Northern short-tailed shrew

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Northern Short-tailed Shrew[1]
Blarina brevicauda.jpg
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Soricomorpha
Family: Soricidae
Genus: Blarina
Species: B. brevicauda
Binomial name
Blarina brevicauda
(Say, 1823)

The Northern Short-tailed Shrew (Blarina brevicauda) is the largest shrew in the genus Blarina,[3] and occurs in the northeastern region of North America.[4] It is a semifossorial, highly active and voracious insectivore and is present in a variety of habitats.[5] It is notable in that it is one of the few venomous mammals. The specific epithet, brevicauda, is a combination of the Latin brevis and cauda, meaning "short tail".[3]

Taxonomy[edit]

Blarina brevicauda is a red-toothed shrew, one of three or four species (depending on the authority)[3] in the genus Blarina. It was formerly considered to be a sister subspecies of the Southern Short-tailed Shrew (Blarina carolinensis).[3] The species has been divided into eleven subspecies based on morphological characteristics, which are grouped into two semi-species: brevicauda and talpoides; these groupings were mirrored by a molecular systematics study of the mitochondrial cytochrome b sequence.[6] The two groups of subspecies are thought to have been kept isolated from each other by Pleistocene glaciers.[6]

Description[edit]

Northern short-tailed shrew

This shrew has a total length of 108 to 140 mm (4.3 to 5.5 inches), of which 18 to 32 mm (0.7 to 1.3 inches) is tail; and weighs 15 to 30 grams (0.5 to 1.1 ounces).[5] The Northern Short-tailed Shrew exhibits slight sexual dimorphism in size, with the male being slightly larger than the female.[3][4] The dorsal fur is thick and velvety, and can be black, brownish black, or silvery gray, with the ventral fur being a bit lighter and grayer.[5] The shrew molts from a summer coat which is shorter and paler than the winter pelage in October and November, and back again sometime in February through July.[3] The tail is quite short, amounting to less than 25% of the total length.[5] The dental formula is I 3-3/1-1, C 1-1/1-1, P 3-3/1-1, M 3-3/3-3 = 16/16 = 32.[3] Three well-developed scent glands are present, one on each side of the animal and one placed ventrally; the scent may be used for marking territories, though the shrew's sense of smell is thought to be poor.[3]

Distribution[edit]

Fossil Record[edit]

Most records of B. brevicauda are from the Pleistocene,[7] though one record from the late Pliocene (Blancan land mammal age) is tentatively attributed to this species.[8] Another source indicates that the earliest record of the genus Blarina is a specimen of the talpoides subspecies of B. brevicauda, from the Blancan (early Pleistocene) in Kansas. The species is thought to have arisen in the middle or late Pliocene.[3] The brevicauda subspecies appeared later.[3]

Range[edit]

This shrew is found throughout central and eastern North America, from southern Saskatchewan to Atlantic Canada and south to Nebraska and Georgia.[4] It is probably the most common shrew in the Great Lakes region.[3][5] Population densities usually range from 5-30 shrews per hectare (2-12 per acre), but rarely exceed 200 per hectare (80 per acre).[5] The typical home range of a shrew is 2.5 hectares, and may overlap slightly with the ranges of other shrews.[3]

Habitat[edit]

Both disturbed and undisturbed habitats are used by the Northern Short-tailed Shrew, including grasslands, old fields, fencerows, marshy areas, deciduous and coniferous forests, and household gardens,[3][4][5] though the preferred habitats are those which are moist with a decent amount of leaf litter or thick plant cover.[3][5] Burned-over forests are not quickly recolonized by B. brevicauda,[4] and shrews quickly depart clear-cuts.[3]

Diet[edit]

The Northern Short-tailed Shrew consumes up to three times its weight in food each day.[4] It will eat small quantities of subterranean fungi and seeds, though it is mostly carnivorous.[5] It prefers insects, earthworms, voles, snails, and other shrews for the bulk of the diet, though salamanders and mice are also eaten.[5] This shrew consumes vertebrates more often than other shrews do.[5] The shrew mostly forages within a few hours after sunset, though it will also be active during cloudy days.[5] High evaporative water loss requires the shrew to have access to a source of water, though it does obtain water from its food as well.[3] The shrew often hoards food, especially in the fall and winter, or during a time of prey abundance;[3] one study found that it caches 87% of the prey it catches, while 9% is eaten immediately and 4% is left where it was killed.[9] They will also chew up anything they can to make their nests in.

Toxin[edit]

The saliva of the Northern Short-tailed Shrew contains a kallikrein-like protease, used to paralyze and subdue its prey.[10] The toxin is strong enough to kill small animals, up to sizes somewhat larger than the shrew itself, and results in painful bites to humans who attempt to handle the shrew.[3] The venomous saliva is secreted from submaxillary glands, through a duct which opens at the base of the lower incisors, where the saliva flows along the groove formed by the two incisors, and into the prey.[3][5] The toxin is very similar in structure to the one used by the Mexican beaded lizard (Heloderma horridum), but evolved independently, however from the same precursor protein.[11]

Physiology[edit]

Senses[edit]

The sense of smell is thought to be poor, and the eyes are degenerate and vision is thought to be limited to the detection of light,[3] but the shrew compensates by using echolocation and a fine sense of touch.[4][5]

Energetics[edit]

Its ability to consume almost anything it can catch allows the Northern Short-tailed Shrew to survive the cold winters of temperate regions.[3] The thermoneutral zone of this species is from 25 to 33 degrees Celsius[3]—meaning that no extra energy must be expended by the animal to maintain its body temperature (which averages 38–38.5 degrees Celsius[3]) when the ambient temperature is within this range. Food consumption is 43% higher in winter than in summer,[3] as the shrew must increase its metabolic rate in order to maintain its body temperature under the cold conditions. Temperatures at or above 35 degrees Celsius are lethal for this shrew.[3] A study of captive shrews found that though they were primarily nocturnal, the degree of nocturnality changed with the season; that is, during the colder winter months the shrews exhibited more out-of-burrow activity earlier in the evening, but were active later in the night during the summer.[12] The study indicated that this seasonal pattern was due to solar radiation and changing daily temperatures, and that it allows the shrews to minimize the energy needed for thermoregulation.[12] Other winter adaptations include the creation of a lined nest which aids the shrew in conserving heat, the caching of food in case of prey shortages, foraging below the leaf litter or snow where the temperature is milder, and decreasing activity levels during cold periods.[13] Along with these behavioral adaptations, the Northern Short-tailed Shrew increases its ability to generate body heat during the winter by nonshivering thermogenesis in brown adipose tissue.[13]

Behavior[edit]

Other shrews spend more time above ground than does the Northern Short-tailed Shrew, which prefers to tunnel along below ground, through the leaf litter, or at the snow/ground interface.[3][5] Bouts of frenzied activity, lasting approximately five minutes, are followed by longer periods of resting, with the total active time amounting to only 16% of a 24-hour day.[3] This animal is capable of digging at a rate of 2.5 centimeters per minute, in-between resting.[3]

The shrew constructs a nest up to 20 centimeters (8 inches) in diameter underground or underneath a log, and lines it with leaves or the fur of the Meadow Vole (Microtus pennsylvanicus).[3][5] This nest is kept clean, with wastes deposited outside the nest in a latrine area.[3][5] Other parts of the burrow system are used for food storage.[5]

Typically solitary,[4] B. brevicauda exhibits several aggressive displays and vocalizations to ward off other members of the species when encounters occur.[14] Pairs of shrews which were introduced to a cage simultaneously coexisted for less than four months before one killed the other, and a new shrew placed in a cage containing an already established shrew will be killed within a few hours.[15]

Reproduction[edit]

Mating occurs from March through September, though most births occur early or late in that period.[5] The Microtus pennsylvanicus is promiscuous[clarification needed]. Male shrews in captivity were observed to make clicking sounds while courting a female.[14] During copulation the male and female are locked together, and the female will drag the male along with her.[3] Gestation lasts 21–24 days, and after birth the 6-8 young are suckled for up to 25 days before the babies are weaned off.[3][5] Two litters per season are typical, though three is possible.[4] The female strengthens the nest when the young are nursing, and is more active to support her increased nutritional needs.[3] The young, which were born hairless and blind, weighing less than a gram, may become sexually mature in as soon as 2–3 months; those born in the spring mature more quickly than those born late in the season, and may themselves reproduce in the same year that they were born.[3][5] The juvenile pelage is pale and quite similar to the adults' summer fur, and is molted when the young reaches adult size.[3]

Predation[edit]

The Northern Short-tailed Shrew has a high mortality rate, though it attempts to escape predation by remaining hidden under vegetation, soil, leaf litter, or snow;[4] in a study only 6% of a marked group of shrews survived to the next year,[5] and winter mortality of 90% has been recorded, probably due to cold stress.[3] This shrew is consumed by many predators: trout, snakes, raptors, canids, cats, mustelids, skunks, raccoons, and opossums,[3][5] though mammalian carnivores appear to be deterred by the musky odor produced by the shrew's scent glands.[5]

Conservation[edit]

The Northern Short-tailed Shrew is considered a species of least concern in the IUCN Red List, as it is widespread, abundant, and its population is not declining.[2]

References[edit]

  1. ^ Hutterer, R. (2005). Wilson, D. E.; Reeder, D. M, eds. Mammal Species of the World (3rd ed.). Johns Hopkins University Press. p. 269. ISBN 978-0-8018-8221-0. OCLC 62265494. 
  2. ^ a b NatureServe (Hammerson, G.) (2008). Blarina brevicauda. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 2008-11-23.
  3. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak "Mammalian Species Accounts: Northern Short-tailed Shrew (Blarina brevicauda)". Retrieved 2008-11-23.  Includes a range map.
  4. ^ a b c d e f g h i j "Animal Diversity Web: Blarina brevicauda". Retrieved 2008-11-23. 
  5. ^ a b c d e f g h i j k l m n o p q r s t u v w x Kurta, Allen (1995). Mammals of the Great Lakes Region. Ann Arbor, MI: The University of Michigan Press. pp. 46–49. ISBN 978-0-472-06497-7. 
  6. ^ a b Brant, Sara V.; Orti, Guillermo (2003), "Phylogeography of the Northern short-tailed shrew, Blarina brevicauda (Insectivora: Soricidae): past fragmentation and postglacial recolonization", Molecular Ecology 12: 1435–1449, doi:10.1046/j.1365-294x.2003.01789.x 
  7. ^ "The Paleobiology Database - Blarina brevicauda". Retrieved 2008-11-23. 
  8. ^ "The Paleobiology Database - Collection 19930". Retrieved 2008-11-23. 
  9. ^ Robinson, Denise E.; Brodie, Edmund D., Jr. (1982), "Food Hoarding Behavior in the Short-tailed Shrew Blarina brevicauda", American Midland Naturalist 108 (2): 369–375, doi:10.2307/2425498 
  10. ^ Kita, Masaki; Nakamura, Yasuo; Ohdachi, Satoshi D.; Oba, Yuichi; Yoshikuni, Michiyasu; Kido, Hiroshi; Uemura, Daisuke (2004), "Blarina toxin, a mammalian lethal venom from the short-tailed shrew Blarina brevicauda: Isolation and characterization", PNAS 101 (20): 7542–7547, doi:10.1073/pnas.0402517101 
  11. ^ Aminetzach et al. 2009
  12. ^ a b Martin, Irwin G. (1983), "Daily Activity of Short-tailed Shrews (Blarina brevicauda) in Simulated Natural Conditions", American Midland Naturalist 109 (1): 136–144, doi:10.2307/2425523 
  13. ^ a b Merritt, Joseph F. (1986), "Winter Survival Adaptations of the Short-tailed Shrew (Blarina brevicauda) in an Appalachian Montane Forest", Journal of Mammalogy 67 (3): 450–464, doi:10.2307/1381276 
  14. ^ a b Martin, Irwin G. (1980), "An Ethogram of Captive Blarina brevicauda", American Midland Naturalist 104 (2): 290–294, doi:10.2307/2424868 
  15. ^ Martin, Irwin G. (1981), "Tolerance of Conspecifics by Short-Tailed Shrews (Blarina brevicauda) in Simulated Natural Conditions", American Midland Naturalist 106 (1): 206–208, doi:10.2307/2425153 
  • Aminetzach, Yael T.; Srouji, John R.; Kong, Chung Yin & Hoekstra, Hopi E. (2009): Convergent Evolution of Novel Protein Function in Shrew and Lizard Venom. Current Biology doi:10.1016/j.cub.2009.09.022