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* ''[[Cephalopachus]]''
* ''[[Cephalopachus]]''
* ''[[Tarsius]]''
* ''[[Tarsius]]''
| range_map = Combined Range of 9 Tarsier Species.jpg
| range_map_caption = Range of all tarsier species
}}
}}


'''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>
'''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==
==Evolutionary history==
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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.
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/>
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 a [[simian]] (anthropoid) primate.<ref name=sirindhornae/>


So far, four fossil species of tarsiers are known from the fossil record:
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>
* ''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 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>
* ''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>
* ''[[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>
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>
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{{see also|List of tarsiiformes}}
{{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>
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 | 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]]
[[File:Bohol Tarsier.jpg|thumb|right|[[Philippine tarsier]] (''Carlito syrichta''), one of the smallest primates]]
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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 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>
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 |article-number=e104340 |doi=10.1371/journal.pone.0104340 |doi-access=free |issn=1932-6203 |pmc=4138104 |pmid=25136854 |bibcode=2014PLoSO...9j4340B }}</ref>


* Infraorder Tarsiiformes
* Infraorder Tarsiiformes
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[[File:Tarsius Syrichta-GG.jpg|thumb|right|Tarsiers tree-climbing]]
[[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>
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.{{pn |date=October 2025}} 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]].
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 |page=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>
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>
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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>
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>
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|url=https://durham-repository.worktribe.com/output/1623678 }}</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>
Tarsiers tend to be extremely shy animals and are sensitive to bright lights, loud noises, and physical contact. They have been reported to behave [[Animal_suicide#Tarsiers|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 |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 |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===
===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.
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>
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" |journal=International Journal of Primatology |volume=26 |issue=1 |pages=207–221 |doi=10.1007/s10764-005-0731-0 |s2cid=21188050 }}</ref>


===Diet===
===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" />
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 |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}}</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 |page=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===
===Reproduction===
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==Conservation==
==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>
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 }}</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 | 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 | 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 | 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>
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 were 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 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 | archive-url = https://web.archive.org/web/20100906171822/http://www.primate-sg.org/siau07.htm | archive-date = 6 September 2010 }}</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>
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>


Latest revision as of 07:59, 2 November 2025

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Tarsiers (Template:IPAc-en Template:Respell) are haplorhine primates of the 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.[1]

File:Tarsier (Inside Philippine National Museum of Natural History).jpg
Tarsier image inside 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.[2]

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,[3] and it is sometimes listed in its own family, Afrotarsiidae, within the infraorder Tarsiiformes,[4] or considered a simian (anthropoid) primate.[5]

So far, four fossil species of tarsiers are known from the fossil record:

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.[9]

Classification

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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 simians (Anthropoidea) in the infraorder Haplorhini. Analysis of 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.[10]

File:Bohol Tarsier.jpg
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,[11] 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).[12] Species level taxonomy is complex, with morphology often being of limited use compared to vocalizations.Script error: No such module "Unsubst". 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.[11]

In 2010, Colin Groves and Myron Shekelle suggested splitting the genus Tarsius into three genera, the Philippine tarsiers (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 mammae, chromosome count, socioecology, vocalizations, and distribution. The senior taxon of the species, T. tarsier was restricted to the population of a Selayar island, which then required the resurrection of the defunct taxon T. fuscus.[13]

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.[14]

Anatomy and physiology

File:Tarsius Syrichta-GG.jpg
Tarsiers tree-climbing

Tarsiers are small animals with enormous eyes; each eyeball is approximately Template:Convert in diameter and is as large as, or in some cases larger than, its entire brain.[16][17] 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.[18] Tarsiers have a strong auditory sense, and their auditory cortex is distinct.[18] Tarsiers also have long hind limbs, owing mostly to the elongated tarsus bones of the feet, from which the animals get their name.Template:Pn The combination of their elongated tarsi and fused tibiofibulae makes them morphologically specialized for vertical clinging and leaping.[19] The head and body range from 10 to 15 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 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.[20]

Tarsier morphology allows for them to move their heads 180 degrees in either direction, allowing for them to see 360 degrees around them.[21] Their dental formula is also unique: Template:DentalFormula[22] Unlike many nocturnal vertebrates, tarsiers lack a light-reflecting layer (tapetum lucidum) of the retina and have a 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 lemurs, lorises, and monkeys, which are all similar in this respect.[23] 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."[24]

Philippine tarsiers are capable of hearing frequencies as high as 91 kHz. They are also capable of vocalizations with a dominant frequency of 70 kHz.[25]

Unlike most primates, male tarsiers do not have bacula.[26]

Behavior

Pygmy tarsiers differ from other species in terms of their morphology, communication, and behavior.[27] The differences in morphology that distinguish pygmy tarsiers from other species are likely based on their high altitude environment.[28]

All tarsier species are 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.[29][30][31][32]

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.[33] For example, the colder climate at higher elevations can influence cranial morphology.[34]

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.[35][36]

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 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.[37][21][38]

Diet

Tarsiers are the only entirely carnivorous extant primates, albeit mainly insectivorous, catching invertebrates by jumping at them. The tarsiers also opportunistically prey on a variety of arboreal and small forest animals, including orthopterans, scarab beetles, small flying frogs, lizards and, occasionally, amphibious crabs that climb into the lower sections of trees.[39][40][41] However, it has been found that their favorite prey are arthropods, beetles, arachnids, cockroaches, grasshoppers, katydids, cicadas, and walking sticks.[21] Tarsiers are, rarely, also known to prey on baby birds, small tree snakes and even baby bats.[20]

Reproduction

Gestation takes about six months,[42] 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.[43][44][45][46][47]

A sanctuary near the town of Corella, on the Philippine island of Bohol, has had some success restoring tarsier populations.[48] 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.[49]

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 were 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.[50]

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.[51] 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.[52]

References

Template:Reflist

External links

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  1. ↑ The Editors of Encyclopædia Britannica. "Tarsier." Encyclopædia Britannica, Encyclopædia Britannica, Inc., 14 April 2019, http://www.britannica.com/animal/tarsier.
  2. ↑ Script error: No such module "Citation/CS1".
  3. ↑ a b c Script error: No such module "citation/CS1".
  4. ↑ McKenna, M.C., and Bell, S.K. 1997. Classification of Mammals Above the Species Level. Columbia University Press, New York, 337–340 pp. Template:ISBN
  5. ↑ a b Script error: No such module "Citation/CS1".
  6. ↑ Script error: No such module "Citation/CS1".
  7. ↑ Script error: No such module "citation/CS1".
  8. ↑ Script error: No such module "Citation/CS1".
  9. ↑ Script error: No such module "citation/CS1".
  10. ↑ Script error: No such module "Citation/CS1".
  11. ↑ a b Cite error: Invalid <ref> tag; no text was provided for refs named MSW3
  12. ↑ Script error: No such module "Citation/CS1".
  13. ↑ Script error: No such module "Citation/CS1".
  14. ↑ Script error: No such module "Citation/CS1".
  15. ↑ a b Script error: No such module "Citation/CS1".
  16. ↑ Script error: No such module "citation/CS1".
  17. ↑ Script error: No such module "citation/CS1".
  18. ↑ a b Script error: No such module "Citation/CS1".
  19. ↑ Script error: No such module "Citation/CS1".
  20. ↑ a b Script error: No such module "citation/CS1".
  21. ↑ a b c Gron KJ. 2010 December 1. Primate Factsheets: Tarsier (Tarsius) Taxonomy, Morphology, & Ecology . http://pin.primate.wisc.edu/factsheets/entry/tarsier . Retrieved 12 November 2019.
  22. ↑ Script error: No such module "citation/CS1".
  23. ↑ Script error: No such module "Citation/CS1".
  24. ↑ Script error: No such module "Citation/CS1".
  25. ↑ Script error: No such module "Citation/CS1".
  26. ↑ Script error: No such module "citation/CS1".
  27. ↑ Script error: No such module "Citation/CS1".
  28. ↑ Script error: No such module "Citation/CS1".
  29. ↑ Script error: No such module "Citation/CS1".
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  31. ↑ 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.
  32. ↑ 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
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  35. ↑ Script error: No such module "citation/CS1".
  36. ↑ Script error: No such module "citation/CS1".
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  40. ↑ Script error: No such module "Citation/CS1".
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  44. ↑ Script error: No such module "citation/CS1".
  45. ↑ Script error: No such module "citation/CS1".
  46. ↑ Script error: No such module "citation/CS1".
  47. ↑ Script error: No such module "citation/CS1".
  48. ↑ Script error: No such module "citation/CS1".
  49. ↑ Script error: No such module "Citation/CS1".
  50. ↑ Script error: No such module "Citation/CS1".
  51. ↑ Script error: No such module "citation/CS1".
  52. ↑ Script error: No such module "citation/CS1".Template:Dead link
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