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{{Short description|Genus of Old World monkeys}}
{{Automatic taxobox
{{Automatic taxobox
|name = Black-and-white colobus<ref name=msw3>{{MSW3 Groves|pages=167–168}}</ref>
|name = Black-and-white colobus<ref name=msw3>{{MSW3 Groves|pages=167–168}}</ref>
Line 5: Line 6:
|taxon = Colobus
|taxon = Colobus
|authority = [[Johann Karl Wilhelm Illiger|Illiger]], 1811
|authority = [[Johann Karl Wilhelm Illiger|Illiger]], 1811
|display_parents = 2
|type_species = ''[[King Colobus|Simia polycomos]]''
|type_species = ''[[King Colobus|Simia polycomos]]''
|type_species_authority = [[Johann Christian Daniel von Schreber|Schreber]], 1800<br>(''Cebus polykomos'' [[Eberhard August Wilhelm von Zimmermann|Zimmermann]], 1780)
|type_species_authority = [[Johann Christian Daniel von Schreber|Schreber]], 1800<br>(''Cebus polykomos'' [[Eberhard August Wilhelm von Zimmermann|Zimmermann]], 1780)
|subdivision_ranks = [[Species]]
|subdivision_ranks = [[Species]]
|subdivision = :{{small|See text}}
|subdivision = ''[[Black colobus|Colobus satanas]]''<br> ''[[Angola colobus|Colobus angolensis]]''<br> ''[[King colobus|Colobus polykomos]]''<br> ''[[Ursine colobus|Colobus vellerosus]]''<br> ''[[Mantled guereza|Colobus guereza]]''
}}
}}

'''Black-and-white colobuses''' (or '''colobi''') are [[Old World monkey]]s of the genus '''''Colobus''''', native to [[Africa]]. They are closely related to the red<!--brown : the supporting reference uses 'red' for every species--> 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=":02" /> 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.
'''Black-and-white colobuses''' (or '''colobi''') are [[Old World monkey]]s of the genus '''''Colobus''''', native to [[Africa]]. They are closely related to the red<!--brown : the supporting reference uses 'red' for every species--> 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==
==Etymology==
The word "colobus" comes from [[Greek language|Greek]] κολοβός ''kolobós'' ("docked"), so-named because the thumb is stump-like.
The word "colobus" comes from [[ancient Greek language|Greek]] {{wikt-lang|grc|κολοβός}} (''kolobós'', "docked", "maimed"), so named because the thumb is stump-like.


==Taxonomy==
==Habitat and food==
{{Species table |no-note=y |genus=[[Colobus]] |authority-name=[[Johann Karl Wilhelm Illiger|Illiger]] |authority-year=1811 |species-count=five|narrow-percent=75}}
Colobuses 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 [[Herbivore|herbivorous]], eating leaves, fruit, flowers, lichen and twigs.
{{Species table/row
|name=[[Angola colobus]] |binomial=C. angolensis
|image=File:Colobus_angolensis.jpg |image-size=180px |image-alt=Gray monkey
|authority-name=[[Philip Sclater|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. palliates'' (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=[[George Robert Waterhouse|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=[[Eberhard August Wilhelm von Zimmermann|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=[[Eduard Rüppell|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=[[Isidore Geoffroy Saint-Hilaire|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'''
Leaf toughness influences colobus foraging efficiency. Tougher leaves correlate negatively with ingestion rate (g/min) as they are costly in terms of [[Chewing|mastication]], but positively with investment (chews/g).<ref name=":0">{{Cite journal|last=Dunham|first=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 |pages=1-24 |doi=10.1007/s10764-020-00143-w |url=https://link.springer.com/article/10.1007/s10764-020-00143-w}}</ref>
*†''[[Colobus flandrini]]''
*†''[[Colobus freedmani]]''{{cn|date=February 2022}}


==Social patterns and morphology==
==Behaviour and ecology==
[[File:Multi-male unit of Rwenzori colobus in Nyungwe .jpg|thumb|A multi-male unit of Angolan colobus in a multilevel society]]
Colobuses live in territorial groups that vary in both size (3-15 individuals) and structure.<ref name=":02">{{Cite journal|last=Kutsukake|first=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|issn=0164-0291}}</ref><ref name=":04">{{cite journal |author1=Miller, A. |author2=Uddin, S. |author3=Judge, D. S. |author4=Kaplin, B. A. |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 |doi=10.1002/ajp.23127 |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajp.23127}}</ref><ref name=":05">{{cite journal |author1=Stead, S. M. |author2=Teichroeb, J. A. |title=A multi-level society comprised of one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii). |journal=PLoS One |date=2019 |doi=10.1371/journal.pone.0217666 |url=https://journals.plos.org/plosone/article/file?type=printable&id=10.1371/journal.pone.0217666}}</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=":02" /> 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|last=Sterck|first=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|hdl-access=free}}</ref> Colobuses do not display any type of seasonal breeding patterns.<ref>{{Cite journal|last=Teichroeb|first=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|issn=0275-2565}}</ref>
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 [[Herbivore|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 [[Chewing|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>
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]] are documented, which means members of the troop other than the infant's biological mother care for it. Possible explanations to this are, increasing inclusive fitness or maternal practice which will benefit future offspring.<ref>{{Cite journal|last=Dunham|first=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|issn=0032-8332}}</ref>


== Social behaviours ==
===Social patterns and morphology===
Many members participate in a greeting ritual when they are reunited with familiar individuals, an act of reaffirming.<ref name=":03">{{Cite journal|last=Kutsukake|first=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|issn=0164-0291}}</ref> The greeting behaviour is generally carried out by the approaching monkey and often times 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" />[[File:Eastern black-and-white colobus (Colobus guereza occidentalis) with juvenile.jpg|thumb|Adult Eastern black-and-white colobus with a juvenile ]]Black-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|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" />
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 |pages=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 |pages=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]] are documented, which means members of the troop other than the infant's biological mother care for it. [[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>
== Ecology ==
Colobuses are important for [[seed dispersal]] through their sloppy eating habits, as well as through their digestive systems.


=== Social behaviours ===
==Predators and threat level==
[[File:Angolan colobus with infant.jpg|thumb|Angolan black-and-white colobus with infant.]]
They are prey for many forest predators, and are threatened by hunting for the [[bushmeat]] trade, logging, and habitat destruction.
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" />[[File:Eastern black-and-white colobus (Colobus guereza occidentalis) with juvenile.jpg|thumb|Adult Eastern black-and-white colobus with a juvenile]]Black-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==
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|last=MacIntosh|first=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}}</ref> There are 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|last=Baldellou|first=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|issn=0003-3472}}</ref>
They are prey for many forest predators such as [[leopard]]s and [[chimpanzee]]s,{{cn|date=May 2024}} and are threatened by hunting for the [[bushmeat]] trade, logging, and habitat destruction.
==Species==

[[File:Trapped-1.JPG|thumb|right|Angolan black-and-white colobus (''Colobus angolensis'') seen in Tanzania.]]
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 are 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>
*'''Genus ''Colobus'' '''
**[[Black colobus]], ''C. satanas''
***Gabon black colobus, ''C. s. anthracinus''
***Bioko black colobus, ''C. s. satanas''
**[[Angola colobus]], ''C. angolensis''
***Sclater's Angola colobus, ''C. a. angolensis''
***Powell-Cotton's Angola colobus, ''C. a. cottoni''
***Adolf Friedrichs's Angola colobus, or Ruwenzori black-and-white colobus, ''C. a. ruwenzorii''
***Cordier's Angola colobus, ''C. a. cordieri''
***Prigogine's Angola colobus, ''C. a. prigoginei''
***Peters's Angola colobus or Tanzanian black-and-white colobus, ''C. a. palliatus''
**[[King colobus]], ''C. polykomos''
**[[Ursine colobus]], ''C. vellerosus''
**[[Mantled guereza]] or Abyssinian black-and-white colobus, ''C. guereza''<ref>Wolfheim, J. H. (1983). ''Primates Of The World: Distribution, Abundance And Conservation''. Routledge. {{ISBN|3-7186-0190-7}}.</ref>''


==References==
==References==
{{Reflist}}
{{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 |page=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 |page=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 |page=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 |page=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 |page=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">[[#CITEREF_AfricanMammals|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==
==External links==
Line 63: Line 150:
* [http://pin.primate.wisc.edu/factsheets/links/colobus Primate Info Net ''Colobus'' Factsheets]
* [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]
* [http://www.colobusconservation.org/index.php/ecotours/primates-category/168-angolan-black-and-white-colobus Angolan Black & White Colobus]
* [http://www.seaworld.org/animal-info/animal-bytes/animalia/eumetazoa/coelomates/deuterostomes/chordata/craniata/mammalia/primates/b-&-w-colobus.htm Animal Bytes]


{{C.Colobinae nav}}
{{C.Colobinae nav}}
{{Haplorhini|C.}}
{{Taxonbar|from=Q358813}}
{{Taxonbar|from=Q358813}}


Latest revision as of 17:05, 28 July 2024

Black-and-white colobus[1]
Mantled guereza (Colobus guereza)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Cercopithecidae
Subfamily: Colobinae
Tribe: Colobini
Genus: Colobus
Illiger, 1811
Type species
Simia polycomos
Schreber, 1800
(Cebus polykomos Zimmermann, 1780)
Species
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.[1] There are five species of this monkey, and at least eight subspecies.[1] 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.[2] Resident-egalitarian and allomothering relationships have been observed among the female population.[3] Complex behaviours have also been observed in this species, including greeting rituals[2] and varying group sleeping patterns.[4] Colobi play a significant role in seed dispersal.

Etymology

[edit]

The word "colobus" comes from Greek κολοβός (kolobós, "docked", "maimed"), so named because the thumb is stump-like.

Taxonomy

[edit]
Genus Colobus – Illiger, 1811 – five species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Angola colobus

Gray monkey

C. angolensis
P. L. Sclater, 1860

Six subspecies
  • C. a. angolensis (Sclater's Angola colobus)
  • C. a. cordieri (Cordier's Angola colobus)
  • C. a. cottoni (Powell-Cotton's Angola colobus)
  • C. a. palliates (Tanzanian black-and-white colobus)
  • C. a. prigoginei (Prigogine's Angola colobus)
  • C. a. ruwenzorii (Ruwenzori colobus)
  • C. a. sharpei (Sharpe's Angola Colobus)
Central Africa
Map of range 		Size: 49–68 cm (19–27 in) long, plus 70–83 cm (28–33 in) tail[5]

Habitat: Forest[6]

Diet: Leaves, as well as stems, bark, flowers, buds, shoots, fruits, and insects[5] 		 VU 


Unknown Population declining[6]

Black colobus

Black-and-white drawing of monkeys

C. satanas
Waterhouse, 1838

Two subspecies
  • C. s. anthracinus (Gabon black colobus)
  • C. s. satanas (Bioko black colobus)
Western Africa
Map of range 		Size: 50–70 cm (20–28 in) long, plus 62–88 cm (24–35 in) tail[7]

Habitat: Forest[8]

Diet: Nuts and seeds, as well as unripe fruit and leaves[7] 		 VU 


Unknown Population declining[8]

King colobus

Gray monkey

C. polykomos
(Zimmermann, 1780) 		Western Africa
Map of range 		Size: 45–72 cm (18–28 in) long, plus 52–100 cm (20–39 in) tail[9]

Habitat: Forest and savanna[10]

Diet: Leaves, as well as fruit and flowers[9] 		 EN 


Unknown Population declining[10]

Mantled guereza

Gray monkey

C. guereza
Rüppell, 1835

Seven subspecies
  • 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)
Central Africa
Map of range 		Size: 45–72 cm (18–28 in) long, plus 52–100 cm (20–39 in) tail[11]

Habitat: Forest[12]

Diet: Leaves, as well as fruit, buds, and blossoms[11] 		 LC 


Unknown Population declining[12]

Ursine colobus

Black-and-white monkey

C. vellerosus
(I. Geoffroy, 1834) 		Western Africa
Map of range 		Size: 60–67 cm (24–26 in) long, plus 73–93 cm (29–37 in) tail[13]

Habitat: Forest[14]

Diet: Leaves and seeds, as well as fruit, insects, and clay[15] 		 CR 


975 Population declining[14]

Fossil species

Behaviour and ecology

[edit]
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).[16] Individuals spend approximately 150 minutes actively feeding each day.[16] In a montane habitat colobus are known to utilise lichen as a fallback food during periods of low food availability.[17]

Social patterns and morphology

[edit]

Colobuses live in territorial groups that vary in both size (3-15 individuals) and structure.[2][18][19] It was originally believed that the structure of these groups consisted of one male and about 8 female members.[20] 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.[18][19] There appears to be a dominant male, whilst there is no clear dominance among female members.[2] 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.[3] Colobuses do not display any type of seasonal breeding patterns.[21]

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 are documented, which means members of the troop other than the infant's biological mother care for it. Allomothering is believed to increase inclusive fitness or maternal practice for the benefit of future offspring.[22]

Social behaviours

[edit]
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.[2] 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.[2] It seems as though these behaviours do not have any relationship with mating or courting.[2]

Adult Eastern black-and-white colobus with a juvenile

Black-and-white colobus have complex sleeping patterns. They sleep in trees near a food source, which may serve to save energy.[4] 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.[4] 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.[4] Although there is no obvious preference for tree type, they have often been observed in Antiaris toxicaria.[4]

Conservation

[edit]

They are prey for many forest predators such as leopards and chimpanzees,[citation needed] 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.[23] There are no clear difference in vigilance between male and females. However, there is a positive correlation between mean monthly vigilance and encounter rates.[23] Male vigilance generally increases during mating.[24]

References

[edit]
  1. ^ a b c Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 167–168. ISBN 0-801-88221-4. OCLC 62265494.
  2. ^ a b c d e f g Kutsukake, Nobuyuki; Suetsugu, Noyuri; Hasegawa, Toshikazu (2006-11-16). "Pattern, Distribution, and Function of Greeting Behavior Among Black-and-White Colobus". International Journal of Primatology. 27 (5): 1271–1291. doi:10.1007/s10764-006-9072-x. ISSN 0164-0291. S2CID 24835234.
  3. ^ a b Sterck, Elisabeth H. M.; Watts, David P.; van Schaik, Carel P. (1997-11-20). "The evolution of female social relationships in nonhuman primates". Behavioral Ecology and Sociobiology. 41 (5): 291–309. doi:10.1007/s002650050390. hdl:1874/1183. ISSN 0340-5443. S2CID 25163826.
  4. ^ a b c d e Von Hippel, Frank A. (1998). "Use of sleeping trees by black and white Colobus monkeys (Colobus guereza) in the Kakamega Forest, Kenya". American Journal of Primatology. 45 (3): 281–290. doi:10.1002/(sici)1098-2345(1998)45:3<281::aid-ajp4>3.0.co;2-s. ISSN 0275-2565. PMID 9651650.
  5. ^ a b Thompson, Brandon (2002). "Colobus angolensis". Animal Diversity Web. University of Michigan. Archived from the original on August 12, 2023. Retrieved July 24, 2023.
  6. ^ a b de Jong, Y. A.; Cunneyworth, P.; Butynski, T. M.; Maisels, F.; Hart, J. A.; Rovero, F. (2020). "Colobus angolensis". IUCN Red List of Threatened Species. 2020: e.T5142A17945007. doi:10.2305/IUCN.UK.2020-2.RLTS.T5142A17945007.en.
  7. ^ a b Lane, Whitney (2011). "Colobus satanas". Animal Diversity Web. University of Michigan. Archived from the original on August 12, 2023. Retrieved July 24, 2023.
  8. ^ a b Maisels, F.; Cronin, D. T. (2020). "Colobus satanas". IUCN Red List of Threatened Species. 2020: e.T5145A17944405. doi:10.2305/IUCN.UK.2020-2.RLTS.T5145A17944405.en.
  9. ^ a b Landes, Devon (2000). "Colobus polykomos". Animal Diversity Web. University of Michigan. Archived from the original on August 12, 2023. Retrieved July 24, 2023.
  10. ^ a b Gonedelé Bi, S.; Koné, I.; Matsuda Goodwin, R.; Alonso, C.; Hernansaiz, A.; Oates, J. F. (2020). "Colobus polykomos". IUCN Red List of Threatened Species. 2020: e.T5144A17944855. doi:10.2305/IUCN.UK.2020-2.RLTS.T5144A17944855.en.
  11. ^ a b Kim, Kenneth (2002). "Colobus guereza". Animal Diversity Web. University of Michigan. Archived from the original on August 12, 2023. Retrieved July 24, 2023.
  12. ^ a b de Jong, Y. A.; Butynski, T. M.; Oates, J. F. (2019). "Colobus guereza". IUCN Red List of Threatened Species. 2019: e.T5143A17944705. doi:10.2305/IUCN.UK.2019-3.RLTS.T5143A17944705.en.
  13. ^ Kingdon 2015, p. 114
  14. ^ a b Matsuda Goodwin, R.; Gonedelé Bi, S.; Nobimè, G.; Koné, I.; Osei, D.; Segniagbeto, G.; Oates, J. F. (2020). "Colobus vellerosus". IUCN Red List of Threatened Species. 2020: e.T5146A169472127. doi:10.2305/IUCN.UK.2020-2.RLTS.T5146A169472127.en.
  15. ^ Walker, Shannon (2009). "Colobus vellerosus". Animal Diversity Web. University of Michigan. Archived from the original on August 4, 2020. Retrieved July 24, 2023.
  16. ^ a b Dunham, Noah T.; Lambert, Alexander L. (2016). "The role of leaf toughness on foraging efficiency in Angola black and white colobus monkeys ( Colobus angolensis palliatus ): Dunham and Lambert". American Journal of Physical Anthropology. 161 (2): 343–354. doi:10.1002/ajpa.23036. PMID 27346431.
  17. ^ Miller, A.; Judge, D.; Uwingeneye, G.; Ndayishimiye, D.; Grueter, C. C. (2020). "Diet and Use of Fallback Foods by Rwenzori Black-and-White Colobus (Colobus angolensis ruwenzorii) in Rwanda: Implications for Supergroup Formation". International Journal of Primatology. 41 (3): 434–457. doi:10.1007/s10764-020-00143-w. S2CID 212731904.
  18. ^ a b Miller, A.; Uddin, S.; Judge, D. S.; Kaplin, B. A.; Ndayishimiye, D.; Uwingeneye, G.; Grueter, C. C. (2020). "Spatiotemporal association patterns in a supergroup of Rwenzori black-and-white colobus (Colobus angolensis ruwenzorii) are consistent with a multilevel society". American Journal of Primatology. 82 (6): e23127. doi:10.1002/ajp.23127. PMID 32249977. S2CID 214808996.
  19. ^ a b Stead, S. M.; Teichroeb, J. A. (2019). "A multi-level society comprised of one-male and multi-male core units in an African colobine (Colobus angolensis ruwenzorii)". PLOS ONE. 14 (10): e0217666. doi:10.1371/journal.pone.0217666. PMC 6795445. PMID 31618212.
  20. ^ Marler, Peter (1972). "Vocalizations of East African Monkeys II: Black and White Colobus". Behaviour. 42 (3–4): 175–197. doi:10.1163/156853972x00266. ISSN 0005-7959.
  21. ^ Teichroeb, Julie A.; Sicotte, Pascale (2009). "Test of the ecological-constraints model on ursine colobus monkeys (Colobus vellerosus) in Ghana". American Journal of Primatology. 71 (1): 49–59. doi:10.1002/ajp.20617. ISSN 0275-2565. PMID 18828147. S2CID 31872244.
  22. ^ Dunham, Noah Thomas; Opere, Paul Otieno (2016-02-12). "A unique case of extra-group infant adoption in free-ranging Angola black and white colobus monkeys (Colobus angolensis palliatus)". Primates. 57 (2): 187–194. doi:10.1007/s10329-016-0515-5. ISSN 0032-8332. PMID 26872896. S2CID 14120148.
  23. ^ a b MacIntosh, Andrew J.J.; Sicotte, Pascale (2009). "Vigilance in ursine black and white colobus monkeys ( Colobus vellerosus ): an examination of the effects of conspecific threat and predation". American Journal of Primatology. 71 (11): 919–927. doi:10.1002/ajp.20730. PMID 19598222. S2CID 205329258.
  24. ^ Baldellou, Maribel; Peter Henzi, S. (1992). "Vigilance, predator detection and the presence of supernumerary males in vervet monkey troops". Animal Behaviour. 43 (3): 451–461. doi:10.1016/s0003-3472(05)80104-6. ISSN 0003-3472. S2CID 53202789.

Sources

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