Chladobates Schinz, 1824
Tupaia is a treeshrew genus in the family Tupaiidae that was first described by Thomas Stamford Raffles in 1821. The name of this genus derives from the Malay word tupai meaning squirrels or small animals that resemble squirrels.
Raffles described the genus as having an elongated snout, eight to 10 incisors, well-developed limbs, five-toed naked feet, and the sole furnished with projecting pads and sharp claws, with a habit and tail of a squirrel.
Marcus Ward Lyon published a revision of the genus in 1913, and also noted the squirrel-like appearance of Tupaia species, which only lack the long black whiskers and have smaller ears. They do not have any markings on the face, the naked area of the nose is finely reticulated, an oblique stripe on the shoulder is more or less distinct, and the tail is haired but not tufted. The braincase is about as wide as the maxillary tooth row is long. The temporal fossa is smaller than the orbit. The dental formula is 188.8.131.52. The first pair of upper incisors is longer than the second, while the second pair of lower incisors is slightly larger than the first and third pairs. The lower canines are better developed than the upper ones and stand high above the adjacent premolars. The size of head, body, and tail varies between species.
Classification and taxonomic history
Subsequent to Raffles' description of the genus, these additional Tupaia species were described:
- Northern treeshrew T. belangeri — by Wagner in 1841
- Golden-bellied treeshrew T. chrysogaster — by Miller in 1903
- Striped treeshrew T. dorsalis — by Schlegel in 1857
- Slender treeshrew T. gracilis — by Thomas in 1893
- Horsfield's treeshrew T. javanica — by Horsfield in 1821
- Long-footed treeshrew T. longipes — by Thomas in 1893
- Pygmy treeshrew T. minor — by Günther in 1876
- Calamian treeshrew T. möllendorffi — by Matschie in 1898
- Mountain treeshrew T. montana — by Thomas in 1892
- Nicobar treeshrew T. nicobarica — by Zelebor in 1868
- Palawan treeshrew T. palawanensis — by Thomas in 1894
- Painted treeshrew T. picta — by Thomas in 1892
- Ruddy treeshrew T. splendidula — by Gray in 1865
- Large treeshrew T. tana — by Raffles in 1821
- †Tupaia miocenica — by Mein and Ginsburg in 1997
In the past, various authors proposed to place treeshrews in the ordinal rank Insectivora, or considered them close relatives of primates. Since 1972, the treeshrew families Tupaiidae and Ptilocercidae are grouped in the order Scandentia.
Distribution and habitat
Tupaia species range from northeastern India, Burma, and Nicobar Islands eastward to members of the Philippine Islands, and from central China south to Java, Borneo, and Sumatra, including islands on the southwest coast. They do not occur on Celebes, nor on islands to the east of Java, with the possible exception of Bali. They inhabit the dense undergrowth of tropical forests. With the exception of T. minor, they are primarily terrestrial and forage on the forest floor, usually below 1.5 m (4.9 ft). Since they are rarely seen crossing wide roads, populations likely are negatively affected by fragmentation of forests caused by logging operations.
Ecology and behaviour
Early naturalists described wild-caught captive Tupaia specimens as restless, nervous, and rapidly reacting to sounds and movements. Their auditory sensitivity is highly developed as the broad frequency range of their hearing reaches far into the ultrasonic.
The shape of the cheek-teeth of Tupaia species indicate they are foremost insectivores. Captive specimens were reported to hunt ants, flies, crickets, grasshoppers, cockroaches, and small beetles. They hold their food between their forelegs while sitting on their haunches. After feeding, they smooth their heads and faces with both forepaws, and lick their lips and palms. They are also fond of water, both to drink and to bathe. They fortify their diet with soft fruits that are mostly dispersed by birds. They swallow the pulp, but reject fibrous components, which they cannot digest due to their long and small intestines and rudimentary ceca.
The well-developed olfactory perception of treeshrews enables them to easily detect food among the leaf litter on the forest floor. Their sensitivity for odours coupled with scent-marking of their territories is important in their interaction with conspecifics.
Observations of Tupaia species in their natural habitats suggest they usually form monogamous pairs. Social behaviour differs between species and the available food resources in their territories. Where food is adequate and sufficient, they tolerate conspecifics without engaging in territorial disputes. Common treeshrews observed in the Bukit Timah Nature Reserve defended diminishing food resources by chasing away residents of adjacent areas.
Birds of prey, snakes, and small carnivores are known to hunt treeshrews. Humans have no interest in killing them for food because of their unpleasant taste, and they are rarely seen as pests.
Reproduction and development
Treeshrews share more similarity with rodents and squirrels than with primates in regards to their reproduction and development. In contrast to primates, which produce one baby with longer gestation periods, treeshrews generally have litters of two or three young and are only in utero about 45 days. Female treeshrews give birth in nests made of many dry leaves, and are known to leave the young unattended while returning occasionally to give them milk. Parental care of Tupaia is relatively limited. The young remain in the nest for 33 days on average, developing gradually before they exit the nest. Ten identified embryonic developmental stages are seen in Tupaia species. Baby treeshrews are sexually immature until about 90 days after birth.
Their close relationship to primates makes treeshrews important model organisms in human medical research. A study investigating the effects of the Borna disease virus on treeshrews has given new insight into neurological disease. Since Tupaia species share so many similarities with primates, yet are more abundant and have more plentiful progeny than them, interest is increasing in using them as an alternative model for use in human medical research. Successful psychosocial studies were carried out, and dramatic behavioral, neuroendocrinal, and physiological changes occurred in subordinate males of Tupaia, similar to depressed human patients. Their susceptibility to viruses has also piqued interest in using them to study immune responses to infections such as hepatitis B. Tupaia species have been used to overcome the limitations of using rodent models in the study of human biology and disease mechanisms, as well as the development of new drugs and diagnostic tools. Recent studies have used treeshrews to study infectious, metabolic, neurological, and psychiatric diseases, as well as cancers.
In 2013, the Virology Journal published an article that documents the use of the northern treeshrew (T. belangeri) as medical models for the H1N1 influenza virus. This was advantageous because other possible candidates such as guinea pigs, rats, mice, and other rodents leave gaps in the information, especially regarding clinical symptoms and transmission. Tupaia, though, displays moderate systemic and respiratory symptoms, as well as pathological changes in the respiratory tract, supporting its use as a beneficial model in H1N1 research.
- Helgen, K.M. (2005). "Tupaia". In Wilson, D.E.; Reeder, D.M. Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. p. 104. ISBN 978-0-8018-8221-0. OCLC 62265494.
- Raffles, T. S. (1821). "Descriptive Catalogue of a Zoological Collection made on account of the Honourable East India Company, in the Island of Sumatra and its Vicinity, under the Direction of Sir Thomas Stamford Raffles, Lieutenant-Governor of Fort Marlborough; with additional Notices illustrative of the Natural History of those Countries.". The Transactions of the Linnean Society of London. Linnean Society of London. XIII: 239–340.
- Wilkinson, R. J. (1901). A Malay-English dictionary Kelly & Walsh Limited, Hongkong, Shanghai and Yokohama.
- Lyon, M. W., Jr. (1913). Tree shrews: An account of the mammalian family Tupaiidae. Proceedings of the United States National Museum, 45:1–188.
- Shriver, J .G., Noback, C. R. (1967). Color Vision in the Tree Shrew (Tupaia glis). Folia Primatologia 6: 161−169.
- Diard, P.M., Duvaucel, A. (1820). "Sur une nouvelle espèce de Sorex — Sorex Glis". Asiatick researches, or, Transactions of the society instituted in Bengal, for inquiring into the history and antiquities, the arts, sciences, and literature of Asia, Volume 14: 470–475.
- Wagner, J. A. (1841). Das peguanische Spitzhörnchen. In: Die Säugethiere in Abbildungen nach der Natur mit Beschreibungen. Supplementband 2. Erlangen: Expedition des Schreber'schen Säugethier- und des Esper'schen Schmetterlingswerkes. Pp. 42–43.
- Miller, G. S. Jr. (1903). Seventy New Malayan Mammals. Smithsonian Miscellaneous Collections 45: 1–73.
- Schlegel, H. (1857). Tana dorsalis. In: Handleiding Tot de Beoefening der Dierkunde, Ie Deel. Boekdrukkerij van Nys, Breda. Page 59.
- Thomas, O. (1893). Description of a new Bornean Tupaia. The Annals and Magazine of Natural History 6 (12): 53–54.
- Horsfield, T. (1824). Zoological researches in Java, and the neighbouring islands. London: Kingsbury, Parbury, & Allen.
- Thomas, O. (1893). On some new Bornean Mammalia. The Annals and Magazine of Natural History, 6 (11): 341–347.
- Günther, A. G. (1876). Remarks on some Indian and, more especially, Bornean Mammals. Proceedings of the general meetings for scientific business of the Zoological Society of London: 424–428.
- Matschie, P. (1898). Säugethiere von den Philippinen. Sitzungsbericht der Gesellschaft Naturforschender Freunde zu Berlin: 38–43.
- Thomas, O. (1892). On some new Mammalia from the East-Indian Archipelago. The Annals and Magazine of Natural History 6 (9): 250–254.
- Zelebor, J. (1868). Cladobates Nicobaricus. In: Reise der österreichischen Fregatte Novara um die Erde. Zoologischer Theil, Band 1 Säugethiere. Wien: Kaiserliche Akademie der Wissenschaften. Pp. 17–19.
- Thomas, O. (1894). On the Palawan Representative of Tupaia ferruginea. The Annals and Magazine of Natural History 6 (13): 367.
- Gray, J. E. (1865). Notice of a Species of Tupaia from Borneo, in the Collection of the British Museum. Proceedings of the general meetings for scientific business of the Zoological Society of London: 322.
- Mein, P. and Ginsburg, L. (1997). Les mammifères du gisement miocène inférieur de Li Mae Long, Thaïlande : systématique, biostratigraphie et paléoenvironnement. Geodiversitas 19(4): 783–844.
- Butler, P. M. (1972). The problem of insectivore classification. In: K. A. Joysey and T. S. Kemp (eds.) Studies in vertebrate evolution. Oliver and Boyd, Edinburgh. Pp. 253−265.
- Martin, R. D. (1968). Reproduction and Ontogeny in tree-shrews (Tupaia belangeri), with reference to their general behaviour and taxonomic relationships. Zeitschrift für Tierpsychologie 25(4): 409–495.
- McKenna, M. C., Bell, S. K. (1997). Classification of mammals above the species level. Columbia University Press, New York.
- Emmons, L. (2000). Tupai: A field study of Bornean treeshrews. Berkeley and Los Angeles: University of California Press.
- Peterson, E. A., Wruble, S. D., Ponzoli, V. I. (1968). Auditory responses in tree shrews and primates. Journal of Auditory Research 8(3): 345–355.
- Emmons, L. H. (1991). Frugivory in Treeshrews (Tupaia). The American Naturalist. 138(3): 642–649.
- Gould, E. (1978). The behavior of the moonrat, Echinosorex gymnurus (Erinaceidae) and the pentail shrew, Ptilocercus lowi (Tupaiidae) with comments on the behavior of other insectivora. Zeitschrift für Tierpsychologie 48(1): 1–27.
- Kawamichi, T. and Kawamichi, M. (1979). Spatial Organization and Territory of Tree Shrews (Tupaia glis). Animal Behavior 27(2): 381–393.
- Cisneros, L. (2005). "Tupaia glis" (On-line), Animal Diversity Web.
- Collins, P. M. and Tsang, W. N. (1987). Growth and reproductive development in the male treeshrew (Tupaia belangeri) from birth to sexual maturity. Biology of reproduction 37(2): 261–267.
- Kuhn, H, and Schwaier, A. (1973). Implantation, early placentation, and the chronology of embryogenesis in Tupaia belangeri. Zeitschrift für Anatomie und Entwicklungsgeschichte 142(3): 315–340.
- Sprankel, H., Richarz, K., Ludwig, H. and Rott, R. (1978). Behavior Alterations in Tree Shrews Induced by Borna Disease Virus. Medical Microbiology and Immunology 165(1): 1–18.
- Cao, J., Yang, E. B., Su, J. J., Li, Y., Chow, P. (2003). The Tree Shrew: Adjuncts and Alternatives to Primates as Models for Biomedical Research. Journal of Medical Primatology. 32(3): 123–130.
- Xu, L., Zhang, Y., Liang, B., Lü, L. B., Chen, C. S., Chen, Y. B., Yao, Y. G. (2013). Tree shrews under the spot light: emerging model of human diseases. Dongwuxue Yanjiu 34(2): 59–69. (in Chinese)
- Yang, Z. F., Zhao, J., Zhu, Y. T., Wang, Y. T., Liu, R., Zhao, S. S., Li, R. F., Yang, C., Li, J., Zhong, N. S. (2013). The tree shrew provides a useful alternative model for the study of influenza H1N1 virus. Virology Journal 10 (1): 111.