5" pageNumber="632">Macaque de Heck</vernacularName>
/
<emphasis id="74C4F613FFC4FFC1FE9F6267FD97F870" bold="true" box="[454,544,1911,1932]" pageId="5" pageNumber="632">German:</emphasis>
<vernacularName id="C8B35A2FFFC4FFC1FD726267FD12F870" box="[555,677,1911,1932]" pageId="5" pageNumber="632">Heck-Makak</vernacularName>
/
<emphasis id="74C4F613FFC4FFC1FDE36267FCA3F870" bold="true" box="[698,788,1911,1932]" pageId="5" pageNumber="632">Spanish:</emphasis>
<vernacularName id="C8B35A2FFFC4FFC1FC466267FC72F870" box="[799,965,1911,1932]" pageId="5" pageNumber="632">Macaco de Heck</vernacularName>
</heading>
</paragraph>
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<paragraph id="460F2A01FFC4FFC1FC5462AEFCE0F7FF" blockId="5.[780,1367,1982,2405]" pageId="5" pageNumber="632">
Taxonomy.
<taxonomicName id="81B05182FFC4FFC1FCE062AEFCE5F7FF" authority="Matschie, 1901" authorityName="Matschie" authorityYear="1901" class="Mammalia" family="Cercopithecidae" genus="Papio" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="hecki" subGenus="Inuus">Papio (Inuus) hecki Matschie, 1901</taxonomicName>
,
</paragraph>
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<paragraph id="460F2A01FFC4FFC1FC3A62FAFAF0F7FF" blockId="5.[780,1367,1982,2405]" box="[867,1351,2026,2051]" pageId="5" pageNumber="632">
<materialsCitation id="F6D8205CFFC4FFC1FC3A62FAFAF4F7FF" ID-GBIF-Occurrence="3864421322" box="[867,1347,2026,2051]" pageId="5" pageNumber="632">Buol, Tengah, Sulawesi, Indonesia</materialsCitation>
.
</paragraph>
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<subSubSection id="0EAA798AFFC4FFC1FC566D1DFC02F563" pageId="5" pageNumber="632" type="discussion">
<paragraph id="460F2A01FFC4FFC1FC566D1DFC02F563" blockId="5.[780,1367,1982,2405]" lastBlockId="5.[159,1368,2419,3465]" pageId="5" pageNumber="632">
M. heckiis a member of the
<taxonomicName id="81B05182FFC4FFC1FBC96D1DFB51F7D6" authorityName="Linnaeus" authorityYear="1758" box="[1168,1254,2061,2090]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="silenus">silenus</taxonomicName>
species-group of macaques, including
<taxonomicName id="81B05182FFC4FFC1FB8A6D29FAE7F7AE" box="[1235,1360,2105,2130]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="siberu">M. siberu</taxonomicName>
,
<taxonomicName id="81B05182FFC4FFC1FC566D70FC03F785" box="[783,948,2144,2169]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="pagensis">M. pagensis</taxonomicName>
,
<taxonomicName id="81B05182FFC4FFC1FC8B6D70FBDBF785" baseAuthorityName="Blyth" baseAuthorityYear="1863" box="[978,1132,2144,2169]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="636" phylum="Chordata" rank="species" species="leonina">M. leonina</taxonomicName>
,
<taxonomicName id="81B05182FFC4FFC1FBD26D70FAE6F785" baseAuthorityName="Linnaeus" baseAuthorityYear="1766" box="[1163,1361,2144,2169]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="nemestrina">M. nemestrina</taxonomicName>
,
<taxonomicName id="81B05182FFC4FFC1FC566D6FFC24F75C" authorityName="Linnaeus" authorityYear="1758" box="[783,915,2175,2208]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="638" phylum="Chordata" rank="species" species="silenus">M. silenus</taxonomicName>
, and the other Sulawesi species. It hybridizes with
<taxonomicName id="81B05182FFC4FFC1FB4B6DB6FB00F73B" baseAuthorityName="Meyer" baseAuthorityYear="1899" box="[1042,1207,2214,2247]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="tonkeana">M. tonkeana</taxonomicName>
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
<taxonomicName id="81B05182FFC4FFC1FC0D6C58FBB6F699" box="[852,1025,2376,2405]" class="Mammalia" family="Cercopithecidae" genus="Papio" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="nigrescens">M. nigrescens</taxonomicName>
just east Gorontalo and in the basins of the Bolango and Bone rivers. In his 1980 review, C. P. Groves considered
<taxonomicName id="81B05182FFC4FFC1FFB76C86FEECF64F" box="[238,347,2454,2483]" class="Mammalia" family="Cercopithecidae" genus="Papio" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="hecki">M. hecki</taxonomicName>
to be a subspecies of
<taxonomicName id="81B05182FFC4FFC1FDD46C86FC87F64F" baseAuthorityName="Meyer" baseAuthorityYear="1899" box="[653,816,2454,2483]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="tonkeana">M. tonkeana</taxonomicName>
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.
</paragraph>
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<caption id="12CF7A89FFC4FFC1FFFD6FB5FE02F511" ID-DOI="http://doi.org/10.5281/zenodo.6867087" ID-Zenodo-Dep="6867087" httpUri="https://zenodo.org/record/6867087/files/figure.png" inLine="true" pageId="5" pageNumber="632" startId="5.[164,336,2725,2758]" targetBox="[168,748,1989,2395]" targetPageId="5">
<paragraph id="460F2A01FFC4FFC1FFFD6FB5FE02F511" blockId="5.[159,1368,2419,3465]" pageId="5" pageNumber="632">Distribution. NW Sulawesi from the base of the N peninsula (Isthmus of Palu) NE to just E of Gorontalo.</paragraph>
</caption>
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<subSubSection id="0EAA798AFFC4FFC1FFFD6FE8F96BFE71" pageId="5" pageNumber="632" type="description">
<paragraph id="460F2A01FFC4FFC1FFFD6FE8F96BFE71" blockId="5.[159,1368,2419,3465]" lastBlockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">
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). Hecks 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
<taxonomicName id="81B05182FFC4FFC1FF846E80FE97F44D" box="[221,288,2960,2993]" pageId="5" pageNumber="744" rank="form">form</taxonomicName>
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 (
<taxonomicName id="81B05182FFC4FFC1FD25691AFC9BF3DB" box="[636,812,3082,3111]" class="Mammalia" family="Cercopithecidae" genus="Papio" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="nigrescens">M. nigrescens</taxonomicName>
) with Hecks Macaque, the Gorontalo Macaque has a baboon-like face, an elongated narrow crest, blackish limbs contrasting with a brown body, and gray ischial callosities. Hecks 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 (
<taxonomicName id="81B05182FFC4FFC1FCBE69E6FB25F2EF" baseAuthorityName="Meyer" baseAuthorityYear="1899" box="[999,1170,3318,3347]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="tonkeana">M. tonkeana</taxonomicName>
) with Hecks 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 Hecks Macaque), cheek whiskers have a pale grayish or brownish tint (not in Hecks Macaque), rump patch is more clearly marked and bushy (not so in Hecks Macaque), and ischial callosities are oval and orange-colored (similar to the Moor Macaque,
<taxonomicName id="81B05182FFC4FFC1F7B1645AF6DDFE9B" baseAuthorityName="Schinz" baseAuthorityYear="1825" box="[2280,2410,330,359]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="maura">M. maura</taxonomicName>
, and the Booted Macaque,
<taxonomicName id="81B05182FFC4FFC1F96A6460F967FE71" baseAuthorityName="Ogilby" baseAuthorityYear="1841" box="[1587,1744,368,397]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="ochreata">M. ochreata</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAF86484F845FDAE" pageId="5" pageNumber="632" type="biology_ecology">
<paragraph id="460F2A01FFC4FFC1FAF86484F845FDAE" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">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 Hecks Macaque can thrive in degraded and secondary forest is not known.</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAF96749F944FCEA" pageId="5" pageNumber="632" type="food_feeding">
<paragraph id="460F2A01FFC4FFC1FAF96749F944FCEA" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">Food and Feeding. The three macaque species of the northern peninsula, including Hecks 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.</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAF96632F61BFBF8" pageId="5" pageNumber="632" type="breeding">
<paragraph id="460F2A01FFC4FFC1FAF96632F61BFBF8" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">Breeding. Reproductive cycle of female Hecks 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.</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAC7611EF99DFB85" pageId="5" pageNumber="632" type="activity">
<paragraph id="460F2A01FFC4FFC1FAC7611EF99DFB85" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">Activity patterns. Hecks 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.</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAC66190F854FAC3" pageId="5" pageNumber="632" type="biology_ecology">
<paragraph id="460F2A01FFC4FFC1FAC66190F854FAC3" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">Movements, Home range and Social organization. Hecks 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?.</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAC6605AF6D0F734" pageId="5" pageNumber="632" type="conservation">
<paragraph id="460F2A01FFC4FFC1FAC6605AF6D0F734" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">
Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Hecks 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, Indonesias 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. Hecks 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 (
<taxonomicName id="81B05182FFC4FFC1FAF563B8F996F935" baseAuthorityName="Desmarest" baseAuthorityYear="1822" box="[1452,1569,1704,1737]" class="Mammalia" family="Cercopithecidae" genus="Macaca" kingdom="Animalia" order="Primates" pageId="5" pageNumber="632" phylum="Chordata" rank="species" species="nigra">M. nigra</taxonomicName>
) 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 Hecks 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 Hecks Macaques at higher elevations is unknown. In 1999, the lack of strictly protected areas in the range of Hecks 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 Hecks Macaque. Since 1999, Nantu Wildlife Reserve and a number of other reserves have been created that protect, or should protect, Hecks Macaque: Dolangan, Mas Popaya Raja, Pangi Binanga, Pinjam/Tanjung Mantop, and Tangale reserves.
</paragraph>
</subSubSection>
<subSubSection id="0EAA798AFFC4FFC1FAC76DC7F8C1F671" pageId="5" pageNumber="632" type="bibRefCitation_list">
<paragraph id="460F2A01FFC4FFC1FAC76DC7F8C1F671" blockId="5.[1437,2646,291,2447]" pageId="5" pageNumber="632">Bibliography. Abegg &amp; Thierry (2002a), Bynum (1995, 1999), Bynum, Bynum, Froehlich &amp; Supriatna (1997), Bynum, Bynum &amp; Supriatna (1997), Bynum, Kohlhaas &amp; Pramono (1999), Cannon et al. (2007), Clayton &amp; Millner-Gulland (2000), Corbet &amp; 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 &amp; Matsumura (1991), Thierry (2000b, 2007, 2011), Watanabe, Lapasere &amp; Tantu (1991).</paragraph>
</subSubSection>
</treatment>
</document>