Black-and-white colobus

{{Short description|Genus of Old World monkeys}} {{Automatic taxobox |name = Black-and-white colobus<ref name=msw3>{{MSW3 Groves|pages=167–168}}</ref> |image = Colubusmonkey.JPG |image_caption = Mantled guereza (''Colobus guereza'') |taxon = Colobus |authority = Illiger, 1811 |display_parents = 2 |type_species = ''Simia polycomos'' |type_species_authority = Schreber, 1800<br />(= ''Cebus polykomos'' Zimmermann, 1780) |subdivision_ranks = Species |subdivision = :{{small|See text}} }}

'''Black-and-white colobuses''' (or '''colobi''') are Old World monkeys of the genus '''''Colobus''''', native to Africa. They are closely related to the red colobus monkeys of genus ''Piliocolobus''.<ref name=msw3/> There are five species of this monkey, and at least eight subspecies.<ref name=msw3/> They are generally found in high-density forests where they forage on leaves, flowers and fruit. Social groups of colobus are diverse, varying from group to group.<ref name=":03"/> Resident-egalitarian and allomothering relationships have been observed among the female population.<ref name=":1" /> Complex behaviours have also been observed in this species, including greeting rituals<ref name=":03" /> and varying group sleeping patterns.<ref name=":3" /> Colobi play a significant role in seed dispersal.

==Etymology== The word "colobus" comes from the Ancient Greek κολοβός (''kolobós''), meaning "docked, maimed" and refers to their stump-like thumb.

==Taxonomy== {{Species table |no-note=y |genus=Colobus |authority-name=Illiger |authority-year=1811 |species-count=five|narrow-percent=75}} {{Species table/row |name=Angola colobus |binomial=C. angolensis |image=File:Colobus_angolensis.jpg |image-size=180px |image-alt=Gray monkey |authority-name=P. L. Sclater |authority-year=1860 |subspecies={{Collapsible list |expand=yes |title=Six subspecies |bullets=on | ''C. a. angolensis'' (Sclater's Angola colobus) | ''C. a. cordieri'' (Cordier's Angola colobus) | ''C. a. cottoni'' (Powell-Cotton's Angola colobus) | ''C. a. palliatus'' (Tanzanian black-and-white colobus) | ''C. a. prigoginei'' (Prigogine's Angola colobus) | ''C. a. ruwenzorii'' (Ruwenzori colobus) | ''C. a. sharpei'' (Sharpe's Angola Colobus) }} |range=Central Africa |range-image=File:Angola_Colobus_area.png |range-image-size=160px |size={{convert|49|–|68|cm|in|0|abbr=on}} long, plus {{convert|70|–|83|cm|in|0|abbr=on}} tail<ref name="ADWAngolacolobus"/> |habitat=Forest<ref name="IUCNAngolacolobus"/> |hunting=Leaves, as well as stems, bark, flowers, buds, shoots, fruits, and insects<ref name="ADWAngolacolobus"/> |iucn-status=VU |population=Unknown |direction={{decrease|Population declining}}<ref name="IUCNAngolacolobus"/> }} {{Species table/row |name=Black colobus |binomial=C. satanas |image=File:1_-_Colobus_polykomos_2_-_Colobus_satanas.jpg |image-size=180px |image-alt=Black-and-white drawing of monkeys |authority-name=Waterhouse |authority-year=1838 |subspecies={{Collapsible list |expand=yes |title=Two subspecies |bullets=on | ''C. s. anthracinus'' (Gabon black colobus) | ''C. s. satanas'' (Bioko black colobus) }} |range=Western Africa |range-image=File:Black_Colobus_area.png |range-image-size=160px |size={{convert|50|–|70|cm|in|0|abbr=on}} long, plus {{convert|62|–|88|cm|in|0|abbr=on}} tail<ref name="ADWBlackcolobus"/> |habitat=Forest<ref name="IUCNBlackcolobus"/> |hunting=Nuts and seeds, as well as unripe fruit and leaves<ref name="ADWBlackcolobus"/> |iucn-status=VU |population=Unknown |direction={{decrease|Population declining}}<ref name="IUCNBlackcolobus"/> }} {{Species table/row |name=King colobus |binomial=C. polykomos |image=File:King_colobus_monkeys.jpg |image-size=180px |image-alt=Gray monkey |authority-name=Zimmermann |authority-year=1780 |authority-not-original=yes |range=Western Africa |range-image=File:King_Colobus_area.png |range-image-size=160px |size={{convert|45|–|72|cm|in|0|abbr=on}} long, plus {{convert|52|–|100|cm|in|0|abbr=on}} tail<ref name="ADWKingcolobus"/> |habitat=Forest and savanna<ref name="IUCNKingcolobus"/> |hunting=Leaves, as well as fruit and flowers<ref name="ADWKingcolobus"/> |iucn-status=EN |population=Unknown |direction={{decrease|Population declining}}<ref name="IUCNKingcolobus"/> }} {{Species table/row |name=Mantled guereza |binomial=C. guereza |image=File:Mantled_Guereza.jpg |image-size=148px |image-alt=Gray monkey |authority-name=Rüppell |authority-year=1835 |subspecies={{Collapsible list |expand= |title=Seven subspecies |bullets=on | ''C. g. caudatus'' (Kilimanjaro guereza) | ''C. g. dodingae'' (Dodinga Hills guereza) | ''C. g. guereza'' (Omo River guereza) | ''C. g. kikuyuensis'' (Eastern black-and-white colobus) | ''C. g. matschiei'' (Mau Forest guereza) | ''C. g. occidentalis'' (Western guereza) | ''C. g. percivali'' (Mt Uaraguess guereza) }} |range=Central Africa |range-image=File:Mapa distribución Colobus guereza.png |range-image-size=162px |size={{convert|45|–|72|cm|in|0|abbr=on}} long, plus {{convert|52|–|100|cm|in|0|abbr=on}} tail<ref name="ADWMantledguereza"/> |habitat=Forest<ref name="IUCNMantledguereza"/> |hunting=Leaves, as well as fruit, buds, and blossoms<ref name="ADWMantledguereza"/> |iucn-status=LC |population=Unknown |direction={{decrease|Population declining}}<ref name="IUCNMantledguereza"/> }} {{Species table/row |name=Ursine colobus |binomial=C. vellerosus |image=File:Colobus vellerosus 12785335.jpg |image-size=180px |image-alt=Black-and-white monkey |authority-name=I. Geoffroy |authority-year=1834 |authority-not-original=yes |range=Western Africa |range-image=File:Colobus_vellerosus_distribution.svg |range-image-size=180px |size={{convert|60|–|67|cm|in|0|abbr=on}} long, plus {{convert|73|–|93|cm|in|0|abbr=on}} tail<ref name="FieldAfrica114"/> |habitat=Forest<ref name="IUCNUrsinecolobus"/> |hunting=Leaves and seeds, as well as fruit, insects, and clay<ref name="ADWUrsinecolobus"/> |iucn-status=CR |population=975 |direction={{decrease|Population declining}}<ref name="IUCNUrsinecolobus"/> }} {{Species table/end}}

'''Fossil species''' *†''Colobus flandrini'' *†''Colobus freedmani''{{cn|date=February 2022}}

==Behaviour and ecology== thumb|A multi-male unit of Angolan colobus in a multilevel society Colobus habitats include primary and secondary forests, riverine forests, and wooded grasslands; they are found more in higher-density logged forests than in other primary forests. Their ruminant-like digestive systems have enabled them to occupy niches that are inaccessible to other primates: they are herbivorous, eating leaves, fruit, flowers, lichen, herbaceous vegetation and bark. Colobuses are important for seed dispersal through their sloppy eating habits, as well as through their digestive systems.

Leaf toughness influences colobus foraging efficiency. Tougher leaves correlate negatively with ingestion rate (g/min) as they are costly in terms of mastication, but positively with investment (chews/g).<ref name=":0">{{Cite journal|last1=Dunham|first1=Noah T.|last2=Lambert|first2=Alexander L.|date=2016|title=The role of leaf toughness on foraging efficiency in Angola black and white colobus monkeys ( Colobus angolensis palliatus ): Dunham and Lambert|journal=American Journal of Physical Anthropology|language=en|volume=161|issue=2|pages=343–354|doi=10.1002/ajpa.23036|pmid=27346431}}</ref> Individuals spend approximately 150 minutes actively feeding each day.<ref name=":0" /> In a montane habitat colobus are known to utilise lichen as a fallback food during periods of low food availability.<ref>{{cite journal |author1=Miller, A. |author2=Judge, D. |author3=Uwingeneye, G. |author4=Ndayishimiye, D. |author5=Grueter, C. C. |title=Diet and Use of Fallback Foods by Rwenzori Black-and-White Colobus (Colobus angolensis ruwenzorii) in Rwanda: Implications for Supergroup Formation. |journal=International Journal of Primatology |date=2020 |volume=41 |issue=3 |pages=434–457 |doi=10.1007/s10764-020-00143-w |s2cid=212731904 }}</ref>

===Social patterns and morphology=== Colobuses live in territorial groups that vary in both size (3-15 individuals) and structure.<ref name=":03">{{Cite journal|last1=Kutsukake|first1=Nobuyuki|last2=Suetsugu|first2=Noyuri|last3=Hasegawa|first3=Toshikazu|date=2006-11-16|title=Pattern, Distribution, and Function of Greeting Behavior Among Black-and-White Colobus|journal=International Journal of Primatology|language=en|volume=27|issue=5|pages=1271–1291|doi=10.1007/s10764-006-9072-x|s2cid=24835234|issn=0164-0291}}</ref><ref name=":04">{{cite journal |author1=Miller, A. |author2=Uddin, S. |author3=Judge, D. S. |author4=Kaplin, B. A.|author4-link=Beth Kaplin |author5=Ndayishimiye, D. |author6=Uwingeneye, G. |author7=Grueter, C. C. |title=Spatiotemporal association patterns in a supergroup of Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) are consistent with a multilevel society. |journal=American Journal of Primatology |date=2020 |volume=82 |issue=6 |article-number=e23127 |doi=10.1002/ajp.23127 |pmid=32249977 |s2cid=214808996 }}</ref><ref name=":05">{{cite journal |author1=Stead, S. M. |author2=Teichroeb, J. A. |title=A multi-level society {{sic|comprised |hide=y|of}} one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii). |journal=PLOS ONE |date=2019 |volume=14 |issue=10 |article-number=e0217666 |doi=10.1371/journal.pone.0217666 |pmid=31618212 |pmc=6795445 |doi-access=free }}</ref> It was originally believed that the structure of these groups consisted of one male and about 8 female members.<ref>{{Cite journal|last=Marler|first=Peter|date=1972|title=Vocalizations of East African Monkeys II: Black and White Colobus|journal=Behaviour|volume=42|issue=3–4|pages=175–197|doi=10.1163/156853972x00266|issn=0005-7959}}</ref> However, more recent observations have shown variation in structure and the number of males within groups, with one species forming multi-male, multifemale groups in a multilevel society, and in some populations supergroups form exceeding 500 individuals.<ref name=":04" /><ref name=":05" /> There appears to be a dominant male, whilst there is no clear dominance among female members.<ref name=":03"/> Relationships among females are considered to be resident-egalitarian, as there is low competition and aggression between them within their own groups. Juveniles are treated as a lower-rank (in regards to authority) than subadults and likewise when comparing subadults to adults.<ref name=":1">{{Cite journal|last1=Sterck|first1=Elisabeth H. M.|last2=Watts|first2=David P.|last3=van Schaik|first3=Carel P.|date=1997-11-20|title=The evolution of female social relationships in nonhuman primates|journal=Behavioral Ecology and Sociobiology|volume=41|issue=5|pages=291–309|doi=10.1007/s002650050390|issn=0340-5443|hdl=1874/1183|s2cid=25163826|hdl-access=free}}</ref> Colobuses do not display any type of seasonal breeding patterns.<ref>{{Cite journal|last1=Teichroeb|first1=Julie A.|last2=Sicotte|first2=Pascale|date=2009|title=Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana|journal=American Journal of Primatology|volume=71|issue=1|pages=49–59|doi=10.1002/ajp.20617|pmid=18828147|s2cid=31872244|issn=0275-2565}}</ref>

As suggested by their name, adult colobi have black fur with white features. White fur surrounds their facial region and a "U" shape of long white fur runs along the sides of their body. Newborn colobi are completely white with a pink face. Cases of allomothering, whereby members of the troop other than the infant's biological mother care for it, are documented. Allomothering is believed to increase inclusive fitness or maternal practice for the benefit of future offspring.<ref>{{Cite journal|last1=Dunham|first1=Noah Thomas|last2=Opere|first2=Paul Otieno|date=2016-02-12|title=A unique case of extra-group infant adoption in free-ranging Angola black and white colobus monkeys (Colobus angolensis palliatus)|journal=Primates|volume=57|issue=2|pages=187–194|doi=10.1007/s10329-016-0515-5|pmid=26872896|s2cid=14120148|issn=0032-8332}}</ref>

=== Social behaviours === thumb|Angolan black-and-white colobus with infant. Many members participate in a greeting ritual when they are reunited with familiar individuals, an act of reaffirming.<ref name=":03"/> The greeting behaviour is generally carried out by the approaching monkey and often is followed with grooming. They participate in three greeting behaviours of physical contact. This includes mounting, head mounting (grasps the shoulders) and embracing.<ref name=":03" /> It seems as though these behaviours do not have any relationship with mating or courting.<ref name=":03" />thumb|Adult Eastern black-and-white colobus with a juvenileBlack-and-white colobus have complex sleeping patterns. They sleep in trees near a food source, which may serve to save energy.<ref name=":3">{{Cite journal|last=Von Hippel|first=Frank A.|date=1998|title=Use of sleeping trees by black and white Colobus monkeys (Colobus guereza) in the Kakamega Forest, Kenya|journal=American Journal of Primatology|volume=45|issue=3|pages=281–290|doi=10.1002/(sici)1098-2345(1998)45:3<281::aid-ajp4>3.0.co;2-s|pmid=9651650|issn=0275-2565}}</ref> Groups seem to regularly switch up sleeping locations (suggested due to reducing risk of parasites and placement prediction) and generally do not sleep near other groups. They also tend to sleep more tightly together on nights with great visibility.<ref name=":3" /> They sleep in mid- to upper sections of tall trees which allows for predator watch as well as protection from ground and aerial predators while they are asleep.<ref name=":3" /> Although there is no obvious preference for tree type, they have often been observed in ''Antiaris toxicaria.''<ref name=":3" />

==Conservation== Colobuses are prey for many forest predators such as leopards and chimpanzees,{{cn|date=May 2024}} and are threatened by hunting for the bushmeat trade, logging, and habitat destruction.

Individuals are more vigilant (conspecific threat) in low canopy, they also spend less time scanning when they are around familiar group members as opposed to unfamiliar.<ref name=":2">{{Cite journal|last1=MacIntosh|first1=Andrew J.J.|last2=Sicotte|first2=Pascale|date=2009|title=Vigilance in ursine black and white colobus monkeys ( Colobus vellerosus ): an examination of the effects of conspecific threat and predation|journal=American Journal of Primatology|language=en|volume=71|issue=11|pages=919–927|doi=10.1002/ajp.20730|pmid=19598222|s2cid=205329258}}</ref> There exists no clear difference in vigilance between male and females. However, there is a positive correlation between mean monthly vigilance and encounter rates.<ref name=":2" /> Male vigilance generally increases during mating.<ref>{{Cite journal|last1=Baldellou|first1=Maribel|last2=Peter Henzi|first2=S.|date=1992|title=Vigilance, predator detection and the presence of supernumerary males in vervet monkey troops|journal=Animal Behaviour|volume=43|issue=3|pages=451–461|doi=10.1016/s0003-3472(05)80104-6|s2cid=53202789|issn=0003-3472}}</ref>

==References== {{Reflist|refs= <ref name="IUCNAngolacolobus">{{cite iucn |last1=de Jong |first1=Y. A. |last2=Cunneyworth |first2=P. |last3=Butynski |first3=T. M. |last4=Maisels |first4=F. |last5=Hart |first5=J. A. |last6=Rovero |first6=F. |title=''Colobus angolensis'' |volume=2020 |article-number=e.T5142A17945007 |date=2020 |doi=10.2305/IUCN.UK.2020-2.RLTS.T5142A17945007.en}}</ref> <ref name="ADWAngolacolobus">{{cite web |first1=Brandon |last1=Thompson |title=''Colobus angolensis'' |url=https://animaldiversity.org/accounts/Colobus_angolensis/ |date=2002 |website=Animal Diversity Web |publisher=University of Michigan |access-date=July 24, 2023 |archive-date=August 12, 2023 |archive-url=https://web.archive.org/web/20230812151146/https://animaldiversity.org/accounts/Colobus_angolensis/ |url-status=live }}</ref> <ref name="IUCNMantledguereza">{{cite iucn |last1=de Jong |first1=Y. A. |last2=Butynski |first2=T. M. |last3=Oates |first3=J. F. |title=''Colobus guereza'' |volume=2019 |article-number=e.T5143A17944705 |date=2019 |doi=10.2305/IUCN.UK.2019-3.RLTS.T5143A17944705.en}}</ref> <ref name="ADWMantledguereza">{{cite web |first1=Kenneth |last1=Kim |title=''Colobus guereza'' |url=https://animaldiversity.org/accounts/Colobus_guereza/ |date=2002 |website=Animal Diversity Web |publisher=University of Michigan |access-date=July 24, 2023 |archive-date=August 12, 2023 |archive-url=https://web.archive.org/web/20230812151658/https://animaldiversity.org/accounts/Colobus_guereza/ |url-status=live }}</ref> <ref name="IUCNKingcolobus">{{cite iucn |last1=Gonedelé Bi |first1=S. |last2=Koné |first2=I. |last3=Matsuda Goodwin |first3=R. |last4=Alonso |first4=C. |last5=Hernansaiz |first5=A. |last6=Oates |first6=J. F. |title=''Colobus polykomos'' |volume=2020 |article-number=e.T5144A17944855 |date=2020 |doi=10.2305/IUCN.UK.2020-2.RLTS.T5144A17944855.en}}</ref> <ref name="ADWKingcolobus">{{cite web |first1=Devon |last1=Landes |title=''Colobus polykomos'' |url=https://animaldiversity.org/accounts/Colobus_polykomos/ |date=2000 |website=Animal Diversity Web |publisher=University of Michigan |access-date=July 24, 2023 |archive-date=August 12, 2023 |archive-url=https://web.archive.org/web/20230812172410/https://animaldiversity.org/accounts/Colobus_polykomos/ |url-status=live }}</ref> <ref name="IUCNBlackcolobus">{{cite iucn |last1=Maisels |first1=F. |last2=Cronin |first2=D. T. |title=''Colobus satanas'' |volume=2020 |article-number=e.T5145A17944405 |date=2020 |doi=10.2305/IUCN.UK.2020-2.RLTS.T5145A17944405.en}}</ref> <ref name="ADWBlackcolobus">{{cite web |first1=Whitney |last1=Lane |title=''Colobus satanas'' |url=https://animaldiversity.org/accounts/Colobus_satanas/ |date=2011 |website=Animal Diversity Web |publisher=University of Michigan |access-date=July 24, 2023 |archive-date=August 12, 2023 |archive-url=https://web.archive.org/web/20230812153212/https://animaldiversity.org/accounts/Colobus_satanas/ |url-status=live }}</ref> <ref name="IUCNUrsinecolobus">{{cite iucn |last1=Matsuda Goodwin |first1=R. |last2=Gonedelé Bi |first2=S. |last3=Nobimè |first3=G. |last4=Koné |first4=I. |last5=Osei |first5=D. |last6=Segniagbeto |first6=G. |last7=Oates |first7=J. F. |title=''Colobus vellerosus'' |volume=2020 |article-number=e.T5146A169472127 |date=2020 |doi=10.2305/IUCN.UK.2020-2.RLTS.T5146A169472127.en}}</ref> <ref name="ADWUrsinecolobus">{{cite web |first1=Shannon |last1=Walker |title=''Colobus vellerosus'' |url=https://animaldiversity.org/accounts/Colobus_vellerosus/ |date=2009 |website=Animal Diversity Web |publisher=University of Michigan |access-date=July 24, 2023 |archive-date=August 4, 2020 |archive-url=https://web.archive.org/web/20200804000004/https://animaldiversity.org/accounts/Colobus_vellerosus/ |url-status=live }}</ref> <ref name="FieldAfrica114">Kingdon 2015, p. 114</ref>}}

==Sources== {{refbegin}} * {{cite book |title=The Kingdon Field Guide to African Mammals |edition=Second |last=Kingdon |first=Jonathan |date=2015 |publisher=Bloomsbury Publishing |isbn=978-1-4729-2531-2 |ref=CITEREF_AfricanMammals}} {{refend}}

==External links== {{Commons category|Colobus|''Colobus''}} {{Wikispecies|Colobus''Colobus''}}

* [http://www.awf.org/wildlives/66 Colobus Monkey: Wildlife summary from the African Wildlife Foundation] * [http://www.colobusconservation.org/ Colobus Conservation] * [http://pin.primate.wisc.edu/factsheets/links/colobus Primate Info Net ''Colobus'' Factsheets] * [http://www.colobusconservation.org/index.php/ecotours/primates-category/168-angolan-black-and-white-colobus Angolan Black & White Colobus] {{Webarchive|url=https://web.archive.org/web/20131224100000/http://www.colobusconservation.org/index.php/ecotours/primates-category/168-angolan-black-and-white-colobus |date=2013-12-24 }}

Category:Colobus Category:Primates of Africa Category:Folivores Category:Taxa named by Johann Karl Wilhelm Illiger