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Squirrels Temporal range: Late Eocene – Recent PreꞒ Ꞓ O S D C P T J K Pg N
Various members of the family Sciuridae Callosciurus prevostii Eutamias sibiricus Tamiasciurus hudsonicus Sciurus niger Urocitellus columbianus Paraxerus cepapi Geosciurus inauris Marmota sp. Cynomys ludovicianus
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Rodentia
Suborder:	Sciuromorpha
Family:	Sciuridae Fischer de Waldheim, 1817
Type genus
Sciurus Linnaeus, 1758
Subfamilies and tribes
Subfamily Ratufinae Subfamily Sciurillinae Subfamily Sciurinae Tribe Sciurini Tribe Pteromyini Subfamily Callosciurinae Tribe Callosciurini Tribe Funambulini Subfamily Xerinae Tribe Xerini Tribe Protoxerini Tribe Marmotini

Squirrels are members of the family Sciuridae (/sɪˈjuːrɪdeɪ, -diː/), a family that includes small or medium-sized rodents. The squirrel family includes tree squirrels, ground squirrels (including chipmunks and prairie dogs, among others), and flying squirrels. Squirrels are indigenous to the Americas, Eurasia, and Africa, and were introduced by humans to Australia. The earliest known fossilized squirrels date from the Eocene epoch, and among other living rodent families, the squirrels are most closely related to the mountain beaver and dormice.

Etymology

The word squirrel, first attested in 1327, comes from the Anglo-Norman esquirel which is from the Old French escurel, the reflex of a Latin word sciurus, which was taken from the Ancient Greek word σκίουρος (skiouros; from σκία-ουρος) 'shadow-tailed', referring to the long bushy tail which many of its members have. Sciurus is also the name of one of its genuses.

The native Old English word for the squirrel, ācweorna, only survived into Middle English (as aquerne) before being replaced. The Old English word is of Common Germanic origin, cognates of which are still used in other Germanic languages, including the German Eichhörnchen (diminutive of Eichhorn, which is not as frequently used); the Norwegian ikorn/ekorn; the Dutch eekhoorn; the Swedish ekorre and the Danish egern.

A group of squirrels is called a "dray" or a "scurry".

Characteristics

Squirrels are generally small animals, ranging in size from the African pygmy squirrel and least pygmy squirrel at 10–14 cm (3.9–5.5 in) in total length and just 12–26 g (0.42–0.92 oz) in weight, to the Bhutan giant flying squirrel at up to 1.27 m (4 ft 2 in) in total length, and several marmot species, which can weigh 8 kg (18 lb) or more. Squirrels typically have slender bodies with very long very bushy tails and large eyes. In general, their fur is soft and silky, though much thicker in some species than others. The coat color of squirrels is highly variable between—and often even within—species.

In most squirrel species, the hind limbs are longer than the forelimbs, while all species have either four or five toes on each foot. The feet, which include an often poorly developed thumb, have soft pads on the undersides and versatile, sturdy claws for grasping and climbing. Tree squirrels, unlike most mammals, can descend a tree headfirst. They do so by rotating their ankles 180 degrees, enabling the hind feet to point backward and thus grip the tree bark from the opposite direction.

Head

As their large eyes indicate, squirrels have excellent vision, which is especially important for the tree-dwelling species. Many also have a good sense of touch, with vibrissae on their limbs as well as their heads.

The teeth of sciurids follow the typical rodent pattern, with large incisors (for gnawing) that grow throughout life, and cheek teeth (for grinding) that are set back behind a wide gap, or diastema. The typical dental formula for sciurids is 1.0.1.31.0.1.3.

Tail

The purposes of squirrels' tails, to benefit the squirrel, include:

• To keep rain, wind, or cold off itself.
• To cool off when hot, by pumping more blood through its tail.
• As a counterbalance when jumping about in trees
• As a parachute when jumping.
• To signal with.

The hairs from squirrel tails are prized in fly fishing when tying fishing flies. Squirrel hair is very fine, making it better for tying fishing flies.

When the squirrel sits upright, its tail folded up its back may stop predators looking from behind from seeing the characteristic shape of a small mammal.

Lifetime

Squirrels live in almost every habitat, from tropical rainforest to semiarid desert, avoiding only the high polar regions and the driest of deserts. They are predominantly herbivorous, subsisting on seeds and nuts, but many will eat insects and even small vertebrates.

Many juvenile squirrels die in the first year of life. Adult squirrels can have a lifespan of 5 to 10 years in the wild. Some can survive 10 to 20 years in captivity. Premature death may occur when a nest falls from the tree, in which case the mother may abandon her young if their body temperature is not correct. Many such baby squirrels have been rescued and fostered by a professional wildlife rehabilitator until they could be safely returned to the wild, although the density of squirrel populations in many places and the constant care required by premature squirrels means that few rehabilitators are willing to spend their time doing this and such animals are routinely euthanized instead.

• Squirrel in sunlight
• Squirrel in Chandigarh
• Squirrel near Chandigarh
• Squirrel on mango tree

Behavior

Squirrels mate either once or twice a year and, following a gestation period of three to six weeks, give birth to a number of offspring that varies by species. The young are altricial, being born naked, toothless, and blind. In most species of squirrel, the female alone looks after the young, which are weaned at six to ten weeks and become sexually mature by the end of their first year. In general, the ground-dwelling squirrel species are social, often living in well-developed colonies, while the tree-dwelling species are more solitary.

Ground squirrels and tree squirrels are usually either diurnal or crepuscular, while the flying squirrels tend to be nocturnal—except for lactating flying squirrels and their young, which have a period of diurnality during the summer.

During hot periods, squirrels have been documented to sploot, or lay their stomachs down on cool surfaces.

Squirrels, like other rodents, employ species-specific strategies to store food, buffering against periods of scarcity. In temperate regions, squirrels commonly cache nuts beneath leaf litter, inside hollow trees, or underground. However, in subtropical and humid environments, traditional caching can lead to mold growth, decomposition, or premature germination. To counteract these challenges, some squirrels, particularly in subtropical zones, hang nuts or mushrooms on tree branches. This behavior, believed to minimize fungal infections and reduce the risk of food loss, also inadvertently aids certain trees, like Cyclobalanopsis, in expanding their range, with forgotten or dislodged nuts sprouting in new locations, influencing forest ecology. Two species of flying squirrel, the particolored flying squirrel and Hainan flying squirrel aid such cacheing by carving grooves into the nuts to fix the nuts tightly between small intersecting twigs, akin to the mortise-tenon joint in carpentry.

Feeding

Because squirrels cannot digest cellulose, they must rely on foods rich in protein, carbohydrates, and fats. In temperate regions, early spring is the hardest time of year for squirrels because the nuts they buried are beginning to sprout (and thus are no longer available to eat), while many of the usual food sources are not yet available. During these times, squirrels rely heavily on tree buds. Squirrels, being primarily herbivores, eat a wide variety of plants, as well as nuts, seeds, conifer cones, fruits, fungi, and green vegetation. Some squirrels, however, also consume meat, especially when faced with hunger. Squirrels have been known to eat small birds, young snakes, and smaller rodents, as well as bird eggs and insects. Some tropical squirrel species have shifted almost entirely to a diet of insects.

Squirrels, like pigeons and other fauna, are synanthropes, in that they benefit and thrive from their interaction in human environments. This gradual process of successful interaction is called synurbanization, wherein squirrels lose their inherent fear of humans in an urban environment. When squirrels were almost completely eradicated during the Industrial Revolution in New York, they were later re-introduced to "entertain and remind" humans of nature. The squirrel blended into the urban environment so efficiently that when synanthropic behavior stops (i.e. people do not leave trash outside during particularly cold winters), they can become aggressive in their search for food.

Aggression and predatory behavior has been observed in various species of ground squirrels, in particular the thirteen-lined ground squirrel. For example, Bernard Bailey, a scientist in the 1920s, observed a thirteen-lined ground squirrel preying upon a young chicken. Wistrand reported seeing this same species eating a freshly killed snake. There has also been at least one report of squirrels preying on atypical animals, such as an incident in 2005 where a pack of black squirrels killed and ate a large stray dog in Lazo, Russia. Squirrel attacks on humans are exceedingly rare, but do occur.

Whitaker examined the stomachs of 139 thirteen-lined ground squirrels and found bird flesh in four of the specimens and the remains of a short-tailed shrew in one; Bradley, examining the stomachs of white-tailed antelope squirrels, found at least 10% of his 609 specimens' stomachs contained some type of vertebrate, mostly lizards and rodents. Morgart observed a white-tailed antelope squirrel capturing and eating a silky pocket mouse.

Taxonomy

The living squirrels are divided into five subfamilies, with about 58 genera and some 285 species. The oldest squirrel fossil, Hesperopetes, dates back to the Chadronian (late Eocene, about 40–35 million years ago) and is similar to modern flying squirrels.

A variety of fossil squirrels, from the latest Eocene to the Miocene, have not been assigned with certainty to any living lineage. At least some of these probably were variants of the oldest basal "protosquirrels" (in the sense that they lacked the full range of living squirrels' autapomorphies). The distribution and diversity of such ancient and ancestral forms suggest the squirrels as a group may have originated in North America.

Apart from these sometimes little-known fossil forms, the phylogeny of the living squirrels is fairly straightforward. The three main lineages are the Ratufinae (Oriental giant squirrels), Sciurillinae and all other subfamilies. The Ratufinae contain a mere handful of living species in tropical Asia. The neotropical pygmy squirrel of tropical South America is the sole living member of the Sciurillinae. The third lineage, by far the largest, has a near-cosmopolitan distribution. This further supports the hypothesis that the common ancestor of all squirrels, living and fossil, lived in North America, as these three most ancient lineages seem to have radiated from there; if squirrels had originated in Eurasia, for example, one would expect quite ancient lineages in Africa, but African squirrels seem to be of more recent origin.

The main group of squirrels can be split into five subfamilies: the Callosciurinae, 60 species mostly found in South East Asia; the Ratufinae, 4 cat-sized species found in south and southeast Asia; the Sciurinae, which contains the flying squirrels (Pteromyini) and the tree squirrels, 83 species found worldwide; Sciurillinae, a single South American species; and Xerinae, which includes three tribes of mostly terrestrial squirrels, including the Marmotini (marmots, chipmunks, prairie dogs, and other Holarctic ground squirrels), Xerini (African and some Eurasian ground squirrels), and Protoxerini (African tree squirrels).

Taxonomy list

• Basal and incertae sedis Sciuridae (all fossil)
  • Hesperopetes
  • Kherem
  • Lagrivea
  • Oligosciurus
  • Plesiosciurus
  • Prospermophilus
  • Sciurion
  • Similisciurus
  • Sinotamias
  • Vulcanisciurus
• Subfamily Cedromurinae (fossil)
• Subfamily Ratufinae – Oriental giant squirrels (1 genus, 4 species)
• Subfamily Sciurillinae – neotropical pygmy squirrel (monotypic)
• Subfamily Sciurinae
  • Tribe Sciurini – tree squirrels (5 genera, about 38 species)
  • Tribe Pteromyini – true flying squirrels (15 genera, about 45 species)
• Subfamily Callosciurinae – Asian ornate squirrels
  • Tribe Callosciurini (13 genera, nearly 60 species)
  • Tribe Funambulini palm squirrels (1 genus, 5 species)
• Subfamily Xerinae – terrestrial squirrels
  • Tribe Xerini – spiny squirrels (3 genera, 6 species)
  • Tribe Protoxerini (6 genera, about 50 species)
  • Tribe Marmotini – ground squirrels, marmots, chipmunks, prairie dogs, etc. (6 genera, about 90 species)

Relationship with humans

See also

• List of animal names#squirrel
• Variant Creutzfeldt–Jakob disease (Kuru) from eating squirrel brains.

References

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17. The Beginning of the Age of Mammals Kenneth D. Rose (2006) ISBN 978-0-801-88472-6 p. 326
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28. HADJ-CHIKH, LEILA Z.; STEELE, MICHAEL A.; SMALLWOOD, PETER D. (1996). "Caching decisions by grey squirrels: a test of the handling time and perishability hypotheses". Animal Behaviour. 52 (5): 941–948. doi:10.1006/anbe.1996.0242. ISSN 0003-3472.
29. ^ Xiao, Zhishu; Gao, Xu; Zhang, Zhibin (5 April 2013). "The combined effects of seed perishability and seed size on hoarding decisions by Pére David's rock squirrels". Behavioral Ecology and Sociobiology. 67 (7): 1067–1075. doi:10.1007/s00265-013-1531-8. ISSN 0340-5443. S2CID 253815798.
30. ^ Xu, Han; Xia, Lian; Spence, John R; Lin, Mingxian; Lu, Chunyang; Li, Yanpeng; Chen, Jie; Luo, Tushou; Li, Yide; Fang, Suqin (13 June 2023). "Flying squirrels use a mortise-tenon structure to fix nuts on understory twigs". eLife. 12: e84967. doi:10.7554/elife.84967. ISSN 2050-084X. PMC 10328505. PMID 37309191.
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41. Whitaker, J.O. (1972). "Food and external parasites of Spermophilus tridecemlineatus in Vigo County, Indiana". Journal of Mammalogy. 53 (3): 644–648. doi:10.2307/1379067. JSTOR 1379067.
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43. Morgart, J. R. (May 1985). "Carnivorous behavior by a white-tailed antelope ground squirrel Ammospermophilus leucurus". The Southwestern Naturalist. 30 (2): 304–305. doi:10.2307/3670745. JSTOR 3670745.
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46. ^ Steppan & Hamm (2006)
47. Steppan, S. J. B. L. Storz, and R. S. Hoffmann. 2004. Nuclear DNA phylogeny of the squirrels (Mammalia: Rodentia) and the evolution of arboreality from c-myc and RAG1. Molecular Phylogenetics and Evolution, 30:703-719.
48. Blakeslee, Sandra (29 August 1997). "Kentucky Doctors Warn Against a Regional Dish: Squirrels' Brains". The New York Times. Retrieved 18 April 2019.
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Sources

• Whitaker, John O. Jr.; Elman, Robert (1980). The Audubon Society Field Guide to North American Mammals (2nd ed.). New York: Alfred A. Knopf, Jr. ISBN 0-394-50762-2.

Further reading

• Milton, Katherine (1984): "Family Sciuridae". In: Macdonald, D. (ed.): The Encyclopedia of Mammals: 612–623. Facts on File, New York. ISBN 0-87196-871-1.
• Steppan, Scott J. and Hamm, Shawn M. (2006): Tree of Life Web Project – "Sciuridae (Squirrels)". Version of 13 May 2006. Retrieved 10 December 2007.
• Steppan, S. J.; Storz, B. L.; Hoffmann, R. S. (2004). "Nuclear DNA phylogeny of the squirrels (Mammalia: Rodentia) and the evolution of arboreality from c-myc and RAG1". Molecular Phylogenetics and Evolution. 30 (3): 703–719. Bibcode:2004MolPE..30..703S. doi:10.1016/S1055-7903(03)00204-5. PMID 15012949.
• Thorington, R.W. and Hoffmann, R.S. (2005): "Family Sciuridae". In: Mammal Species of the World: A Taxonomic and Geographic Reference: 754–818. Johns Hopkins University Press, Baltimore.

External links

• Quotations related to Squirrel at Wikiquote
• Tree of Life: Sciuridae
• Squirrel Tracks: How to identify squirrel tracks in the wild
• National Geographic link on Squirrels
• List of names of squirrel taxa

v t e Extant species of family Sciuridae (subfamily Sciurinae, Sciurini tribe)
Kingdom: Animalia Phylum: Chordata Class: Mammalia Order: Rodentia Suborder: Sciuromorpha
Microsciurus (Dwarf squirrels)	Central American dwarf squirrel (Microsciurus alfari) Amazon dwarf squirrel (Microsciurus flaviventer) Western dwarf squirrel (Microsciurus mimulus) Santander dwarf squirrel (Microsciurus santanderensis)
Rheithrosciurus	Tufted ground squirrel (Rheithrosciurus macrotis)
Sciurus	Subgenus Guerlinguetus Brazilian squirrel (Sciurus aestuans) South Yungas red squirrel (Sciurus argentinius) Yellow-throated squirrel (Sciurus gilvigularis) Red-tailed squirrel (Sciurus granatensis) Bolivian squirrel (Sciurus ignitus) Atlantic Forest squirrel (Sciurus ingrami) Andean squirrel (Sciurus pucheranii) Richmond's squirrel (Sciurus richmondi) Sanborn's squirrel (Sciurus sanborni) Guayaquil squirrel (Sciurus stramineus) Subgenus Hadrosciurus Fiery squirrel (Sciurus flammifer) Junín red squirrel (Sciurus pyrrhinus) Subgenus Hesperosciurus Western gray squirrel (Sciurus griseus) Subgenus Otosciurus Abert's squirrel (Sciurus aberti) Subgenus Sciurus Allen's squirrel (Sciurus alleni) Arizona gray squirrel (Sciurus arizonensis) Mexican gray squirrel (Sciurus aureogaster) Eastern gray squirrel (Sciurus carolinensis) Collie's squirrel (Sciurus colliaei) Deppe's squirrel (Sciurus deppei) Japanese squirrel (Sciurus lis) Calabrian black squirrel (Sciurus meridionalis) Mexican fox squirrel (Sciurus nayaritensis) Fox squirrel (Sciurus niger) Peters's squirrel (Sciurus oculatus) Variegated squirrel (Sciurus variegatoides) Red squirrel (Sciurus vulgaris) Yucatan squirrel (Sciurus yucatanensis) Subgenus Tenes Caucasian squirrel (Sciurus anomalus) Subgenus Urosciurus Northern Amazon red squirrel (Sciurus igniventris) Southern Amazon red squirrel (Sciurus spadiceus)
Syntheosciurus	Bangs's mountain squirrel (Syntheosciurus brochus)
Tamiasciurus (Pine squirrels)	Douglas squirrel (Tamiasciurus douglasii) Southwestern red squirrel (Tamiasciurus fremonti) American red squirrel (Tamiasciurus hudsonicus)
Category

v t e Extant species of family Sciuridae (subfamily Sciurinae, Pteromyini (Flying squirrels) tribe)
Kingdom Animalia Phylum Chordata Class Mammalia Order: Rodentia Suborder: Sciuromorpha
Aeretes	Groove-toothed flying squirrel (Aeretes melanopterus)
Aeromys (Large black flying squirrels)	Black flying squirrel (Aeromys tephromelas) Thomas's flying squirrel (Aeromys thomasi)
Belomys	Hairy-footed flying squirrel (Belomys pearsonii)
Biswamoyopterus	Namdapha flying squirrel (Biswamoyopterus biswasi) Mount Gaoligong flying squirrel (Biswamoyopterus gaoligongensis) Laotian giant flying squirrel (Biswamoyopterus laoensis)
Eoglaucomys	Kashmir flying squirrel (Eoglaucomys fimbriatus)
Eupetaurus	Western woolly flying squirrel (Eupetaurus cinereus) Yunnan woolly flying squirrel (Eupetaurus nivamons) Tibetan woolly flying squirrel (Eupetaurus tibetensis)
Glaucomys (New World flying squirrels)	Humboldt's flying squirrel (Glaucomys oregonensis) Northern flying squirrel (Glaucomys sabrinus) Southern flying squirrel (Glaucomys volans)
Hylopetes	Particolored flying squirrel (Hylopetes alboniger) Bartel's flying squirrel (Hylopetes bartelsi) Hainan flying squirrel (Hylopetes electilis) Palawan flying squirrel (Hylopetes nigripes) Indochinese flying squirrel (Hylopetes phayrei) Jentink's flying squirrel (Hylopetes platyurus) Arrow flying squirrel (Hylopetes sagitta) Sipora flying squirrel (Hylopetes sipora) Red-cheeked flying squirrel (Hylopetes spadiceus) Sumatran flying squirrel (Hylopetes winstoni)
Iomys	Javanese flying squirrel (Iomys horsfieldi) Mentawi flying squirrel (Iomys sipora)
Petaurillus (Pygmy flying squirrels)	Lesser pygmy flying squirrel (Petaurillus emiliae) Hose's pygmy flying squirrel (Petaurillus hosei) Selangor pygmy flying squirrel (Petaurillus kinlochii)
Petaurista	Red and white giant flying squirrel (Petaurista alborufus) Spotted giant flying squirrel (Petaurista elegans) Japanese giant flying squirrel (Petaurista leucogenys) Hodgson's giant flying squirrel (Petaurista magnificus) Mechuka giant flying squirrel (Petaurista mechukaensis) Mishmi giant flying squirrel (Petaurista mishmiensis) Bhutan giant flying squirrel (Petaurista nobilis) Red giant flying squirrel (Petaurista petaurista) Indian giant flying squirrel (Petaurista philippensis) Mebo giant flying squirrel (Petaurista siangensis) Chinese giant flying squirrel (Petaurista xanthotis)
Petinomys	Basilan flying squirrel (Petinomys crinitus) Travancore flying squirrel (Petinomys fuscocapillus) Whiskered flying squirrel (Petinomys genibarbis) Hagen's flying squirrel (Petinomys hageni) Siberut flying squirrel (Petinomys lugens) Mindanao flying squirrel (Petinomys mindanensis) Arrow flying squirrel (Petinomys sagitta) Temminck's flying squirrel (Petinomys setosus) Vordermann's flying squirrel (Petinomys vordermanni)
Pteromys (Old World flying squirrels)	Japanese dwarf flying squirrel (Pteromys momonga) Siberian flying squirrel (Pteromys volans)
Pteromyscus	Smoky flying squirrel (Pteromyscus pulverulentus)
Trogopterus	Complex-toothed flying squirrel (Trogopterus xanthipes)
Category

• Squirrels
• Extant Eocene first appearances
• Rodents by common name
• Taxa named by Gotthelf Fischer von Waldheim