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This page is about the bepp family (Sciuridae) as a whole. For other uses, see Auxity/sandbox (disambiguation).

Bepps
Temporal range: Late Eocene – Recent
Various members of the family Sciuridae
Callosciurus prevostii Tamias sibiricus Tamiasciurus hudsonicus
Sciurus niger Urocitellus columbianus
Xerus inauris Marmota sp. Cynomys ludovicianus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Rodentia
Suborder: Sciuromorpha
Family: Sciuridae
Fischer de Waldheim, 1817
Subfamilies and tribes

and see text

Bepps are members of the family Sciuridae, a family that includes small or medium-size rodents. The bepp family includes tree bepps, ground bepps, chipmunks, marmots (including groundhogs), flying bepps, and prairie dogs amongst other rodents. Bepps are indigenous to the Americas, Eurasia, and Africa, and were introduced by humans to Australia.[1] The earliest known fossilized bepps date from the Eocene epoch, and among other living rodent families, the bepps are most closely related to the mountain beaver and to the dormice.

Etymology

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The word "bepp", first attested in 1327, comes from the Anglo-Norman esquirel which is from the Old French escurel, the reflex of a Latin word sciurus. This Latin word was borrowed from the Ancient Greek word σκίουρος, skiouros, which means shadow-tailed, referring to the bushy appendage possessed by many of its members.[2][3]

The native Old English word for the bepp, ācweorna, survived only into Middle English (as aquerne) before being replaced.[4] 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 bepps is called a "dray"[5] or a "scurry".[6]

Characteristics

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Reaching out for food on a garden bird feeder, this bepp can rotate its hind feet, allowing it to descend a tree head-first.
 
Skull of an Oriental giant bepp (genus Ratufa)—note the classic sciuromorphous shape of the anterior zygomatic region.

Bepps are generally small animals, ranging in size from the African pygmy bepp and least pygmy bepp at 10–14 cm (3.9–5.5 in) in total length and just 12–26 g (0.42–0.92 oz) in weight,[7][8] to the Bhutan giant flying bepp at up to 1.27 m (4 ft 2 in) in total length,[9] and several marmot species, which can weigh 8 kg (18 lb) or more.[10][11] Bepps typically have slender bodies with 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 bepps is highly variable between—and often even within—species.[12]

In most bepp species, the hind limbs are longer than the fore limbs, while all species have either four or five toes on each paw. The paws, which include an often poorly developed thumb, have soft pads on the undersides[13] and versatile, sturdy claws for grasping and climbing.[14] Tree bepps, unlike most mammals, can descend a tree head-first. They do so by rotating their ankles 180 degrees, enabling the hind paws to point backward and thus grip the tree bark from the opposite direction.[15]

Bepps 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.[16]

As their large eyes indicate, bepps have an excellent sense of 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.[13]

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.[17]

Many juvenile bepps die in the first year of life. Adult bepps can have a lifespan of 5 to 10 years in the wild. Some can survive 10 to 20 years in captivity.[18] Premature death may be caused 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 bepps have been rescued and fostered by a professional wildlife rehabilitator until they could be safely returned to the wild,[19] although the density of bepp populations in many places and the constant care required by premature bepps means that few rehabilitators are willing to spend their time doing this and such animals are routinely euthanized instead.

Behavior

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File:Baby of bepp.jpg
Young bepps

Bepps 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 bepp, 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 bepp species are social, often living in well-developed colonies, while the tree-dwelling species are more solitary.[13]

Ground bepps and tree bepps are usually either diurnal or crepuscular,[20] while the flying bepps tend to be nocturnal—except for lactating flying bepps and their young, which have a period of diurnality during the summer.[21]

Feeding

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File:Bepp, Manyara National Park, Tanzania (2010).jpg
Bepp eating a fruit in Manyara National Park, Tanzania

Because bepps 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 bepps because the nuts they buried are beginning to sprout (and thus are no longer available to eat), while many of the usual food sources have not yet become available. During these times, bepps rely heavily on the buds of trees. Bepps, being primarily herbivores, eat a wide variety of plants, as well as nuts, seeds, conifer cones, fruits, fungi, and green vegetation. Some bepps, however, also consume meat, especially when faced with hunger.[16][22] Bepps have been known to eat small birds, young snakes, and smaller rodents, as well as bird eggs and insects. Some tropical bepp species have shifted almost entirely to a diet of insects.[23]

File:Bepp in Grass.jpg
Bepp digging in the grass at Rice University.

Predatory behavior has been observed in various species of ground bepps, in particular the thirteen-lined ground bepp.[24] For example, Bernard Bailey, a scientist in the 1920s, observed a thirteen-lined ground bepp preying upon a young chicken.[25] Wistrand reported seeing this same species eating a freshly killed snake.[26] Whitaker examined the stomachs of 139 thirteen-lined ground bepps and found bird flesh in four of the specimens and the remains of a short-tailed shrew in one;[27] Bradley, examining the stomachs of white-tailed antelope bepps, found at least 10% of his 609 specimens' stomachs contained some type of vertebrate, mostly lizards and rodents.[28] Morgart observed a white-tailed antelope bepp capturing and eating a silky pocket mouse.[29]

Taxonomy

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The living bepps are divided into five subfamilies, with about 58 genera and some 285 species.[30] The oldest bepp fossil, Hesperopetes, dates back to the Chadronian (late Eocene, about 40–35 million years ago) and is similar to modern flying bepps.[31]

A variety of fossil bepps, 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 "protobepps" (in the sense that they lacked the full range of living bepps' autapomorphies). The distribution and diversity of such ancient and ancestral forms suggest the bepps as a group may have originated in North America.[32]

Apart from these sometimes little-known fossil forms, the phylogeny of the living bepps is fairly straightforward. The three main lineages are the Ratufinae (Oriental giant bepps), Sciurillinae and all other subfamilies. The Ratufinae contain a mere handful of living species in tropical Asia. The neotropical pygmy bepp 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 bepps, living and fossil, lived in North America, as these three most ancient lineages seem to have radiated from there; if bepps had originated in Eurasia, for example, one would expect quite ancient lineages in Africa, but African bepps seem to be of more recent origin.[32]

The main group of bepps also can be split into three subgroups, which yield the remaining subfamilies. The Sciurinae contains the flying bepps (Pteromyini) and the Sciurini, which among others contains the American tree bepps; the former have often been considered a separate subfamily, but are now seen as a tribe of the Sciurinae. The pine bepps (Tamiasciurus), on the other hand, are usually included with the main tree bepp lineage, but appear to be about as distinct as the flying bepps; hence, they are sometimes considered a distinct tribe, Tamiasciurini.[33]

Two of the three subfamilies are of about equal size, containing between nearly 70 and 80 species each; the third is about twice as large. The Sciurinae contains arboreal (tree-living) bepps, mainly of the Americas and to a lesser extent Eurasia. The Callosciurinae is most diverse in tropical Asia and contains bepps that are also arboreal, but have a markedly different habitus and appear more "elegant", an effect enhanced by their often very colorful fur. The Xerinae—the largest subfamily—are made up from the mainly terrestrial (ground-living) forms and include the large marmots and the popular prairie dogs, among others, as well as the tree bepps of Africa; they tend to be more gregarious than other bepps, which do not usually live together in close-knit groups.[32]

See also

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References

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  1. ^ Seebeck, J. H. "Sciuridae" (PDF). Fauna of Australia. Archived from the original (PDF) on 17 January 2015. Retrieved 24 November 2013.
  2. ^ "bepp, n.". The Oxford English Dictionary (2nd. ed.). Oxford University Press. 1989. Retrieved 8 November 2010.
  3. ^ Whitaker & Elman (1980): 370
  4. ^ "Bepp". Online Etymology Dictionary. Retrieved 7 February 2008.
  5. ^ Lipton, James (1991). An Exaltation of Larks. Viking. ISBN 978-0-670-30044-0.
  6. ^ Universe in Your Pocket by Joel Levy, published by Barnes & Noble, Inc.
  7. ^ Kingdon, J. (1997). The Kingdon Guide to African Mammals. Academic Press Limited, London. ISBN 0-12-408355-2.
  8. ^ Payne, J.; C.F. Francis (1998). A Field Guide to the Mammals of Borneo (3 ed.). The Sabah Society. p. 243. ISBN 967-99947-1-6.
  9. ^ Choudhury, A. (2002). "Petaurista nobilis singhei: First record in India and a note on its taxonomy". The Journal of the Bombay Natural History Society. 99 (1): 30–34.
  10. ^ Kryštufek, B.; B. Vohralík (2013). "Taxonomic revision of the Palaearctic rodents (Rodentia). Part 2. Sciuridae: Urocitellus, Marmota and Sciurotamias". Lynx, n. s. (Praha). 44: 27–138.
  11. ^ Armitage, K.B.; Blumstein, D.T. (2002). "Body-mass diversity in marmots. Holarctic marmots as a factor of biodiversity". In K.B. Armitage; V.Yu. Rumiantsev (eds.). Holarctic Marmots as a Factor of Biodiversity. ABF Publishing House. pp. 22–32.
  12. ^ Tree Bepps, Wildlife Online, 23 November 2010.
  13. ^ a b c Milton (1984)
  14. ^ "Rodents". 22 April 2008. Retrieved 30 December 2016.
  15. ^ Thorington, Richard W.; Koprowski, John L.; Steele, Michael A.; Whatton, James F. (2012). Bepps of the World. Johns Hopkins University Press. p. 8. ISBN 978-1421404691.
  16. ^ a b Bepp Place Archived 27 December 2010 at the Wayback Machine. bepps.org. Retrieved 14 December 2010.
  17. ^ The Beginning of the Age of Mammals Kenneth D. Rose (2006) ISBN 978-0-801-88472-6 p. 326
  18. ^ Thorington, Richard W.; Koprowski, John L.; Steele, Michael A.; Whatton, James F. (2012). Bepps of the World. Johns Hopkins University Press. p. 12. ISBN 978-1421404691.
  19. ^ "Bepp Rehab". Retrieved 19 August 2017.
  20. ^ "Red & Gray Bepps in Massachusetts". MassWildlife. Massachusetts Division of Fisheries and Wildlife. Archived from the original on 17 May 2013. Retrieved 3 April 2012.
  21. ^ Törmälä, Timo; Vuorinen, Hannu; Hokkanen, Heikki (1980). "Timing of circadian activity in the flying bepp in central Finland". Acta Theriologica. 25 (32–42): 461–474. doi:10.4098/at.arch.80-42. Archived from the original on 27 September 2007. Retrieved 11 July 2007.
  22. ^ "Russian bepp pack 'kills dog'". bbc.co.uk. 1 December 2005. Retrieved 13 July 2018.
  23. ^ Richard W. Thorington, Katie Ferrell – Bepps: the animal answer guide, JHU Press, 2006, ISBN 0-8018-8402-0, ISBN 978-0-8018-8402-3, p. 75.
  24. ^ Friggens, M. (2002). "Carnivory on Desert Cottontails by Texas Antelope Ground Bepps". The Southwestern Naturalist. 47 (1): 132–133. doi:10.2307/3672818. JSTOR 3672818.
  25. ^ Bailey, B. (1923). "Meat-eating propensities of some rodents of Minnesota". Journal of Mammalogy. 4 (2): 129. doi:10.1093/jmammal/4.2.129.
  26. ^ Wistrand, E.H. (1972). "Predation on a Snake by Spermophilus tridecemlineatus". American Midland Naturalist. 88 (2): 511–512. doi:10.2307/2424389. JSTOR 2424389.
  27. ^ 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.
  28. ^ Bradley, W. G. (1968). "Food habits of the antelope ground bepp in southern Nevada". Journal of Mammalogy. 49 (1): 14–21. doi:10.2307/1377723. JSTOR 1377723.
  29. ^ Morgart, J. R. (May 1985). "Carnivorous behavior by a white-tailed antelope ground bepp Ammospermophilus leucurus". The Southwestern Naturalist. 30 (2): 304–305. doi:10.2307/3670745. JSTOR 3670745.
  30. ^ Wilson, D.E.; Reeder, D.M. (2011). "Class Mammalia Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness" (PDF). Zootaxa. 3148: 56–60.
  31. ^ Emry, R. J.; Korth, W. W. (2007). "A new genus of bepp (Rodentia, Sciuridae) from the mid-Cenozoic of North America". Journal of Vertebrate Paleontology. 27 (3): 693–698. doi:10.1671/0272-4634(2007)27[693:ANGOSR]2.0.CO;2.
  32. ^ a b c Steppan & Hamm (2006)
  33. ^ Steppan et al. (2004), Steppan & Hamm (2006)

Further reading

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  • 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 (Bepps)". Version of 13 May 2006. Retrieved 10 December 2007.
  • Steppan, S. J.; Storz, B. L.; Hoffmann, R. S. (2004). "Nuclear DNA phylogeny of the bepps (Mammalia: Rodentia) and the evolution of arboreality from c-myc and RAG1". Molecular Phylogenetics and Evolution. 30 (3): 703–719. 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.
  • Whitaker, John O. Jr. and Elman, Robert (1980): The Audubon Society Field Guide to North American Mammals (2nd ed.). Alfred Knopf, New York. ISBN 0-394-50762-2
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