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The treeshrews (or tree shrews or banxrings) are small
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s native to the
tropical forest Tropical forests (a.k.a. jungle) are forested landscapes in tropical regions: ''i.e.'' land areas approximately bounded by the tropic of Cancer and Capricorn, but possibly affected by other factors such as prevailing winds. Some tropical fores ...
s of South and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
. They make up the entire order Scandentia, which split into two
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
: the
Tupaiidae Tupaiidae is one of two families of treeshrews, the other family being Ptilocercidae. The family contains three living genera and 19 living species. The family name derives from ''tupai'', the Malay word for treeshrew and also for squirrel which ...
(19 species, "ordinary" treeshrews), and the Ptilocercidae (one species, the
pen-tailed treeshrew The pen-tailed treeshrew (''Ptilocercus lowii'') is a treeshrew of the family Ptilocercidae native to southern Thailand, the Malay Peninsula, Borneo, and some Indonesian islands. It is the only living species in the genus ''Ptilocercus''. All ...
). Though called 'treeshrews', and despite having previously been classified in
Insectivora The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, w ...
, they are not true
shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...
s, and not all species live in trees. They are
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
s; among other things, treeshrews eat fruit. Treeshrews have a higher
brain to body mass ratio A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
than any other mammal, including
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s, but high ratios are not uncommon for animals weighing less than . Among orders of mammals, treeshrews are closely related to
primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
, and have been used as an alternative to primates in experimental studies of
myopia Near-sightedness, also known as myopia and short-sightedness, is an eye disease where light focuses in front of, instead of on, the retina. As a result, distant objects appear blurry while close objects appear normal. Other symptoms may include ...
, psychosocial stress, and
hepatitis Hepatitis is inflammation of the liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes (jaundice), poor appetite, vomiting, tiredness, abdominal pa ...
.


Name

The name '' Tupaia'' is derived from ''
tupai Tupai ( ty, TÅ«pai), also called Motu Iti, is a low-lying atoll in Society Islands, French Polynesia. It lies 19 km to the north of Bora Bora and belongs to the western Leeward Islands ( French: ''ÃŽles Sous-le-vent''). This small atoll is ...
'', the
Indonesian Indonesian is anything of, from, or related to Indonesia, an archipelagic country in Southeast Asia. It may refer to: * Indonesians, citizens of Indonesia ** Native Indonesians, diverse groups of local inhabitants of the archipelago ** Indonesian ...
word for
squirrel Squirrels are members of the family Sciuridae, a family that includes small or medium-size rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squ ...
, and was provided by Sir
Stamford Raffles Sir Thomas Stamford Bingley Raffles (5 July 1781 – 5 July 1826) was a British statesman who served as the Lieutenant-Governor of the Dutch East Indies between 1811 and 1816, and Lieutenant-Governor of Bencoolen between 1818 and 1824. He is ...
.


Description

Treeshrews are slender animals with long tails and soft, greyish to reddish-brown fur. The terrestrial species tend to be larger than the arboreal forms, and to have larger claws, which they use for digging up insect prey. They have poorly developed canine teeth and unspecialised molars, with an overall
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
of Treeshrews have good
vision Vision, Visions, or The Vision may refer to: Perception Optical perception * Visual perception, the sense of sight * Visual system, the physical mechanism of eyesight * Computer vision, a field dealing with how computers can be made to gain un ...
, which is binocular in the case of the more arboreal species.


Reproduction

Female treeshrews have a gestation period of 45–50 days and give birth to up to three young in nests lined with dry leaves inside tree hollows. The young are born blind and hairless, but are able to leave the nest after about a month. During this period, the mother provides relatively little maternal care, visiting her young only for a few minutes every other day to suckle them. Treeshrews reach sexual maturity after around four months, and breed for much of the year, with no clear breeding season in most species.


Behavior

Treeshrews live in small family groups, which defend their
territory A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or a ...
from intruders. Most are diurnal, although the
pen-tailed treeshrew The pen-tailed treeshrew (''Ptilocercus lowii'') is a treeshrew of the family Ptilocercidae native to southern Thailand, the Malay Peninsula, Borneo, and some Indonesian islands. It is the only living species in the genus ''Ptilocercus''. All ...
is
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
. They mark their territories using various
scent gland Scent gland are exocrine glands found in most mammals. They produce semi-viscous secretions which contain pheromones and other semiochemical compounds. These odor-messengers indicate information such as status, territorial marking, mood, and sexu ...
s or
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excretion, excreted from the body through the urethra. Cel ...
, depending on the particular species.


Diet

Treeshrews are
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutri ...
, feeding on insects, small vertebrates, fruit, and seeds. Among other things, treeshrews eat ''
Rafflesia ''Rafflesia'' () is a genus of parasitic flowering plants in the family Rafflesiaceae. The species have enormous flowers, the buds rising from the ground or directly from the lower stems of their host plants; one species has the largest flowers i ...
'' fruit. The pen-tailed treeshrew in Malaysia is able to consume large amounts of naturally fermented nectar from flower buds of the bertam palm '' Eugeissona tristis'' (with up to 3.8% alcohol content) the entire year without it having any effects on behaviour. Treeshrews have also been observed intentionally eating foods high in
capsaicin Capsaicin (8-methyl-''N''-vanillyl-6-nonenamide) ( or ) is an active component of chili peppers, which are plants belonging to the genus ''Capsicum''. It is a chemical irritant for mammals, including humans, and produces a sensation of burning ...
and Cap2, a behavior unique among mammals other than humans. A single
TRPV1 The transient receptor potential cation channel subfamily V member 1 (TrpV1), also known as the capsaicin receptor and the vanilloid receptor 1, is a protein that, in humans, is encoded by the ''TRPV1'' gene. It was the first isolated member of th ...
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mi ...
reduces their pain response to capsaicinoids, which scientists believe is an evolutionary adaptation to be able to consume spicy foods in their natural habitats.


Taxonomy

They make up the entire order Scandentia, split into the
families Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
Tupaiidae Tupaiidae is one of two families of treeshrews, the other family being Ptilocercidae. The family contains three living genera and 19 living species. The family name derives from ''tupai'', the Malay word for treeshrew and also for squirrel which ...
, the treeshrews, and Ptilocercidae, the pen-tailed treeshrew. The 20 
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are placed in five
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
. Treeshrews were moved from the order
Insectivora The order Insectivora (from Latin ''insectum'' "insect" and ''vorare'' "to eat") is a now-abandoned biological grouping within the class of mammals. Some species have now been moved out, leaving the remaining ones in the order Eulipotyphla, w ...
into the order
Primates Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
because of certain internal similarities to primates (for example, similarities in the
brain A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a v ...
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having its ...
, highlighted by Sir
Wilfrid Le Gros Clark Sir Wilfrid Edward Le Gros Clark (5 June 1895 – 28 June 1971) was a British anatomist, surgeon, primatologist and palaeoanthropologist, today best remembered for his contribution to the study of human evolution. He was Dr Lee's Professor of ...
), and classified as a "primitive
prosimian Prosimians are a group of primates that includes all living and extinct Strepsirrhini, strepsirrhines (lemurs, Lorisoidea, lorisoids, and Adapiformes, adapiforms), as well as the Haplorhini, haplorhine tarsiers and their extinct relatives, the O ...
", however they were soon split from the primates and moved into their own
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
. Taxonomists continue to refine the treeshrews' relations to primates and to other closely related clades.
Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies have suggested that the treeshrews should be given the same rank ( order) as the primates and, with the primates and the flying lemurs (colugos), belong to the
grandorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Euarchonta The Euarchonta are a proposed grandorder of mammals: the order Scandentia (treeshrews), and its sister Primatomorpha mirorder, containing the Dermoptera or colugos and the primates (Plesiadapiformes and descendents). The term "Euarchonta" (mea ...
. According to this classification, the Euarchonta are sister to the
Glires Glires (, Latin ''glīrēs'' 'dormice') is a clade (sometimes ranked as a grandorder) consisting of rodents and lagomorphs ( rabbits, hares, and pikas). The hypothesis that these form a monophyletic group has been long debated based on morp ...
(
lagomorphs The lagomorphs are the members of the taxonomic order Lagomorpha, of which there are two living families: the Leporidae ( hares and rabbits) and the Ochotonidae (pikas). The name of the order is derived from the Ancient Greek ''lagos'' (λαΠ...
and
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
s), and the two groups are combined into the
superorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Euarchontoglires Euarchontoglires (synonymous with Supraprimates) is a clade and a superorder of mammals, the living members of which belong to one of the five following groups: rodents, lagomorphs, treeshrews, colugos, and primates. Evolutionary affinities wit ...
. However, the alternative placement of treeshrews as sister to both Glires and
Primatomorpha The Primatomorpha are a proposed mirorder of mammals containing the flying lemurs (order Dermoptera or colugos) and lemurs (Strepsirrhini, adapiformes and lemuriformes). However, notably, the haplorhini are sister to the lemurs, together form ...
cannot be ruled out. Recent studies place Scandentia as sister of the Glires, invalidating Euarchonta: It is this organization that is shown in the tree diagram below. Several other arrangements of these orders have been proposed in the past, and the above tree is only a well-favored proposal. Although it is known that Scandentia is one of the most basal Euarchontoglire clades, the exact phylogenetic position is not yet considered resolved: It may be a sister of Glires, Primatomorpha, or Dermoptera, or separate from and sister to all other Euarchontoglires. Short interspersed elements (SINEs) strongly shows scandentia belong to the euarchonta group:


Order Scandentia

The 23 
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are placed in four
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, which are divided into two families. The majority are in the "ordinary" treeshrew family,
Tupaiidae Tupaiidae is one of two families of treeshrews, the other family being Ptilocercidae. The family contains three living genera and 19 living species. The family name derives from ''tupai'', the Malay word for treeshrew and also for squirrel which ...
, but one species, the
pen-tailed treeshrew The pen-tailed treeshrew (''Ptilocercus lowii'') is a treeshrew of the family Ptilocercidae native to southern Thailand, the Malay Peninsula, Borneo, and some Indonesian islands. It is the only living species in the genus ''Ptilocercus''. All ...
, is different enough to warrant placement in its own family, Ptilocercidae; the two families are thought to have separated 60 million years ago. The former Tupaiidae genus ''
Urogale The Mindanao treeshrew (''Tupaia everetti''), also called the Philippine tree shrew, is a species of treeshrew endemic to the Mindanao region in the Philippines. It was formerly considered the only member of the genus ''Urogale'', but that genus ...
'' was disbanded in 2011 when the Mindanao treeshrew was moved to ''Tupaia'' based on a molecular phylogeny. ;Family
Tupaiidae Tupaiidae is one of two families of treeshrews, the other family being Ptilocercidae. The family contains three living genera and 19 living species. The family name derives from ''tupai'', the Malay word for treeshrew and also for squirrel which ...
: *Genus ''
Anathana The Madras treeshrew (''Anathana ellioti''), also known as the Indian treeshrew, is a species of treeshrew in the monotypic genus ''Anathana'' found in the hill forests of central and southern India. The genus name is derived from the Tamil name ...
'' **
Madras treeshrew The Madras treeshrew (''Anathana ellioti''), also known as the Indian treeshrew, is a species of treeshrew in the monotypic genus ''Anathana'' found in the hill forests of central and southern India. The genus name is derived from the Tamil name ...
, ''A. ellioti'' *Genus ''
Dendrogale ''Dendrogale'' is a genus of treeshrew in the family Tupaiidae found in Southeast Asia and Borneo. It contains these species: * Bornean smooth-tailed treeshrew (''D. melanura'') * Northern smooth-tailed treeshrew (''D. murina'') References

...
'' **
Bornean smooth-tailed treeshrew The Bornean smooth-tailed treeshrew (''Dendrogale melanura'') is a species of treeshrew in the family Tupaiidae. It is endemic to Borneo. Its natural habitat is subtropical or tropical dry forests. It is threatened by habitat loss. Description ...
, ''D. melanura'' **
Northern smooth-tailed treeshrew The northern smooth-tailed treeshrew (''Dendrogale murina'') is a species of treeshrew in the family Tupaiidae found in Cambodia, Thailand, and Vietnam. Their diet primarily consists of invertebrates but rarely includes fruits and plants. Refere ...
, ''D. murina'' *Genus '' Tupaia'' **
Northern treeshrew The northern treeshrew (''Tupaia belangeri'') is a treeshrew species native to Southeast Asia. In 1841, the German zoologist Johann Andreas Wagner first used the specific name ''Cladobates belangeri'' for treeshrews that had been collected in Pe ...
, ''T. belangeri'' **
Golden-bellied treeshrew The golden-bellied treeshrew (''Tupaia chrysogaster'') is a treeshrew species within the Tupaiidae. It is also called Mentawai treeshrew as it is endemic to the Indonesian Mentawai islands of Sipora, North and South Pagai. It lives in forests, an ...
, ''T. chrysogaster'' **
Bangka Island treeshrew The Bangka Island treeshrew (''Tupaia discolor''), also known as the Bangkan treeshrew, is a treeshrew species within the Tupaiidae. It was previously listed as a junior synonym to ''Tupaia glis'', but was raised up to species status in 2013. It ...
, ''T. discolor'' **
Striped treeshrew The striped treeshrew (''Tupaia dorsalis'') is a treeshrew species within the Tupaiidae. It is endemic to Borneo and known only from a few individuals in Sabah, Sarawak, Brunei and Kalimantan Kalimantan () is the Indonesian portion of the isla ...
, ''T. dorsalis'' ** Mindanao treeshrew, ''T. everetti'' ** Sumatran treeshrew, ''T. ferruginea'' **
Common treeshrew The common treeshrew (''Tupaia glis'') is a small mammal in the treeshrew family Tupaiidae, and is native to Thailand, Malaysia, and Indonesia. It has been listed as Least Concern by IUCN as it remains common and displays some adaptability to on ...
, ''T. glis'' ** Slender treeshrew, ''T. gracilis'' ** Javan treeshrew, ''T. hypochrysa'' **
Horsfield's treeshrew Horsfield's treeshrew (''Tupaia javanica''), also called Javan treeshrew, is a treeshrew species within the Tupaiidae. It is endemic to the Indonesian islands of Sumatra, Bali, Java and Nias where it inhabits foremost primary forest. It was firs ...
, ''T. javanica'' **
Long-footed treeshrew The long-footed treeshrew (''Tupaia longipes'') is a treeshrew species within the Tupaiidae. It is endemic to Borneo and threatened due to deforestation Deforestation or forest clearance is the removal of a forest or stand of trees fro ...
, ''T. longipes'' **
Pygmy treeshrew The pygmy treeshrew (''Tupaia minor'') is a treeshrew species within the family Tupaiidae. It is native to Thailand, Malaysia and Indonesia. The generic name is derived from the Malay word ''tupai'' meaning squirrel or small animals that resemb ...
, ''T. minor'' **
Mountain treeshrew The mountain treeshrew (''Tupaia montana'') is a treeshrew species within the Tupaiidae. It is endemic to Borneo and inhabits montane forests in Sarawak and Sabah. The first specimen was described by Oldfield Thomas and was part of a zoological ...
, ''T. montana'' ** Nicobar treeshrew, ''T. nicobarica'' ** Palawan treeshrew, ''T. palawanensis'' ** Painted treeshrew, ''T. picta'' ** Kalimantan treeshrew, ''T. salatana'' ** Ruddy treeshrew, ''T. splendidula'' **
Large treeshrew The large treeshrew (''Tupaia tana'') is a treeshrew species within the Tupaiidae. It is native to Sumatra and adjacent small islands, as well as in the lowlands and hills of Borneo.Payne, J., Francis, C. M., Phillipps, K. (1985). ''Field guide ...
, ''T. tana'' ;Family Ptilocercidae: *Genus ''
Ptilocercus ''Ptilocercus'' is a genus of treeshrew and the sole member of the family Ptilocercidae. Taxonomy Today the genus (and family) is represented by a single species, the pen-tailed treeshrew ''(Ptilocercus lowii)''. However, this genus is very anc ...
'' **
Pen-tailed treeshrew The pen-tailed treeshrew (''Ptilocercus lowii'') is a treeshrew of the family Ptilocercidae native to southern Thailand, the Malay Peninsula, Borneo, and some Indonesian islands. It is the only living species in the genus ''Ptilocercus''. All ...
, ''P. lowii''


Fossil record

The fossil record of treeshrews is poor. The oldest putative treeshrew, '' Eodendrogale parva'', is from the
Middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "dawn ...
of
Henan Henan (; or ; ; alternatively Honan) is a landlocked province of China, in the central part of the country. Henan is often referred to as Zhongyuan or Zhongzhou (), which literally means "central plain" or "midland", although the name is al ...
, China, but the identity of this animal is uncertain. Other fossils have come from the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
of Thailand, Pakistan, India, and
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
, China, as well as the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58 Named fossil species include '' Prodendrogale yunnanica'', '' Prodendrogale engesseri'', and '' Tupaia storchi'' from Yunnan, ''
Tupaia miocenica ''Tupaia miocenica'' is a fossil treeshrew from the Miocene of Thailand. Known only from a single tooth, an upper first or second molar, it is among the few known fossil treeshrews. With a length of 3.57 mm, the tooth is large for a treeshr ...
'' from Thailand, '' Palaeotupaia sivalicus'' from India and ''
Ptilocercus kylin ''Ptilocercus'' is a genus of treeshrew and the sole member of the family Ptilocercidae. Taxonomy Today the genus (and family) is represented by a single species, the pen-tailed treeshrew ''(Ptilocercus lowii)''. However, this genus is very anc ...
'' from Yunnan.


References

{{Taxonbar, from=Q231550 Fauna of Southeast Asia Extant Eocene first appearances