Editing Tarsier

{{Short description|Family of 'dry nosed' primates}}
{{Cleanup rewrite|date=November 2024}}
{{Use dmy dates|date=February 2025}}
{{Automatic taxobox
| name                 = Tarsiers<ref name=MSW3>{{MSW3 Groves|pages=127–128}}</ref><ref name=2010_Groves_Shekelle/>
| fossil_range         = {{Fossil range|45|0}}Middle [[Eocene]] to recent
| image                = Tarsier-GG.jpg
| image_caption        = [[Philippine tarsier]] (''Carlito syrichta'')
| taxon                = Tarsiidae 
| authority            = [[John Edward Gray|Gray]], 1825
| type_genus           = ''[[Tarsius]]''
| type_genus_authority = [[Gottlieb Conrad Christian Storr|Storr]], 1780
| subdivision_ranks    = [[Genus|Genera]]
| subdivision          =
* ''[[Carlito (genus)|Carlito]]''
* ''[[Cephalopachus]]''
* ''[[Tarsius]]''
}}

'''Tarsiers''' ({{IPAc-en|ˈ|t|ɑr|s|i|ər|z}} {{respell|TAR|see|ərz}}) are [[Haplorhini|haplorhine]] [[primate]]s of the [[family (biology)|family]] '''Tarsiidae''', which is the lone extant family within the infraorder [[Tarsiiformes]]. Although the group was prehistorically more globally widespread, all of the existing [[species]] are restricted to [[Maritime Southeast Asia]], predominantly in [[Brunei]], [[Indonesia]], [[Malaysia]] and the [[Philippines]].<ref>The Editors of Encyclopædia Britannica. “Tarsier.” ''Encyclopædia Britannica'', Encyclopædia Britannica, Inc., 14 April 2019, http://www.britannica.com/animal/tarsier.</ref>[[File:Tarsier (Inside Philippine National Museum of Natural History).jpg|thumb|Tarsier image inside [[National Museum of the Philippines|Philippine National Museum]] of Natural History]]They are found primarily in forested habitats, especially forests that have [[liana]], since the vine gives tarsiers vertical support when climbing trees.<ref>{{cite journal |last1=Simeon |first1=S.G.F. |last2=Duya |first2=M.R.M |last3=Duya |first3=M.V. |last4=Galindon |first4=J.M.M. |last5=Pasion |first5=B.O. |last6=Ong |first6=P.S. |title=Living in small spaces: Forest fragment characterization and its use by Philippine tarsiers (Tarsius syrichta Linnaeus, 1758) in Mindanao Island, Philippines |journal=Primates |date=2020 |volume=61 |issue=3 |pages=529–542 |doi=10.1007/s10329-020-00798-2|pmid=32043166 |s2cid=211075031 }}</ref>

==Evolutionary history==
===Fossil record===
Fossils of [[tarsiiform]] primates have been found in Asia, Europe, and North America (with disputed fossils from Northern Africa), but extant tarsiers are restricted to several Southeast Asian islands. The fossil record indicates that their [[dentition]] has not changed much, except in size, over the past 45 million years.

Within the family Tarsiidae, there are two extinct genera—''[[Xanthorhysis]]'' and ''[[Afrotarsius]]''; however, the placement of ''Afrotarsius'' is not certain,<ref name=Hartwig1>{{cite book | last1 = Gunnell | first1 = G. | last2 = Rose | first2 = K. | editor1-last = Hartwig | editor1-first = W.C. | title = The Primate Fossil Record | publisher = Cambridge University Press | year = 2002 | isbn = 978-0-521-66315-1 | chapter = Tarsiiformes: Evolutionary History and Adaptation}}</ref> and it is sometimes listed in its own family, [[Afrotarsiidae]], within the infraorder Tarsiiformes,<ref name=McKenna_Bell>McKenna, M.C., and Bell, S.K. 1997. ''Classification of Mammals Above the Species Level.'' Columbia University Press, New York, 337–340 pp. {{ISBN|0-231-11013-8}}</ref> or considered an [[Simian|anthropoid]] primate.<ref name=sirindhornae/>

So far, four fossil species of tarsiers are known from the fossil record:
* ''[[Tarsius eocaenus]]'' is known from the [[Middle Eocene]] in [[China]].<ref name=Hartwig1/><ref name=2006Rossie>{{cite journal | last1 = Rossie | first1 = J.B. | last2 = Ni | first2 = X. | last3 = Beard | first3 = K.C. | title = Cranial remains of an Eocene tarsier | year = 2006 | url = http://www.pnas.org/content/103/12/4381.full.pdf | journal = PNAS | volume =  103| issue = 12 | pages = 4381–4385 | pmid = 16537385|doi=10.1073/pnas.0509424103 | pmc=1450180| doi-access = free }}</ref>
* ''[[Hesperotarsius|Hesperotarsius thailandicus]]'' lived during the [[Early Miocene]] in northwestern [[Thailand]].<ref name=Hartwig1/><ref name=Nowak>{{cite book | last = Nowak | first = R.M. | publisher = Johns Hopkins University Press | title = Walker's Mammals of the World | url = https://archive.org/details/walkersmammalsof0002nowa | url-access = registration | edition = 6th | year = 1999 | isbn = 978-0-8018-5789-8 | pages = 94–97}}</ref>
* ''[[Hesperotarsius|Hesperotarsius sindhensis]]'' lived during the [[Miocene]] in [[Pakistan]].<ref>{{cite journal | last1 = Zijlstra | first1 = Jelle S. | last2 = Flynn | first2 = Lawrence J. | last3 = Wessels | first3 = Wilma | year = 2013 | title = The westernmost tarsier: A new genus and species from the Miocene of Pakistan | journal = Journal of Human Evolution | volume = 65 | issue = 5| pages = 544–550 | doi = 10.1016/j.jhevol.2013.06.015 | pmid=23928350| bibcode = 2013JHumE..65..544Z }}</ref>
* ''[[Tarsius sirindhornae]]'' lived during the [[Middle Miocene]] in northern Thailand.<ref name=sirindhornae>{{cite journal | author = Chiamanee, Y., Lebrun, R., Yamee, C., and Jaeger, J.-J. | year = 2010 | title = A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric–morphometric method | journal = Proceedings of the Royal Society B: Biological Sciences | volume = 278| issue = 1714| pages = 1956–1963| doi = 10.1098/rspb.2010.2062 | pmid=21123264 | pmc=3107645}}</ref>

The genus ''Tarsius'' has a longer fossil record than any other primate genus, but the assignment of the Eocene and Miocene fossils to the genus is dubious.<ref>{{cite book | last = Simons | first = E.L. | year = 2003 | title = Tarsiers: past, present, and future | editor1-last = Wright | editor1-first = P.C. | editor2-last = Simons | editor2-first = E.L. | editor3-last = Gursky | editor3-first = S. | chapter = The Fossil Record of Tarsier Evolution | publisher = Rutgers University Press | isbn = 978-0-8135-3236-3}}</ref>

===Classification===
{{see also|List of tarsiiformes}}

The [[phylogenetic]] position of extant tarsiers within the order Primates has been debated for much of the 20th century, and tarsiers have alternately been classified with [[strepsirrhine]] primates in the suborder Prosimii, or as the sister group to the [[simian]]s (Anthropoidea) in the infraorder [[Haplorrhini|Haplorhini]]. Analysis of [[Retrotransposon#SINEs|SINE]] insertions, a type of macromutation to the DNA, is argued to offer very persuasive evidence for the [[monophyly]] of Haplorhini, where other lines of evidence, such as [[DNA sequence]] data, remain ambiguous. Thus, some systematists argue the debate is conclusively settled in favor of a monophyletic Haplorrhini. In common with simians, tarsiers have a mutation in the [[L-gulonolactone oxidase]] (GULO) gene, which prevents their bodies from synthesizing vitamin C so they must find it in the diet. Since the strepsirrhines do not have this mutation and have retained the ability to make vitamin C, the genetic trait that confers the need for it in the diet would tend to place tarsiers with haplorhines.<ref>{{cite journal | title = Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of ''Tarsius'' | author1 = Pollock, J. I. | author2 = Mullin, R. J. | name-list-style = amp | journal = American Journal of Physical Anthropology | year = 1986 | volume = 73 | issue = 1 | pages = 65–70 | doi = 10.1002/ajpa.1330730106 | url = http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 | archive-url = https://archive.today/20120628232930/http://www3.interscience.wiley.com/journal/110488482/abstract?CRETRY=1&SRETRY=0 | url-status = dead | archive-date = 28 June 2012 | pmid = 3113259 | access-date = 16 March 2010 | url-access = subscription }}</ref>

[[File:Bohol Tarsier.jpg|thumb|right|[[Philippine tarsier]] (''Carlito syrichta''), one of the smallest primates]]

At a lower phylogenetic level, the tarsiers have, until recently, all been placed in the genus ''Tarsius'',<ref name=MSW3/> while it was debated whether the species should be placed in two (a [[Sulawesi]] and a [[Philippine]]-western group) or three separate genera (Sulawesi, Philippine and western groups).<ref name="Asian">{{cite journal |last=Brandon-Jones |first=D. |year=2004 |title=Asian primate classification |journal=International Journal of Primatology |volume=25 |issue=1 |pages=97–164 |doi=10.1023/B:IJOP.0000014647.18720.32 |s2cid=29045930 |display-authors=etal}}</ref> Species level [[Taxonomy (biology)|taxonomy]] is complex, with [[Morphology (biology)|morphology]] often being of limited use compared to vocalizations.{{Citation needed|date=March 2012}} Further confusion existed over the validity of certain names. Among others, the widely used ''T. dianae'' has been shown to be a [[junior synonym]] of ''T. dentatus'', and comparably, ''T. spectrum'' is now considered a junior synonym of ''T. tarsier''.<ref name=MSW3/>
<!--Anything to save in here? Text is uncited (although a ref is given for part). The recent taxonomy changes address these issues
Several "vocal morphs" may represent undescribed [[taxa]], taxonomically separable from ''T. tarsier'' (=''spectrum'') (such as [[Minahasa]] and a tarsier from the [[Togian Islands]]), and many others from Sulawesi and surrounding islands (Shekelle & Leksono 2004). This may also be the case for a number of poorly known, isolated Philippine populations (such as the [[Basilan]], [[Leyte (island)|Leyte]] and [[Dinagat Islands|Dinagat]] populations of the ''T. syrichta'' group).
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In 2010, [[Colin Groves]] and Myron Shekelle suggested splitting the genus ''Tarsius'' into three genera, the Philippine tarsiers (genus ''[[Carlito (genus)|Carlito]]''), the western tarsiers (genus ''[[Cephalopachus]]''), and the eastern tarsiers (genus ''Tarsius''). This was based on differences in [[dentition]], eye size, limb and hand length, tail tufts, tail sitting pads, the number of [[Mammary gland|mammae]], chromosome count, [[socioecology]], vocalizations, and distribution. The senior taxon of the species, ''T. tarsier'' was restricted to the population of a [[Selayar Islands|Selayar]] island, which then required the resurrection of the defunct taxon ''T. fuscus''.<ref name=2010_Groves_Shekelle>{{cite journal | last1 = Groves | first1 = C. | last2 = Shekelle | first2 = M. | title = The Genera and Species of Tarsiidae | journal = International Journal of Primatology | volume = 31 | issue = 6 | pages = 1071–1082 | year = 2010 | doi = 10.1007/s10764-010-9443-1| s2cid = 21220811 }}</ref>

In 2014, scientists published the results of a genetic study from across the range of the [[Philippine tarsier]], revealing previously unrecognised genetic diversity. Three subspecies are recognised in the established taxonomy: ''Carlito syrichta syrichta'' from [[Leyte]] and [[Samar]], ''C. syrichta fraterculus'' from [[Bohol]], and ''C. syrichta carbonarius'' from [[Mindanao]]. Their analysis of mitochondrial and nuclear DNA sequences suggested that ssp. ''syrichta'' and ''fraterculus'' may represent a single lineage, whereas ssp. ''carbonarius'' may represent two lineages – one occupies the majority of Mindanao while the other is in northeastern Mindanao and the nearby [[Dinagat Island]], which the authors termed the 'Dinagat-Caraga tarsier'. More detailed studies that integrate morphological data will be needed to review the taxonomy of tarsiers in the Philippines.<ref>{{Cite journal |last1=Brown |first1=Rafe M. |last2=Weghorst |first2=Jennifer A. |last3=Olson |first3=Karen V. |last4=Duya |first4=Mariano R. M. |last5=Barley |first5=Anthony J. |last6=Duya |first6=Melizar V. |last7=Shekelle |first7=Myron |last8=Neri-Arboleda |first8=Irene |last9=Esselstyn |first9=Jacob A. |last10=Dominy |first10=Nathaniel J. |last11=Ong |first11=Perry S. |last12=Moritz |first12=Gillian L. |last13=Luczon |first13=Adrian |last14=Diesmos |first14=Mae Lowe L. |last15=Diesmos |first15=Arvin C. |date=19 August 2014 |title=Conservation Genetics of the Philippine Tarsier: Cryptic Genetic Variation Restructures Conservation Priorities for an Island Archipelago Primate |journal=PLOS ONE |language=en |volume=9 |issue=8 |pages=e104340 |doi=10.1371/journal.pone.0104340 |doi-access=free |issn=1932-6203 |pmc=4138104 |pmid=25136854 |bibcode=2014PLoSO...9j4340B }}</ref>

* Infraorder Tarsiiformes
**'''Family Tarsiidae''': tarsiers
***'''Genus ''[[Carlito (genus)|Carlito]]'''''
**** [[Philippine tarsier]], ''Carlito syrichta''
***** ''C. s. syrichta''
***** ''C. s. fraterculus'' (to be combined into ''C. s. syrichta''?)
***** ''C. s. carbonarius''
*** '''Genus ''[[Cephalopachus]]'''''
**** [[Horsfield's tarsier]], ''Cephalopachus bancanus''
***** ''C. b. bancanus''
***** ''C. b. natunensis''
***** ''C. b. boreanus''
***** ''C. b. saltator''
*** '''Genus ''[[Tarsius]]'''''
****[[Dian's tarsier]], ''T. dentatus''
****[[Makassar tarsier]] ''T. fuscus'' 
****[[Lariang tarsier]], ''T. lariang''
****[[Niemitz's tarsier]], ''T. niemitzi''
****[[Peleng tarsier]], ''T. pelengensis''
****[[Sangihe tarsier]], ''T. sangirensis''
****[[Gursky's spectral tarsier]], ''T. spectrumgurskyae''<ref name="Shekelle">{{cite journal | title=Two New Tarsier Species (Tarsiidae, Primates) and the Biogeography of Sulawesi, Indonesia | first1=Myron |last1=Shekelle |first2=Colin P |last2=Groves |author-link2=Colin Groves |first3=Ibnu |last3=Maryanto |first4=Russell A |last4=Mittermeier |author-link4=Russell A Mittermeier |date=May 2017 |journal=Primate Conservation |volume=31 |issue=1 |pages=1–9 |url=https://www.researchgate.net/publication/316714884}}</ref>
****[[Jatna's tarsier]], ''T. supriatnai''<ref name=Shekelle/>
****[[Spectral tarsier]], ''T. tarsier''
****[[Siau Island tarsier]], ''T. tumpara''
****[[Pygmy tarsier]], ''T. pumilus''
****[[Wallace's tarsier]], ''T. wallacei''

==Anatomy and physiology==
[[File:Tarsius Syrichta-GG.jpg|thumb|right|Tarsiers tree-climbing]]

Tarsiers are small animals with enormous eyes; each eyeball is approximately {{convert|16|mm}} in diameter and is as large as, or in some cases larger than, its entire brain.<ref>{{cite book |last= Soluri |first= K. Elizabeth |author2= Sabrina C. Agarwal |title=The Laboratory Manual and Workbook for Biological Anthropology |publisher= W.W. Norton |year=2016 |isbn= 978-0-393-91291-3}}</ref><ref>{{cite book |last= Shumaker |first= Robert W. |author2=Benjamin B. Beck |title= Primates in Question |url= https://archive.org/details/primatesinquesti00shum |url-access= registration |publisher= Smithsonian Books |year= 2003 |isbn= 978-1-58834-151-8 }}</ref> The unique cranial anatomy of the tarsier results from the need to balance their large eyes and heavy head so they are able to wait silently for nutritious prey.<ref name="ShekelleMyronGursky2010">{{cite journal | last1 = Shekelle | first1 = Myron | last2 = Gursky | year = 2010 | title = Why tarsiers? Why now? An introduction to the special edition on tarsiers | journal = International Journal of Primatology | volume = 31 | issue = 6| pages = 937–940 | doi=10.1007/s10764-010-9459-6| s2cid = 326565 }}</ref> Tarsiers have a strong auditory sense, and their auditory cortex is distinct.<ref name="ShekelleMyronGursky2010" /> Tarsiers also have long hind limbs, owing mostly to the elongated [[tarsus (skeleton)|tarsus]] bones of the feet, from which the animals get their name. The combination of their elongated tarsi and fused tibiofibulae makes them morphologically specialized for vertical clinging and leaping.<ref>{{cite journal | last1 = Rasmussen | first1 = D. T. | last2 = Conroy | first2 = G. C. | last3 = Simons | first3 = E. L. | year = 1998 | title = Tarsier-like locomotor specializations in the Oligocene primate Afrotarsius | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 95 | issue = 25| pages = 14848–14850 | doi=10.1073/pnas.95.25.14848| pmid = 9843978 | pmc = 24538 | bibcode = 1998PNAS...9514848T | doi-access = free }}</ref> The head and body range from 10 to 15&nbsp;cm in length, but the hind limbs are about twice this long (including the feet), and they also have a slender tail from 20 to 25&nbsp;cm long. Their fingers are also elongated, with the third finger being about the same length as the upper arm. Most of the digits have nails, but the second and third toes of the hind feet bear claws instead, which are used for grooming. Tarsiers have soft, velvety fur, which is generally buff, beige, or ochre in color.<ref name=EoM>{{cite book|editor= Macdonald, D.|author= Niemitz, Carsten|author-link= Carsten Niemitz|year= 1984|title= The Encyclopedia of Mammals|publisher= Facts on File|location= New York|pages= [https://archive.org/details/encyclopediaofma00mals_0/page/338 338–339]|isbn= 978-0-87196-871-5|url-access= registration|url= https://archive.org/details/encyclopediaofma00mals_0/page/338}}</ref>

Tarsier morphology allows for them to move their heads 180 degrees in either direction, allowing for them to see 360 degrees around them.<ref name="RefA">Gron KJ. 2010 December 1. Primate Factsheets: Tarsier (Tarsius) Taxonomy, Morphology, & Ecology . http://pin.primate.wisc.edu/factsheets/entry/tarsier . Retrieved 12 November 2019.</ref> Their [[Dentition#Dental formula|dental formula]] is also unique: {{DentalFormula|upper=2.1.3.3|lower=1.1.3.3}}<ref>{{Cite book |last1=Simons |first1=Elwyn L. |url=https://books.google.com/books?id=9g_TwVzDOI0C |title=Tarsiers: Past, Present, and Future |last2=Wright |first2=Patricia C. |last3=Gursky |first3=Sharon |date=2003 |isbn=0-8135-3236-1 |pages=65 |publisher=Rutgers University Press |author-link=Elwyn L. Simons |author-link2=Patricia Wright (primatologist)}}</ref> Unlike many nocturnal vertebrates, tarsiers lack a light-reflecting layer ([[tapetum lucidum]]) of the retina and have a [[Fovea centralis|fovea]].

The tarsier's brain is different from that of other primates in terms of the arrangement of the connections between the two eyes and the [[lateral geniculate nucleus]], which is the main region of the [[thalamus]] that receives visual information. The sequence of cellular layers receiving information from the ipsilateral (same side of the head) and contralateral (opposite side of the head) eyes in the lateral geniculate nucleus distinguishes tarsiers from [[lemur]]s, [[lorises]], and [[monkey]]s, which are all similar in this respect.<ref name="Rosa1996">{{cite journal |last=Rosa |first=M. G. |author2=Pettigrew J. D. |author3=Cooper H. M. |year=1996 |title=Unusual pattern of retinogeniculate projections in the controversial primate ''Tarsius'' |journal=Brain, Behavior and Evolution |volume=48 |issue=3 |pages=121–129 |doi=10.1159/000113191 |pmid=8872317}}</ref> Some neuroscientists suggested that "this apparent difference distinguishes tarsiers from all other primates, reinforcing the view that they arose in an early, independent line of primate evolution."<ref name="Collins2005">{{cite journal |last=Collins |first=C. E. |author2=Hendrickson, A. |author3=Kaas, J. H. |year=2005 |title=Overview of the visual system of ''tarsius'' |journal=The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology|volume=287 |issue=1 |pages=1013–1025 |doi=10.1002/ar.a.20263 |pmid=16200648 |doi-access= |s2cid=21448186 }}</ref>

[[Philippine tarsier]]s are capable of hearing frequencies as high as 91&nbsp;kHz. They are also capable of vocalizations with a dominant frequency of 70&nbsp;kHz.<ref name="Ramsier2012">{{cite journal |last=Ramsier |first=Marissa A. |author2=Cunningham A.J. |author3=Moritz G.L. |author4=Finneran J.J. |author5=Williams C.V. |author6=Ong P.S. |author7=Gursky-Doyen S.L. |author8= Dominy N.J. |year=2012 |title=Primate communication in the pure ultrasound |journal=Biology Letters |doi=10.1098/rsbl.2011.1149 |pmid=22319094 |volume=8 |issue=4 |pages=508–11 |pmc=3391437}}</ref>

Unlike most primates, male tarsiers do not have [[baculum|bacula]].<ref name="Ankel-Simons2010">{{cite book|author=Friderun Ankel-Simons|title=Primate Anatomy: An Introduction|url=https://books.google.com/books?id=Mwl3M6c5KzoC|date=27 July 2010|publisher=Academic Press|isbn=978-0-08-046911-9|pages=442, 521 }}</ref>

==Behavior==
[[Pygmy tarsier]]s differ from other species in terms of their morphology, communication, and behavior.<ref>{{cite journal |last1=Grow |first1=Nanda |last2=Gursky-Doyen |first2=Sharon |year=2010 |title=Preliminary Data On The Behavior, Ecology, And Morphology Of Pygmy Tarsiers ( Tarsius Pumilus) |journal=International Journal of Primatology |volume=31 |issue=6 |pages=1174–1191 |doi=10.1007/s10764-010-9456-9 |s2cid=23939646}}</ref> The differences in morphology that distinguish pygmy tarsiers from other species are likely based on their high altitude environment.<ref>{{cite journal |last1=Musser |first1=G. G. |last2=Dagosto |first2=M. |year=1987 |title=The identity of Tarsius pumilus, a pygmy species endemic to the montane mossy forests of Central Sulawesi |journal=American Museum Novitates |issue=2867 |pages=1–53}}</ref>

All tarsier species are [[Nocturnality|nocturnal]] in their habits, but like many nocturnal organisms, some individuals may show more or less activity during the daytime. Based on the anatomy of all tarsiers, they are all adapted for leaping even though they all vary based on their species.<ref>{{cite journal |last1=Dagosto |first1=M. |last2=Gebo |first2=D. L. |last3=Dolino |first3=C. |year=2001 |title=Positional behavior and social organization of the Philippine tarsier (''Tarsius syrichta'') |journal=Primates |volume=42 |issue=3 |pages=233–243 |doi=10.1007/bf02629639 |s2cid=41499839}}</ref><ref>{{cite journal |last1=Niemitz |first1=C |year=1977 |title=Zur funktionsmorphologie und biometrie der gattung Tarsius, Storr, 1780 |journal=Courier Forschungsinstitut Senckenberg |volume=25 |pages=1–161}}</ref><ref>Niemitz, C. (1979). Relationships among anatomy, ecology, and behavior: A model developed in the genus Tarsius, with thoughts about phylogenetic mechanisms and adaptive interactions. In S. 1190 N. Grow, S. Gursky-DoyenMorbeck, H. Preuschoft, & N. Gomberg (Eds.), Environment, behavior, and morphology: Dynamic interactions (pp. 119–138). New York: Gustav Fischer.</ref><ref>Niemitz, C. (1984). An investigation and review of the territorial behaviour and social organization of the genus Tarsius. In C. Niemitz (Ed.), Biology of tarsiers (pp. 117–128). New York: Gustav Fischer</ref>

Ecological variation is responsible for differences in morphology and behavior in tarsiers because different species become adapted to local conditions based on the level of altitude.<ref>{{cite journal |last1=Körner |first1=C |year=2007 |title=The use of 'altitude' in ecological research |journal=Trends in Ecology and Evolution |volume=22 |issue=11 |pages=569–574 |doi=10.1016/j.tree.2007.09.006 |pmid=17988759|bibcode=2007TEcoE..22..569K }}</ref> For example, the colder climate at higher elevations can influence cranial morphology.<ref>{{cite journal |last1=Rae |first1=T. C. |last2=Hill |first2=R. I. |last3=Hamada |first3=Y. |last4=Koppe |first4=T. |year=2003 |title=Clinal variation of maxillary sinus volume in Japanese macaques (Macaca fuscata) |journal=American Journal of Primatology |volume=59 |issue=4 |pages=153–158 |doi=10.1002/ajp.10072 |pmid=12682923 |s2cid=12290499}}</ref>

Tarsiers tend to be extremely shy animals and are sensitive to bright lights, loud noises, and physical contact. They have been reported to behave suicidally when stressed or kept in captivity.<ref>{{Cite news |last=Jarosz |first=Andy |date=20 May 2019 |title=The Philippines: The Tarsier Man |work=[[National Geographic]] |url=https://www.nationalgeographic.co.uk/travel/2019/05/philippines-tarsier-man |url-status=dead |access-date=28 September 2019 |archive-url=https://web.archive.org/web/20190521012214/https://www.nationalgeographic.co.uk/travel/2019/05/philippines-tarsier-man |archive-date=21 May 2019}}</ref><ref>{{Cite news |last=Sinclair |first=Joe |date=12 December 2011 |title=Tourism threatens tiny Philippine primate |work=My SinChew |publisher=[[Agence France-Presse|AFP]] |url=http://mysinchew.sinchew.com.my/node/67669 |url-status=dead |access-date=28 September 2019 |archive-url=https://web.archive.org/web/20190928114156/http://mysinchew.sinchew.com.my/node/67669 |archive-date=28 September 2019}}</ref>

===Predators===
Due to their small size, tarsiers are prey to various other animals. Tarsiers primarily inhabit the lower vegetation layers as they face threats from both terrestrial predators such as cats, lizards, and snakes, and aerial predators such as owls and birds. By residing in these lower layers, they can minimize their chances of being preyed upon by staying off the ground and yet still low enough to avoid birds of prey.

Tarsiers, though known as being shy and reclusive, are known to mob predators. In nature, [[Mobbing_(animal_behavior)|mobbing]] is the act of harassing predators to reduce the chance of being attacked. When predators are near, tarsiers will make a warning vocalization. Other tarsiers will respond to the call, and within a short period of time, 2-10 tarsiers will show up to mob the predator. The majority of the group consists of adult males, but there will occasionally be a female or two. While tarsier groups only contain one adult male, males from other territories will join in the mob event, meaning there are multiple alpha male tarsiers attacking the predator.<ref>{{cite journal |last1=Řeháková-Petrů |first1=M. |last2=Peške |first2=L. |date=2012 |title=Predation on a wild Philippine tarsier (Tarsius syrichta) |journal=Acta Ethologica |volume=15 |issue=2 |pages=217–220 |doi=10.1007/s10211-011-0096-7 |s2cid=254163428}}</ref><ref name="RefA" /><ref>{{Cite journal |last=Gursky |first=Sharon |date=February 2005 |title="Predator Mobbing in Tarsius Spectrum" |url=https://doi.org/10.1007/s10764-005-0731-0 |journal=International Journal of Primatology |volume=26 |issue=1 |pages=207–221 |doi=10.1007/s10764-005-0731-0 |s2cid=21188050 |via=EBSCOhost|url-access=subscription }}</ref>

===Diet===
Tarsiers are the only entirely [[Carnivore#Obligate carnivores|carnivorous]] [[Extant taxon|extant]] primates, albeit mainly [[insectivore|insectivorous]], catching invertebrates by jumping at them. The tarsiers also opportunistically prey on a variety of arboreal and small forest animals, including [[orthopterans]], [[scarab beetle]]s, small [[flying frogs]], [[lizards]] and, occasionally, amphibious [[crab]]s that climb into the lower sections of trees.<ref>{{Cite journal |last1=Crompton |first1=Robin Huw |last2=Blanchard |first2=Mary L. |last3=Coward |first3=Sam |last4=Alexander |first4=R. McNeill |last5=Thorpe |first5=Susannah K. |date=1 December 2010 |title=Vertical Clinging and Leaping Revisited: Locomotion and Habitat Use in the Western Tarsier, Tarsius bancanus Explored Via Loglinear Modeling |url=https://doi.org/10.1007/s10764-010-9420-8 |journal=International Journal of Primatology |language=en |volume=31 |issue=6 |pages=958–979 |doi=10.1007/s10764-010-9420-8 |s2cid=45884124 |issn=1573-8604|url-access=subscription }}</ref><ref>{{Cite journal |last1=Crompton |first1=Robin Huw |last2=Savage |first2=Russell |last3=Spears |first3=Iain R. |date=14 February 1998 |title=The Mechanics of Food Reduction in Tarsius bancanus |url=https://brill.com/view/journals/ijfp/69/7/article-p41_6.xml |journal=Folia Primatologica |volume=69 |issue=7 |pages=41–59 |doi=10.1159/000052698 |pmid=9595687 |s2cid=24464173 |issn=1421-9980|url-access=subscription }}</ref><ref>{{Cite journal |last1=Syahrullah |first1=Fakhri Naufal |last2=Maddus |first2=Un |last3=Mustari |first3=Abdul Haris |last4=Gursky |first4=Sharon |last5=Indrawan |first5=Mochamad |date=15 July 2023 |title=Distribution and abundance of Peleng Tarsier (Tarsius pelengensis) in Banggai Island group, Indonesia |journal=Scientific Reports |language=en |volume=13 |issue=1 |pages=11445 |doi=10.1038/s41598-023-30049-5 |issn=2045-2322 |pmc=10349819 |pmid=37454197|bibcode=2023NatSR..1311445S }}</ref> However, it has been found that their favorite prey are [[arthropods]], [[beetles]], [[arachnids]], [[cockroaches]], [[grasshoppers]], [[katydid]]s, [[cicadas]], and [[Phasmatodea|walking sticks]].<ref name="RefA" /> Tarsiers are, rarely, also known to prey on baby birds, small tree snakes and even baby bats.<ref name="EoM" />

===Reproduction===
[[Gestation]] takes about six months,<ref name="Izard et al., 1985">{{cite journal |last=Izard |first=Kay M. |author2=Wright, Simons |year=1985 |title=Gestation length in Tarsius bancanus |journal=Am J Primatol |volume=9 |issue=4 |pages=327–331 |doi=10.1002/ajp.1350090408 |pmid=31979510 |s2cid=83711759 }}</ref> and tarsiers give birth to single offspring. Young tarsiers are born furred, and with open eyes, and are able to climb within a day of birth. They reach sexual maturity by the end of their second year. Sociality and mating system varies, with tarsiers from Sulawesi living in small family groups, while Philippine and western tarsiers are reported to sleep and forage alone.

==Conservation==
Tarsiers have never formed successful breeding colonies in captivity; this may be due in part to their special feeding requirements.<ref name="1993Roberts_and_Kohn">{{cite journal | doi = 10.1002/zoo.1430120207 | title = Habitat Use, Foraging Behavior, and Activity Patterns in Reproducing Western Tarsiers, ''Tarsius bancanus'', in Captivity: A Management Synthesis | last1 = Roberts | first1 = M. | last2 = Kohn | first2 = F. | journal = Zoo Biology | year = 1993 | volume = 12 | issue = 2 | pages = 217–232 | url = http://si-pddr.si.edu/dspace/bitstream/10088/503/1/Roberts1993.pdf | access-date = 19 November 2010 | archive-date = 12 March 2012 | archive-url = https://web.archive.org/web/20120312045122/http://si-pddr.si.edu/dspace/bitstream/10088/503/1/Roberts1993.pdf | url-status = dead }}</ref><ref name="2008aSevern_etal">{{cite book | editor1-last = Shekelle | editor1-first = M. | editor2-last = Maryano | editor2-first = T. | editor3-last = Groves | editor3-first = C. | editor4-last = Schulze | editor4-first = H. | editor5-last = Fitch-Snyder | editor5-first = H. | title = Tarsier Longevity: Data from a Recapture in the Wild and from Captive Animals | publisher = LIPI Press | year = 2008 | isbn = 978-979-799-263-7 | last1 = Shekelle | first1 = M. | last2 = Nietsch | first2 = A. | pages = 85–89 | url = http://tarsier.org/products_files/2008-POTON-p85-ShekelleNietsch-Longevity.pdf | url-status = dead | archive-url = https://web.archive.org/web/20110728081843/http://tarsier.org/products_files/2008-POTON-p85-ShekelleNietsch-Longevity.pdf | archive-date = 28 July 2011 }}</ref><ref name="2008bSevern_etal">{{cite book | editor1-last = Shekelle | editor1-first = M. | editor2-last = Maryano | editor2-first = T. | editor3-last = Groves | editor3-first = C. | editor4-last = Schulze | editor4-first = H. | editor5-last = Fitch-Snyder | editor5-first = H. | publisher = LIPI Press | year = 2008 | isbn = 978-979-799-263-7 | last1 = Severn | first1 = K. | last2 = Dahang | first2 = D. | last3 = Shekelle | first3 = M. | title = Eastern Tarsiers in Captivity, Part I: Enclosure and Enrichment | pages = 91–96 | url = http://www.tarsier.org/products_files/2008-POTON-p91-Severnetal-CaptivityEnclosure.pdf | url-status = dead | archive-url = https://web.archive.org/web/20110724183841/http://www.tarsier.org/products_files/2008-POTON-p91-Severnetal-CaptivityEnclosure.pdf | archive-date = 24 July 2011 }}</ref><ref name="2008cSevern_etal">{{cite book | editor1-last = Shekelle | editor1-first = M. | editor2-last = Maryano | editor2-first = T. | editor3-last = Groves | editor3-first = C. | editor4-last = Schulze | editor4-first = H. | editor5-last = Fitch-Snyder | editor5-first = H. | publisher = LIPI Press | year = 2008 | isbn = 978-979-799-263-7 | last1 = Severn | first1 = K. | last2 = Dahang | first2 = D. | last3 = Shekelle | first3 = M. | title = Eastern Tarsiers in Captivity, Part II: A Preliminary Assessment of Diet | pages = 97–103 | url = http://www.tarsier.org/products_files/2008-POTON-p97-Dahangetal-CaptiveDiet.pdf | url-status = dead | archive-url = https://web.archive.org/web/20110728081958/http://www.tarsier.org/products_files/2008-POTON-p97-Dahangetal-CaptiveDiet.pdf | archive-date = 28 July 2011 }}</ref><ref name="2003Fitch-Snyder">{{cite book | last = Fitch-Snyder | first = H. | chapter = History of Captive Conservation of Tarsiers | pages = 227–295 | title = Tarsiers: Past, Present, and Future | editor1-last = Wright | editor1-first = P.C. | editor2-last = Simons | editor2-first = E.L. | editor3-last = Gursky | editor3-first = S. | year = 2003 | publisher = Rutgers University Press | isbn = 978-0-8135-3236-3}}</ref>

A sanctuary near the town of [[Corella, Bohol|Corella]], on the Philippine island of Bohol, has had some success restoring tarsier populations.<ref>{{cite web|url=http://www.stephenmbland.com/#!the-man-trying-to-save-the-philippine-ta/zoom/c11sk/cqq5|title=StephenMBland|access-date=18 October 2016}}</ref> The Philippines Tarsier Foundation has developed a large, semi-wild enclosure known as the Tarsier Research and Development Center. Carlito Pizarras, also known as the "Tarsier man", founded the sanctuary, where visitors can observe tarsiers in the wild. The trees in the sanctuary are populated with nocturnal insects that make up the tarsier's diet.<ref name="Jachowski2005">{{cite journal |last=Jachowski |first=David S. |author2=Pizzaras, Carlito |year=2005 |title=Introducing an innovative semi-captive environment for the Philippine tarsier (''Tarsius syrichta'') |journal=Zoo Biology |volume=24 |issue=1 |pages=101–109 |doi=10.1002/zoo.20023 }}</ref>

The conservation status of all tarsiers is vulnerable to extinction. Tarsiers are a conservation-dependent species, meaning that they need to have more and improved management of protected habitats, or they will become extinct in the future.<ref name="ShekelleMyronGursky2010" />

The first quantitative study on the activity patterns of captive Philippine tarsiers (''Tarsius syrichta'') was conducted at the Subayon Conservation Centre for the Philippine Tarsier in [[Bilar, Bohol]]. From December 2014 to January 2016, female and male ''T. syrichta'' were observed based on their time apportioned to normal activities during non-mating versus mating seasons. During the non-mating season, a significant amount of their waking hours was spent scanning prior to resting, foraging, and traveling. Feeding, scent-marking, self-grooming, social activities, and other activities were minimal. Scanning was still a common activity among the paired sexes during mating season. However, resting markedly decreased, while increases in travel and foraging were evident.<ref>{{cite journal |last1=Wojciechowski |first1=Filip J. |last2=Kaszycka |first2=K.A. |last3=Wielbass |first3=A.M. |last4=Řeháková |first4=M. |date=2019 |title=Activity Patterns of Captive Philippine Tarsiers (Tarsius Syrichta): Differences Related to Sex and Social Context. |journal=Folia Primatologica |volume=90 |issue=2 |pages=109–23 |doi=10.1159/000495612|pmid=30826810 |s2cid=73491766 }}</ref> 

The 2008-described [[Siau Island tarsier]] in Indonesia is regarded as [[critically endangered]] and was listed among [[The World's 25 Most Endangered Primates]] by [[Conservation International]] and the IUCN/SCC Primate Specialist Group in 2008.<ref>{{cite web | title = Siau Island Tarsier | url = http://www.primate-sg.org/siau07.htm | first1 = Myron | last1 = Shekelle | first2 = Agus | last2 = Salim | publisher = IUCN/SSC Primate Specialist Group | access-date = 1 January 2010 | url-status = dead | archive-url = https://web.archive.org/web/20100906171822/http://www.primate-sg.org/siau07.htm | archive-date = 6 September 2010 }}</ref>
The Malaysian government protects tarsiers by listing them in the Totally Protected Animals of [[Sarawak]], on the island of Borneo, where they are commonly found.<ref name="srwkprtct">{{cite web | url = http://www.forestry.sarawauk.gov.my/forweb/wildlife/mgmt/tpa/tprotsp.htm | title = Totally Protected Animals of Sarawak | publisher = Forestry Department of Sarawak | access-date = 1 January 2010 }}{{Dead link|date=February 2020 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

==References==
{{Reflist}}

==External links==
{{Wikispecies|Tarsiidae}}
{{Wiktionary|Tarsiiformes|Tarsiidae|tarsier}}
{{Commons|Tarsius}}
* [http://www.tarsier.org Tarsier.org] {{Webarchive|url=https://web.archive.org/web/20220116110244/http://tarsier.org/ |date=16 January 2022 }}, an international research and conservation project
* {{webarchive |url=https://web.archive.org/web/20041015083950/http://www.szgdocent.org/pp/p-tarsir.htm |date=15 October 2004 |title=Tarsiers (Tarsiidae) }}, Singapore Zoological Gardens Docents, 2000
* {{cite web | url = http://pin.primate.wisc.edu/factsheets/entry/tarsier | title = Primate Factsheets: Tarsier (Tarsius) Taxonomy, Morphology, & Ecology | publisher = National Primate Research Center, University of Wisconsin–Madison | first = Kurt J. | last = Gron |date=July 2008}}
* [http://www.nomadicnotes.com/travel-blog/aww-tarsiers-the-unofficial-mascot-of-bohol/ Tarsiers – Visiting the two Tarsier sanctuaries in Bohol, Philippines]
* [http://eskeletons.org/boneviewer/nid/12544/region/skull/bone/cranium Tarsier skeleton] {{Webarchive|url=https://web.archive.org/web/20170818033649/http://eskeletons.org/boneviewer/nid/12544/region/skull/bone/cranium |date=18 August 2017 }} – Skeleton from the University of Texas at Austin

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[[Category:Mammals of Southeast Asia]]
[[Category:Tarsiers| ]]
[[Category:Extant Lutetian first appearances]]