Tarsier
Tarsiers[1][2] Temporal range: Eocene to Recent
Middle | |
---|---|
Philippine tarsier (Carlito syrichta) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Tarsiiformes |
Family: | Tarsiidae Gray, 1825 |
Type genus | |
Tarsius | |
Genera | |
Tarsiers are haplorrhine primates of the family Tarsiidae, which is itself the lone extant family within the infraorder Tarsiiformes. Although the group was once more widespread, all of its species living today are found in the islands of Southeast Asia, specifically the Philippines, Malaysia, Indonesia, and Brunei.[3]
They are found primarily in forested habitats, especially forests that have liana, since they give them vertical support when climbing trees.[4]
Evolutionary history
Fossil record
Fossils of tarsiiform primates are found in Asia, Europe, and North America, with disputed fossils from Africa, but extant tarsiers are restricted to several Southeast Asian islands in Indonesia, Philippines, and Malaysia, The fossil record indicates that their dentition has not changed much, except in size, in 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,[5] and it is sometimes listed in its own family, Afrotarsiidae, within the infraorder Tarsiiformes,[6] or considered an anthropoid primate.[7]
So far, four fossil species of tarsiers are known from the fossil record:
- Tarsius eocaenus is known from the Middle Eocene in China.[5][8]
- Hesperotarsius thailandicus lived during the Early Miocene in northwestern Thailand.[5][9]
- Hesperotarsius sindhensis lived during the Miocene in Pakistan.[10]
- Tarsius sirindhornae lived during the Middle Miocene in northern Thailand.[7]
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 questionable.[11]
Classification
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 Haplorrhini. Analysis of SINE insertions, a type of macromutation to the DNA, is argued to offer very persuasive evidence for the monophyly of Haplorrhini, 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 confers the need for vitamin C 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 haplorrhines.[12]
At a lower phylogenetic level, the tarsiers have, until recently, all been placed in the genus Tarsius,[1] 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).[13] Species level taxonomy is complex, with morphology often being of limited use compared to vocalizations.[citation needed] 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.[1]
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.[2]
In 2014, scientists from the University of the Philippines (Diliman Campus) – Institute of Biology in partnership with the University of Kansas have discovered a distinct genus of Philippine tarsier. The genetically distinct populations are found in the Dinagat Islands, Surigao del Norte, and probably Siargao Islands in Mindanao Island's northeast portion. Isolation is the key to the population's distinctiveness. Prior to the study, scientists generally accepted three subspecies of Philippine tarsier: the large island of Mindanao contained one subspecies, Tarsius syrichta carbonarius; while the islands of Samar and Leyte sported another, Tarsius syrichta syrichta; and Bohol held the third, Tarsius syrichta fraterculus. However the new genetic research found the relationships among the Philippine tarsier populations was even messier. Looking at mitochondrial and nuclear DNA, Brown's team uncovered three different evolutionary lineages: one lineage of tarsier makes their home on Bohol, Samar, and Leyte Islands (putting two presently accepted Philippine tarsier subspecies into a single subspecies); another has conquered the vast majority of Mindanao; while a long-cryptic branch has evolved in northeastern Mindanao and Dinagat Island (the new subspecies). For the purposes of the paper, the scientists refer to this as the Dinagat-Caraga tarsier. Rafe Brown of the University of Kansas' Biodiversity Institute, an author of the study, also said that through a more keen study, the only current Philippine tarsier species, Carlito syrichta, could be split into three distinct full species in the future.[14][15]
- Infraorder Tarsiiformes
- Family Tarsiidae: tarsiers
- Genus Carlito
- Philippine tarsier, Carlito syrichta
- C. s. syrichta (to be combined with C. s. fraterculus)
- C. s. fraterculus (to be combined with C. s. syrichta)
- C. s. carbonarius
- Philippine tarsier, Carlito syrichta
- Genus Cephalopachus
- Horsfield's tarsier, Cephalopachus bancanus
- C. b. bancanus
- C. b. natunensis
- C. b. boreanus
- C. b. saltator
- Horsfield's tarsier, Cephalopachus bancanus
- Genus Tarsius
- Dian's tarsier, T. dentatus
- Makassar tarsier T. fuscus
- Lariang tarsier, T. lariang
- Niemitz’s tarsier, T. niemitzi
- Peleng tarsier, T. pelengensis
- Sangihe tarsier, T. sangirensis
- Gursky's spectral tarsier, T. spectrumgurskyae[16]
- Jatna's tarsier, T. supriatnai[16]
- Spectral tarsier, T. tarsier
- Siau Island tarsier, T. tumpara
- Pygmy tarsier, T. pumilus
- Wallace's tarsier, T. wallacei
- Genus Carlito
- Family Tarsiidae: tarsiers
Anatomy and physiology
Tarsiers are small animals with enormous eyes; each eyeball is approximately 16 millimetres (0.63 in) in diameter and is as large as, or in some cases larger than, its entire brain.[17][18] 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.[19] Tarsiers have a strong auditory sense, and their auditory cortex is distinct.[19] Tarsiers also have long hind limbs, owing mostly to the elongated 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.[20] 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.[21]
Tarsiers morphology allows for them to move their heads 180 degrees in either direction, allowing for them to see 360 degrees around them.[22]
Their dental formula is also unique: 2.1.3.31.1.3.3
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]
Behavior
Tarsiers are the only extant entirely carnivorous primates: they are primarily insectivorous, and catch insects by jumping at them. Their favorite prey are arthropods like beetles, spiders, cockroaches, grasshoppers, and walking sticks.[22] They are also known to prey on birds, snakes, lizards, and bats.[21]
Pygmy tarsiers differ from other species in terms of their morphology, communication, and behavior.[26] The differences in morphology that distinguish pygmy tarsiers from other species are likely based on their high altitude environment.[27]
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.[28][29][30][31]
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.[32] For example, the colder climate at higher elevations can influence cranial morphology.[33]
Gestation takes about six months,[34] 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.
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]
Due to their small size, tarsiers are prey to snakes, owls, lizards, and cats. When a predator is present the tarsiers surround the threat vocalizing and attacking it. While tarsier groups only contain one male, when confronting a threat other groups will join, meaning there are multiple alpha male tarsiers attacking.[22]
Conservation
Tarsiers have never formed successful breeding colonies in captivity. This may be due in part to their special feeding requirements.[37][38][39][40][41]
A sanctuary near the town of Corella, on the Philippine island of Bohol, is having some success restoring tarsier populations.[42] The Philippines Tarsier Foundation (PTFI) has developed a large, semi-wild enclosure known as the Tarsier Research and Development Center. Carlito Pizarras, also known as the "Tarsier man", founded this sanctuary where visitors can observe tarsiers in the wild. As of 2011, the sanctuary was maintained by him and his brother.[citation needed] The trees in the sanctuary are populated with nocturnal insects that make up the tarsier's diet.[43]
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 definitely become extinct in the future.[19]
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.[44] The Malaysian government protects tarsiers by listing them in the Totally Protected Animals of Sarawak, the Malaysian state in Borneo where they are commonly found.[45]
A new scheme to conserve the tarsiers of Mount Matutum near Tupi in South Cotabato on the island of Mindanao is being organised by the Tupi civil government and the charity Endangered Species International (ESI). Tarsier UK are also involved on the margins helping the Tupi Government to educate the children of Tupi about the importance of the animal. ESI are hoping to build a visitor centre on the slopes of Mount Matutum and help the local indigenous peoples to farm more environmentally and look after the tarsiers. The first stage in this is educating the local peoples on the importance of keeping the animal safe and secure. A number of native tarsier-friendly trees have been replanted on land which had been cleared previously for fruit tree and coconut tree planting.[citation needed]
References
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- ^ 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.
- ^ The Editors of Encyclopædia Britannica. “Tarsier.” Encyclopædia Britannica, Encyclopædia Britannica, Inc., 14 Apr. 2019, http://www.britannica.com/animal/tarsier.
- ^ Simeon, S.G.F.; Duya, M.R.M; Duya, M.V.; Galindon, J.M.M.; Pasion, B.O.; Ong, P.S. (2020). "Living in small spaces: Forest fragment characterization and its use by Philippine tarsiers (Tarsius syrichta Linnaeus, 1758) in Mindanao Island, Philippines". Primates. 61 (3): 529–542. doi:10.1007/s10329-020-00798-2. PMID 32043166. S2CID 211075031.
- ^ a b c Gunnell, G.; Rose, K. (2002). "Tarsiiformes: Evolutionary History and Adaptation". In Hartwig, W.C. (ed.). The Primate Fossil Record. Cambridge University Press. ISBN 978-0-521-66315-1.
- ^ 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
- ^ a b Chiamanee, Y., Lebrun, R., Yamee, C., and Jaeger, J.-J. (2010). "A new Middle Miocene tarsier from Thailand and the reconstruction of its orbital morphology using a geometric–morphometric method". Proceedings of the Royal Society B: Biological Sciences. 278 (1714): 1956–1963. doi:10.1098/rspb.2010.2062. PMC 3107645. PMID 21123264.
{{cite journal}}
: CS1 maint: multiple names: authors list (link) - ^ Rossie, J.B.; Ni, X.; Beard, K.C. (2006). "Cranial remains of an Eocene tarsier" (PDF). PNAS. 103 (12): 4381–4385. doi:10.1073/pnas.0509424103. PMC 1450180. PMID 16537385.
- ^ Nowak, R.M. (1999). Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. pp. 94–97. ISBN 978-0-8018-5789-8.
- ^ Zijlstra, Jelle S.; Flynn, Lawrence J.; Wessels, Wilma (2013). "The westernmost tarsier: A new genus and species from the Miocene of Pakistan". Journal of Human Evolution. 65 (5): 544–550. doi:10.1016/j.jhevol.2013.06.015. PMID 23928350.
- ^ Simons, E.L. (2003). "The Fossil Record of Tarsier Evolution". In Wright, P.C.; Simons, E.L.; Gursky, S. (eds.). Tarsiers: past, present, and future. ISBN 978-0-8135-3236-3.
- ^ Pollock, J. I. & Mullin, R. J. (1986). "Vitamin C biosynthesis in prosimians: Evidence for the anthropoid affinity of Tarsius". American Journal of Physical Anthropology. 73 (1): 65–70. doi:10.1002/ajpa.1330730106. PMID 3113259. Archived from the original on 2012-06-28.
- ^ Brandon-Jones, D.; et al. (2004). "Asian primate classification". International Journal of Primatology. 25 (1): 97–164. doi:10.1023/B:IJOP.0000014647.18720.32. S2CID 29045930.
- ^ Jeremy Hance (2014-08-21). "Have scientists discovered a new primate in the Philippines?". Mongabay Environmental News.
- ^ "We have a new tarsier species – UP biologists". Archived from the original on 2015-07-08. Retrieved 2015-07-07.
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- ^ Soluri, K. Elizabeth; Sabrina C. Agarwal (2016). The Laboratory Manual and Workbook for Biological Anthropology. W.W. Norton. ISBN 978-0-393-91291-3.
- ^ Shumaker, Robert W.; Benjamin B. Beck (2003). Primates in Question. Smithsonian Books. ISBN 978-1-58834-151-8.
- ^ a b c Shekelle, Myron; Gursky (2010). "Why tarsiers? Why now? An introduction to the special edition on tarsiers". International Journal of Primatology. 31 (6): 937–940. doi:10.1007/s10764-010-9459-6. S2CID 326565.
- ^ Rasmussen, D. T.; Conroy, G. C.; Simons, E. L. (1998). "Tarsier-like locomotor specializations in the Oligocene primate Afrotarsius". Proceedings of the National Academy of Sciences of the United States of America. 95 (25): 14848–14850. doi:10.1073/pnas.95.25.14848. PMC 24538. PMID 9843978.
- ^ a b Niemitz, Carsten (1984). Macdonald, D. (ed.). The Encyclopedia of Mammals. New York: Facts on File. pp. 338–339. ISBN 978-0-87196-871-5.
- ^ a b c Gron KJ. 2010 December 1. Primate Factsheets: Tarsier (Tarsius) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/tarsier>. Accessed 2019 November 12.
- ^ Rosa, M. G.; Pettigrew J. D.; Cooper H. M. (1996). "Unusual pattern of retinogeniculate projections in the controversial primate Tarsius". Brain, Behavior and Evolution. 48 (3): 121–129. doi:10.1159/000113191. PMID 8872317.
- ^ Collins, C. E.; Hendrickson, A.; Kaas, J. H. (2005). "Overview of the visual system of tarsius". The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology. 287 (1): 1013–1025. doi:10.1002/ar.a.20263. PMID 16200648.
- ^ Ramsier, Marissa A.; Cunningham A.J.; Moritz G.L.; Finneran J.J.; Williams C.V.; Ong P.S.; Gursky-Doyen S.L.; Dominy N.J. (2012). "Primate communication in the pure ultrasound". Biology Letters. 8 (4): 508–11. doi:10.1098/rsbl.2011.1149. PMC 3391437. PMID 22319094.
- ^ Grow, Nanda; Gursky-Doyen, Sharon (2010). "Preliminary Data On The Behavior, Ecology, And Morphology Of Pygmy Tarsiers ( Tarsius Pumilus)". International Journal of Primatology. 31 (6): 1174–1191. doi:10.1007/s10764-010-9456-9. S2CID 23939646.
- ^ Musser, G. G.; Dagosto, M. (1987). "The identity of Tarsius pumilus, a pygmy species endemic to the montane mossy forests of Central Sulawesi". American Museum Novitates. 2867: 1–53.
- ^ Dagosto, M.; Gebo, D. L.; Dolino, C. (2001). "Positional behavior and social organization of the Philippine tarsier (Tarsius syrichta)". Primates. 42 (3): 233–243. doi:10.1007/bf02629639. S2CID 41499839.
- ^ Niemitz, C (1977). "Zur funktionsmorphologie und biometrie der gattung Tarsius, Storr, 1780". Courier Forschungsinstitut Senckenberg. 25: 1–161.
- ^ 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.
- ^ 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
- ^ Körner, C (2007). "The use of 'altitude' in ecological research". Trends in Ecology and Evolution. 22 (11): 569–574. doi:10.1016/j.tree.2007.09.006. PMID 17988759.
- ^ Rae, T. C.; Hill, R. I.; Hamada, Y.; Koppe, T. (2003). "Clinal variation of maxillary sinus volume in Japanese macaques (Macaca fuscata)". American Journal of Primatology. 59 (4): 153–158. doi:10.1002/ajp.10072. PMID 12682923. S2CID 12290499.
- ^ Izard, Kay M.; Wright, Simons (1985). "Gestation length in Tarsius bancanus". Am J Primatol. 9 (4): 327–331. doi:10.1002/ajp.1350090408. PMID 31979510. S2CID 83711759.
- ^ Jarosz, Andy (2019-05-20). "The Philippines: The Tarsier Man". National Geographic. Archived from the original on 2019-05-21. Retrieved 2019-09-28.
- ^ Sinclair, Joe (2011-12-12). "Tourism threatens tiny Philippine primate". My SinChew. AFP. Archived from the original on 2019-09-28. Retrieved 2019-09-28.
- ^ Roberts, M.; Kohn, F. (1993). "Habitat Use, Foraging Behavior, and Activity Patterns in Reproducing Western Tarsiers, Tarsius bancanus, in Captivity: A Management Synthesis" (PDF). Zoo Biology. 12 (2): 217–232. doi:10.1002/zoo.1430120207.
- ^ Shekelle, M.; Nietsch, A. (2008). Shekelle, M.; Maryano, T.; Groves, C.; Schulze, H.; Fitch-Snyder, H. (eds.). Tarsier Longevity: Data from a Recapture in the Wild and from Captive Animals (PDF). LIPI Press. pp. 85–89. ISBN 978-979-799-263-7. Archived from the original (PDF) on 28 July 2011.
{{cite book}}
:|work=
ignored (help) - ^ Severn, K.; Dahang, D.; Shekelle, M. (2008). Shekelle, M.; Maryano, T.; Groves, C.; Schulze, H.; Fitch-Snyder, H. (eds.). Eastern Tarsiers in Captivity, Part I: Enclosure and Enrichment (PDF). LIPI Press. pp. 91–96. ISBN 978-979-799-263-7. Archived from the original (PDF) on 24 July 2011.
{{cite book}}
:|work=
ignored (help) - ^ Severn, K.; Dahang, D.; Shekelle, M. (2008). Shekelle, M.; Maryano, T.; Groves, C.; Schulze, H.; Fitch-Snyder, H. (eds.). Eastern Tarsiers in Captivity, Part II: A Preliminary Assessment of Diet (PDF). LIPI Press. pp. 97–103. ISBN 978-979-799-263-7. Archived from the original (PDF) on 28 July 2011.
{{cite book}}
:|work=
ignored (help) - ^ Fitch-Snyder, H. (2003). "History of Captive Conservation of Tarsiers". In Wright, P.C.; Simons, E.L.; Gursky, S. (eds.). Tarsiers: Past, Present, and Future. Rutgers University Press. pp. 227–295. ISBN 978-0-8135-3236-3.
- ^ "StephenMBland". Retrieved 18 October 2016.
- ^ Jachowski, David S.; Pizzaras, Carlito (2005). "Introducing an innovative semi-captive environment for the Philippine tarsier (Tarsius syrichta)". Zoo Biology. 24 (1): 101–109. doi:10.1002/zoo.20023.
- ^ Shekelle, Myron; Salim, Agus. "Siau Island Tarsier". IUCN/SSC Primate Specialist Group. Archived from the original on 6 September 2010. Retrieved 1 January 2010.
- ^ "Totally Protected Animals of Sarawak". Forestry Department of Sarawak. Retrieved 1 January 2010.[permanent dead link ]
External links
- Philippine Tarsier Foundation
- Tarsier.org, an international research and conservation project
- Tarsiers (Tarsiidae) at the Wayback Machine (archived October 15, 2004), Singapore Zoological Gardens Docents, 2000
- Gron, Kurt J. (July 2008). "Primate Factsheets: Tarsier (Tarsius) Taxonomy, Morphology, & Ecology". National Primate Research Center, University of Wisconsin–Madison.
- Tarsier UK, Charitable Organisation for the Philippine Tarsier
- Tarsiers – Visiting the two Tarsier sanctuaries in Bohol, Philippines
- Tarsier skeleton – Skeleton from the University of Texas at Austin