Colugo

{{Short description|Family of mammals}} {{redirect-distinguish|Cynocephalidae|Cybocephalidae}} {{redirect-distinguish|Dermoptera|Dermaptera}} {{use dmy dates|date=May 2026}} {{Automatic taxobox | name = Colugos<ref name="msw3">{{MSW3 Stafford|pages=110|id=12000002}}</ref> | fossil_range = <br/>Eocene-Present, {{Fossil range|37|0}} | image = Colugo (Galeopterus variegatus, adult female), Central Catchment Area, Singapore - 20060618.jpg | image_caption = Sunda flying lemur | display_parents = 4 | parent_authority = Illiger, 1811 | taxon = Cynocephalidae | authority = Simpson, 1945 | type_genus = ''Cynocephalus'' | type_genus_authority = Boddaert, 1768 | subdivision_ranks = Genera | subdivision = : {{leftlegend|#00FF00|''Cynocephalus''|outline=gray}} : {{leftlegend|#ff3131|''Galeopterus''|outline=gray}} : {{spaces}}† {{spaces}}{{font|size=90%|''Dermotherium''}} | range_map = DistributionDermopteraCorrecting.png | synonyms = * Colugidae * Galeopithecidae * Galeopteridae * Galeopithidae }}

'''Colugos''' ({{IPAc-en|k|ə|ˈ|l|uː|ɡ|oʊ}}<ref>{{Cite dictionary |title=Colugo |dictionary=Lexico UK English Dictionary |publisher=Oxford University Press |url=http://www.lexico.com/definition/Colugo |archive-url=https://web.archive.org/web/20210214184126/https://www.lexico.com/definition/colugo |url-status=dead |archive-date=14 February 2021 }}</ref><ref>{{MerriamWebsterDictionary |Colugo |access-date=2016-01-21}}</ref>), '''flying lemurs''', or '''cobegos''' ({{IPAc-en|k|ə|ˈ|b|iː|ɡ|oʊ}}{{refn|{{MerriamWebsterDictionary |access-date=2024-11-06 |Cobego}}}}), are arboreal gliding primatomorphs that are native to Southeast Asia. Their evolutionary closest relatives are primates. There are just two living species of colugos: the Sunda flying lemur (''Galeopterus variegatus'') and the Philippine flying lemur (''Cynocephalus volans''). These two species make up the entire family '''Cynocephalidae''' ({{IPAc-en|ˌ|s|aɪ|n|oʊ-|ˌ|s|ɛ|f|ə|ˈ|l|aɪ|d|i|,_|-|ˌ|k|ɛ|-}}){{efn|Cf. words with analogous pronunciations{{or|date=November 2024}} such as Meningoencephalitis, see {{Cite dictionary |title=Meningoencephalitis |dictionary=Lexico UK English Dictionary UK English Dictionary |publisher=Oxford University Press |url=http://www.lexico.com/definition/Meningoencephalitis |archive-url=https://web.archive.org/web/20200322185903/https://www.lexico.com/definition/meningoencephalitis |url-status=dead |archive-date=March 22, 2020}}}} and order '''Dermoptera''',<ref name="msw3"/><ref name="TNHC"/> from Ancient Greek {{math|δέρμα}} (''dérma''), meaning "skin", and {{math|πτερόν}} (''pterón''), meaning "wing".<ref>{{cite book |last1=Klein |first1=Jan |last2=Takahata |first2=Naoyuki |year=2013 |title=Where Do We Come From?: The molecular evidence for human descent |page=182 |publisher=Springer |place=Berlin, DE / Heidelberg, DE |isbn=978-3-66204-847-4 |url=https://books.google.com/books?id=0FvxCAAAQBAJ&pg=PA182 |access-date=16 December 2024 }}</ref>

== Characteristics == Colugos are nocturnal, tree-dwelling, gliding mammals.

=== Appearance and anatomy === They reach lengths of {{cvt|35|to|40|cm}} and weigh {{cvt|1|to|2|kg}}.<ref>{{cite book |last1=Lim |first1=Norman |title=Colugo: The flying lemur of South-East Asia |date=2007 |publisher=Draco Publishing and Distribution Pte Ltd |location=Singapore}}</ref> They have long, slender front and rear limbs, a medium-length tail, and a relatively light build. The head is small, with large, front-focused eyes for excellent binocular vision, and small rounded ears.

The incisor teeth of colugos are highly distinctive; they are comb-like in shape with up to 20&nbsp;tines on each tooth. The incisors are analogous in appearance and function to the incisor suite in strepsirrhines, which is used for grooming. The second upper incisors have two roots, another unique feature among mammals.<ref name=EoM/> The dental formula of colugos is: {{DentalFormula|upper=2.1.2.3|lower=3.1.2.3}} thumbnail|right|Feet of Philippine colugo

=== Movement === Colugos are proficient gliders, and thought better adapted for flight than any other gliding mammal. They can travel as far as {{cvt|70|m}} from one tree to another without losing much altitude,<ref>{{cite book |last=Dawkins |first=Richard |author-link=Richard Dawkins |year=2004 |title=The Ancestor's Tale |publisher=Phoenix |isbn=978-0-7538-1996-8 |url-access=registration |url=https://archive.org/details/ancestorstalepil0000dawk }}</ref> with a Malayan colugo (''Galeopterus variegatus'') individual having been observed traveling about {{cvt|150|m}} in one glide.<ref name="pmid18252673">{{cite journal |last1=Byrnes |first1=G. |last2=Lim |first2=N.T. |last3=Spence |first3=A.J. |title=Take-off and landing kinetics of a free-ranging gliding mammal, the Malayan colugo (''Galeopterus variegatus'') |year=2008 |journal=Proceedings of the Royal Society B: Biological Sciences |volume=275 |issue=1638 |pages=1007–1013 |pmid=18252673 |doi=10.1098/rspb.2007.1684 |pmc=2600906 }}</ref>

Their ability to glide is possible because of a large membrane of skin that extends between their paired limbs. This gliding membrane, or patagium, runs from the shoulder blades to the fore paws, from the tip of the rear-most fingers to the tip of the toes, and from the hind legs to the tip of the tail.<ref name=EoM>{{cite book |editor-last=Macdonald |editor-first=D. |last=MacKinnon |first=Kathy |year=1984 |title=The Encyclopedia of Mammals |publisher=Facts on File |location=New York, NY |isbn=978-0-87196-871-5 |pages=[https://archive.org/details/encyclopediaofma00mals_0/page/446 446–447] |url-access=registration |url=https://archive.org/details/encyclopediaofma00mals_0/page/446 }}</ref> Colugos' gliding ability is aided by specialized muscles that control the tension and movement of the gliding membrane. The platysma myoides and platysma cervicale help adjust the patagium’s shape for better control during flight. The sternomastoideus and cleidomastoideus stabilize the head and neck, improving balance and maneuverability mid-air.<ref>{{Cite journal |last=Diogo |first=Rui |year=2009 |title=The head and neck muscles of the Philippine colugo (Dermoptera: ''Cynocephalus volans''), with a comparison to tree-shrews, primates, and other mammals |journal=Journal of Morphology |lang=en |volume=270 |issue=1 |pages=14–51 |doi=10.1002/jmor.10666 |pmid=18798250 |issn=1097-4687 |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.10666 |url-access=subscription }}</ref> The spaces between the colugo's fingers and toes are webbed. As a result, colugos were once considered to be close relatives of bats. Today, on account of genetic data, they are considered to be more closely related to primates.<ref name="Janeckaetal2007">{{cite journal |first1=Jan E. |last1=Janecka |first2=Webb |last2=Miller |first3=Thomas H. |last3=Pringle |first4=Frank |last4=Wiens |first5=Annette |last5=Zitzmann |first6=Kristofer M. |last6=Helgen |first7=Mark S. |last7=Springer |first8=William J. |last8=Murphy |display-authors=6 |year=2007 |title=Molecular and genomic data identify that their closest living relative non-colugo relatives are primates |journal=Science |volume=318 |issue=5851 |pages=792–794 |bibcode=2007Sci...318..792J |doi=10.1126/science.1147555 |pmid=17975064 |s2cid=12251814 }}</ref>

[[File:Dermoptera - toothcomb 01.jpg|150px|thumbnail|right|Lower jaw (''Galeopterus'')]] Colugos are unskilled climbers; they lack opposable thumbs.<ref name="Piotrowski 2015">{{cite web |last=Piotrowski |first=Jan |date=2015-05-15 |series=Zoologger |title=The clumsy tree-dweller transforms into a gliding ace |website=New Scientist |url=https://www.newscientist.com/article/dn27539-zoologger-the-clumsy-tree-dweller-transforms-into-a-gliding-ace/ |url-status=live |access-date=2021-11-23 |archive-url=https://web.archive.org/web/20211123163640/https://www.newscientist.com/article/dn27539-zoologger-the-clumsy-tree-dweller-transforms-into-a-gliding-ace/ |archive-date=2021-11-23 }}</ref> They progress up trees in a series of slow hops, gripping onto the bark with their small, sharp claws. They spend most of the day resting. At night, colugos spend most of their time up in the trees foraging, with gliding being used to either find another foraging tree or to find possible mates and protect territory.<ref>{{cite journal |last1=Spence |first1=Andrew J. |last2=Yeong |first2=Charlene |last3=Lim |first3=Norman T.-L. |last4=Byrnes |first4=Greg |date=2011-04-15 |title=Sex differences in the locomotor ecology of a gliding mammal, the Malayan colugo (''Galeopterus variegatus'') |journal=Journal of Mammalogy |lang=en |volume=92 |issue=2 |pages=444–451 |doi=10.1644/10-MAMM-A-048.1 |s2cid=84539846 |issn=0022-2372 |url=https://academic.oup.com/jmammal/article/92/2/444/869618 |url-status=live |access-date=2018-12-05 |archive-url=https://web.archive.org/web/20181205130744/https://academic.oup.com/jmammal/article/92/2/444/869618 |archive-date=2018-12-05 }}</ref>

=== Behavior and diet === Colugos are shy, nocturnal, solitary animals found in the tropical forests of Southeast Asia. Consequently, very little is known about their behavior. They are herbivorous and eat leaves, shoots, flowers, sap, and fruit. They have well-developed stomachs and long intestines capable of extracting nutrients from leaves and other fibrous material.

As part of colugos' evolution into nocturnal species, they developed night vision.<ref>{{cite journal |last1=Moritz |first1=G.L. |last2=Lim |first2=N.T. |last3=Neitz |first3=M. |last4=Peichl |first4=L. |last5=Dominy |first5=N.J. |year=2013 |title=Expression and evolution of short wavelength sensitive opsins in colugos: A nocturnal lineage that informs debate on primate origins |journal=Evolutionary Biology |volume=40 |issue=4 |pages=542–553 |doi=10.1007/s11692-013-9230-y |pmid=24293738 |pmc=3832777 |url=https://doi.org/10.1007/s11692-013-9230-y |archive-url=https://web.archive.org/web/20231116074310/https://link.springer.com/article/10.1007/s11692-013-9230-y |archive-date=2023-11-16 }}</ref> Colugos spend their days resting in tree holes and are active at night time; traveling around {{Convert|1.7|km|mi|1|abbr=off}} at night. Colugos may also be a territorial species.<ref>{{cite web |title=Flying lemur |date=25 May 2021 |website=A-Z Animals |url=https://a-z-animals.com/animals/flying-lemur/ |url-status=live |archive-url=https://web.archive.org/web/20231224100805/https://a-z-animals.com/animals/flying-lemur/ |archive-date=24 December 2023 }}</ref>

=== Life cycle === Although they are placentals, colugos raise their young in a manner similar to marsupials. Newborn colugos are underdeveloped and weigh only {{cvt|35|g}}.<ref>{{cite book |editor-last=Macdonald |editor-first=David W. |title=The Encyclopedia of Mammals |publisher=Oxford University Press |year=2006 |isbn=978-0-19-920608-7}}</ref> They spend the first six months of life clinging to their mother's belly. The mother colugo curls her tail and folds her patagium into a warm, secure, quasi-pouch to protect and transport her young. The young do not reach maturity until they are two to three years old.<ref name=EoM/> In captivity, they live up to 15 years, but their lifespan in the wild is unknown.<ref>{{cite web |url=https://www.exploringnature.org/db/view/Colugo |title=Colugo |last=Amsel |first=Sheri |website=Exploring Nature Educational Resource |access-date=2018-12-23 |archive-date=2018-12-23 |archive-url=https://web.archive.org/web/20181223121153/https://www.exploringnature.org/db/view/Colugo |url-status=live }}</ref>

== Status == Both species are threatened by habitat destruction, and the Philippine flying lemur was once classified by the IUCN as vulnerable. In 1996, the IUCN declared the species vulnerable owing to destruction of lowland forests and hunting. It was downlisted to least-concern status in 2008 but still faces the same threats. In addition to the ongoing clearing of its rainforest habitat, it is hunted for its meat and fur. It is also a favorite prey item for the critically endangered Philippine eagle; some studies suggest colugos account for 90% of the eagle's diet.<ref name=Chandler>{{Cite book |last1 =Chandler |first1 = David |last2 = Couzens |first2 = Dominic |year = 2008 |title = 100&nbsp;Birds to See Before You Die |place = London, UK |publisher = Carleton Books |isbn = 978-1-84442-019-3 |page = 171 }}</ref>

== Taxonomy == Their family name Cynocephalidae comes from the Ancient Greek κύων (''kúon''), meaning "dog", and κεφαλή (''kephalḗ''), meaning "head", because their heads are broad with short snouts like dogs.<ref name="TNHC">{{cite web |last=Baines |first=Elizabeth<!-- fr Acknowledgements --> |year=2001<!-- fr Acknowledgements --> |title=Colugos or 'flying lemurs' |website=The Natural History Collections |publisher=University of Edinburgh |url=http://www.nhc.ed.ac.uk/index.php?page=493.494 |url-status=live |access-date=30 August 2023 |archive-url=https://web.archive.org/web/20230830142003/http://www.nhc.ed.ac.uk/index.php?page=493.494 |archive-date=30 August 2023 }}</ref>

==Summary of extant species== {| class="wikitable sortable" |- ! width="15%"|Common name ! width="15%"|Binomial name ! data-sort-type="number" width="15%"|Population ! width="5%"|Status ! width="5%"|Trend ! class="unsortable"|Notes ! width="15%" class="unsortable"|Image |- |'''Sunda flying lemur''' |''Galeopterus variegatus'' |unknown<ref name="G.variegatus IUCN">{{Cite IUCN|title=''Galeopterus variegatus''|article-number=e.T41502A10479343|last=Boeadi|year=2008|doi=10.2305/IUCN.UK.2008.RLTS.T41502A10479343.en|last2=Steinmetz, R.}}</ref> |{{IUCN-LC}}<ref name="G.variegatus IUCN"/> |align="center"|{{decrease}}<ref name="G.variegatus IUCN"/> | |175px |- |'''Philippine flying lemur''' |''Cynocephalus volans'' |unknown<ref name="C.volans IUCN">{{cite iucn |author1=Gonzalez, J.C. |author2=Custodio, C. |author3=Carino, P. |author4=Pamaong-Jose, R. |year=2008 |title=''Cynocephalus volans'' |volume=2008 |article-number=e.T6081A12410826 |doi=10.2305/IUCN.UK.2008.RLTS.T6081A12410826.en |language=en |access-date=3 May 2026}}</ref> |{{IUCN-LC}}<ref name="C.volans IUCN"/> |align="center"|{{steady}}<ref name="C.volans IUCN"/> | |175px |}

== Classification and evolution == The ancestors of the colugos are estimated to split from other mammals about 80 million years ago, leading to the present-day forms, which may include up to 14 extant species.<ref>[https://phys.org/news/2016-08-dna-analysis-colugos-primate-sister.html DNA analysis suggests colugos are part of a primate sister group]</ref> Mixodectidae and Plagiomenidae are two extinct families of North American mammals which may represent fossil dermopterans or at least close relatives.<ref name="Rose-2006">{{cite book |last=Rose |first=Kenneth D. |year=2006 |title=The Beginning of the Age of Mammals |publisher=Johns Hopkins University Press |isbn=978-0-8018-8472-6 |pages=164–165|lang=en |doi=10.56021/9780801884726 |url=https://books.google.com/books?id=3bs0D5ix4VAC&pg=PA164 }}</ref> However, some studies doubt a close relationship between these families and modern dermopterans (Cynocephalidae).<ref name="MacPhee-1989">{{cite journal |last1=MacPhee |first1=R.D.E. |last2=Cartmill |first2=M. |last3=Rose |first3=K.D. |date=1989-09-28 |title=Craniodental morphology and relationships of the supposed Eocene dermopteran Plagiomene (Mammalia) |journal=Journal of Vertebrate Paleontology |lang=en |volume=9 |issue=3 |pages=329–349 |doi=10.1080/02724634.1989.10011766 |issn=0272-4634 |jstor=4523269 |url=http://www.tandfonline.com/doi/abs/10.1080/02724634.1989.10011766 }}</ref><ref name="Chester-2025">{{Cite journal |last1=Chester |first1=Stephen G. B. |last2=Williamson |first2=Thomas E. |last3=Crowell |first3=Jordan W. |last4=Silcox |first4=Mary T. |last5=Bloch |first5=Jonathan I. |last6=Sargis |first6=Eric J. |date=2025-03-11 |title=New remarkably complete skeleton of Mixodectes reveals arboreality in a large Paleocene primatomorphan mammal following the Cretaceous-Paleogene mass extinction |journal=Scientific Reports |lang=en |volume=15 |issue=1 |pages=8041 |doi=10.1038/s41598-025-90203-z |pmid=40069232 |pmc=11897203 |issn=2045-2322 }}</ref> A complete skeleton of ''Mixodectes'' shows that, though it was well-adapted for an arboreal lifestyle, it had no adaptations for gliding.<ref name="Chester-2025" /> Although other Paleogene mammals have been interpreted as related to dermopterans, the evidence for this association is uncertain and many of the fossils are no longer interpreted as being gliding mammals.<ref>{{cite conference |title=The first dentally associated skeleton of Plagiomenidae (Mammalia, ?Dermoptera) from the late Paleocene of Wyoming |date=November 2011 |doi=10.13140/2.1.1302.4322 |conference=Society of Vertebrate Paleontology 71st Annual Meeting |location=Las Vegas, NV }}</ref> At present, the fossil record of definitive dermopterans is limited to two species of the Eocene and Oligocene cynocephalid genus ''Dermotherium''.<ref name=Marivauxetal06>{{cite journal |first1=L. |last1=Marivaux |first2=L. |last2=Bocat |first3=Y. |last3=Chaimanee |first4=J.-J. |last4=Jaeger |first5=B. |last5=Marandat |first6=P. |last6=Srisuk |first7=P. |last7=Tafforeau |first8=C. |last8=Yamee |first9=J.-L. |last9=Welcomme |display-authors=6 |name-list-style=amp |year=2006 |title=Cynocephalid dermopterans from the Palaeogene of South Asia (Thailand, Myanmar and Pakistan): Systematic, evolutionary and palaeobiogeographic implications |journal=Zoologica Scripta |volume=35 |issue=4 |pages=395–420 |doi=10.1111/j.1463-6409.2006.00235.x |s2cid=86572864}}</ref>

Molecular phylogenetic studies have demonstrated that colugos emerged as a basal Primatomorpha clade – which, in turn, is a basal Euarchontoglires clade. Scandentia are widely considered to be the closest relatives of Primatomorpha, within Euarchonta. Some studies, however, place Scandentia as sister of Glires (lagomorphs and rodents), in an unnamed sister clade of the Primatomorpha.<ref>{{cite journal |last1=Meredith |first1=Robert W. |last2=Janečka |first2=Jan E. |last3=Gatesy |first3=John |last4=Ryder |first4=Oliver A. |last5=Fisher |first5=Colleen A. |last6=Teeling |first6=Emma C. |last7=Goodbla |first7=Alisha |last8=Eizirik |first8=Eduardo |last9=Simão |first9=Taiz L.L. |date=2011-10-28 |title=Impacts of the cretaceous terrestrial revolution and KPg extinction on mammal diversification |journal=Science |volume=334 |issue=6055 |pages=521–524 |doi=10.1126/science.1211028 |issn=0036-8075 |pmid=21940861 |bibcode=2011Sci...334..521M |s2cid=38120449 |url=https://zotero.org/groups/5435545/items/UZ7N5QRK }}</ref><ref>{{Cite journal |last1=Zhou |first1=Xuming |last2=Sun |first2=Fengming |last3=Xu |first3=Shixia |last4=Yang |first4=Guang |last5=Li |first5=Ming |date=2015-03-01 |title=The position of tree shrews in the mammalian tree: Comparing multi-gene analyses with phylogenomic results leaves monophyly of Euarchonta doubtful |journal=Integrative Zoology |lang=en |volume=10 |issue=2 |pages=186–198 |doi=10.1111/1749-4877.12116 |issn=1749-4877 |pmid=25311886 }}</ref>

{{Clade | label1=Euarchontoglires | 1={{Clade |1={{clade | 1=Scandentia (treeshrews) | label2=Glires | 2={{Clade | 1=Lagomorpha (rabbits, hares, pikas) | 2=Rodentia (rodents) }} }}

| label2=Primatomorpha | 2={{Clade | 1='''Dermoptera''' (colugos) | 2={{Clade | 1=†Plesiadapiformes |state1=double | 2=Primates}} }} }} }}

* '''Order Dermoptera''' ** †'''Family Plagiomenidae'''? ** †'''Family Mixodectidae'''? ** '''Family Cynocephalidae''' *** ''Cynocephalus'' **** Philippine flying lemur, ''Cynocephalus volans'' *** ''Galeopterus'' **** Sunda flying lemur, ''Galeopterus variegatus'' *** †''Dermotherium'' **** †''Dermotherium major'' **** †''Dermotherium chimaera''

== Gallery == <gallery> File:Colugo's claw.jpg|Claw of Sunda flying lemur File:Colugo (Cynocephalus variegatus) " red " morph (8064072543).jpg|A "red" morph of a colugo (''Cynocephalus variegatus'') File:Cynocephalus volans 58339590.jpg|Black morph of a Sunda flying colugo (''Cynocephalus variegatus'') File:Sunda Flying Lemur.jpg|Colugo in a tree (''Cynocephalus variegatus'') </gallery>

== Notes == {{notelist|128em}}

== References == {{reflist|25em}}

== External links == {{EB1911 poster|Galeopithecus}} {{Commons}}

* {{cite press release |first=Barbara K. |last=Kennedy |date=2007-11-01 |title=Flying lemurs are the closest relatives of primates |website=Eureka Alert |publisher=Penn State |url=https://www.eurekalert.org/news-releases/688278 }}

{{Mammals}} {{Colugo nav}} {{Euarchontoglires|E.}} {{Taxonbar|from=Q183383|from2=Q11762767}} {{Authority control}}

Category:Colugos Category:Priabonian first appearances Category:Extant Eocene first appearances