Editing Colobinae

{{Short description|Subfamily of Old World monkeys}}
{{Redirect|Langur|other uses}}
{{Automatic taxobox
| name = Colobine monkeys<ref name=msw3>{{MSW3 Primates|pages=167–178|id=12100596 |heading=SUBFAMILY '''Colobinae'''}}</ref>
| fossil_range = Late Miocene-recent {{fossilrange|12.5|0}}
| image = Gray langur (Semnopithecus) by Shantanu Kuveskar.jpg
| image_caption = [[Gray langur]] in Maharashtra, India
| taxon = Colobinae
| authority = [[Thomas C. Jerdon|Jerdon]], 1867
| subdivision_ranks = [[Genus|Genera]]
| subdivision = ''[[Black-and-white colobus|Colobus]]''<br />
''[[Red colobus|Piliocolobus]]''<br />
''[[Olive Colobus|Procolobus]]''<br />
''[[Trachypithecus]]''<br />
''[[Surili|Presbytis]]''<br />
''[[Gray langur|Semnopithecus]]''<br />
''[[Douc|Pygathrix]]''<br />
''[[Snub-nosed monkey|Rhinopithecus]]''<br />
''[[Proboscis monkey|Nasalis]]''<br />
''[[Pig-tailed langur|Simias]]''<br />
†''[[Cercopithecoides]]''<br />
†''[[Dolichopithecus]]''<br />
†''[[Mesopithecus]]''<br />
†''[[Microcolobus]]''<br />
†''[[Paracolobus]]''<br />
†''[[Parapresbytis]]''<br />
†''[[Rhinocolobus]]''
}}

The '''Colobinae''' or '''leaf-eating monkeys''' are a [[family (biology)|subfamily]] of the [[Old World monkey]] family that includes 61 [[species]] in 11 [[genus|genera]], including the [[black-and-white colobus]], the large-nosed [[proboscis monkey]], and the [[gray langur]]s. Some classifications split the '''colobine''' monkeys into two tribes, while others split them into three groups. Both classifications put the three African genera ''[[Black-and-white colobus|Colobus]]'', ''[[Red colobus|Piliocolobus]]'', and ''[[Olive colobus|Procolobus]]'' in one group; these genera are distinct in that they have stub thumbs (Greek κολοβός ''kolobós'' = "docked"). The various Asian genera are placed into another one or two groups. Analysis of [[Mitochondrial DNA|mtDNA]] confirms the Asian species form two distinct groups, one of langurs and the other of the "odd-nosed" species, but are inconsistent as to the relationships of the gray langurs; some studies suggest that the gray langurs are not closely related to either of these groups,<ref name=Sterner2006/> while others place them firmly within the langur group.<ref name=Osterholz2008>{{cite journal | last1=Osterholz | first1=Martin | last2=Walter | first2=Lutz | last3=Roos | first3=Christian | title=Phylogenetic position of the langur genera Semnopithecus and Trachypithecus among Asian colobines, and genus affiliations of their species groups | journal=BMC Evolutionary Biology | publisher=Springer Nature | volume=8 | issue=1 | year=2008 | issn=1471-2148 | doi=10.1186/1471-2148-8-58 | page=58|pmid=18298809|pmc=2268674 | doi-access=free | bibcode=2008BMCEE...8...58O }}</ref>

== Characteristics ==
[[File:NILGIRI LANGUR Trachypithecus johnii.jpg|thumb|250px|Colobines are mainly [[arboreal]] and [[folivorous]] primates. Adult [[Nilgiri Langur]] pictured.]]

Colobines are medium-sized [[primate]]s with long tails (except for the [[pig-tailed langur]]) and diverse colorations. The coloring of nearly all young animals differs remarkably from that of the adults.

Most species are [[arboreal]], although some live a more [[terrestrial animal|terrestrial]] life. They are found in many different habitats of different climate zones ([[rainforest]]s, [[mangrove]]s, mountain forests, and [[savannah]]), but not in deserts and other dry areas. They live in groups, but in social forms vary.

Colobines are [[folivore|folivorous]], though their diet may be supplemented with flowers, fruits and the occasional insect. To aid in digestion, particularly of hard-to-digest leaves, they have multichambered, complex stomachs, making them the only primates with [[foregut fermentation]]. Foregut fermenters use bacteria to detoxify plant compounds before reaching the intestine, where toxins can be absorbed. Foregut fermentation is also associated with higher protein extraction and efficient digestion of fiber;<ref name=":0">{{Cite journal|last1=Edwards|first1=Mark S.|last2=Ullrey|first2=Duane E.|date=1999|title=Effect of dietary fiber concentration on apparent digestibility and digesta passage in non-human primates. II. Hindgut- and foregut-fermenting folivores|journal=Zoo Biology|language=en|volume=18|issue=6|pages=537–549|doi=10.1002/(SICI)1098-2361(1999)18:6<537::AID-ZOO8>3.0.CO;2-F|issn=1098-2361|doi-access=free}}</ref><ref>{{Cite book|last=Feldhamer, George A.; Lee C. Drickamer; Stephen H. Vessey|url=http://worldcat.org/oclc/1097452280|title=Mammalogy : Adaptation, Diversity, Ecology|year=2007|isbn=978-1-4214-3652-4|pages=128–130|publisher=JHU Press |oclc=1097452280}}</ref> it is the dominant form of digestions in diverse herbivore taxa, including most Artiodactyla (e.g., deer, cattle, antelope), sloths, and kangaroos. In contrast, lower diversity howler monkeys in the New World rely on hindgut fermentation – occurring lower in the colon or cecum – much like horses and elephants.<ref name=":0" /> Unlike the other subfamily of Old World monkeys, the [[Cercopithecinae]], they do not possess cheek pouches.

In the [[silvery lutung]] (''Trachypithecus cristatus'') and the [[Northern plains gray langur]] (''Semnopithecus entellus'') females begin reproducing after around 3 years of age.<ref>{{Cite journal |last=Dirks |first=Wendy |last2=Bowman |first2=Jacqui E. |date=September 2007 |title=Life history theory and dental development in four species of catarrhine primates |url=https://linkinghub.elsevier.com/retrieve/pii/S0047248407001029 |journal=Journal of Human Evolution |language=en |volume=53 |issue=3 |pages=309–320 |doi=10.1016/j.jhevol.2007.04.007|url-access=subscription }}</ref>

== Classification and evolution ==
Colobinae is split into two tribes: [[Colobini]], found in Africa, and [[Presbytini]], found in Asia. Based on fossil records, the tribes split between 10 and 13 million years ago.<ref name="XiaoPing2013">{{cite journal |last1=XiaoPing |first1=Wang |title= Summary of phylogeny in subfamily Colobinae (Primate: Cercopithecidae) |journal= [[Chinese Science Bulletin]] |date=2013 |volume=58 |issue=18 |pages=2097–2103 | doi = 10.1007/s11434-012-5624-y|bibcode=2013ChSBu..58.2097W |doi-access=free }}</ref><ref>{{cite journal |title=A Molecular Phylogeny of Living Primates |first1=Polina |last1=Perelman |first2=Warren E. |last2=Johnson |first3=Christian |last3=Roos |first4=Hector N. |last4=Seuánez |first5=Julie E. |last5=Horvath |first6=Miguel A. M. |last6=Moreira |first7=Bailey |last7=Kessing |first8=Joan |last8=Pontius |first9=Melody |last9=Roelke |first10=Yves |last10=Rumpler |first11=Maria Paula C. |last11=Schneider |first12=Artur |last12=Silva |first13=Stephen J. |last13=O'Brien |first14=Jill |last14=Pecon-Slattery |journal=PLOS Genetics |display-authors=6 |date=17 March 2011 |volume=7 |issue=3 |pages=e1001342 |doi=10.1371/journal.pgen.1001342|pmid=21436896 |pmc=3060065 |doi-access=free }}</ref> The Colobini tribe contains three genera, black-and-white colobuses, red colobuses, and the olive colobus, all of whom are found in Africa. The Asian Presbytini comprises seven genera split into two clades, the odd-nosed group and the langur group.<ref name="XiaoPing2013" /><ref name=Sterner2006>{{cite journal | author1 = Sterner, Kirstin N. | author2 = Raaum, Ryan L. | author3 = Zhang, Ya-Ping | author4 = Stewart, Caro-Beth | author5 = Disotell, Todd R. | year = 2006 | title = Mitochondrial data support an odd-nosed colobine clade | journal = Molecular Phylogenetics and Evolution | volume = 40 | issue = 1 | pages = 1–7 | doi = 10.1016/j.ympev.2006.01.017 | url = http://www.raaum.org/uploads/1/1/7/5/1175545/sterner2006p1532.pdf | pmid = 16500120 | bibcode = 2006MolPE..40....1S | access-date = 2021-06-19 }}</ref> The discordant gene tree topologies and divergence age estimates suggest that hybridization, particularly involving female introgression from ''Piliocolobus''/''Procolobus'' into ''Colobus'' and male introgression from ''Semnopithecus'' into ''Trachypithecus'', played a prominent role in shaping the phylogenetic relationships of African and Asian colobine monkeys during their evolutionary history.<ref name=Osterholz2008/>

The earliest remains of Colobinae are known from the Tugen Hills of Kenya, dating to 12.5 million years ago. The earliest fossils of the genus in Eurasia are those of ''[[Mesopithecus]]'' found in Greece, dating to around 8.2 million years ago.<ref>{{Citation |last1=Frost |first1=Stephen R. |title=The Colobine Fossil Record |date=2022-02-28 |work=The Colobines |pages=13–31 |editor-last=Matsuda |editor-first=Ikki |url=https://www.cambridge.org/core/product/identifier/9781108347150%23CN-bp-3/type/book_part |access-date=2024-08-25 |edition=1 |publisher=Cambridge University Press |doi=10.1017/9781108347150.004 |isbn=978-1-108-34715-0 |last2=Gilbert |first2=Christopher C. |last3=Nakatsukasa |first3=Masato |editor2-last=Grueter |editor2-first=Cyril C. |editor3-last=Teichroeb |editor3-first=Julie A.|url-access=subscription }}</ref>

* Family [[Old World monkey|Cercopithecidae]]<ref name=msw3/><ref name=Sterner2006 />
** Subfamily [[Cercopithecinae]]
** '''Subfamily Colobinae'''
*** Tribe [[Colobini]]
**** Genus ''[[Black-and-white colobus|Colobus]]'' - black-and-white colobus monkeys
**** Genus ''[[Red colobus|Piliocolobus]]'' - red colobus monkeys
**** Genus ''[[Olive colobus|Procolobus]]'' - olive colobus
**** {{extinct}}Genus ''[[Cercopithecoides]]''
*** Tribe [[Presbytini]]
**** Langur (leaf monkey) group
***** Genus ''[[Trachypithecus]]'' - lutungs
***** Genus ''[[Surili|Presbytis]]'' - surilis
***** Genus ''[[Gray langur|Semnopithecus]]'' - gray langurs
**** Odd-nosed group
***** Genus ''[[Douc|Pygathrix]]'' - doucs
***** Genus ''[[Snub-nosed monkey|Rhinopithecus]]'' - snub-nosed monkeys
***** Genus ''[[Proboscis monkey|Nasalis]]'' - proboscis monkey
***** Genus ''[[Pig-tailed langur|Simias]]'' - pig-tailed langur
***** {{extinct}}Genus ''[[Mesopithecus]]''

===Hybrids===
Intergeneric [[Hybrid (biology)|hybrids]] are known to occur within the subfamily Colobinae. In India, [[gray langur]]s (''Semnopithecus'' spp.) are known to hybridize with [[Nilgiri langur]]s (''Trachypithecus johnii'').<ref name=Rowe1996>{{cite book|title=The Pictorial Guide to the Living Primates|url=https://archive.org/details/pictorialguideto0000rowe|url-access=registration|author=Rowe, N.|year=1996|publisher=Pogonias Press|isbn=978-0-9648825-0-8|pages=[https://archive.org/details/pictorialguideto0000rowe/page/139 139], 143, 154, 185, 223}}</ref>

==References==
{{Commons category|Colobinae}}
{{Wikispecies|Colobinae}}
{{Reflist}}
Roos, Christian, et al. “Nuclear Versus Mitochondrial DNA: Evidence for Hybridization in Colobine Monkeys.” BMC Evolutionary Biology, vol. 11, no. 1, 2011, pp. 77–77, https://doi.org/10.1186/1471-2148-11-77.
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{{Haplorhini|C.}}
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[[Category:Colobinae| ]]
[[Category:Mammal subfamilies]]
[[Category:Taxa named by Thomas C. Jerdon]]
[[Category:Taxa described in 1867]]