MAMMALIAN SPECIES wo. 261, pp. 1-9, 3 fas Blarina brevicauda. By sorah B. George, Jerry R. Choate, and Hugh H. Genoways Published 16 June 1986 by The American Society of Mammalogists Blarina Gray, 1838 Blarina Gray, 1838:124. Type species Corsa (Blavina) ral- ipoides Gray (=Sorex talpoides Gapper ~ Sorex brevicaudus Say), by eriginal designation. Elevated to generic rank by Lesson, 1842:89, Brachysorex Duvervoy, 1842:37~41. Type species Brachysorex brevicaudus Duvernoy (=Sorex brevicaudus Say), by orginal designation, Talposorex Pomel, 1848:248, Type species Talposorex platyurus ‘Pomel (=Sorex brevicaudus Say), by erginal designation. Anotus Wagner, 1855:550-551. Type spocies Sores carolinensis Bachman, by orginal designation CONTEXT AND CONTENT. Order Insetivora, Family Sericidae, Subfamily Sorcinee, Tribe Blinini (Repenning, 1967) ‘The genus Blarina includes three species, Key characters used herein were adapted primarily from George el. (1981, 1982) but with modifeation based on data presented by Braun and Kennedy (1983), Cenoways and Goeate (1972), Monerief et el. (1982), and Tate etal (1980) 1 Size large (total length usually greater than 110 mm; ‘eciptopremaxilary length (Choate, 1972a) usually treater than 20,5 mm if eat of Mississippi River, usu ally greater than 21.5 if west of Misinippi River karyotype 2n = 48 t0 50, FN = 48. B. brevicauda Size smaller (total length usually less than 110 am: oc ‘pite-premaxilary length sully les than 20.0 sna if east of Misssippt River, usualy less than 21.5 if west fof Mississippi River karyotype other than indicated [tors = aaeaee aaa 2 (1) Sie smallest in genus (eraial breadth as small os 9.6 ‘mum, usually less than 11.0); karyotype 2a = 50 to 52, FN = 52 (in the subspecies B. ¢. peninsulae) or 2a’ 46, 39, 38, of 37, FN ~ 48 or 44 (i B. c carolinensis) ——— B. carolinensis ‘Sae larger (canal breadth as great a2 12.2 mm, ually (greater than 10.5} karyotype 2n = 52, FN = 60 1 o —B. hylophaga Blarina brevicauda (Say, 1823) Northern Short-tailed Shrew Sores brevicauds Say, 1823:164, Type locality west bank of ‘incur River, approximately 2 mi & Fe. Calhoun, former Engineer Cantonment, Westingtn Co, Nessa Joes, 196% 6, larina’brevicaud: — . . Sores talpaides Capper, 1830:202, Type lcaty between Yor ‘nd Lake Sines, Ontario, Soren dekoyi Bachman, 1887:372, Type locality New Jerey. The nomenclatoral history of this nae sod "Sorex dekayt De Kay” were reviewed by Handley and Choate (1970) Golemys micruras Pomel, 1848:249, A new name propored for "Sorex dekeyi De Kay” (Handley and Choate, 1970). Blarina angusticeps Bair, 1858-34, Type locality Burington, Chistenien Cry Vermant. Regarded by Meriams (1898) 8 based ona deformed sll (Bale and Moultrop, 1942) Blarina cotaricenss JA. Allen, 1891:205-208. Type laity ‘ippsely La Carpstra, Conta Ric, but acured by Mere tian (1895) to have been somentere tn Upper Missi Vale, proebly lowe (Alen, 1897; Ble and Mautrep. 1942). Barina tlaleses Merit, 1895:15. Type loalty Lake Drum ‘nond, Diamal Samp, Notfols Co, Vegi, Boird, 1858:42; fist use of current name Blarina simplicidens Cope, 1899:219. Type locality Port Kenoedy Cave (pre Wiacotinan lace fauna), Montgomery Ca, Penn saan Blarina brevicouda ozarkensis Brown, 1908170. Type losly Conard Fisore (a pre-Wisconsinan lot] fauna), Newton Co. Arkansas, Elevated to wpecie rank by Graham snd Semken, 1976484 lari fossils Hibbard, 1943:258. Type locality Reusbek gravel ‘te pre-Wiconsian lel fauna}, Lineal Ca. Kanss CONTEXT AND CONTENT. Context is given above in the generic account. Eleven Recent subspecies of B. brevicauda (exclusive of B. carolinensis and B. hylophaga), referable to two semispecies Jones et al 1984), currently are recognized (Hall, 1981; Handley, 1979} B. b.aloga Bangs, 1902:76. Type locality West Tisbury, Martha’ Vineyard, Dukes Co., Massachosetts B. b. angusta Anderson, 1943:52. Type locality Kelly's Camp, Berry Mountain Brook, near head of Grand Cascapedia River, Gaspe Co., Quebec, about 1,600 f Bb, brevicauda (Say, i Long, 1823:164), see above, BB. church: Bele and Moulthrop, 1942:109. Type locality Roen ‘Mountain, Mitchell Co., North Caroline Bb. compacta Bangs, 1902:77. Type locality Nantucket, Nam B. B. tucket Co., Messachuses. b- hooperi Bole and Moulthrop, 1942:110. Type locality Lyn- don, Caledonia Co., Vermont .irdlands Bole and Moulthzop, 1942:99. Type locality Holden Arboretum, Lake and Geauge counties (the county line bisects the type locality), Ohio, Bb. manitobensis Anderson, 1947:23. Type locality Max Lake, Turtle Mountains, Manitoba, “latitude a lide north of 49th parallel, longitude about 100 degrees west tude about 2,100, foot.” b, pallida R. W. Smith, 1940:223. Type locality WolWville, Kings Co., Nova Scotia . talpodes (Gapper. 1830:202), see above. : telmalestes Merriam, 1895:15, see above. DIAGNOSIS. The Nearetic genus Blarina includes the nearly uniformly slver to black (often with brown tips on baits, short- taled shrews having five unieuspidate teeth in each upper jaw (Fig. 1). The dental formula is as in the genus Sorex: faletorm incisor, five unicurpis, the fourth premolar, and three molars in each uppee twothrow: procumbent incor, one unieuspid, the fourth premolar, and three molars in each lower toothrow, total 82 (Choate, 1968, 1970, 1975). The genera Blarina and Sorex realy canbe distin guished externally bythe relatively much shorter tail ofthe former (209% of total length is typical for Blarina,wheress mare than 40% fs usual for Sores). Blarina can be distinguished from Cryptots in thatthe letter lacks one unicuspid (30 total teeth in Cryprots, 32 in Blarina; Hall, 1981). Blaring brevicauda is the largest species in the genus (Cen corays and Choate, 1972: George et al, 1981; Graham and Semen, 1976; Monerief et al, 1982). Its geographic range (Fig- 2) hes north of the ranges of B. ylophaga (inthe west) and B. caro rensis (in the eat), from which it usually can be distinguished by its greater sie. In southern lowa, northern Missouri, and mort astern Kansas, however, small individuals of B. brevicauda may fall within the range of measurements of B. Aylophage (Moncriel totaly 1982). Therefore, the most diagnostic character of B. brew “cauda is its karyotype: EN = 48, 20 = 48 to 50 (Ceorge etal 1982). GENERAL CHARACTERS. Blarina brevicauda iss rel. sively large, robust shrew (Fig. 3). Its external ears are inconspic- Uuous and concealed by pelage and its eyes are minute. The snout = Betrim ventral, and lateral views of cranium and lateral view of loner ja of Blorina brevicauda (CM 50584, male, fom Round Hill Regional Park, 13.7 miS, 9.1 ai E Pittsburgh, Allegheny Co, Penneylrania). Drawn by N. J. Perkins is pointed and somewhat proboscislike but is comparstively shorter fand heavier than in ober shrews. The tal is noticeably hairy and in adults is faintly to indtintly bicolored. Feet are pentadactylous tnd are relatively broader and stronger than those of all but the ‘most fostoral of other American shrews (Choete, 1970). Dorsal plage is short, sot, and molelike in winter, when it often has 3 fark slate color: vental pelage sometimes appears paler, atleast in part because ventral far is shorter and denser. Summer pelage is shorter and slightly paler than winter pelage and sometimes in realy indisingusbable from the short fury, juvenal peage. The skull is more massive and angular (Fig. 1) than those of other ‘American shrews end is charactriad (jn adult) by prominent edges tnd processes (Jackson, 1961:43}. Teeth are pigmented (deep chestutin color) and exhibit a relatively unepecialized sorcid con guration: “frst upper incisors incumbent with tips curved or hooked ventrad and... posesing a 2nd... unicuspdlike conule.. all other incisors, canines, and all [bu the fourth] premolars uni usp: crowns of upper molars W-shaped” (Hal, 1981:24), Other dental and mandibular characters of shrews were described and ihstrated by Repenning (1967). [Nesling short-sailed shrews grow rapidly, end they attain es MAMMALIAN SPECIES 261 Fic. 2. Mep showing the geographic range of Blarina bree fcauda. Subnpecies ae: ly B. be loge: 2, B.b. angusta; 3, B bbrevicauda: 4, B. b. churehi: 8, B. b. compacta; 6, B. b hhooperi 7, B. 6. krdlandi 8, B. . manitobensis; 9, Bb pallida; 10, Bb. folpoides: 11, B. . telmatestes. The dot in northeastern Kanes represents an apparently iclated population of B. b. rt lands sentilly adult sie but not necessrily mass; Dapson, 1968) before they become susceptible to trapping (Guilay, 1957). Accordingly, ‘most authors have opted to disregard variation with age in analyse of morphometric variation (Choate, 1972). Certain authors (Beum fardver and McPherson, 1985; Cuilday, 1957; Jackson, 1961) Fave assorted that males average sbghtly larger than females in external, cranial, and_postranial meatirements, whereas others (Choste, 19724) have found litle or no secondary sexual rariation in the apecis, anges ard means (in parentheses) of selected exteenal and cranial measurements (ia mm) of samples from Nebraska (Jones, 1964), Towa (Bowles, 1975), Iinois (Elis et al, 1978), Pennsy- ‘ania (males only, Guilday, 1957), and Connecticut (msles onl, Ghoate, 1972e) are as follows: total longi 125 to 141 (132.8) 120 v0 138 (125.3), 106 to 126 (117.2), length of hindfoct, 16 to 18 (17.0) 14 to 17 (15.8), Beis (14.0), + cranial eadth, 12.8 t9 14.0 (13.5), 13.3 10 18.8 (13.1), 115 w 126 (2.1), 11.8 to 13.4 (125), 11.9 to 13.2 (12.6) maxilary breadth, 8.2 to 9.1 (8.7), 8.2 to 8.8 (8.5) 7.3, 10 8.0 (7.6), 7.0 to 8:4 (7.8), 7.2 to 8.1 (7.7), Ranges and means of cranial measurements (in mim) of 25 specimens of B. b. brevi- ‘cauda from Nebraska and 23 specimens of B. 6, kirtlandi from Obio (Monerif et al, 1982) are as follows: ocepitopremaxilary length, 23.1 to 25.2 (24.0), 20.8 to 23.0 (21.5), PA-M3 length, 6.3 10 6.8 (6.6), 5.6 to 6.0 (5.9 cranial breath, 12.6 10 14.0 (13.5), 11.5 to 12.6 (12.0f breadth of zygomatic plate, 2.3 to 3.1 (2:7), 20 to 26 (2.3 maxilary breadth, 8.0 t0 9.1 (8.7), 7.1 0 7.8 (7.6 intororbital breadth, 5.9 to 6.7 (6.2), 5.3 t0 5.8 (5.5% length of mandible, 13.3 to 14.4 (13.8), 11.6 10 13.0 (12.1); height of mandible, 7-4 to 8.2 (7.9), 6.3 10 7-1 (6.6) articular breadth, 28 to 32 (30), 23 t0.2:7 (2.4). DISTRIBUTION. Blerine brevicauda occurs throughout ‘uch of the north-central nd northeastern United States and roth frm regions of adjacent Canadian provinces (Fig. 2). Marginal ree fords were identified by Hal (1981), but he regarded B. carolinensis ‘only subepecifeeliy distinct from B, brevicauda, B. hylophag fea synonym of B, carolinensis, and Bb. telmalestes ea distnet specie, The datribution of B. brevicauda overlaps with that of B. Fylophaga in couthern lowa, northern Misoui, northeastern Kam sas, and possibly southeastern Nebraska (Moncraf t al, 1982), land with that of B- carolinensis in central Tennessee (Braun and Kennedy, 1983), southern Inois (Els etal, 1978), eastern Vir- ina (Tate etal, 1980), western North Csrolina and Georgia, and eastern Alabama (French, 1981), FOSSIL RECORD. Blarina brevicauda ot an ancestral specin probably arose from the blriaine stem in the idle or late Phocene, ‘The earliest record of the genus is represented by spec mens of the falpoides semispeciee of B. brevicauda from late Blancan (early Pleistocene) fsunas in Kansas, The brevicauda semiMAMMALIAN SPECIES 261, Fic. 3, Photograph of « feeding Blarina breviceuda. Pho- tograph by Roger W. Barbour. species of B. brevicauda eppesred later inthe early Pleistocene, perhaps during the Kansan glacial, but before the origin af the two father species of Blarina (B. carolinensis in the mid.lrvingconian tnd B. hylophaga after the Wisconsinan glacistion). Continity of gene How between the two semispecies of B, Brevicauda during the Preistocene and Holocene and the absence of fixed chromosomal Aiferences between the semispecies apparently prevented speciation af these phena (Jones et al, 1988), FORM. Merriam (1895:11) described the pelage of B. brev ‘cauda as "sooty plmheous above, becoming ashy-plumbeous be Tow, verying withthe light.” Findley and Jones (1956) described ‘three malts: The first, postuvenal molt docs not occur ntl the shrew is ewentially of adult sae, The fuzzy juvenal pelage sheds inilly inthe head region and proceeds easdad; the new pelage (whether long, silky winter furor shorter summer fu) i determined by the time of year in which the sbrew is born, Molt from summer to winter pelage proceeds in a tailtorhead direction in both first and second:year animals and most often is seen in Octeber and November. Spring molt can occur at any time between February and Jul in females iin head-to-tail Section, whereas in males itis more irregular (Findley and Jones, 1956). Albiniste Blorina have been reported from Delaware, Indiana, New York, Perey vania, Vermont (Willams, 1962), and Ohio (Svendson and Srend- son, 1975), Blarina brevicanda possesses thee dermal scent glands, one ventral nd one on each flank. Sweat glands seem tobe the principle ‘component of the lateral glands, whereas sebaceous tise i more Pronounced inthe ventral gland (Pearson, 1946). Scent glands are well developed in both males and female; they inerease in size in ‘males concomitant wit testicular enlargement in spring end autumn (Eadie, 1938) and hecome smaller in females duting estrus, preg nancy, and leetation (Pearson, 1946). Eade (1938) suggested that Scent glands might serve as a means of protection; the musky odor thay produce is distasteful 19 many carnivores, and the abdominal skin is associated closely withthe underlying muscles, posi to facilitate secretion when a shrew becomes excited or upset. The lands alo might be used for marking territories, ths serving to Separate shrews at times other than during the breeding secson (Pearson, 1946); however, the sense of smell is considered to be poorly developed (Sehwarte and Schwartz, 1981) "The eye of B. brevicauda is degenerate (Gaughran, 1954; Ryder, 1890), It and the optic nerve are diminutive and, although slight motion is possible, the reduced ocular muscles donot arise Azectly from the skull. Vision probably is limited to pereeption of light (Schwartz and Sehwartz, 1981). The lachrymal gland = much larger than the eyeball, and its duct opens inte the conjunctival cavity. The gland has its own investment of striated voluntary mus- les, suggesting thatthe shrew can compres th gland volutariy, Secret over the eyeball, and wash away dirt that might recumulate ‘here during burrowing activity. There is controversy about the dental formal of short-iled shrews. Mersiam (1895) described it ax 14/2, ¢ 1/0, p 2/1, m 3/3, total 32. Ambich-Christie-Liede (1912) opined that the dental formula actualy is 13, 1d, 15, Pl. P2, P3, Pa, MI, M2, M3/4, Pl, pd ml, m2, m3, and suggested that two adltonal rudimentary 3 incisors are present early in embryonic development. Kinds (1960) found no evidence for these rudimentary inciwors and stated that the dental formula is 1,12, 13 C, P2, P3, Pa, M1, M2, M3/. 3. James (1963) reinterpreted Kindabl's results the dental formula is, in fact, 11,12, 13, C, P2, PS, PS, M1, M2, MB/i3, c, pf ml, m2, m3. Choate (1968) eserned derial abnormalities in. brevicauda and followed James’ (1963) interpretation of the dental formula. Because of uncertainty regarding dental homologies, most authors employ the formula given in GENERAL CHARACTERS. Repenning (1967) described the anatomy of individual teeth, Blarina.brevicauda possesses “a pair of extrapulmonary bronchial diverticula emerging from the dorsal eaudal margin of the ‘ight posterior lobe of the lung," which may be a morphological ‘adaptation to the environment in whieh the shrews live (Parke, 19562236). Dust collects n the diverticula, balls wp, ix ejected into the lungs by the muscular ection of the diverticula and removed by peristalsis (Parke and Wetzel, 1968). ‘The ventral aspect ofthe brain of B. brevicauda was figured by Hyde (1950), and the lateral aspect by Le Gros Clark (1932) The secondary optic tracts of the brain are largo relative to the primary visual clement (Cillan, 1941 the optic, oculomotor, ro: thler, and abducens nerves are small for mammals andere dwarfed by the trigeminal nerve (Hyde, 1959). With respect tothe trigem- inus, Hyde (1959:845) stated that Blarina “probably represents the greatest reduction in neuronal population that ie consistent with adequate bran function.” The olfactory tubercle and bubs ee large (Le Gros Clark, 1952). The relative size ofthe amygdaloid complex is simlar to tht of other mammals (Crosby and Humphrey, 1944). The average hematocrit for B. brevicauda was 45% and the average hemoglobin was 16.5 4/100 ml, well within the range for ‘other mammals. The average red blood cell count wae 18 milion’ mm, very high for mammals, whereas the average white blood cell ‘souat af 2,730/mm was lowe (Doremus and Jaffe, 1964) ‘Alen (1894:270) commented on the sll end skeleton of B. brevicauda and concluded thst the "posterior extremity is of low specialization, and one which supports the trunk imperfectly.” Caughran (1954) described and contrasted the estoology and myol ‘ogy of the erenial and cervial regions of B. brevicauda and the teastern mole, Scalopus aquaticus. He concluded that their crania Ihave sinilar proportions and features, but that Blarina is speciale ‘more for marication and Sealopus more for burrowing. ‘estes are situated inside the abdominal cavity: there is no serotum (Pearson, 1944), Testes in winter vary in length from 2.5 to 3.5 mm, whereas in the breeding season they usually ate longer than 9 mun. An erect pens in the breeding season measures ebout ‘30 mam, the terminal thied to half of which i glans (Martin, 19675, Pearson, 1944). Male accossory reproductive glands include pros tate and bulbo-urethral glands (Eadie, 1947). The vagina during estrus is bent in an Shape (Pearson, 1944). The uterus is bicorn- late: the distance fom the base of the bladder to the junetion of the uterine horns in ‘winter was as litle as 2 mm but in the breeding Season increased to more than 12 mm (Pearson, 1944). Ovaries ace completely enclosed in ovarian capsules end consist primarily Of folie, with elatively fee interstitial cells (Pearson, 1948). The ‘horiellantoie placenta has been deserted as endotheliochorial in orgsnization (Wimsatt etal 1973). Females have three pairs of ‘mammae in the groin cegion (ehwarte and Schwartz, 1981), FUNCTION. The metabolic rete of B.brevicauda is chac- acterized by shor periods of activity with intervening pero of inactivity Pearson," 1987). The pattern i netiy connnaoun a though there ie dsinet tendency for shorted shrew to be rove ecive in the ely moming (Morton, 1948) Periods of fctvigy average about 4.5 min (Bucker, 1968), The shrews ae fesive for only 165 of a 24h period (Marien, 1969), the fe Stunde bing nent st «lower, resting metab rte (Ralph, 1973), Maransen (1969) bypassed hat th, topes wh he proclivity of thee shrews to eat eary any source of energy, te oun for the ability ofthe species to survive in coldtemperure slime. Within the range of OF to 250, metabo very oportonal to ambient temperature (Randalph, 1973) ood comumptin averages about 0.36°¢ §* day” in B. breviaude (Morrison #1957) betes onygen consumption averages about 5.2 ce gh (Pearvn, T947), Food consumption in'winer ie about 43% higher than in turner Randolph, 1979) (Clclted values for besa metab rate range fom 218 fo 3.44 c= O, g bot (Martincen, 1969; Neal end Lustick, 1973: Pearson, 1947}. Because of its bigh evaporative water loss, B. brevicauda requires free water even though it drives fre water from food ad rmetabalie water from oxidation of food (Chew, 1951). Jackson (1961) itutrated and dereribed shrew feces ax being about 2.5 em long, dark green in color, and twisted into corkserow shape. Computed from oxygen consumption, the thermoneutrl zone of B. brevicauda extends from 25°C to 33°C. Minimal oxygen ‘consumption occurs at 30°C (Neal and Lustick, 1973). The upper lethal ambient temperature appears to be 35°C, et which no amount Of evaporative water loss is effective in reversing hyperthermia Mean body temperature ranges from 38°C to 38.5°C (Chew, 19515, oremus, 1963), but body temperature elevates apprecshly during periods of activity (Kendeigh, 1945). “The poisonous nature of the saliva of B. brevicauda was suspected ts early a8 1889, and several authors devcribed the se- ‘ous effects experienced after they were bitten by shrews (Krosch, 1973; Maynard, 1889). The poison is secreted {rom the submax. ilary glands through a duct atthe base of the lower incisors; when ' shorttaled shrew bites another animal, the tore saliva probably flows along the groove between the two teeth into the wound (Pear son, 1942). In small mammals, the toxin ean lead to death from separ flr accnpanid by sever peripheral vacation (Elis and Krayer, 1955; Pearson, 1942; DeMeules (1958) abo demonstrated posible antnadrenalin action of the venom, The toxin ise protein with several histiine-ike features (Elis and Kray. fr, 1955} The LD,, of crudely purfed toxin is approximately 3.4 ‘g/kg in mice and'eats and 0.6 to 1.2 mg/kg in rabbits (Elis and Krayer, 1955). Lawrence (1945) compared shrew venom to snake venom and found that, in is neurototic und hemotoxe elect, it is most comparable to elapine venom, She suggested thet, in aditon to its role in predation, it may aid in the breakdown of protein ddring digestion. Tomasi (1978) opined that one function of the ‘venom is to stan or paralyze its prey, thereby allowing short-tailed shrews to take advantage of prey even though it might not actually be eaten until later. Martin (19814) described the immabiling sffect of the venom on insets and suggested that the venom fac itates food hoarding by Blarina Evidence that shorttaled shrews employ echolocation to ex- plore their environment ws frst presented by Gould e al. (1968) Subsequently, Tomasi (1979) investigated the echolocating ability of B. brevicauda by testing the ability of individuals to discriminate between openended and closed tubes simulating burrows. Lacking other sensory input, ultrasonic “clicks” emitted by the shrews were ‘used to ditinguich between open and clred tubes up to 61 em in length, and at 30.5 em this distinction was possible for openings ax final as 0.63 em in deter and around corners up to 90". The shrews alko could distinguish among diferent kinds of meterials Locking the tubes. Echolocation “licke” were recorded from 30 to 55 KH (Gould etal, 1964; Tomas, 1979), ONTOGENY AND REPRODUCTION. The. breeding season of B. brevicauda lasts from early February to September (Christian, 1969; Pearson, 1944). Females in estrus were caught ‘nearly January, when sexual maturation of males wes ony begin ning (Christan, 1969} conversely, males in heeding conion were ‘aught in mid October (Pearson, 1944), Dapeon (1968) recorded the capture in February of shortsiled shrew that must have been born in January or December. Two peaks of breeding, in spring tnd late summer or early autumn, have been noted (Blair, 1940: Hamilton, 1929), Athough Pearson (1944) assumed that there was ‘po postpartum estrus in his captive females, lus (1971) concluded that evidence from wild-caught shrews was too indirect to conclude that it never oceurs, Both authors documented instances of ests foeurring after the desth of « iter. CCopslation in B. Brevicauda lasts ae long as 25 min and averages 5 min. The shrews are locked together, probably hy “the penis becoming rigid after it has passed around one or more sharp bends inthe vagina” (Pearson, 1944:77). Erections were observed in sleeping shrews, andthe rigid glans took on a Ast, leak sppearance in an Shape, caused parly by erectile tisues and parlly by s pai of retractor pens muscles (Gibbs, 1955; Pearson, 1944). During copulation, the female usualy is active and drags the inactive male behind her. No posteopalatory plug is formed in the vagina, After copulation, the male hed to use his mouth to retract the peis. Twenty or more matings in 1 day wore observed: tt Teast six matings/day are required to induce ovulation, which MAMMALIAN SPECIES 261 usually occurs from 85 to 71 h after the fist copulation and never faccurs inthe absence of copulation. Receptvity ofthe female de creases {ovulation has occurred but may last for a long at 1 ‘month if thas not. Daring pregnancy, the corpora ates regress (Pearson, 1944). Asdell (1965) suggested that the placenta pro- {uces enough progesterone to continue pregnancy ‘Gestation lasts 21 or 22 days (Hanson, 1925; Pearson, 1944) Average liter sizes of “ix or seven™ (Hamilton, 1929:134), 4.7 (Bus, 1971), and 4.5 (Pearson, 1944) have been reported. Harn iton (1929) described a liter of seven neonates as being naked (except for vibriske, which averaged 1 mm i lengthy dark pink, land about “honeybee size,” ard as having closed eyes and ears. At 2 days, external measurement ir mum) were total length, 31; length of tall 4: and length of hinfoot, 4.5: weight averaged 1.38 g. At “days of age they weighed an average of 3.8 and were 48 mam in total length. At 8 days they weighed 6.2 g and fad standard reasarements of 61, 9.3, and 9. BY that time, hair had appeared But teeth had not yet erupted; the shrews were noted to emit a sucking sound and” were able to crawl. At 13. days, the young weighed 9'g and measured 73, 12, 16: sex was dcernabe by the {ppesrance of mame in females, At 19 days, the upper incisors had appeared through the gum, weight was 99 g, and total length ‘vas 81 mm, On day 22, when the last ofthe litter died, the eyes tad not opened although the external ear was prominent. Weaning accurs by 25 days of age (Bhs, 1971). Pearson (1944) mentioned {captive female that demonstrated receptivity at 47 days of age fn noted that captive males had spermetono in ther teste, which were nealy of adult sae, at 50 days of age. The earest succesful Trceding of « male recorded by Poeran (1944) was at 83 days of age: Bl (1971) obverved a male to breed succesfully at 65 days B brevicauda born spring mature more rapily than those born in outumn, and some breed inthe rame season in which they are Born (Blus, 1971; Dapson, 1966; Pearson, 1944), In mark and release experiments by Pearson (1945), 6% of ‘the orginally marked population war recaptured in the second sun rer. One wi-eaught, captive female lived to atleast 30 onthe ‘of age, and one captivetorn male lived 33 mnths. Blas (1971) Studied morality in a captive colony and found that 11.1% lived ‘hore than 1 year. The number of young that survived from bith to weaning was 72.6%, Average minimal survival for females and male waa 4-4 and 46 mont, respectively. Age may be deter- tine from the degree of totinear, mith maximum wear indicating tan age of shout 18 months (Pearson, 1945). ECOLOGY. Ferthworms (Oligochaeta) (Munford and Whi kee, 1982; Whitaker and Ferrazo, 1963) or milipedes (Dplopoda) (Lingey and Linzey, 1973) make up © major portion ofthe det of B. brevicauda. Hamilton (1941) analyzed 460 stomachs and found that the majority contained insects and annelids and that (in de creasing order of frequency) plant material, centipedes (Chilopede), trachnids, molluscs, and vertebrates also were represented. He as Serted that shorted shrews were not heavy predators on field tice, 2s they hed the eputatio of being (Merriam, 1684). Eade (198%, 1948) analyeed Blarina feces during high and low popula: Sion ejeles of Mierotus pennsyleanicus and found that, Whereas insocts predominated in the diet of B. Brevicauda even when voles ‘rere most numerous, the dct included more voles during periods ot high vole density than during low vole density. Eadie (1944, 1952) estimated that three shrews consumed 14 to 27 mice per 25 ba during the winter months, thereby acting as an effective ‘control on microtine populations. Allen (1938) and Platt and Blake ley (1973) thought that mice might become important in the diet of Blarina when insects are relatively unavailable. In adtion, B. Srevicauda reportedly has preyed on Sorex (Eadie, 1940; Ham ton, 1940), « young Lepus americanus (Rongstad, 1965), a ribbon snake, Thamnophis =p. (O'Reily, 1949), 2 6Q:em water snake, Nerodia sp. (Cope, 1873), and slimy silamander, Plethodon alutinosis (Hamilton, 1936). Endogone and otber fungi (Diehl, 1939; Whitsker, 1962) sometimes are inchided in their det. Short. tailed shrews store food for fre use (Hamilton, 1930; Robinson sand Brodie, 1982), expecially sails (Gastropoda) (Cleneh, 1925: Ingram, 1942). Martin (1984) found that food-hoarding by short tailed shrews occurred primarty in auturin and winter although it could be induced in suramer by «sudden abundance of prey ‘Species predatory on B. brevicauda include: owls dewolius acadicus, Asio otus, A. flammeus, Bubo virginianus, Otis asio,MAMMALIAN SPECIES 261 and Stris varia (Chose, 1972b; Dexter, 1978; Gets, 1961; Kike patiek and Conway, 1947; Mumford and Whitaker, 1982; Pearson End Pearson, 1987; Willams, 1936} hawks Bute lagopus, Cir us cyaneus, and Falco spaeeras(Mrnford and Whitaker, 1982} Striker Lantus ap. Jackaoo, 1961) snakes Nevada ep, kis trodon contort, Pitwophis melanoleucus, and members ofthe Grotalinae Gckeon, 1901); fede Felis cotus and F rufes (Es Fingtom, 1936; Story ly 1982} canids Canis domeateus, C latrans, Vulpes vulpes, and Urocyon cinercoargenteus (Andrews sd Bogges, 1978: Fowie ae awards, 1955: Hamiton, 1935: Muraford ana Whitaker, 1982 mosteds— Mustela ermine, Mf {frenata, vison, a Mephitis mephtie Haman, 1928, 1959; Mumford and Whitaker, 1982 ctoon, Procyon lotr Hamilton, 1936} oporsum, Didelphisvirginianus (Blumenthal snd Kilian, 1976), Shrews aio have been scovered in the stomachs of lke two, Salvelinus namayeush (Fowe and Edwards, 1955) and green suns, Lepomis eyanelus (ish and Hoffmeister, 1947) ‘A lierature starch on parasites of B. brevicauda produced 127 cations, mos of which were orginal doerptions of [44 eto and endoparstes. Witrock snd Hendrickson (1979) lated 18 be ‘ithe tat occurred in B. brevicouda in Toma, and Mumford and ‘Whitaker (1982) lnted 32 specie of ectoparaites and thre orders of endoperastes (Nematoda, Trematoda, and Cetod) that occurred Ona aborted henna Non Win er os) site examining our biography, reported tht the papers reer to tbe folowing ectoparaae. 2 species of Anoplura; 2 Coleoptera {both leptin ipteran (e citerebrd); 23 Sipbonapteret 3 eat ‘Miter and Gets (1977) clelated that short-tied shrews have Ira habitat requirements but were mort comnten in arese wih ‘ore than 50% herbaceous cover Conversely, Dueser and Shugat (1979) iterated that shorsaied threws in eastern Tennessee have ‘narrow, somewhat epeciald iche, Get (1961) fund that food ves the limiting Factor n wooded habitats type of voetation, cover Temperature, and moisture hed lite effect on local dsuibaon, tlthough shrews avoided arets with ite cover and wi exteres of temperatore and moisture Pruitt (1983, 1939) suggested that deep ter protected shrews in hardwood forest fom tenperatre i npitre extremes. . revicauda was the most ubigutous snd Stunden of Sve specie of mammals wed in farmstead shelter Init in wouthern Minnesota, sed on eaptres in both wooded and ssnwooded habitats (Yahner, 1982, 1983). brevicauda moved between sbeterbels more oiten than other species stlid. In Towa, B. brevionuda was asoiated vith big hiestem. Andropogon ger dards (Plat, 1973} in Quebec, thy oovurredprmariy sn mate deestuow onferous fret and secondary in elds of tall rases Stn edges (Wrigley, 1969). Since (1967) found shorted Shrew: asociated with stone walls in eelatively dry situations in fatern decihuous forest, they suggested that humidity might be Liger near he stone walls than in adjacent microhabtats, thereby nabing shortaled shrews to inhabit otbervize dry areas In ast frm Tennesses, Blarnn consistently oecupiol ares of high sump Sd og density, hard ground, few abrabs, and dense oversor, and they fed on larval insets found inthe stump and loge (Kitcings tnd Levy, 191). Tae subepeciee B. .tlmalestes coours primary in marshy babites in and around the Dial Swamp of Virginia sk North Cazola (Handy, 1979) Plat and Blakeley (1973) investigated the interepecie rela tionship between B.brevicauda and Sorex cinereus, an suggested {hat Sorex populations might be somewhst negatively correlated with density of Blarina. Harson (1940) thought B. Brevicauda Inight have am advere eect on 5 fumeus popsatons, but Jemeson (1949) found the opposite to be rue: Lindeborg (1941) found 4 Poitve correltion between Buctuntions in Peromyscus leucopus nd B. brevicauda, and Calhoun (1963) found evidence thet the presence of P. leucopus on the surface ofthe ground might force Sirewe to remain underground. Zegers and Ha (1981) petulated tha P. leucopus used arborea! abiats to srnimiae competition tril Blarinas In Towa, Heider ot al (1983) found that members the gens Peromyscus reinvadedSooded seas much more quick Iy thm Blarina "Winter mortaty of upto 90% of populations of B. brevicaw dla has boen Jocumetel,prcbably related to stress rom cold (Bar behenn, 1958; Cotwcbang, 1965: Jackson, 1961). Population den sy varies from year to year Jackion, 1961; Plt, 1968), and popustions of shoretaled news occainaly cra, requiring 2 tral years to recover (Onoge and Verme, 1968). Christian (1963) 5 found that mean adrenal weight was related directly to population size. Brenner etal. (1983) concluded that reproduction in B. bree ‘cauda may not be affected adversely by behavioral interaction as itis in microtines, Estimates of population density range from 1.67 hha to nearly 121/ba (Jackson, 1961; Willams, 1936). Estimates of homeange ste usually average about 2.5 he (Bhi, 1940, 941; Buckner, 1966), and the range of each shrew usualy over laps with the range of one or more other shrews, Blair (1940) thought that B. Brevienuda did not have territories defended from other shrews In abandoned strip-mines,B. brevicauda is found ony in older areas with stable, mott environmental conditions (Jones, 1874 Kirkland, 1976; Wetzel, 1958), After a timbered azea is clear-cut, populations of shrews decline abruptly (Kirkland, 1977). Powerline fotrdore seem to be a dispersal barrier for short-taed shrews (Schreier and Graves, 1977), Beeause shrews are predstors, they ‘concentrate DDT residues at levels 10 times those found in Pero imyscus and Clethrionomys (Dimord and Sherburne, 1969). Steho tal. (1976) found that shows significantly increased their con: fumption of arthropods after an area was sprayed with orthene, ‘hus increasing thei intake of pesticide residues. Gets etal (1977) ‘documented concentrations of lead in Blarina adjacent te highways, BEHAVIOR. Martin (1980) described 54 behavioral pt tera in captive shor-taied crews,B.Srevionuda is semifoeoria Tearamal sith rotwaye woul inthe top 10 em of tll bot wth tome 1 deep as 50 cm Lelow the sl surface aaikon, 1981 Jameson, 1943). Runways unslly parallel the srtace bit occa sional’ ascend vertically to it (meson, 1943). Srews dig along fd through old rotten fogs and often ite runways of aerotines a rales (Hamiton, 1981), although there india variation in digging behavior, shrews generally dig with their front fect an, shen ennogh soi) accumulates, hick it fromthe tunnel entrance wrth ther hindee (Rood, 1958). If the dtance to the entrance {reat enough, sbrews o's aidenays somerset and push the dirt tut with thet noses. They dig at rate of approximately 2-5 cm! tin mith Irequent ope for short naps. Shll (1907) and Brooks (1908) found burrow systems of B.brevcoude Bera o honey. comb an aces. Shrews pen relatively litle tie on the surface of the ground (Food, 1988) but bave been reported to climb tees (Carer, 1986). Gets (19618) found them fo be more active on cloudy days then on sunny or rainy day Nests are underground ard spherical in shape, and may be lined with vegetation anderen fur of meadow vl, rots pean. ‘yloanicus (amt, 1929, Rapp and Rapp, 1945; Stal, 1907) However, eaves and grass provided by Rood (1958) for hit captive sbrors were ignored. ‘During lnctaton, male shrews constrict the openings of nests and retnfree the ning materia (Martin, 982)" Aetety of fe tualenincrouss during pregnancy tod lactation, pow s« the Tent of erated utiaonl needs, Female seve retire tt pups hy dragging snd by « behavior simiae to caravanning. The Inter, However, involves ony the female and one young. Alla ternal boavior ceases when the young ace weaned (Martin, 1982). eoos rarely are found inthe nest amon, 1925: Rood 1958; Shull, 1907} they usually are deposited neatly the side ofa runway, outsie the entrance tothe nest or. by expe shrews, inthe corners ofthe cage. B. brevicnud twitches rele when Sheping. The mom cettmon postion swith the ove and pews tucked ander the beliy (Rood, 1958). These shrews rarely stay io one poston for more than a ow mines sd often aroue fo yan seth, and clean themestves before going back to ep If sver familiar inviduals share «cage, they seep together and contatiy try to get tothe bottom of the ple Allen et 1977) quanted Ata ot seep in B. brevicuda Many tutors consider Barina tobe altary and untiendly (ackson, 1961; Metin, 19816: Shal, 1907). Rood (1958), how ‘ver, found that the sciaity of short sae shows depend greatly Sn inlividual dispositions wilh age and sex playing 8 ese role ‘eb ander amas tended tobe les frien than females ad $ounger animal: Dapeion alan sere to have beating on pre {tory pelestions. Some individuals seemed "tered of mice pst into tir cage, others stacked the tice balGbenrtelly, and some stacked without hesitation. Philips (1956:543) noted that shrew frst "faster is teeth ont behind the let ear ofthe vole aed tegen to gnaw atthe bab ofthe bul. I sequred 11 mince for the shrew 0 kilt prey, and dating that ime it was dragged6 oughly and rapidly about the cage, as the vole attempted to shake loose.” Olten (1969) analyzed the agonistic behavior of shorted shrews and recogniaed four setion patterns ard five postures. He Suggested thatthe postures were used in species recognition, thus ‘minimizing the amount of energy wasted in competition for food, space, and social poston GENETICS. Standard karyotypes of B. brevicauda are ccharscteraed by # iplid number of 50, 49, of 48 and a funds rental number of 48 (George et al, 1982). Meylan (1967) con- ‘lided that variation in diploid numbers isthe result of a Bsion~ fusion event between a pair of large acrocentric autoeomes and a pair of small acrocentric sutosomes. The totally acrocentic diploid ‘number of 50 is most common (Genoveays etal, 1977; George et al, 1982}, and » diploid number of 48 bas been found in oly one specimen from central Tinos (Lee snd Zimmerman, 1969). The Xechromoaome is a large metacentic (Genoways etal, 1977; Mey lan, 1967). Gonoways etal. (1977) reported the Y-chromosome to bbe a small acrocentric in B. b, breviewuda and B. b. birtlandi, \nhereas Meylan (1967) reported it tobe a small meiacentric in B. 5 talpoides: George et al. (1982) demonstrated that each specia tion event inthe genus Barina has been accompanied by fixation ‘of chromosomal diferences Examination (using starch gel electrophoresis) of 18 prevump- tive lock in individuals of B. Brevioauda from Massachusers and Pennsylvania revealed that mean beterorygosty was nil and the percent of loci pelymorphic was 11.1 (George, 1984). Brenner and ‘Atno (1983) found seven distinct protein fractions, cepresenting six autosomal genetic tats, in the lens of the eye of B. brevicauda. REMARKS. Based on the revisonary studies of Merriam (1895) and Bole and Moultrop (1042), the genus Blarina was thought to contin two species. brevicauda and B. telmalestes ‘The latter subsequently war shown by Handley (1979) to be a subspecies of B. brevicauda, However, series of papers published Since 1972 (Braun and Kennedy, 1983: French, 1981: Genoways find Choate, 1972; Genoways et al, 1977; George et al, 1981, 1982; Moncref et al 1982; Tate ot al, 1980) has demorsteated {hat the genus consists of atleast three species-—B. brevicauda, B. carolinensis, and B. hylophoga. Moreover, the nominal sub- species B. carolinensis peninsulae may repreeenta fourth species, Finally, the taxon B. carolinensis shermani may prove to be an isolated subspecies of B. Brevirauda or sil another species of Blarina (Jones ets, 1984). ‘The generic name Blarina is « coined name that hae 10 Aecivaton. The specie pit is combisation of two Latin words — brevis and cauda—meening short tal. “There are literally hundreds of citations that pertain to B. revicaude. We have not been able to inchude them all bere, but have attempted to eite those that give the moet complete informa- tion or that summarize other authors work, LITERATURE CITED ALLEN, D. L. 1938, Ecological studies on the vertebrate fauna ‘of a 500acre farm in Kalamazoo County, Michigan. Eco Monogr 8347-836. Auses, H.” 1894. Observations on Blarina brevicauda. Proc. ‘Acad, Nat. Sei, Philadelphia, pp. 269-270 Auten, I-A. 1891,” Notes on colection of mammals from Costa Tica, Bull Amer. Mus. Nat. Hist, 3:203-218, 1897, “Additional notes on Gosta Rican mammals with Aescriptions of new species. Bull. Amer. Mis. Nat. Hist, 9 31-44. Aso, TS. D. Grn, S. M. Busxouove, ax G. L. Devoe. 1977." A behavioral and polygraphic stady of sleep in the shrews of Suncus murinus, Blerina brevicauda, and Cryp {otis parva. Behav. Bil, 20:354-366, AnoERSON, RLM. “1943. Nine additions to the lst of Quebec ‘mammals with descriptions of ix new forms. Ann. Rept. for 1942, Provander Soc. Nat. Hist, Canada, pp. 49-62. 1947. “Catalogue of Canadian Recent mammals, Bull Nal, Mus. Canada, 102:1-288. ANDREWS, RAL, AND E. K. Boccss. 1978, Eoology of coyotes in Towa. Pp. 249-265, in Coyotes: biology, behavior, and ‘management (M. Bekoff, ed.. Academic Press, New Yerk, sent PP nwa CRasre-Linor, A. 1912, Der Bau der Soriiden und ‘hve Berichungen 2 andern Saugetioren. I Zur Entice MAMMALIAN SPECIES 261 Tungsgeschichte der Zahne. Ontogenie. Morpholog. Jahrbuch. ‘44:201-296. ASDELL, &. A. 1965. Reproduction and development. Pp. 1—41 in Physiological mammalogy (W. V. Mayer and R. C. Van Gelder, es.) Academie Press, New York, 2:1-326. Bacawan, J. 1837. Some remarks on the genus Sorex, with a ‘monograph ofthe North American species. J. Acad. Nat. Soi, 362-102, ‘Mammals. In Reports of explorations and ‘to ascertain the most practicable and economic route oad from the Missiseppi River to the Pace Ocean, fora Alpart 11-757. Bayes, 0.1902, Descriptions of two new, insular Harinas from ‘eastern Massachusetts. Proc. New England Zool. Chub 8:75~ 7A Banaeuenn, K. R. 1958. Spatial and population relationships ‘between Merotus and Blarina, Ecology, 39:293-304, BavweauDnen, G.D., ANDI. M. McPutison. 1985. Inraspeciic ‘arahlty in cranial and port-cranil features of Blarina brev- Teauda in Tennesce. J. Tennesse Acad. Se, 60:16-20 Buait, W. F. 1940, Notes on home ranges and populations of ‘the shor-tiled threw. Ecology, 21:284-288, T941. Some dats on the home ranges and general life history of the shortaled shrew, red:backed vole, and wood land jumping moure in northern Michigan. Amer. Midland Nat. 25:681-685. BuuMeNTiaL, E. M., AND G, L. KIRKLAND, JR. 1976, The biology ofthe opossum, Didephis virginiana, in soutacentral Pennsyl- ‘ania, Proe. Pennsylvania Acad. So, 50:B1-B5. Bus, LJ. 1971. Reproduction and survival of shar-ailed shrews (Blarina brevicauda) in eaptwvity. Lab. Anim. Sei, 21:084— B91. Bout, B.P., Jn ano P. N. Moutrinor. 1942. ‘The Ohio Recent ‘mammal colletion in the Cleveland Museum of Natal Hie tory. Sei Publ, Cleveland Mus. Nat. Hit, 5:83-181 Bowtts, J. B._1973. Distribution and biogeography of mammals of Towa. Spee. Publ. Mus., Texas Tech Univ., 9:1-184 Brawn, JK. AND M. L. Kewvepy. 1983. Systematics of the ‘genie Blarina in Tennessee and adjacent areas. J. Mazam. O44 4-425, Busnsen, F. Jn aN D. K. ATN0. 1983. Morphological and eye lens protein characters of three species of small mammal, Proc. Pennsylvania Acad, Sci, 97:173-176, Buren, FJ, RB. Ket, xD J. KFLLY. 1983. Physiological ‘characteristics of three species of sal mammals on surface ‘mined lands, Proc. Pennsylvania Acad. Sei, 57:41~44 Buooks, F. E.. 1908. Notes on the habits of mice, moles, and ‘htews, West Virgina Univ. Agric. Exp. Sta, Buy, 113:89- 158. Bow, B. 1908. The Conard Fissure, a Plistocene bone deposit, in northern Arkansas: with a description of two new genera fand twenty new species of mammals. Mem. Amer. Mus. Nat. His, 9:157-208, Buckner, C. H. 1964, Metabolism, food capacity, and feeding Toehavior in four specien of shrews. Canadian J. Zool, 42: 259-279, 1966, Populations and ecological relationships of shrews in tamarack bogs of southeastern Manitoba. J. Maran., 47 Tal-195, CALHOUN, J. B. 1963. The socal use of space. Pp, 1-187, in Physialogical mamsmslogy (W. V. Mayer and R.C. Van Cali er, eds) Academie Press, New York, 1:1-381 antes, T. D. 1936. The shortaied shrew as a tee climber. J. Maun. 17:285. Cues, RM. 1951. The water exchanges of some small mam: nals, Ecol, Monogr, 21:215-225, Cwoxre, J. R. 1968. Dental abnormalities in the shortailed threw, Blaring brevicaud, J. Mar, 49:251-258, 1970. "Systematic end aoogeography of Middle Amer. jean shrews of the genus Crypto. Univ. Kanses Pub, Mus. Nat. Hist, 19:195-317, 1972a. Variation within and among populations of the shorttailed shrew in Connecticut. J. Mam, 53:116-128. T9T2b, Notes on geographic distribution and habitats ‘of mammals eaten by owls in southern New England. Trans Kansas Acad, Sei, 74:212-216, 1975. [Review of] A manual of mammalogy: with keysMAMMALIAN SPECIES 261 to families of the worl, by A. F. DeBlase and R. F. Martin. 1. Mamm., 56:281-283 unistian, J.J. 1963. Endocrine adaptive mechanisms and the ‘pbsologic regulation of population growth. Pp. 189-353, cm Physiologic mammalogy (MV. Mayer and R.G. Van Geer, wis). Academic Press, New York, 1:1-381 1969. "Maturetion and breeding of Blarina brevicauda in winter. I. Mamm,, 50:272-276. Gurncn, W. J.” 1925. Snails eaten by shrews. Nautilus, 39:28, Core, ED. 1873. Ons habit of species of Blarina. Amer. Nat, 7490-491 1899, Vertebrate remains from Port Kennedy bone deposit. J. Acad. Nat. Se, Philadelphia, 11:193-207. Crosoy, E-C., an T. Huwuicy. 1044. "Studies of the vere rate telencephalon IL, The amygdaloid complex in the shrew (Blarina brevicouda). j. Comp. Neurol, 81:285~305, Darsox, R. W. 1968. Reproduction and age structure in «pop: lsion of shorttaled shrews, Blarina Brevieauda. J. Maryn, 149:205-214, DeMevtes, D. H. 1954, Possible antiadrenalin action of shrew ‘venom. J. Mamm., 35:425. Dexren, RoW. 1978, Mammals utilized as food by owls in ‘reference tothe local fauna of northesstern Ohio. Kiland, 2ad6. Dieu, W. W. 1959. Endogone as enimal food. Science, 90: daa DWoNd, J.B. AND J. A. SHERBURNE, 1969, Persistence of DDT in wld populations of small mammals. Nature, 221:486—487, DoneMes, HM. “1965. Heart rate, temperature and respiration ate of the short-talled shrew in captivity J. Mami, 46:424— 35. DoneMvs, H. M., AND J.J. JArv=. 1964. Hematologic values for the shortialed shrew, Blarina brevicauda. J. Mamm., 45 648-699. Dusen, RD., ano H. H. SkucaRT, JR. 1979. Niche patterns in's forestloor stallmamma fauna. Ecology, 60:108-118. Duvesnoy, G. L. 1842. Notices pour Servir& la Monographie au Genre Musaraigne, Mag. Anat, Comp. Palsontol, sar. 2, 1-48 Exois, W.R. 1938, The dermal glands of shrews. J. Mamm., igs71-174, 1914._ The shorted shrew and feld mouse predation. 1. Mamm., 25:389-368. 1947, Homologiee of the male accessory reproductive lands in Sorex and Blarina, Anat, Rec., 98:347~354, 1948. Shrew-moute predation daring low mouse abun. dance. J. Mamm., 29:3 1949.” Predation on Sorex by Blarina, J. Mamm., 30: 308-309, 1952. Shrew predation and vole populations on a lo: alized area. J. Mara, $3:185~189, Buss, LS, V. E. Dietsine, aN D. F. Horrweisten, 1978 "Pazonomie status of shorttaled shrews (Blarina) in Minis J. Mamm., 59:305-311 Fuss, S., an 0. Knaven. 1955. Properties ofa toxin fom the “alivery gland of the shrew, Blarina brevicaida. J. Pharmacol, Exp. Therap., 114127137 Ennincton, P. L. 1936. Notes an food habits of southern Wis- ‘onsin house eats, J. Mamm., 17:68-65, Fivouy, JS. AND J. K. JONES, TR. 1956. Molt of the short tailed shrew, Blarina brevieauda. Amer. Midland Nat, 36: 246-240, Fow1s, C.D.,aXpR. Y. Enwanos, 1953. An unusual abundance of shorttaled shrews, Blarina brevicauda. J. Mamm., 36: 36-41, France, T. W. 1981. Notes on the distribution and taxonomy of short.taled shrews (gems Blarina) inthe Southeast. Bem leyana, 6:101-110, Garren. 1830. Observations on the quadrupeds found in the distiet of Upper Canada extending between York and Lake Simcoe, with the view of ilustreting their geographical distri. bution, a6 well as of describing some species hitherto unno teed. Zook. J, $:201-207. Gavetitan, G. R. L. 1954, A comparative study of the osteclogy ‘and myology ofthe cranial and cervieal regions ofthe shrew, Blarina brevicauda, ad the tole, Scalopus aquaticus, Mie Publ. Mus. Zool, Univ. Michigan, 80:1-82. Graoways, Ho HL, ano J. R Cvoxre, 1972. A mubiva ‘analysis of systematic relationships among populations of the shortaled shrew (genus Blarina) in Nebraska. Syst. Zoo 21:106-116. Gesowars, HHL, J. C. Parton, IL, ano J. R. Cuoate. 1977, ‘Karyotypes of shrews of the genera Cryptotis and Blarina (Mammalia: Soricdae). Experientia, $3:1294-1295. Gronce, §.B. 1984. Systematics, evolution, and historical bio- ‘geography ofthe Soricinae with epecal reference to the gens Sorex. Unpubl. Ph.D. Disert., Univ. New Mexico, Albaquer. ‘ae, 78 pp, Gvonct, 8. BJ. R. Cuosre, AND H. HL Gavoways. 1981. Dis: ‘tribution and taxonomic tatus of Blarina hylophaga Flot (Insectivora: Sorieidae). Ann. Carnegie Mus., 50:493~513. Goonce, SB. H. H. Gexoways, J. R. Coste, aNo R. J. Bakes. 1982. _Karyotypic relationships within the short-tailed shrews, genus Blarina. J. Mam, 63:639-645. Ger, LL 1961a. Factors influencing the local distribution of ‘shrews, Amer. Midland Nat, 65:67-88. 1961. 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