Horsfield's tarsier: Difference between revisions
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{{Short description|Species of primate}} |
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{{Taxobox |
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{{Speciesbox |
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| name = Horsfield's Tarsier<ref name=msw3>{{MSW3 Groves|pages=127|id=12100167}}</ref> |
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| name = Western tarsier<ref name=msw3>{{MSW3 Groves|pages=127|id=12100167}}</ref> |
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| image = Stavenn Tarsius bancanus 00.jpg |
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| image = Western Tarsier.jpg |
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| image_caption = ''Tarsius bancanus'', specimen - [[AMNH]] |
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| image_caption = ''C. bancanus'' in [[Borneo]]. Note the [[eyeshine]]. |
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| status = VU |
| status = VU |
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| status_system = |
| status_system = IUCN3.1 |
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| status_ref = <ref name=iucn>{{ |
| status_ref = <ref name="iucn status 13 November 2021">{{cite iucn |author=Shekelle, M. |author2=Yustian, I. |date=2020 |title=''Cephalopachus bancanus'' |volume=2020 |page=e.T21488A17976989 |doi=10.2305/IUCN.UK.2020-3.RLTS.T21488A17976989.en |access-date=13 November 2021}}</ref> |
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| parent_authority = [[William John Swainson|Swainson]], 1835 |
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| regnum = [[Animal]]ia |
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| taxon = Cephalopachus bancanus |
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| phylum = [[Chordate|Chordata]] |
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| authority = ([[Thomas Horsfield|Horsfield]], 1821) |
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| classis = [[Mammal]]ia |
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| range_map = Horsfield's Tarsier area.png |
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| ordo = [[Primate]]s |
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| range_map_caption = Western tarsier range |
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| familia = [[Tarsiidae]] |
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| genus = ''[[Tarsius]]'' |
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| species = '''''T. bancanus''''' |
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| binomial = ''Tarsius bancanus'' |
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| binomial_authority = [[Thomas Horsfield|Horsfield]], 1821 |
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| synonyms = |
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''T. b. boreanus'': |
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*''natunensis'' <small>Chasen, 1940</small> |
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}} |
}} |
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'''Horsfeld's tarsier''' ('''''Cephalopachus bancanus''''') is the [[Monotypic taxon|only species]] of [[tarsier]] in the genus '''''Cephalopachus'''''. Named by American [[Natural history|naturalist]] [[Thomas Horsfield]], it is also referred to as '''western tarsier'''. The species occurs on [[Borneo]], [[Sumatra]] and nearby islands and is, like other members of the group, entirely nocturnal. |
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==Taxonomy== |
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'''Horsfield's Tarsier''' ('''''Tarsius bancanus'''''), also known as the '''Western Tarsier''', is a species of [[tarsier]]. |
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Although Horsfeld's tarsier was usually placed in the genus ''[[Tarsius]]'' with all other living tarsiers, it is quite distinct from the [[Philippine tarsier]] and the various tarsiers of [[Sulawesi]] and nearby islands; therefore, scientists have placed it in a separate genus, ''Cephalopachus''.<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 | pages = 1071–1082 | year = 2010 | issue = 6 | doi = 10.1007/s10764-010-9443-1| s2cid = 21220811 }}</ref> |
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The taxonomy of this species is in doubt, with some subspecies considered unsure.<ref>{{cite journal | doi = 10.1023/B:IJOP.0000014647.18720.32 | author = Brandon-Jones, D. | author2 = Eudey, A.A. | author3 = Geissmann, T. | author4 = Groves, C.P. | author5 = Melnick D.J., Morales | author6 = J.C., Shekelle, M. | author7 = Stewart, C.B | name-list-style = amp | year = 2004 | title = Asian Primate Classification | journal = International Journal of Primatology | volume = 25 | issue = 1 | pages = 97–164| s2cid = 29045930 }}</ref> In fact, over 20 years few studies have been done on ''C. bancanus'' and a taxonomic revision based upon intensive and systematic field surveys is overdue. The [[IUCN]] believes that these subspecies should be treated as distinct and named as separate taxa until more definitive evidence is available.<ref>{{cite iucn |author=Shekelle, M. |author2=Yustian, I. |date=2021 |title=''Cephalopachus bancanus'' ssp. ''bancanus'' |volume=2021 |page=e.T39762A17992163 |doi=10.2305/IUCN.UK.2021-1.RLTS.T39762A17992163.en |access-date=13 November 2021}}</ref> When splitting the species into its own genus, [[Colin Groves]] and Myron Shekelle recognized the ''natunensis'' population as a distinct subspecies.<ref name=2010_Groves_Shekelle/> |
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==Taxonomic notes== |
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There are four recognized subspecies of Horsfeld's tarsier: |
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The taxonomy of this species is in doubt, with some subspecies considered unsure.<ref>{{cite journal | author = Brandon-Jones, D., Eudey, A.A., Geissmann, T., Groves, C.P., Melnick D.J., Morales, J.C., Shekelle, M., and Stewart, C.B | year = 2004 | title = Asian Primate Classification | journal = International Journal of Primatology | volume = 25 | issue = 1 | pages = 97-164}}</ref> In fact, over 20 years few studies have been done on ''T. bancanus'' and a taxonomic revision based upon intensive and systematic field surveys is overdue. These subspecies should be treated as distinct and named as separate taxa until more definitive evidence is available.<ref>{{IUCN2008|assessors=Shekelle, M. & Yustian, I. | year = 2008 | title = Tarsius bancanus'' ssp. ''bancanus | downloaded = 25 January 2009|id=39762}}</ref> |
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* ''Cephalopachus bancanus bancanus'' — [[Bangka Island]] tarsier |
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* ''Cephalopachus bancanus saltator'' — [[Belitung Island]] tarsier |
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* ''Cephalopachus bancanus borneanus'' — [[Borneo|Bornean]] tarsier |
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* ''Cephalopachus bancanus natunensis'' — [[Natuna Islands]] tarsier |
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==Habitat and distribution== |
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[[Image:Kera hantu Sarawak.jpg|thumb|left|Horsfield's Tarsier in Sarawak]] |
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There are three recognized subspecies of Horfield's Tarsier: |
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* ''Tarsius bancanus bancanus'' |
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* [[Belitung Island Tarsier]], ''Tarsius bancanus saltator'' |
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* [[Bornean Tarsier]], ''Tarsius bancanus borneanus'' |
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Horsfield's tarsier is found in Southern [[Sumatra]], [[Borneo]] and [[List of islands of Indonesia|nearby islands]].<ref name=field/> The Bornean subspecies, ''C. b. borneanus'', is known from many [[lowland]] sites in [[Sabah]], [[Brunei]], [[Sarawak]] and [[West Kalimantan]] and above {{convert|900|m|abbr=on}} in the [[Kelabit Highlands|Kelabit uplands]] in Northern [[Sarawak]]. Other records show it from [[Kutai]] and Peleben in [[East Kalimantan]] and [[Tanjung Maruwe]] in [[Central Kalimantan]].<ref name=field/> This species can live in both [[primary forest|primary]] and [[secondary forest]]s, and it also lives in forests along the coasts or on the edge of [[plantation]]s.<ref name=Niemitz1979/> |
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==Morphology== |
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==Physical description== |
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The [[pelage]] coloration range from pale-olive or reddish brown to pale or dark grey-brown, possibly varying with age. Based on 12 collected specimens,<ref name=field>{{cite book | author = Payne, J., Francis, C. M., & Phillipps, K | year = 1994 | title = A Field guide to the Mammals of Borneo | location = Petaling Jaya | publisher = The Sabah Society}}</ref> the range measurement from head to body is 121-154 mm. ''T. bancanus'' has an extremely long tail which can reach 181 to 224 mm and is hairless except for tufts of hair at the end.<ref name=Sussman>{{cite book | author = Sussman, R.W | year = 1999 | title = Primate Ecology and Social Structure: Vol. 1 Lorises, Lemurs and Tarsiers | publisher = Pearson Custom Publishing}}</ref> This species has two [[Personal grooming|grooming]] [[claws]] on each foot. The fingers are very long and have pads on the tips. The toes have flattened nails except for the second and third toes on hind feet, which bear claw-like nails.<ref name=field/> It has large eyes which do not reflect light. The membranous ears are slender and almost bare. The [[molar (tooth)|molar]]s of this species have high-cusps and are almost [[tritubercular]].<ref name=food>{{cite journal | author = Crompton, R.H., Savage, R., and Spears, I.R | year = 1998 | title = The mechanics of food reduction in Tarsius bancanus: Hard-object feeder, soft-object feeder or both? | journal = Folia Primatologica | volume = 69 | issue = suppl 1 | pages = 41-59}}</ref> The dental formula of ''T. bancanus'' is 2:1:3:3 on the upper jaw and 1:1:3:3 on the lower jaw.<ref name=Nowak>{{cite book | author = Nowak, R.M | year = 1999 | title = Walker's Primates of the World | publisher = Johns Hopkins University Press | location = Baltimore}}</ref> |
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<!--What differentiates this genus from ''[[Tarsius]]''? What are its [[Autapomorphy|diagnostic trait]]s?--> |
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The [[pelage]] coloration ranges from pale-[[Olive (color)|olive]] or reddish brown to pale or dark grey-brown, possibly varying with age. Based on 12 collected specimens,<ref name=field>{{cite book | author = Payne, J. | author2 = Francis, C. M. | author3 = Phillipps, K | name-list-style = amp | year = 1994 | title = A Field guide to the Mammals of Borneo | location = Petaling Jaya | publisher = The Sabah Society}}</ref> the head to body measurement range from {{convert|121|–|154|mm|abbr=on}}. Horsfield's tarsier has an extremely long tail which can reach {{convert|181|to|224|mm|abbr=on}} and is hairless except for tufts of hair at the end.<ref name=Sussman>{{cite book | author = Sussman, R.W | year = 1999 | title = Primate Ecology and Social Structure: Vol. 1 Lorises, Lemurs and Tarsiers | publisher = Pearson Custom Publishing}}</ref> This species has two [[grooming claw]]s on each foot. The fingers are very long and have pads on the tips. The toes have flattened [[Nail (anatomy)|nail]]s except for the second and third toes on hind feet, which bear claw-like nails.<ref name=field/> It has large eyes which do not reflect light.{{Inconsistent|reason=The Infobox Image}} The membranous ears are slender and almost bare. The [[molar (tooth)|molar]]s of this species have high-cusps and are almost [[wikt:tritubercular|tritubercular]].<ref name=food>{{cite journal | author = Crompton, R.H. | author2 = Savage, R. | author3 = Spears, I.R | name-list-style = amp | year = 1998 | title = The mechanics of food reduction in Tarsius bancanus: Hard-object feeder, soft-object feeder or both? | journal = Folia Primatologica | volume = 69 | issue = suppl 1 | pages = 41–59 | doi=10.1159/000052698| pmid = 9595687 | s2cid = 24464173 }}</ref> The dental formula of Horsfield's tarsier is 2:1:3:3 on the upper jaw and 1:1:3:3 on the lower jaw.<ref name=Nowak>{{cite book | author = Nowak, R.M | year = 1999 | title = Walker's Primates of the World | publisher = Johns Hopkins University Press | location = Baltimore | isbn = 0-8018-6251-5 | url-access = registration | url = https://archive.org/details/walkersprimateso0000nowa }}</ref> |
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==Behaviour and ecology== |
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==Distributions== |
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Horsfield's tarsier is a [[nocturnal]] species. It sleeps alone during the day in a tangle of [[vine]]s or [[liana]]s at a height of {{convert|3.5|to|5|m}}.<ref name=Crompton1986>{{cite journal | doi = 10.1007/BF02382075 |author1=Crompton, R.H. |author2=Andau, P.M | year = 1986 | title = Locomotion and Habitat Utilization in Free-ranging Tarsius bancanus: A Preliminary Report | journal = Primates | volume = 27 | issue = 3 | pages = 337–355|s2cid=25836867 }}</ref> This species prefers to sleep, rest, or remain stationary on perches that are angled 5 degrees from the vertical tree trunks, {{convert|1|to|4|cm|abbr=on}} in diameter,<ref name=Niemitz1984a>{{cite book | author = Niemitz, C |author-link=Carsten Niemitz| year = 1984a | chapter = Locomotion and posture of Tarsius bancanus | title = Biology of Tarsiers | editor = C. Niemitz |editor-link=Carsten Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart | isbn = 0-89574-182-2}}</ref> and it sleeps solitarily.<ref name=Crompton1987>{{cite journal | doi = 10.1007/BF02737113 |author1=Crompton, R.H. |author2=Andau, P.M | year = 1987 | title = Ranging, activity rhythms, and sociality in free-ranging Tarsius bancanus: A preliminary report | journal = International Journal of Primatology | volume = 8 | issue = 1 | pages = 43–71|s2cid=39920648 }}</ref> Before sunset, Horsfield's tarsier will wake up and wait 10 to 20 minutes before moving around the understory and spending 1.5 to 2 hours of the night foraging for food.<ref name=Niemitz1979>{{cite book | author = Niemitz, C. |author-link=Carsten Niemitz | year = 1979 | chapter = Outline of the behavior of Tarsius bancanus | title = The Study of Prosimian Behavior |editor1=G.A. Doyle |editor2=R.D. Martin | publisher = Academic Press | location = New York}}</ref> Horsfield's tarsier can be found from ground level up to a height of {{convert|7|m|abbr=on}} or more in the understory.<ref name = field/> |
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''T. bancanus'' is found in Southern [[Sumatra]], [[Borneo]] and nearby islands.<ref name=field/> The Bornean subspecies, ''T. b. borneanus'', is known from many lowland sites in [[Sabah]], [[Brunei]], [[Sarawak]] and West [[Kalimantan]] and above 900m in the Kelabit uplands in Northern [[Sarawak]]. Other records show it from Kutai and Peleben in East [[Kalimantan]] and Tanjung Maruwe in Central [[Kalimantan]].<ref name=field/> |
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[[File:Calliophis intestinalis 1916.jpg|thumb|right|''Calliophis intestinalis'' is preyed on by the western tarsier]] |
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This species is [[carnivorous]]. It mainly eats [[arthropod]]s such as [[beetle]]s, [[orthoptera]]ns like [[grasshopper]]s, [[katydids]], and [[Cricket (insect)|cricket]]s, [[butterflies]], [[moth]]s, [[cockroach]]es, [[praying mantis]], [[ant]]s, [[Phasmatodea|phasmids]], [[cicada]]s, [[dragonflies]], [[freshwater crab]]s, and [[spider]]s,<ref name=Niemitz1979/><ref name="Crompton2010">{{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. |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 |date=2010 |volume=31 |pages=958-979 |doi=10.1007/s10764-010-9420-8 |url=https://link.springer.com/article/10.1007/s10764-010-9420-8 |access-date=21 November 2024}}</ref> but also will eat small [[vertebrates]] such as [[flying frog]]s,<ref name="Crompton2010" /> bats ([[Chiroptera]]) including members of the genus ''[[Taphozous]]'', the lesser short-nosed fruit bat (''[[Cynopterus brachyotis]]''), and the spotted-winged fruit bat (''[[Balionycteris maculata]]''), and snakes, of which [[venomous snake]]s have been found to be consumed, such as the [[Elapidae|elapid]] ''[[Calliophis intestinalis]]''. This species was also found to consume [[bird]]s, including [[spiderhunter]]s, [[warbler]]s, [[kingfisher]]s, and [[pitta]]s. It locates prey primarily by sound and catches the prey with its hands when foraging. Prey is killed through bites to the [[nape|back of the neck]]<ref name=Niemitz1979/> and the tarsier's eyes are shut when attacking.<ref name=Fogden>{{cite book | author = Fogden, M.P.L | year = 1974 | chapter = A preliminary field study of the western tarsier, Tarsius bancanus Horsefield | title = Prosimian Biology |editor1=R.D. Martin |editor2=G.A. Doyle |editor3=A.C. Walker | publisher = University of Pittsburgh Press | location = Pittsburgh}}</ref> It will consume the prey starting with the head and working its way down the body.<ref name=Niemitz1979/> This species [[Drinking|gets water]] both by drinking from a pool or stream, and by licking drops from [[bamboo]] leaves or from water running down the trunks of trees.<ref name = Niemitz1984>{{cite book | author = Niemitz, C |author-link=Carsten Niemitz | year = 1984d | chapter = Synecological relationships and feeding behaviour of the genus Tarsius | title = Biology of Tarsiers | editor = C. Niemitz |editor-link=Carsten Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart | isbn = 0-89574-182-2}}</ref> Horsfield's tarsier is a host of the [[acanthocephala]]n intestinal parasite ''[[Moniliformis tarsii]]''.<ref name="Deveaux1988">{{Cite journal |doi = 10.2307/3282462|pmid = 3128654|jstor = 3282462|title = Two New Species of Moniliformis (Acanthocephala: Moniliformidae) from Malaysia|journal = The Journal of Parasitology|volume = 74|issue = 2|pages = 322–5|year = 1988|last1 = Deveaux|first1 = Timothy P.|last2 = Schmidt|first2 = Gerald D.|last3 = Krishnasamy|first3 = M.}}</ref> |
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Like all tarsiers, Horsfield's tarsiers are vertical clingers and [[Jumping|leaper]]s, known for extraordinary leaping abilities. An individual will mainly support itself with its feet and the tail, both exerting enough force to hold the individual in place. The hands are not always used because of the pads located on the feet provide adequate grip. The hands are usually placed no higher than its nose except when resting; the hands are only placed higher up to maintain the position of the individual.<ref name=Niemitz1984a/> Other modes of locomotion used by the species include [[Arboreal locomotion|climbing]], quadrupedal walking, hopping and "[[cantilever]]ing."<ref name=Crompton1986/><ref name="Crompton2010" /> |
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==Ecology== |
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Horsfield's tarsiers are [[Monogamy in animals|monogamous]], with a copulation frequency during [[Estrous cycle|estrus]] of once per night.<ref name=Wright1986a/> [[Courtship (animals)|Courtship]] [[animal communication|call]]s are performed by the male and he emits 2–3 chirrups while opening and closing the mouth.<ref name=Wright1986a/> This call happens within 5 minutes of looking at the female. Once the male gives his courtship call, if the female is receptive, she will perform genital displays to him.<ref name=Wright1986a>{{cite journal | author = Wright, P.C. | author2 = Toyama, L.M. | author3 = Simons, E.L | name-list-style = amp | year = 1986 | title = Courtship and Copulation in Tarsius bancanus | journal = Folia Primatologica | volume = 46 | issue = 3 | pages = 142–148 | pmid=3100401 | doi=10.1159/000156247}}</ref> If the female is not in estrus, she will emit an [[Agonistic behaviour|agonistic call]] which is often followed by biting and pushing the male away.<ref name=Wright1986a/> Both sexes' calls last on average for 1 second, and the interval between calls is on average 3 seconds.<ref name=Wright1986a/> |
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Horsfield's Tarsier is a [[nocturnal]] species. It sleeps alone during the day in a tangle of vines or creepers at a height of 3.5 to 5 meters.<ref name=Crompton1986>{{cite journal | author = Crompton, R.H. and Andau, P.M | year = 1986 | title = Locomotion and Habitat Utilization in Free-ranging Tarsius bancanus: A Preliminary Report | journal = Primates | volume = 27 | issue = 3 | pages = 337-355}}</ref> This species prefers to sleep, rest, or remain stationary on perches that are angled 5 degrees from the vertical tree trunks, 1 to 4 cm in diameter,<ref name=Niemitz1984a>{{cite book | author = Niemitz, C | year = 1984a | chapter = Locomotion and posture of Tarsius bancanus | title = Biology of Tarsiers | editor = C. Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart}}</ref> and it sleeps [[solitarily]].<ref name=Crompton1987>{{cite journal | author = Crompton, R.H. and Andau, P.M | year = 1987 | title = Ranging, activity rhythms, and sociality in free-ranging Tarsius bancanus: A preliminary report | journal = International Journal of Primatology | volume = 8 | issue = 1 | pages = 43-71}}</ref> Before sunset, ''T. bancanus'' will wake up and wait 10 to 20 minutes before moving around the understory and spends 1.5 to 2 hours of the night foraging for food.<ref name=Niemitz1979>{{cite book | author = Niemitz, C. | year = 1979 | chapter = Outline of the behavior of Tarsius bancanus | title = The Study of Prosimian Behavior | editor = G.A. Doyle and R.D. Martin | publisher = Academic Press | location = New York}}</ref> Horsfield's Tarsier can be found from ground level up at least 7m in the understory.<ref name = field/> |
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Infants are born with their eyes open and fully furred and able to groom themselves.<ref name=Robert1994>{{cite journal | doi = 10.1007/BF02735232 | author = Roberts, M | year = 1994 | title = Growth, development, and parental care in the Western tarsier (Tarsius bancanus) in captivity: Evidence for a "slow" life-history and nonmonogamous mating system | journal = International Journal of Primatology | volume = 15 | issue = 1 | pages = 1–28| s2cid = 7462413 }}</ref> The mother will carry her infant in her mouth and when she forages for food the mother will park the infant on a branch.<ref name=Haring>{{cite journal | author = Haring, D.M. | author2 = Wright, P.C. | author3 = Simons, E.L | name-list-style = amp | year = 1985 | title = Social behaviors of Tarsius syrichta and Tarsius bancanus | journal = American Journal of Physical Anthropology | volume = 66 | pages = 179|doi= 10.1002/ajpa.1330660204}}</ref> Infant sounds are mostly clicks: "k", "tk", "ki", or a rapid "kooih"<ref name=Niemitz1984b>{{cite book | author = Niemitz, C |author-link=Carsten Niemitz | year = 1984b | chapter = Vocal communication of two tarsier species (Tarsius bancanus and Tarsius spectrum) | title = Biology of Tarsiers | editor = C. Niemitz |editor-link=Carsten Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart | isbn = 0-89574-182-2}}</ref> and can be heard when the infant is left alone or is cold.<ref name=Niemitz1984b/> The mother stays in contact with their infants using high-pitched calls. Infants were found to first use the tails as support during resting at 7–10 days.<ref name=Robert1994/> |
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This species is a carnivorous species. It mainly eats [[insects]] such as [[beetle]]s, [[grasshopper]]s, [[cockroach]]es, [[butterflies]], [[moth]]s, [[praying mantis]], [[ant]]s, [[Phasmatodea|phasmids]], and [[cicada]]s,<ref name=Niemitz1979/> but also will eat small [[vertebrates]] such as bats ([[Chiroptera]]) including [[horseshoe bat]]s of the ''[[Taphozus]]'' genus, the Lesser Short-nosed Fruit Bat (''[[Cynopterus brachyotis]]''), and the Spotted-winged Fruit Bat (''[[Balionycteris maculata]]''),<ref name=Niemitz1979/> and snakes, of which poisonous snakes have been found to be consumed. For example the poisonous snake ''[[Manticora intestinalis]]'' was found to be hunted for by this species.<ref name=Niemitz1979/> This species was also found to consume [[bird]]s, including: [[spider-hunters]], [[warbler]]s, [[kingfisher]]s, and [[pitta]]s.<ref name=Niemitz1979/> It locates prey primarily by sound and catches the prey with its hands when foraging.<ref name=Niemitz1979/> The prey items get killed by bites to the back of the neck<ref name=Niemitz1979/> and the eyes are shut when attacking.<ref name=Fogden>{{cite book | author = Fogden, M.P.L | year = 1974 | chapter = A preliminary field study of the western tarsier, Tarsius bancanus Horsefield | title = Prosimian Biology | editor = R.D. Martin, G.A. Doyle, A.C. Walker | publisher = University of Pittsburgh Press | location = Pittsburgh}}</ref> It will consume the prey starting with the head and working its way down the body.<ref name=Niemitz1979/> This species gets water both by drinking from a pool or steam, and by licking drops from bamboo leaves or from trunks of trees when water is running down the bark.<ref name = Niemitz1984>{{cite book | author = Niemitz, C | year = 1984d | chapter = Synecological relationships and feeding behaviour of the genus Tarsius | title = Biology of Tarsiers | editor = C. Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart}}</ref> |
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[[Social grooming]] in this species only occurs between mothers and infants,<ref name=Niemitz1984c>{{cite book | author = Niemitz, C |author-link=Carsten Niemitz | year = 1984c | chapter = An investigation and review of the territorial behaviour and social organisation of the genus Tarsius | title = Biology of Tarsiers | editor = C. Niemitz |editor-link=Carsten Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart | isbn = 0-89574-182-2}}</ref> removing dead skin and [[Ectoparasite|parasite]]s by scratching with their toe claws and licking their fur, avoiding their faces. Faces are cleaned by rubbing on branches and it is to reinforce social bonds.<ref name=Niemitz1984c/> |
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Young leave their mother's range at the onset of [[puberty]], and find their own [[Territory (animal)|territory]].<ref name=Robert1994/> Horsfield's tarsiers mark their territory with scents from urine and glandular secretions on a substrate while scratching the surface with its hind-limb toe claws.<ref name=Wright1986a/> |
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''T. bancanus'', like all tarsiers, is a vertical clinger and leaper known for its extraordinary leaping abilities. An individual will mainly support itself with its feet and the tail exerts enough force to hold the individual in place without using the hands much because of the pads located on the feet. Except when resting, the hands are usually placed no higher than nose.<ref name=Niemitz1984a/> The hands are only placed higher up to maintain the position of the individual.<ref name=Niemitz1984a/> Other modes of locomotion of this species are climbing, quadrupedal, walking, hopping and "cantilevering"<ref name=Crompton1986/> |
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Horsfield's Tarsier is [[monogamous]], with acopulation frequency during estrus of once per night.<ref name=Wright1986a/> [[Courtship]] [[animal communication|call]]s are performed by the male and he emits 2-3 chirrups while opening and closing the mouth.<ref name=Wright1986a/> This call happens within 5 minutes of looking at the female. Once the male gives his courtship call, if the female is receptive, she will perform genital displays to him.<ref name=Wright1986a>{{cite journal | author = Wright, P.C., Toyama, L.M., and Simons, E.L | year = 1986 | title = Courtship and Copulation in Tarsius bancanus | journal = Folia Primatologica | volume = 46 | pages = 142-148}}</ref> If the female is not in estrus, she will emit an [[agonistic]] call which is often followed by biting and pushing the male away.<ref name=Wright1986a/> Both of each call lasts on average for 1 second and the interval between each call is on average 3 seconds.<ref name=Wright1986a/> |
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Infants are born with their eyes open and fully furred and are able to groom themselves.<ref name=Robert1994>{{cite journal | author = Roberts, M | year = 1994 | title = Growth, development, and parental care in the Western tarsier (Tarsius bancanus) in captivity: Evidence for a "slow" life-history and nonmonogamous mating system | journal = International Journal of Primatology | volume = 15 | issue = 1 | pages = 1-28}}</ref> The mother will carry her infant in her mouth and when she forages for food the mother will park the infant on a branch.<ref name=Haring>{{cite journal | author = Haring, D.M., Wright, P.C., and Simons, E.L | year = 1985 | title = Social behaviors of Tarsius syrichta and Tarsius bancanus | journal = American Journal of Physical Anthropology | volume = 66 | pages = 179}}</ref> Infant sounds are mostly clicks: "k", "tk", "ki", or a rapid "kooih"<ref name=Niemitz1984b>{{cite book | author = Niemitz, C | year = 1984b | chapter = Vocal communication of two tarsier species (Tarsius bancanus and Tarsius spectrum) | title = Biology of Tarsiers | editor = C. Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart}}</ref> and can be heard when the infant is left alone or is cold.<ref name=Niemitz1984b/> The mother stays in contact with their infants using high-pitched calls. Infants were found to first use the tails as support during resting at 7–10 days.<ref name=Robert1994/> Young leave their range at the onset of puberty, and find their own territory.<ref name=Robert1994/> |
|||
Social grooming in this species only occurs between mothers and infants,<ref name=Niemitz1984c>{{cite book | author = Niemitz, C | year = 1984c | chapter = An investigation and review of the territorial behaviour and social organisation of the genus Tarsius | title = Biology of Tarsiers | editor = C. Niemitz | publisher = Gustav Fischer Verlag | location = Stuttgart}}</ref> removing dead skin and parasites by scratching with their toe claws and licking their fur, avoiding their faces. Faces are cleaned by rubbing on branches and it is to reinforce social bonds.<ref name=Niemitz1984c/> |
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Horsfield's Tarsier marks its territory with scents from urine and glandular secretions on a substrate while scratching the surface with its hindlimb toe claws.<ref name=Wright1986a/> |
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==Habitat== |
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This species can live in both primary and secondary forests, and it also lives in forests along the coasts or on the edge of plantations.<ref name=Niemitz1979/> |
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==Conservation status== |
==Conservation status== |
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[[File:Kera hantu Sarawak.jpg|thumb|right|150px|upright|Horsfield's tarsier in Sarawak, wearing a metal collar.]] |
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The rapid loss of habitat due to [[forest conversion]], [[Palm oil|oil palm plantation]]s, [[Wildfire|fire]], and [[logging]] is cause for concern.<ref>{{cite journal | author = Yustian, I. | author2 = Merker, S. | author3 = Supriatna, J | name-list-style = amp | year = 2008 | title = RELATIVE POPULATION DENSITY OF Tarsius dianae IN MAN-INFLUENCED HABITATS | journal = Asian Primates Journal | volume = 1 | issue = 1}}</ref> Additionally, the species is also collected for the [[illegal pet trade]] and wrongly considered a [[Agricultural pest|pest to agricultural crops]]. It can suffer, directly and indirectly, from the use of agricultural [[pesticide]]s. |
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Horsfield's tarsier is listed as [[vulnerable species|vulnerable]] in the 2008 IUCN Red List of Threatened Species,<ref name="iucn status 13 November 2021" /> listed in [[CITES]] Appendix II, and protected by law in Indonesia and in Malaysia. |
|||
The rapid loss of habitat due to forest conversion, oil palm plantations, fire and logging is cause for concern.<ref>{{cite journal | author = Yustian, I., Merker, S., & Supriatna, J | year = 2008 | title = RELATIVE POPULATION DENSITY OF Tarsius dianae IN MAN-INFLUENCED HABITATS | journal = Asian Primates Journal | volume = 1 | issue = 1}}</ref> Additionally, the species is also collected for the illegal pet trade and wrongly considered a pest to agricultural crops. It can suffer, directly and indirectly, from the use of agricultural pesticides. |
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In February 2007, the governments of [[Brunei]], [[Malaysia]], and [[Indonesia]] agreed to protect roughly {{convert|220,000|km2|sqmi|abbr=on}} of tropical forest in the "[[Heart of Borneo]]" region. Environmental group [[World Wide Fund for Nature|WWF]] was particularly active in the establishment of the protected area.<ref>{{cite web | author = Butler, R. A | year = 2006 | title = Borneo | access-date = January 27, 2009 | url = http://www.mongabay.com/borneo.html}}</ref> In the "Heart of Borneo" project, [[non-governmental organization]]s (NGOs) have played a role in promoting the critical initiative and in assisting the neighboring nations in its conceptualization, design, and implementation. The countries are to improve biodiversity conservation in Bornean production forests, and to ensure that such forests are not simply converted to agricultural land-uses such as oil-palm plantations after logging.<ref>World Wildlife Fund. (2007). Resolution concerning the ‘Heart of Borneo’. Association for Tropical Biology and Conservation.</ref> |
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Horsfield's Tarsier is listed as [[vulnerable species|vulnerable]] in the 2008 IUCN Red List of Threatened Species,<ref name=iucn/> listed in [[CITES]] Appendix II, and protected by law in Indonesia and in Malaysia. |
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==Gallery== |
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In February 2007, the governments of [[Brunei]], [[Malaysia]], and [[Indonesia]] agreed to protect roughly 220,000 square kilometers (85,000 square miles) of tropical forest in the so-called "Heart of Borneo". Environmental group [[World Wide Fund for Nature|WWF]] was particularly active in the establishment of the protected area.<ref>{{cite web | author = Butler, R. A | year = 2006 | title = Borneo | accessdate = January 27, 2009 | url = http://www.mongabay.com/borneo.html}}</ref> In the "Heart of Borneo" project, [[non-governmental organization]]s (NGOs) have played role in promoting the critical initiative and in assisting the transboundary nations in its conceptualization, design, and implementation. The transboundary nations are to improve biodiversity conservation in Bornean production forests, and to ensure that such forests are not simply converted to agricultural land-uses such as oil-palm plantations after logging.<ref>World Wildlife Fund. (2007). Resolution concerning the ‘Heart of Borneo’. Association for Tropical Biology and Conservation.</ref> |
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<gallery class="center" mode="nolines" widths="250" noborder="no"> |
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File:Tarsier - Tarsius tarsius - NHMI.jpg|''Cephalopachus bancanus bancanus'' from [[Bangka Island|Bangka]] in [[National Museum of Ireland – Natural History|National Museum of Ireland]], Dublin |
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File:Tarsius di Belitung 06.jpg|''Cephalopachus bancanus saltator'' from [[Belitung]] |
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File:Bornean Tarsier (Cephalopachus bancanus borneanus) (8067063777).jpg|''Cephalopachus bancanus borneanus'' |
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File:Stamp of Indonesia - 2010 - Colnect 248262 - Belitung Island Tarsier Tarsius bancanus saltator.jpeg|[[Postage stamp]] |
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File:Sofyan Efendi Tarsius Mencari Mangsa DSC 0837.jpg |
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File:Stamp of Sarawak - 1950 - Colnect 306198 - Western Tarsier Cephalopachus bancanus.jpeg|Postage stamp featuring the western tarsier and [[George VI]] (top left) |
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File:Tarsius Fransiskus Simbolon.jpg |
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File:Cephalopachus bancanus borneanus 49161649.jpg|In darkness |
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</gallery> |
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==References== |
==References== |
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{{Wikispecies|Tarsius bancanus |
{{Wikispecies|Tarsius bancanus}} |
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{{Commons category|Cephalopachus bancanus}} |
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{{reflist}} |
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{{Reflist|25em}} |
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{{Tarsiidae nav}} |
{{Tarsiidae nav}} |
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{{Haplorhini|Ha.}} |
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{{Taxonbar|from1=Q14518750|from2=Q22113538|from3=Q846009}} |
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[[Category:Mammals of Indonesia]] |
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[[Category:Tarsiers]] |
[[Category:Tarsiers]] |
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[[Category:Primates of Indonesia]] |
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[[Category:Primates of Borneo]] |
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[[de:Sunda-Koboldmaki]] |
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[[Category:Mammals of Brunei]] |
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[[es:Tarsius bancanus]] |
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[[Category:Mammals of Malaysia]] |
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[[eo:Okcidentsunda tarsio]] |
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[[Category:Vulnerable fauna of Asia]] |
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[[eu:Tarsius bancanus]] |
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[[Category:Mammals described in 1821]] |
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[[fr:Tarsius bancanus]] |
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[[Category:Taxa named by Thomas Horsfield]] |
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[[it:Tarsius bancanus]] |
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[[he:קופיף מערבי]] |
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[[hu:Nyugati koboldmaki]] |
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[[nl:Soendaspookdier]] |
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[[pt:Tarsius bancanus]] |
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[[sv:Västligt spökdjur]] |
Latest revision as of 15:39, 22 December 2024
Western tarsier[1] | |
---|---|
C. bancanus in Borneo. Note the eyeshine. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Family: | Tarsiidae |
Genus: | Cephalopachus Swainson, 1835 |
Species: | C. bancanus
|
Binomial name | |
Cephalopachus bancanus (Horsfield, 1821)
| |
Western tarsier range |
Horsfeld's tarsier (Cephalopachus bancanus) is the only species of tarsier in the genus Cephalopachus. Named by American naturalist Thomas Horsfield, it is also referred to as western tarsier. The species occurs on Borneo, Sumatra and nearby islands and is, like other members of the group, entirely nocturnal.
Taxonomy
[edit]Although Horsfeld's tarsier was usually placed in the genus Tarsius with all other living tarsiers, it is quite distinct from the Philippine tarsier and the various tarsiers of Sulawesi and nearby islands; therefore, scientists have placed it in a separate genus, Cephalopachus.[3]
The taxonomy of this species is in doubt, with some subspecies considered unsure.[4] In fact, over 20 years few studies have been done on C. bancanus and a taxonomic revision based upon intensive and systematic field surveys is overdue. The IUCN believes that these subspecies should be treated as distinct and named as separate taxa until more definitive evidence is available.[5] When splitting the species into its own genus, Colin Groves and Myron Shekelle recognized the natunensis population as a distinct subspecies.[3]
There are four recognized subspecies of Horsfeld's tarsier:
- Cephalopachus bancanus bancanus — Bangka Island tarsier
- Cephalopachus bancanus saltator — Belitung Island tarsier
- Cephalopachus bancanus borneanus — Bornean tarsier
- Cephalopachus bancanus natunensis — Natuna Islands tarsier
Habitat and distribution
[edit]Horsfield's tarsier is found in Southern Sumatra, Borneo and nearby islands.[6] The Bornean subspecies, C. b. borneanus, is known from many lowland sites in Sabah, Brunei, Sarawak and West Kalimantan and above 900 m (3,000 ft) in the Kelabit uplands in Northern Sarawak. Other records show it from Kutai and Peleben in East Kalimantan and Tanjung Maruwe in Central Kalimantan.[6] This species can live in both primary and secondary forests, and it also lives in forests along the coasts or on the edge of plantations.[7]
Physical description
[edit]The pelage coloration ranges from pale-olive or reddish brown to pale or dark grey-brown, possibly varying with age. Based on 12 collected specimens,[6] the head to body measurement range from 121–154 mm (4.8–6.1 in). Horsfield's tarsier has an extremely long tail which can reach 181 to 224 mm (7.1 to 8.8 in) and is hairless except for tufts of hair at the end.[8] This species has two grooming claws on each foot. The fingers are very long and have pads on the tips. The toes have flattened nails except for the second and third toes on hind feet, which bear claw-like nails.[6] It has large eyes which do not reflect light.[inconsistent] The membranous ears are slender and almost bare. The molars of this species have high-cusps and are almost tritubercular.[9] The dental formula of Horsfield's tarsier is 2:1:3:3 on the upper jaw and 1:1:3:3 on the lower jaw.[10]
Behaviour and ecology
[edit]Horsfield's tarsier is a nocturnal species. It sleeps alone during the day in a tangle of vines or lianas at a height of 3.5 to 5 metres (11 to 16 ft).[11] This species prefers to sleep, rest, or remain stationary on perches that are angled 5 degrees from the vertical tree trunks, 1 to 4 cm (0.39 to 1.57 in) in diameter,[12] and it sleeps solitarily.[13] Before sunset, Horsfield's tarsier will wake up and wait 10 to 20 minutes before moving around the understory and spending 1.5 to 2 hours of the night foraging for food.[7] Horsfield's tarsier can be found from ground level up to a height of 7 m (23 ft) or more in the understory.[6]
This species is carnivorous. It mainly eats arthropods such as beetles, orthopterans like grasshoppers, katydids, and crickets, butterflies, moths, cockroaches, praying mantis, ants, phasmids, cicadas, dragonflies, freshwater crabs, and spiders,[7][14] but also will eat small vertebrates such as flying frogs,[14] bats (Chiroptera) including members of the genus Taphozous, the lesser short-nosed fruit bat (Cynopterus brachyotis), and the spotted-winged fruit bat (Balionycteris maculata), and snakes, of which venomous snakes have been found to be consumed, such as the elapid Calliophis intestinalis. This species was also found to consume birds, including spiderhunters, warblers, kingfishers, and pittas. It locates prey primarily by sound and catches the prey with its hands when foraging. Prey is killed through bites to the back of the neck[7] and the tarsier's eyes are shut when attacking.[15] It will consume the prey starting with the head and working its way down the body.[7] This species gets water both by drinking from a pool or stream, and by licking drops from bamboo leaves or from water running down the trunks of trees.[16] Horsfield's tarsier is a host of the acanthocephalan intestinal parasite Moniliformis tarsii.[17]
Like all tarsiers, Horsfield's tarsiers are vertical clingers and leapers, known for extraordinary leaping abilities. An individual will mainly support itself with its feet and the tail, both exerting enough force to hold the individual in place. The hands are not always used because of the pads located on the feet provide adequate grip. The hands are usually placed no higher than its nose except when resting; the hands are only placed higher up to maintain the position of the individual.[12] Other modes of locomotion used by the species include climbing, quadrupedal walking, hopping and "cantilevering."[11][14]
Horsfield's tarsiers are monogamous, with a copulation frequency during estrus of once per night.[18] Courtship calls are performed by the male and he emits 2–3 chirrups while opening and closing the mouth.[18] This call happens within 5 minutes of looking at the female. Once the male gives his courtship call, if the female is receptive, she will perform genital displays to him.[18] If the female is not in estrus, she will emit an agonistic call which is often followed by biting and pushing the male away.[18] Both sexes' calls last on average for 1 second, and the interval between calls is on average 3 seconds.[18]
Infants are born with their eyes open and fully furred and able to groom themselves.[19] The mother will carry her infant in her mouth and when she forages for food the mother will park the infant on a branch.[20] Infant sounds are mostly clicks: "k", "tk", "ki", or a rapid "kooih"[21] and can be heard when the infant is left alone or is cold.[21] The mother stays in contact with their infants using high-pitched calls. Infants were found to first use the tails as support during resting at 7–10 days.[19] Social grooming in this species only occurs between mothers and infants,[22] removing dead skin and parasites by scratching with their toe claws and licking their fur, avoiding their faces. Faces are cleaned by rubbing on branches and it is to reinforce social bonds.[22]
Young leave their mother's range at the onset of puberty, and find their own territory.[19] Horsfield's tarsiers mark their territory with scents from urine and glandular secretions on a substrate while scratching the surface with its hind-limb toe claws.[18]
Conservation status
[edit]The rapid loss of habitat due to forest conversion, oil palm plantations, fire, and logging is cause for concern.[23] Additionally, the species is also collected for the illegal pet trade and wrongly considered a pest to agricultural crops. It can suffer, directly and indirectly, from the use of agricultural pesticides.
Horsfield's tarsier is listed as vulnerable in the 2008 IUCN Red List of Threatened Species,[2] listed in CITES Appendix II, and protected by law in Indonesia and in Malaysia.
In February 2007, the governments of Brunei, Malaysia, and Indonesia agreed to protect roughly 220,000 km2 (85,000 sq mi) of tropical forest in the "Heart of Borneo" region. Environmental group WWF was particularly active in the establishment of the protected area.[24] In the "Heart of Borneo" project, non-governmental organizations (NGOs) have played a role in promoting the critical initiative and in assisting the neighboring nations in its conceptualization, design, and implementation. The countries are to improve biodiversity conservation in Bornean production forests, and to ensure that such forests are not simply converted to agricultural land-uses such as oil-palm plantations after logging.[25]
Gallery
[edit]-
Cephalopachus bancanus bancanus from Bangka in National Museum of Ireland, Dublin
-
Cephalopachus bancanus saltator from Belitung
-
Cephalopachus bancanus borneanus
-
Postage stamp featuring the western tarsier and George VI (top left)
-
In darkness
References
[edit]- ^ Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 127. ISBN 0-801-88221-4. OCLC 62265494.
- ^ a b Shekelle, M.; Yustian, I. (2020). "Cephalopachus bancanus". IUCN Red List of Threatened Species. 2020: e.T21488A17976989. doi:10.2305/IUCN.UK.2020-3.RLTS.T21488A17976989.en. Retrieved 13 November 2021.
- ^ a b Groves, C.; Shekelle, M. (2010). "The Genera and Species of Tarsiidae". International Journal of Primatology. 31 (6): 1071–1082. doi:10.1007/s10764-010-9443-1. S2CID 21220811.
- ^ Brandon-Jones, D.; Eudey, A.A.; Geissmann, T.; Groves, C.P.; Melnick D.J., Morales; J.C., Shekelle, M. & Stewart, C.B (2004). "Asian Primate Classification". International Journal of Primatology. 25 (1): 97–164. doi:10.1023/B:IJOP.0000014647.18720.32. S2CID 29045930.
{{cite journal}}
: CS1 maint: multiple names: authors list (link) - ^ Shekelle, M.; Yustian, I. (2021). "Cephalopachus bancanus ssp. bancanus". IUCN Red List of Threatened Species. 2021: e.T39762A17992163. doi:10.2305/IUCN.UK.2021-1.RLTS.T39762A17992163.en. Retrieved 13 November 2021.
- ^ a b c d e Payne, J.; Francis, C. M. & Phillipps, K (1994). A Field guide to the Mammals of Borneo. Petaling Jaya: The Sabah Society.
- ^ a b c d e Niemitz, C. (1979). "Outline of the behavior of Tarsius bancanus". In G.A. Doyle; R.D. Martin (eds.). The Study of Prosimian Behavior. New York: Academic Press.
- ^ Sussman, R.W (1999). Primate Ecology and Social Structure: Vol. 1 Lorises, Lemurs and Tarsiers. Pearson Custom Publishing.
- ^ Crompton, R.H.; Savage, R. & Spears, I.R (1998). "The mechanics of food reduction in Tarsius bancanus: Hard-object feeder, soft-object feeder or both?". Folia Primatologica. 69 (suppl 1): 41–59. doi:10.1159/000052698. PMID 9595687. S2CID 24464173.
- ^ Nowak, R.M (1999). Walker's Primates of the World. Baltimore: Johns Hopkins University Press. ISBN 0-8018-6251-5.
- ^ a b Crompton, R.H.; Andau, P.M (1986). "Locomotion and Habitat Utilization in Free-ranging Tarsius bancanus: A Preliminary Report". Primates. 27 (3): 337–355. doi:10.1007/BF02382075. S2CID 25836867.
- ^ a b Niemitz, C (1984a). "Locomotion and posture of Tarsius bancanus". In C. Niemitz (ed.). Biology of Tarsiers. Stuttgart: Gustav Fischer Verlag. ISBN 0-89574-182-2.
- ^ Crompton, R.H.; Andau, P.M (1987). "Ranging, activity rhythms, and sociality in free-ranging Tarsius bancanus: A preliminary report". International Journal of Primatology. 8 (1): 43–71. doi:10.1007/BF02737113. S2CID 39920648.
- ^ a b c Crompton, Robin Huw; Blanchard, Mary L.; Coward, Sam; Alexander, R. Mcneill; Thorpe, Susannah K. (2010). "Vertical Clinging and Leaping Revisited: Locomotion and Habitat Use in the Western Tarsier, Tarsius bancanus Explored Via Loglinear Modeling". International Journal of Primatology. 31: 958–979. doi:10.1007/s10764-010-9420-8. Retrieved 21 November 2024.
- ^ Fogden, M.P.L (1974). "A preliminary field study of the western tarsier, Tarsius bancanus Horsefield". In R.D. Martin; G.A. Doyle; A.C. Walker (eds.). Prosimian Biology. Pittsburgh: University of Pittsburgh Press.
- ^ Niemitz, C (1984d). "Synecological relationships and feeding behaviour of the genus Tarsius". In C. Niemitz (ed.). Biology of Tarsiers. Stuttgart: Gustav Fischer Verlag. ISBN 0-89574-182-2.
- ^ Deveaux, Timothy P.; Schmidt, Gerald D.; Krishnasamy, M. (1988). "Two New Species of Moniliformis (Acanthocephala: Moniliformidae) from Malaysia". The Journal of Parasitology. 74 (2): 322–5. doi:10.2307/3282462. JSTOR 3282462. PMID 3128654.
- ^ a b c d e f Wright, P.C.; Toyama, L.M. & Simons, E.L (1986). "Courtship and Copulation in Tarsius bancanus". Folia Primatologica. 46 (3): 142–148. doi:10.1159/000156247. PMID 3100401.
- ^ a b c Roberts, M (1994). "Growth, development, and parental care in the Western tarsier (Tarsius bancanus) in captivity: Evidence for a "slow" life-history and nonmonogamous mating system". International Journal of Primatology. 15 (1): 1–28. doi:10.1007/BF02735232. S2CID 7462413.
- ^ Haring, D.M.; Wright, P.C. & Simons, E.L (1985). "Social behaviors of Tarsius syrichta and Tarsius bancanus". American Journal of Physical Anthropology. 66: 179. doi:10.1002/ajpa.1330660204.
- ^ a b Niemitz, C (1984b). "Vocal communication of two tarsier species (Tarsius bancanus and Tarsius spectrum)". In C. Niemitz (ed.). Biology of Tarsiers. Stuttgart: Gustav Fischer Verlag. ISBN 0-89574-182-2.
- ^ a b Niemitz, C (1984c). "An investigation and review of the territorial behaviour and social organisation of the genus Tarsius". In C. Niemitz (ed.). Biology of Tarsiers. Stuttgart: Gustav Fischer Verlag. ISBN 0-89574-182-2.
- ^ Yustian, I.; Merker, S. & Supriatna, J (2008). "RELATIVE POPULATION DENSITY OF Tarsius dianae IN MAN-INFLUENCED HABITATS". Asian Primates Journal. 1 (1).
- ^ Butler, R. A (2006). "Borneo". Retrieved January 27, 2009.
- ^ World Wildlife Fund. (2007). Resolution concerning the ‘Heart of Borneo’. Association for Tropical Biology and Conservation.