HARVARD UNIVERSITY Ernst Mayr Library of the Museum of Comparative Zoology r r r- OCCASIONAL PAPERS ^^^ of the bcu ^ 1 1995 NATURAL HISTORY MUSEUM The University of Kansas | y Lawrence, Kansas NUMBER 175, PAGES 1-49 20 DECEMBER 1995 A Key to Recent Soricidae of the Western United States and Canada Based Primarily on Dentaries Leslie N. Carraway Department of Fisheries and Wildlife Nash 104 Oregon State University Corvallis, Oregon 97331-3308. USA ABSTRACT Dentaries of Notiosorex crawfordii, Ciyptotis parva, three species of Blarina. and 26 species of Sorex from west of the Mississippi River (U.S.) and 94°W (Canada) were examined for distinguishing characters. Additionally, subspe- cies of four species of Sorex that exhibited sufficient morphological differences were considered separately. Twenty quantitative characters and states for 12 quali- tative characters were used to develop a key that can resolve the identifications of the 35 taxa under consideration primarily by use of the dentaries. Key words: Soricidae; Blarina: Ciyptotis; Notiosorex: Sorex: Dentary; Mandible; Identification; Western North America. In the western United States and Canada, most soricid remains in fossil matrix, raptor pellets, and carnivore droppings are dentaries (and some rostral elements). However, few species-specific characters for dentaries have been described, and skin characteristics often are the primary basis by which some species of shrews are distinguished. Therefore, when the skin is missing or deteriorated beyond use, identification to species is difficult. Ognev (1962) andGureev (1971) used a combination of nearly equal num- bers of dentary and cranial characters to describe and distinguish taxa of fossil and Recent Palearctic soricids. In describing the subfamilies and © Natural History Museum, The University of Kansas. Lawrence. 1SSN:009 1-7958 2 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 genera of soricids, Repenning (1967) considered a combination of charac- ters of the mandible and rostrum. Subsequently, application of dentary characters to separate species of soricids was limited and sporadic (Butler et al., 1989; Carraway, 1990; Diersing, 1980; Foresman and Jensen, 1992; Handwerk, 1987; Harris and Carraway, 1993; Hausser and Jammot, 1974; Jammot, 1972a,/7;Junge and Hoffmann, 1981; Jungeet al., 1983; Mullican and Carraway, 1990; Ruprecht, 1971; Schaefer, 1975; Vesmanis et al., 1980; Wilson, 1985; Woodman andTimm, 1992). The purpose of this paper is to provide a means of identifying soricids from western North America, north of Mexico, to species based primarily on features of the dentary. Identification of soricid species should facilitate a better understanding of the diet composition of predators and the structure of fossil and Recent small-mamiTial communities. MATERIALS AND METHODS Specimens of Notiosorex crawfordi , Cryptotis parva, three species of Blarina, and 26 species ofSorex from west of the Mississippi River (United States) and 94°W (Canada) were identified by use of published cranial and skin characteristics (Carraway, 1987, 1990; Diersing, 1980; Hall, 1981; Hennings and Hoffmann, 1977; van Zyll de Jong, 1980, 1982). Blarina hrevicauda also was distinguished by use of morphological characters, but specimens of B. carolinensis (2n = 37-39, 46, FN = 44^5) and B. hylophaga (2n = 52, FN = 60-62) were those for which diploid number and fundamental number had been determined (George et al., 1982). Subspe- cies of S. hendirii, S. monticolus, S. sonomae, and S. trowbridgii that exhibited sufficient morphological differences were considered separately. Complete dentaries for each taxon were examined for distinguishing characters. Specimens of taxa examined were selected from throughout their western distributions and included all age classes (Jackson, 1928). Nevertheless, it is possible to encounter individuals that exceed the range of variation of these samples. Only one karyotyped Blarina carolinensis from west of the Mississippi River was available; consequently, 15 from elsewhere in the range were used to verify identifying characteristics. For each dentary, states for 12 qualitative characters (Fig. 1 ) were evalu- ated and 20 quantitative characters (Fig. 2; Tables 1, 2) were measured by use of an ocular micrometer and ocular protractor mounted in a Bausch and Lomb binocular microscope. Terminology for tooth morphology follows that of Butler et al. (1989), Choate (1970), Dannelid (1989), and Hershkovitz (1971). Dentary tooth nomenclature follows that of Hall (1981): //, cl,p4,ml,m2, and wi. Some characters in the key were used previously (Butler et al., 1989; Carraway, 1987; Carraway and Verts, 1994; Dannelid, 1989; Harris and A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE n Fig. 1. Labial (top) and lingual (middle) views of dentary, and occlusal (bot- tom) view of mandibular tooth row of Sore.x to illustrate qualitative characters examined. Key to characters: a = denticle; b = interdenticular space; c = paraconid; d = hypoconid; e = coronoid spicule; f = mental foramen; g = superior condylar process; h = interarticular area; i = inferior condylar process; j = lower sigmoid notch; k = superior opening of temporal fossa; 1 = internal temporal fossa; m = inferior opening of temporal fossa; n = paraconid; o = protoconid; and p = hypoconid. Scales in this and all other figures are 1 mm. UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 Fig. 2. Labial (top) and occlusal (bottom) views ofaSorex dentary to illustrate quantitative characters measured. Key to characters: a = angle e (i.e., il set at an angle e from the horizontal ramus of the dentary); b = length of iJ (used only with Sorex hendirii); c = length of mandibular tooth row (i.e., cl-fn3)\ d = length of cl ; e = length of p4; f = length of/?//; g = length of m2: h = length of nd: i = height of pigmented portion of tfil ; j = height of unpigmented portion of nil : k = height of ml :\ = distance between il ax\Ap4; m = depth of dentary below /;;/; n = length of dentary; o = height of coronoid process; p = length of coronoid-condyloid pro- cesses; q = width of (7; r = width of p4; s = width of ml ; t = width of m2; and u = width of m3. Note that the length of the dentary is measured from the pericentral margin of the alveolus of il to the labial edge of the inferior condylar process, and that the horizontal line at the base of the dentary presents the orientation for the determination of angle e. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 5 Carraway, 1993: Handwerk, 1987; Hausser. 1984; Repenning, 1967). For dimensions in which tooth designations are separated by a dash (e.g., cl- m3), the specified teeth were measured as a unit from the most anterior point to the most posterior point. For dimensions in which tooth designa- tions are separated by a plus (+), the length of each tooth was measured individually and the values summed. This distinction is necessary, because unlike the mandibular teeth of most mammals, soricid mandibular teeth overlap one another to a considerable extent. The key is dichotomous. Because some species are composed of geo- graphic races of disparate sizes, it is possible for a species to be identified in more than one couplet. Camera-lucida tracings illustrate states of the qualitative characters examined and various views of the dentaries. Figures of the dentaries are drawn to the same scale and in the same orientation; however, they are not all printed to the same scale because of publishing constraints. All scale bars represent 1 mm. Species comparisons of taxa of similar size or close geographic proxim- ity either now or possibly during the Pleistocene follow the key. In a few instances in which no quantitative or qualitative dentary characters could be discerned, rostral characters were used. Species are listed in alphabetical order. KEY TO THE DENTARIES 1. In labial view, alveolus of il extending posteriorly beneath at least part of paraconid of ml (Fig. 3, left) 2 In labial view, alveolus of il not extending posteriorly beneath paraconid of ml (Fig. 3, right) 7 S. hoyi Fig. 3. Labial view of the anterior half of dentaries to illustrate Sorex hoyi (KU 37149) with the alveolus of // (lower arrow) extending beneath ml (upper arrow), and S. monticolus (OSUFW 4747) with the alveolus of // not extending beneath ml . 6 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 2(1). Internal temporal fossa composed of an inferior triangular open- ing leading directly into a superior depression; il with three denticulations (Fig. 4); length of dentary usually < 6. 1 mm; length of cl-m3 < 4.2 mm; height of coronoid process usually < 3.4 mm Sorex hoyi. Pygmy shrew (Fig. 4) Internal temporal fossa composed of an inferior round opening separated from a superior depression by a bar (Fig. 4); // with one or two denticulations (Fig. 4); length of dentary > 6.1 mm; length of cl-m3 > 4.2 mm; height of coronoid process > 3.4 mm 3 3(2). Area between condylar processes deeply emarginate — i.e., breadth of interarticular area markedly less than that of superior condylar process (Fig. 5); internal temporal fossa characterized by small, inferior round opening separated from a superior depression by a wide bar (Fig. 4); no pigment present on m2 and m3; height of unpigmented portion of ml > 0.9 mm Notiosore.x Crawford i, Desert shrew (Fig. 4) Area between condylar processes shallowly emarginate — i.e., breadth of interarticular area approximately equal to that of supe- rior condylar process (Fig. 5); internal temporal fossa character- ized by a large, inferior round opening separated from a superior depression by a narrow bar (Fig. 6); pigment present on m2 and m3; height of unpigmented portion of ml < 0.8 mm 4 4(3). Inferior condylar process not completely obscured in labial view (no groove separating process from lower sigmoid notch); coronoid spicule small, not extending beyond posterior edge of coronoid process in lingual view (Fig. 6); no basin in lingual side of interarticular area (Fig. 5); length of dentary < 7. 1 mm; length of cl-m3 <4.9 mm; length of coronoid-condyloid processes < 3.5 mm; height of coronoid process < 4. 1 mm; width of m2 < 1 .0 mm Cryptotis parva. Least shrew (Fig. 6) Inferior condylar process obscured in labial view (Figs. 7); coronoid spicule large, usually extending beyond posterior edge of coronoid process in lingual view; basin may be present in lingual side of interarticular area (Fig. 5); length of dentary > 8.3 mm; length of cl-m3 > 5.8 mm; length of coronoid-condyloid processes > 4.3 mm; height of coronoid process > 5.0 mm; width ofw2> 1.1 mm 5 A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE ingual occlusa Sorex hoyi lingual occlusal Notiosorex crawfordi Fig. 4. Dentaries of 5o7-ev hoyi (ROM 10736) and Notiosorex crawfordi (MSB 47947). In both taxa, note number of denticulations, relationship of posterior edge of il to ml , and structure of internal temporal fossa. Sore.x hoyi has pigment on posterior portion of ventromedial edge of// and one posterolingually directed ridge on p4. Notiosorex crawfordi lacks pigment on m2 and m3. Blarina brevicauda Sorex Notiosorex Cryptotis & ^ & Blarina carolinensis I I Fig. 5. Posterior views (labial to left) of condylar processes of soricids. Note the differences in the relative widths of the superior and inferior condylar pro- cesses, the amount of emargination in the lingual side of interarticular areas, and the presence or absence of a basin in the lingual side of interarticular areas. UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 occlusal Cryptotis parva Fig. 6. Dentary of Cryptotis parva (OSUFW 4475). Note the numbers of denticulations, the relationship of the posterior edge of // to ml , size of the coronoid spicule, and structure of the internal temporal fossae. 5(4). Height of coronoid process usually > 6.0 mm; length of cl-rn3 usually > 6.5 mm; length of coronoid-condyloid processes usually > 5.2 mm; mental foramen positioned beneath hypoconid of ml ; basin in lingual side of interarticular area (Fig. 5) Blarina brevicauda, Northern short-tailed shrew (Fig. 7) Height of coronoid process < 6.0 mm; length of cl-rn3 < 6.5 mm; length of coronoid-condyloid processes usually < 5.0 mm; mental foramen positioned beneath midpoint between protoconid and hypoconid of /;//; no basin in lingual side of interarticular area (Fig. 5) 6 6(5). First lower incisor set at an angle < 17° from the horizontal ramus of the dentary (Fig. 7) Blarina carolinensis, Southern short-tailed shrew (Fig. 7) First lower incisor set at an angle > 18° from the horizontal rainus of the dentary (Fig. 7) Blarina hylophaga, Elliot's short-tailed shrew (Fig. 7) 7( 1 ). Length of dentary usually > 7.7 mm; height of coronoid process > 4.0 mm; length of cl-m3 > 4.9 min 8 Length of dentary usually < 7.7 mm; height of coronoid process < 4.0 mm; length of cl-m3 < 5.0 inm 22 A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE labial lingual occlusal Blarina brevicauda lingual occlusal Blarina carolinensis occlusal Blarina hylophaga Fig. 7. Dentaries of Blarina brevicauda (OSUFW 1111), B. carolinensis (OSUFW 3325). and 5. hylophaga (FHSU 10166). In all taxa. note the numbers of denticulations, the relationship of the posterior edge of // to wz/, the sizes of the coronoid spicules, the structures of the internal temporal fossae, and the obscured inferior condylar process in labial view. The mental foramen is located beneath the hypoconid oiml in B. brevicauda, whereas it lies beneath the midpoint between the protoconid and hypoconid oi ml in B. carolinensis and B. hylophaga. Note angle of il relative to horizontal ramus of dentary in B. carolinensis and B. hylophaga. 10 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 S. bendirii S. sonomae occlusal surface S. bendirii S. monticolus lateral-oblique surface Fig. 8. Views of c] and p4 (anterior to left) to show posterolingually directed ridges on the occlusal surface of p4 of So re.\ bendirii (OSUFW 4873), S. sonomae (KU 145866), 5. I^endirii (OSUFW 4873), and 5. monticolus (OSUFW 4815). Note the presence of two ridges in S. bendirii and only one ridge in the other Sore.x. 8(7). Two posterolingually directed ridges on occlusal surface of p4 (Fig. 8); length of cl > length of p4 9 One posterolingually directed ridge on occlusal surface of p4 (Fig. 8); length of cl < length of p4 10 9(8). Width of cl + width of p4 usually > 1 .8 mm; length of /7 usually > 5.4 mm (J = 5.85, SE = 0.07, // = 29, range 5.2-6.5, CV = 0.06) Sore.x bendirii palmeri. Pacific water shrew (Fig. 9) Width of c! + width of p4 usually < 1.8 mm; length of il usually < 5.4 miTi (.f = 5.16, SE = 0.05, n = 29, range 4.7-5.7, CV = 0.05) all other Sore.x bendirii. Pacific water shrew (Fig. 9) 10(8). No strip of pigment on inside of ventromedial edge of iJs where they meet (Fig. 10) 11 Strip of pigment on inside of ventromedial edge of iJs where they meet (Fig. 10) 13 A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 11 labial occlusal Sorex bendirii palmerii labial occlusal Sorex bendirii (other subspecies) Fig. 9. Dentaries of Sorex bendirii palmeri (OSUFW 1483) and other subspe- cies of 5. hendirii (represented by USNM 233594). Note the relationship of the posterior edge of il to ml , the presence of pigment on the posterior portion of the ventromedial edge of //, and the two posterolingually directed ridges on p4. S. monticolus S. arcticus Fig. 10. Lingual views (anterior to the left) of il of Sorex monticolus (OSUFW 4747) and S. arcticus (KU 45299) to show differences in the posterior extent of the pigment on the ventromedial edge of //. 12 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 1 1(10). Length of cl-ni3 < 2.7 mm; length of dentary < 6.0 mm; depth of dentary below nil < 0.6 mm; height of coronoid process < 2.7 mm Sore.x minutissimus, Holarctic least shrew (Fig. 1 1) Length of cl-m3 > 4.5 mm; length of dentary > 6.8 mm; depth of dentary below ml > 1 .0 mm; height of coronoid process > 3.7 mm 12 12(11). Pigment on /7 in two or three sections (Fig. 1 1); height of coronoid process > 4.4 mm; length of coronoid-condyloid processes > 3.8 mm; length of dentary > 7.9 mm; depth of dentary below /;/7 + height of coronoid process > 5.6 mm Sore.x arcticus, Arctic shrew (Fig. 11) Pigment on // in one section (Fig. 1 1) may be obscured in old adults; height of coronoid process < 4.2 mm; length of coronoid- condyloid processes < 3.6 mm; length of dentary < 7.9 mm; depth of dentary below ml + height of coronoid process < 5.2 mm Sore.x timclrensis. Tundra shrew (Fig. 1 1) 13(10). Height of coronoid process < 4.2 mm; length of coronoid-condy- loid processes usually < 3.6 mm; depth of dentary below mJ + height of coronoid process usually < 5.2 mm 14 Height of coronoid process > 4.3 mm; length of coronoid-condy- loid processes > 3.6 mm; depth of dentary below ml + height of coronoid process usually > 5.3 mm 16 14(13). Mental foramen usually positioned beneath posterior half of /;// (Fig. 12) Sore.x monticolus. Dusky or Montane shrew (Fig. 12) Mental foramen usually positioned beneath anterior half of ml (Fig. 12) 15 15(14) First lower incisor set at an angle 8-10° from the horizontal ramus of the dentary (Fig. 12); distance between il and p4 < 0.3 mm. . ..Sore.x trowhriclgii montereyensis, Trowbridge's shrew (Fig. 12) First lower incisor set at an angle 10-15° from the horizontal ra- mus of the dentary (Fig. 12); distance between il and p4 > 0.4 mm all other Sore.x trowhriclf^ii. Trowbridge's shrew (Fig. 12) A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 13 labial JV^ t occlusal Sorex minutissimus lingual occlusal Sorex arcticus lingual occlusal Sorex tundrensis Fig. 11. Dentaries of Sore.x minutissimus (UAM 19268), 5. arcticus (ROM 1 1622), and 5. tundrensis (UAM 13561) showing relationship of posterior edge of /'/ to »;/. the lack of pigment on the posterior portion of the ventromedial edge of /7, and one posterolingually directed ridge on p4. Note number of pigment sections on il of 5. arcticus and S. tundrensis. 14 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 labial lingual occlusal Sorex monticolus (other subspecies) labia occlusal Sorex trowbridgii montereyensis labial lingual occlusal Sorex trowbridgii (other subspecies) Fig. 12. Dentaries of Sore.x monticolus (OSUFW 6055). S. trowbridgii montereyensis (KU 68098), and other subspecies of 5. trowlvidgii (represented by OSUFW 6111). Note the relationship of the posterior edge of /'/ to /;;/ , the position of the mental foramen relative of a///, one posterolingiially directed ridge on/?-/, and the presence of pigment on the posterior portion of the ventromedial edge of //. In S. monticolus and other subspecies of 5. trowbridgii note the very deep interdenticular space. In all 5. trowbridgii note the angle of // relative to the horizontal ramus of the dentary. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 15 S. monticolus S. bairdii Fig. 13. Labial views (anterior to the left) of the ils of Sorex monticolus (OSUFW 4747) to show the deep interdenticulations and of S. hairdii (OSUFW 7261) to show the shallow interdenticulations. 16(13). Interdenticular space very deep (Fig. 13), may be obscure in old adults; length of coronoid-condyloid processes usually < 4.1 mm 17 Interdenticular space shallow (Fig. 13); length of coronoid-condy- loid processes > 4.1 mm 19 17(16). Height of coronoid process usually >4.5 mm; width of r7 -i- width of p4 usually > 1.4 mm Sorex monticolus neomexicanus Dusky or Montane shrew (Fig. 14) Height of coronoid process usually < 4.5 mm; width of r/ -i- width of p4 usually < 1.4 mm 18 labial occlusal Sorex monticolus neomexicanus Fig. 14. Dentary of Sorex monticolus neomexicanus (MSB 37345) showing very deep interdenticular space, relationship of posterior edge of // to ml , one posterolingually directed ridge on p4, and presence of pigment on the posterior portion of ventromedial edge of //. 16 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 18(17). No dentary characters discerned, but prominent inframaxillary ridge (Fig. 15) Sorex alaskanus, Glacier Bay water shrew (Fig. 15) No inframaxillary ridge, only a depression (Fig. 15) Sorex palustris. Water shrew (Fig. 15) labial lingual occlusal Sorex alaskanus oblique rostrum labial lingual occlusal Sorex palustris oblique rostrum Fig. 15. Dentaries and rostrums (in oblique view at lower right) of Sorex alaskanus (USNM 97712) and S. palustris (OSUFW 4857). Note in the dentaries of both taxa the very deep interdenticular space, the relationship of the posterior edge of // to /;;/, and one posterolingually directed ridge on p4, and presence of pigment on the posterior portion of the ventromedial edge of //. The rostrum of 5. alaskanus has a prominent inframaxillary ridge, whereas that of S. palustris has a depression. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 17 19(16). No dentary characters discerned, but no projection on medial edge of lis (Fig. 16); body of lis completely abutting for length of pigment or at least at dorsal aspect of pigment (Fig. 16); maxillary and mandibular unicuspids robust for skull size; zygomatic pro- cesses of maxilla rounded 20 Projection present within the pigment on the medial edge of lis (Fig. 18); maxillary and mandibular unicuspids quite small for skull size; zygomatic processes of maxilla pointed 21 20(19). Length ml usually > 1.8 mm; width off/ + width of p4 usually > 1.9 mm Sorex sonomae sonomae. Fog shrew (Fig. 16) Length of ml usually < 1.7 mm; width of r7 + width of /;4 usually < 1.8 mm Sorex sonomae teneUiodus. Fog shrew (Fig. 17) labial occlusal Sorex sonomae sonomae Fig. 16. Dentary and anterior view of // (at lower right) of Sorex sonomae sonomae (OSUFW 8876). In the dentary note the shallow interdenticular space, the relationship of the posterior edge of z7 to ml, one posterolingually directed ridge on p4. and presence of pigment on the posterior portion of the ventromedial edge of //. The medial edge of each // lacks a projection. UNIV. KANSAS NAT. fflST. MUS. OCC. PAP. No. 175 labial occlusal Sorex sonomae tenelliodus Fig. 17. Dentary of So re.\ sonomae tenelliodus (OSUFW 7206) showing the shallow interdenticular space, the relationship of the posterior edge of il to m] , one posterolingually directed ridge on p4. and presence of pigment on the posterior portion of the ventromedial edge of il . 21(19). No dentary characters discerned, but small median tine present on lis (Fig. 18) Sorex hairdii, Baird's shrew (Fig. 18) Ridge present at posteromedial edge of Us (Fig. 18) Sorex pacificus, Pacific shrew (Fig. 18) 22(7). Length of dentary usually <6.5 mm; length of cl-m3 usually < 4. 1 mm; height of coronoid process < 3.2 mm 23 Length of dentary usually > 6.5 mm; length of cl-ni3 > 4.1 mm; most with height of coronoid process > 3.2 mm 25 23(22). Pigment on // in one or two sections (Fig. 19), rarely three as a result of wear; dentary below ml shallower than height of ml ... Sorex nanus. Dwarf shrew (Fig. 19) Pigment on // usually in three sections (Fig. 19); dentary below ml as deep as or deeper than height of ml 24 24(23). First lower incisor set at an angle < 8° from the horizontal ramus of the dentary (Fig. 19); width of ml < 0.8 mm; width of w2 < 0.8 mm Sorex prehlei, Preble's shrew (Fig. 19) First lower incisor set at an angle > 8° from the horizontal ramus of the dentary (Fig. 19); width of ml > 1.1 mm; width of /?/2 >0.9 mm Sorex longirostris. Southeastern shrew (Fig. 19) A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 19 occlusal Sorex bairdii labial occlusal Sorex pacificus Fig. 18. Dentaries and anterior views of ils (at lower right) of 5o/-ev bairdii (OSUFW 61 1 1) and 5. pacificus (OSUFW 8306). In the dentaries of both taxa note the shallow interdenticular space, the relationship of the posterior edge of // to ml , one posterolingually directed ridge on/74, and presence of pigment on the posterior portion of the ventromedial edge of //. Each // of 5. bairdii bears a small median tine, whereas each il of 5. pacificus bears a ridge along the posteromedial edge. 20 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 occlusal Sorex nanus Sorex preblei occlusal labial occlusal Sorex longirostris Fig. 19. Dentaries of Sorex nanus (MSB 59339), S. prehlei (OSUFW 4435), and S. longirostris (MSUMZ 24168). Note the number of pigment sections on il, the relationship of the posterior edge of // to ml , and one posterolingually directed ridge on p4, and presence of pigment on the posterior portion of the ventromedial edge of il in all taxa. In S. prehlei and S. longirostris note the angle of // relative to the horizontal ramus of the dentary. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 21 25(22). Height of coronoid process < 3.2 mm; cl with two cusps Sorex iigyunak, Barren ground shrew (Fig. 20) Height of coronoid process usually > 3.2 mm; cl usually with one cusp, but if with two cusps, height of coronoid process > 3.4 mm 26 26(25). Length of coronoid-condyloid processes usually < 3. 1 mm; length of/;// + length of /?/2 usually < 2.4 mm 27 Length of coronoid-condyloid processes usually > 3.1 mm; length of m7 + length of m2 usually > 2.4 mm 32 27(26). Pigment on il distributed in one section (Fig. 20) Sorex tenellus, Inyo shrew (Fig. 20) Pigment on // distributed in two or three sections (Figs. 20, 21, 22) 28 28(27). Present geographic range limited to Sierra Nevada mountains, Mono Co., California (Williams, 1984); pelage bicolored Sorex lyelli, Mt. Lyell shrew (Fig. 20) Present geographic range not including California; pelage bicol- ored or tricolored 29 29(28). Present geographic range limited to Pribliof Islands, Alaska; pel- age tricolored 30 *to^ Present geographic range other than Pribliof Islands, Alaska; pel- age bicolored 31 30(29). Pigment on il distributed in two sections (Fig. 21); il set at an angle > 8° from the horizontal ramus of the dentary (Fig. 21) .... Sorex hydrodromiis , Pribilof Island shrew (Fig. 21) Pigment on // distributed in three sections (Fig. 21); il set at an angle < 8° from the horizontal ramus of the dentary (Fig. 21) .... Sorex jacksoni, St. Lawrence Island shrew (Fig. 21) 3 1 (29). No dentary characters discerned, but maxillary breadth < 4. 1 5 mm (Jackson, 1928; zygomatic processes commonly broken in speci- mens of old adults) Sorex cinereus. Masked shrew (Fig. 21) Maxillary breadth > 4.15 mm (Jackson, 1928; zygomatic pro- cesses commonly broken in specimens of old adults) Sorex haydeni. Hayden's shrew (Fig. 22) 97 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 labial Sorex ugyunak occlusal occlusal Sorex tenellus labial \ ^^*- 13° from the horizontal ramus of the dentary; interdenticular space very shallow (Fig. 13).... 33 First lower incisor set at an angle < 13° from the horizontal ramus of the dentary; interdenticular space very deep (Fig. 13), may be obscured in old individuals 34 33(32). Height of coronoid process < 3.8 mm; ratio of height of coronoid processilength of dentary < 0.54; ratio of distance between il and p4 : length of dentary > 0.04 Sorex arizonae, Arizona shrew (Fig. 22) Height of coronoid process usually > 3.9 mm; ratio of height of coronoid processilength of dentary > 0.55; ratio of distance be- tween II and p4 : length of dentary < 0.04 Sorex merriami , Merriam's shrew (Fig. 22) 34(32). No dentary characters discerned, but median tines on lis set at or above the pigment on the body of the Us (Fig. 23) 35 Median tines on lis set within the pigment on the body of the lis (Fig. 23) 36 S. trowbridgii S. vagrans S. monticolus Fig. 23. Anterior views of ils of Sore.x trowbridgii (PSM 5892), S. vagrans (SDMNH 16971), and S. monticolus (KU 146134). Notice that in S. trowbridgii and S. vagrans the tines are above the level of the pigment; however, in S. trowbridgii the medial edges of the ils are curved and flaring, whereas they are straight and minimally divergent in S. vagrans. In S. monticolus the median tines are set within the pigment. 26 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 labial occlusal Sorex vagrans Fig. 24. Dentary of Sore.x vagrans (OSUFW 6821) showing the very deep interdenticular space, the angle of /7 relative to the horizontal ramus of the dentary, the relationship of the posterior edge of // to ml , one posterolingually directed ridge on p4, and presence of pigment on the posterior portion of the ventromedial edge of //. 35(34). No dentary characters discerned, but medial edge of Us curved and flaring (Fig. 23); least interorbital breadth > 3.5 mm (Carraway, 1987) Sorex trowhridgii, Trowbridge's shrew (Fig. 12) Medial edge of IJs straight and minimally divergent (Fig. 23); least interorbital breadth < 3.5 mm (Carraway, 1987) Sorex vagrans. Vagrant shrew (Fig. 24) 36(34). No dentary characters discerned, but paracrista not pigmented on P4 (Fig. 25) Sorex ornatus. Ornate shrew (Fig. 26) Paracrista partially to completely pigmented on P4 (Fig. 25) Sorex monticolus, Dusky or Montane shrew (Fig. 12) S. ornatus S. monticolus S. monticolus S. monticolus Fig. 25. Occlusal views (lingual to the left) of upper P4 of Sorex ornatus (KU 502 1 8) and three specimens of S. monticolus ( left to right: OSUFW 4798, 4799, and 4815) to show the absence of pigment on the paracrista in S. ornatus and the variable amounts of pigment on the paracrista in S. nionlicolus. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 27 labial So rex 0 mat us occlusal Fig. 26. Dentary of Sorex ornatus (UCD 4688) showing the very deep interdenticular space, the angle of il relative to the horizontal ramus of the dentary. the relationship of the posterior edge of // to ml . one posterolingually directed ridge on p4. and pcesence of pigment on the posterior portion of the ventromedial edge of il . COMPARISONS OF SOME SPECIES OF SORICIDAE Selected species not in direct line in the key are compared in the following annotations of aspects of their morphology. Notiosore.x crawfordi (Desert shrew) Notisorex crawfordi can be distinguished from all other soricids in the western United States and Canada by possession of the following coinbina- tion of characters: alveolus of il extending posteriorly beneath at least part of ml (Figs. 3, 4); no pigment on ml and m3 (Fig. 4); and deeply emargin- ate interarticular area (Fig. 5). However, these three characters may not be sufficient to identify to species old individuals with excessively worn teeth or a damaged dentary. The desert shrew can be separated from Sorex hoyi, S. merriami, S. ornatus, and S. preblei by height of the unpigmented portion oiml > 0.9 mm; from S. hoyi, S. nanus, and S. preblei by height of the coronoid process > 3.5 mm and length of cl-m3 > 4.5 mm: and from S. hoyi and S. ornatus by // set at an angle > 15° from the horizontal ramus of the dentary. Additionally, A^. crawfordi differs from S. hoyi by the width of ml > 0.8 mm, width of m2 > 0.8 mm, and length of the dentary > 6. 1 mm, and from S. preblei by length of the coronoid-condyloid processes > 3.0 mm. Sorex alaskanus (Glacier Bay water shrew) The Glacier Bay water shrew differs from Sorex arcticus, S. monticolus, and S. ugyunak by length of cl-m3 > 5.4 mm. Also, it differs from S. arcticus by presence of a strip of pigment on the inside ventroinedial edge 28 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 of the Us (Figs. 10, 15) and by the length of the coronoid-condyloid processes < 3.7 mm, and from S. ugyunak > 3.5 mm. Sorex alaskaims is distinguished from S. monticolus and S. ugyunak by length of the dentary > 8.6 mm. Additionally, it can be distinguished from S. ugyunak by height of the coronoid process > 4.2 mm, depth of the mandible at //// > 1 .3 mm, by pigment on il in one section, and cl with one cusp. Sorex arcticus (Arctic shrew) Sorex arcticus differs from S. alaskanus, S. cinereus, S. jacksoni, S. hydrodromus, S. monticolus. and S. ugyunak by length of the coronoid- condyloid processes > 3.8 mm and lack of a strip of pigment on the inside ventromedial edge of the Us (Figs. 10, 1 1 ). Additionally, it can be distin- guished from S. alaskanus by length of cl-m3 < 5.3 mm and from S. jacksoni. S. hydrodromus. and S. ugyunak > 4.8 mm; from S. cinereus, S. jacksoni. S. hydrodromus, S. monticolus. and S. ugyunak by height of the coronoid process > 4.4 mm; from S. cinereus, S. jacksoni. S. hydrodromus, and S. ugyunak by length of the dentary > 7.9 mm; from S. jacksoni, S. hydrodromus, and S. ugyunak by width of p4 > 0.7 mm; from S. cinereus and S. ugyunak by width of cl > 0.6 mm; and from S. jacksoni and S. ugyunak by depth of the dentary below mJ > 1.1 mm. Also, from S. jacksoni by width of m2 > 0.7 mm and from S. ugyunak by cl with one cusp. Sorex arizonae (Arizona shrew) This species can be distinguished from Sorex hoyi and S. nanus by height of the coronoid process > 3.5 mm, and from S. merriami < 3.8 mm, and from S. hoyi and S. nanus by length of cJ-m3 > 4.7 mm and length of the dentary > 7.1 mm. Additionally, S. arizonae differs from S. hoyi by length of m3 > 0.8 mm and the alveolus of // not extending posteriorly beneath part of ml (Figs. 3, 22), and from S. nanus by depth of the dentary below ml > 1 .0 mm, length of the coronoid-condyloid processes > 3. 1 mm, length of r7 > 0.9 mm, length of p4 > 1.0 mm, length of ni2 > 1.2 mm, and width of p4 > 0.7 mm. Sorex bairdii (Baird's shrew) Sorex bairdii can be distinguished from S. monticolus by height of the coronoid process usually > 4.3 mm and length of cl-m3 usually > 5.2 mm. Sorex bendirii (Pacific water shrew) The condition of two posterolingually directed ridges (Fig. 8) on the occlusal surface of p4 of the Pacific water shrew is unique among shrews in the western United States and Canada. Also, Sorex bendirii can be distin- A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 29 guished from all other Sorex by length of cl greater than length of p4. Differences in pattern of pigmentation on the iJs of specimens of S. h. bendirii and S. b. palmeri in Figure 9 are the result of individual variation and are not of taxonomic value. Sorex cinereiis (Masked shrew) Sorex cinereus differs from S. arcticus, S. monticolus, S. m. neomexicamis, S. palustris, S. tundrensis, and S. vagrans by length of the coronoid-condyloid processes usually < 3.2 mm; from S. hydrodromus and S. palustris by il set at an angle < 8° from the horizontal ramus of the dentary; and from S. arcticus and S. tundrensis by presence of a strip of pigment on the inside ventromedial edge of the ils (Figs. 10, 21). No single qualitative or quantitative character, or combination of characters recorded for this study, distinguish S. cinereus from S. jacksoni. However, van Zyll de Jong (1982; irt litt., 30 January 1995) could distinguish cinereus and jacksoni by the following discriminant-function equation: discriminant score = 2.75 1 (length of mandible) + 3.633 (height of the coronoid process) + 6.189 (length of coronoid-condyloid processes) -i- 1 1.68 (greatest condy- lar depth) -I- 14.53 (width of lower condylar facet) - 5.923 (width of upper condylar facet) - 3.874 (length of cl-m3) - 17.54496. Those specimens with a score < - 0.293 are S. cinereus. Additionally, the masked shrew can be separated from S. arcticus and S. palustris by length of the dentary < 7.6 mm and height of the coronoid process < 3.9 mm and from S. ugyunak usually > 3.2 mm; from S. arcticus by width of cl < 0.5 mm; from S. palustris by length of c I -mS < 4.9 mm; from S. monticolus by length of ml < 1.4 mm; from S. m. neomexicamis by length of ml < 1.4 mm, length of m2 < 1.2 mm, and height of the unpigmented portion of ml < 0.6 mm; from S. palustris by length of m3 < 0.9 mm; and from S. ugyunak by cl usually with one cusp. Sorex haydeni (Hayden's shrew) Hay den's shrew differs from Sorex hoyi by the alveolus of il not extending posteriorly beneath at least part of ml (Figs. 3, 22) and length of the dentary > 6.2 mm; from S. monticolus by length of ml usually < 1.3 mm and length of cl-m3 < 4.4 mm; and from S. preblei by height of the coronoid process > 3.2 mm and length of cl-m3 usually > 4.1 mm. Sorex hoyi (Pygmy shrew) This species is unique within the genus Sorex because the alveolus of il extends posteriorly beneath at least part of ml (Figs. 3, 4). Sorex hoyi can be distinguished from Notiosorex crawfordi, S. arizonae, and S. merriami by height of the coronoid process < 3.4 mm and from S. nanus > 3.1 mm; 30 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 from N. crawfordi and S. preblei by il set at an angle < 13° and > 10° from the horizontal ramus of the dentary, respectively; from N. crawfordi and S. arizonae by width of ml < 0.8 mm and width of m2 < 0.8 mm; and from S. haydeni and S. merriami by length of the dentary < 6.1 mm. Additionally, it differs from A^. crawfordi by height of the unpigmented portion of ml < 0.7 mm; from A^. crawfordi, S. arizonae, and 5. merriami by length of cl- mS < 4.2 mm; from S. arizonae and S. merriami by length of m3 > 0.9 mm and < 0.8 mm, respectively; and from S. merriami by length of the coronoid- condyloid processes < 3.1 mm. Sore.x hydrodromus (Pribilof Island shrew) The Pribilof Island shrew differs from Sorex arcticus, S. monticolus, and S. tundrensis by length of the coronoid-condyloid processes usually < 3.1 mm and length of c I -m3 < 4.5 mm, and from S. cinereus, S.jacksoni, and S. ugyunak by il set at an angle > 8° from the horizontal ramus of the dentary. Additionally, S. hydrodromus can be distinguished from S. arcticus and S. tundrensis by presence of a strip of pigment on the inside ventrome- dial edge of the ils (Figs. 10, 21) and height of the coronoid process < 3.6 mm and from S. ugyunak > 3.3 mm; from S. arcticus by width of p4 < 0.6 mm and length of the dentary < 7.7 mm; from S. monticolus by length of ml < 1.3 mm; and from S. ugyunak by presence of pigment on // in two sections and cl usually with one cusp. Sorex jacksoni (St. Lawrence Island shrew) Sorex jacksoni can be distinguished from S. arcticus, S. monticolus, and S. tundrensis by length of cl-m3 < 4.4 mm; from S. arcticus, S. monticolus, and S. tundrensis by length of the coronoid-condyloid processes < 3.0 mm; from S. hydrodromus and S. monticolus by il set at an angle < 7° from the horizontal ramus of the dentary; and from S. ugyunak by cl usually with one cusp. No single qualitative or quantitative character, or combination of characters recorded for this study, separates S. jacksoni from S. cinereus. However, van Zyll de Jong (1982; in litt., 30 January 1995) could distin- guish yacAr^o/z/ and cinereus by the following discriminant-function equa- tion: discriminant score = 2.751 (length of mandible) + 3.633 (height of the coronoid process) -i- 6.189 (length of coronoid-condyloid processes) -i- 1 1.68 (greatest condylar depth) -i- 14.53 (width of lower condylar facet) - 5.923 (width of upper condylar facet) - 3.874 (length of r/-mi) - 1 7.54496. Those specimens with a score > 0.460 are S. jacksoni. Further, it differs from S. arcticus and S. tundrensis by height of the coronoid process < 3.5 mm and from S. ugyunak > 3.2 mm; from S. arcticus and S. monticolus by width of p4 < 0.5 mm and width of m2 < 0.6 mm; from S. arcticus by depth of the dentary below ml <1 .0 mm and length of the dentary < 7.0 mm; and from S. arcticus and S. tundrensis by presence of a strip of pigment on the inside of the ventromedial edge of the ils (Figs. 10, 21). A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 31 Sorex longiwsths (Southeastern shrew) This species can be distinguished from Sorex hoyi, S. merriami. and S. nanus by width of ml > 1 . 1 mm and from S. hoyi and S. nanus by width of m2 > 0.9 mm. Also, it differs from S. hoyi by the alveolus of iJ not extending posteriorly beneath part of ml (Figs. 3, 19), and from S. merriami by length ofcl-mS < 4.2 mm, height of the coronoid process < 3.3 mm, and length of the coronoid-condyloid processes < 2.9 mm. Sorex lyelli (Mt. Lyell shrew) The Mt. Lyell shrew differs from Sorex ornatus and S. vagrans by length of the coronoid-condyloid processes < 3.0 mm; from S. nanus and S. ornatus by height of the coronoid process > 3.1 mm and < 3.4 mm, respectively; and from S. tenellus and S. vagrans by il set at an angle < 6° from the horizontal ramus of the dentary. Additionally, S. lyelli can be distinguished from S. nanus by length of ml > 1.3 mm, length of cl-m3 > 4.2 mm, and length of dentary > 6.5 mm. Sorex merriami (Merriam's shrew) Sorex merriami can be distinguished from S. arizonae, S. hoyi, S. longirostris, S. nanus, and S. preblei by height of the coronoid process usually > 3.9 mm; from Notiosorex crawfordi by height of the unpigmented portion of ml < 0.7 mm; and from S. vagrans by the interdenticular space very shallow (Figs. 13, 22) and il set at an angle > 13^ from the horizontal ramus of the dentary (Fig. 22). It differs from S. hoyi, S. longirostris, S. nanus, and S. preblei by length of cl-m3 > 4.3 mm and length of the coronoid-condyloid processes > 3.2 mm, and from S. nanus and S. preblei by depth of the dentary below ml > 0.95 mm. Additionally, Merriam's shrew can be distinguished from 5". hoyi by length of m3 > 0.9 mm, length of the dentary > 6.4 mm, and the alveolus of il not extending posteriorly beneath part of ml (Figs. 3, 22); from S. longirostris by width of /?;/ < 1.0 mm; and from S. preblei by length of ml > 1.3 mm and length of the dentary usually > 6.6 mm. Sorex monticolus (Dusky or Montane shrew) Sorex monticolus can be distinguished from S. arcticus and S. m. neomexicanus by length of the coronoid-condyloid processes < 3.6 mm and from 5'. cinereus, S. jacksoni, S. hydrodromus, and S. ugyunak > 3.2 mm; from S. alaskanus, S. hairdii, and S. m. neomexicanus by length of cl-m3 < 5.2 mm and from S. jacksoni, S. hydrodromus, and S. monticolus > 4.5 mm; and from S. arcticus and S. tundrensis by presence of a strip of pigment on the inside of the ventromedial edge of the ils (Figs. 10, 12). 32 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 Additionally, it differs from S. alaskamis by length of the dentary < 8.1 mm; from S. arcticiis, S. bairdii, and S. m. neomexicanus by height of the coronoid process < 4.2 mm and from S. ugyunak > 3.5 mm; from S. cinereus, S. haydeni, S. hydwdromus , S.jacksoni, and S. ugyunak by length of mi > 1.4 mm; from S.jacksoni by il set at an angle usually > 8° from the horizontal ramus of the dentary, width of p4 > 0.6 mm, and width of m2 > 0.7 mm; and from S. ugyunak by iJ set at an angle usually > 8° from the horizontal ramus of the dentary, pigment on il usually in one section, and cl usually with one cusp. Sorex monticolus neomexicanus (Dusky or Montane shrew) This taxon can be distinguished from Sorex cinereus, other 5. monticolus, and S. palustris by height of the coronoid process usually > 4.5 mm. Additionally, this taxon differs from S. cinereus and other S. monticolus by length of cl-m3 > 5.0 mm and usually > 5.3 mm, respectively, and length of the coronoid-condyloid processes > 3.7 mm; from S. cinereus and S. palustris by height of the unpigmented portion of ml > 0.65 mm; from S. cinereus by length of ml > 1.6 mm, length of m2 > 1.3 mm, and length of the dentary > 7.6 mm; and from S. palustris by widths of cl + p4 usually > 1.4 mm. Sorex nanus (Dwarf shrew) With exception of Sorex minutissimus, the dwarf shrew is the smallest shrew in North America (Tables 1,2). Sorex nanus can be distinguished from Notiosorex crawfordi, S. hoyi, S. lyelli. S. merriami, and S. arizonae by height of the coronoid process usually < 3.1 mm. Also, it differs from S. hoyi by the alveolus of // not extending posteriorly beneath part of ml (Figs. 3, 19); from S. prehlei by il set at an angle usually > 8° from the horizontal ramus of the dentary; from S. longirostris by width of ml < 0.8 mm and width of m2 < 0.8 mm; and from 5. tenellus by length of the dentary < 6.5 mm and length of cl-m3 usually < 4. 1 mm. Additionally, it can be distinguished from N. crawfordi by length of cl-m3 < 4.2 mm and pigment present on m2 and mS; from S. merriami and S. arizonae by length of the coronoid-condyloid processes < 3.0 mm, depth of the dentary below ml < 0.95 mm. and length of cl-m3 < 4.1 mm; from S. arizonae by length of r7 < 0.8 mm, length of p4 < 0.9 mm, length of m2 < 1.1 mm, width of p4 < 0.6 mm, and length of the dentary < 6.7 mm; and from S. lyelli by length of ml < 1.3 mm, length of cl-m3 < 4.1 mm, and length of the dentary usually < 6.5 mm. Sorex ornatus (Ornate shrew) Sorex ornatus can be distinguished from Notiosorex crawfordi by il set at an angle usually < 13° from the horizontal ramus of the dentary, height A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 33 of the unpigmented portion of ml < 0.7 mm, and pigment present on m2 and mS; from S. lyelli by height of the coronoid process usually > 3.4 mm and length of the coronoid-condyloid processes > 3.0 mm; and from S. trowbridgii by length of cl-m3 usually < 4.8 mm. No quantitative or qualitative characters of the dentary could be discerned to distinguish S. ornatiis from S. vagrans. Sorex pacificus (Pacific shrew) This species can be distinguished from Sorex bendirii by one posterolingually directed ridge (Figs. 8, 18) on the occlusal surface oi p4 and length of cl < length of p4. Sorex palustris (Water shrew) This species differs from Sorex bendirii by one posterolingually di- rected ridge (Figs. 8, 15) on the occlusal surface of p4 and length of cl < length of p4; from S. cinereus and S. iigyunak by height of the coronoid process > 4.0 mm and from 5. m. neomexicanus usually < 4.5 mm. Addi- tionally, the water shrew differs from S. cinereus and S. ugyunak by length of m3 > 0.95 mm, length of cl-m3 > 5.2 mm, il set at an angle usually > 8° from the horizontal ramus of the dentary, length of the coronoid-condyloid processes > 3.2 mm, and length of the dentary > 8.0 mm; from S. m. neomexicanus by height of the unpigmented portion of ml < 0.65 mm and widths of cl + p4 < 1 .4 mm; and from S. ugyunak by pigment on il in one section, length of p4 > 1 . 1 mm, width of m2 > 0.8 mm, and cl usually with one cusp. Sorex preblei (Preble's shrew) Sorex preblei can be distinguished from Notiosorex crawfordi and S. merriami by height of the coronoid process < 3.3 mm, length of the coronoid-condyloid processes < 2.9 mm, and length of cl-m3 < 4.2 mm, and from S. hoyi and S. nanus by il set at an angle < 8° from the horizontal ramus of the dentary. Additionally, Preble's shrew can be distinguished from N. crawfordi by height of the unpigmented portion of n^l < 0.65 mm and pigment present on m2 and m3; from S. hoyi by the alveolus of il not extending posteriorly beneath part of ml (Figs. 3, 19); from S. merriami by depth of dentary below ml < 0.9 mm, length of ml < 1.3 mm, and length of dentary usually < 6.6 mm; and from S. haydeni by height of the coronoid process usually < 3.2 mm and length of cl-m3 usually < 4.1 mm. Sorex sonomae (Fog shrew) This species can be distinguished from Sorex bendirii by one posterolingually directed ridge (Fig. 8) on the occlusal surface of p4 and length of cl < length of p4. 34 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 Sorex tenelhis (Inyo shrew) The Inyo shrew can be distinguished from Sorex lyelli by iJ set at an angle > 8° from the horizontal ramus of the dentary (Fig. 20), and from S. nanus by length of cl-m3 > 4.2 mm and length of the dentary usually > 6.4 mm. Sorex trowbridgii (Trowbridge's shrew) This species can be distinguished from Sorex ornatus by length of cl- m3 usually > 5.0 mm. Sorex timdrensis (Tundra shrew) Sorex timdrensis differs from S. hydrodromus. S. jacksoni, and S. ugyunak by height of the coronoid process > 3.7 mm, length of the coronoid- condyloid processes > 3.1 mm, and length of cJ-mJ > 4.5 mm; from S. cinereus, S. jacksoni, S. monticolus, and S. ugyunak by lack of a strip of pigment on the inside of the ventromedial edge of the ils (Figs. 10, 1 1). Additionally, it differs from S. cinereus by length of the coronoid-condy- loid processes usually > 3.3 mm and from S. ugyunak by pigment on il in one or two sections, cl with one cusp, and depth of the dentary below ml + height of the coronoid process > 4.7 mm. Sorex ugyunak (Barren ground shrew) The ban^en ground shrew can be distinguished from all other Sorex by a combination of height of the coronoid process < 3.2 mm and cl with two cusps. Additionally, it differs from Sorex alaskanus, S. arcticus, and S. palustris by length of cl-m3 < 4.5 mm, length of the coronoid-condyloid processes < 2.9 mm, and length of the dentary < 6.9 mm; from S. alaskanus and S. arcticus by depth of the dentary below nil < 1.0 mm; from S. alaskanus. S. hydrodromus, S. palustris, and S. tundrensis by pigment on il in three sections; and from S. hydrodromus, S. monticolus, and S. palustris by il set at an angle < 5° from the horizontal ramus of the dentary. Additionally, S. ugyunak can be distinguished from S. arcticus and S. tundrensis by presence of a strip of pigment on the inside ventromedial edge of the ils (Figs. 10, 20); from S. arcticus and S. palustris by length of p4 < 0.9 mm; from S. monticolus and S. tundrensis by length of the coronoid-condyloid processes < 2.9 mm; and from S. tundrensis by length of cl-m3 < 4.5 mm. Sorex vagrans (Vagrant shrew) Sorex vagrans can be distinguished from S. cinereus and S. lyelli by length of the coronoid-condyloid processes > 3.1 mm; from S. lyelli by /'/ set at an angle > 8° from the horizontal ramus of the dentary (Fig. 24); and A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 35 from S. meiriami by the interdenticular space very deep (Figs. 13, 24) and il set at an angle < 13° from the horizontal ramus of the dentary (Fig. 24). No quantitative or qualitative dentary characters could be discerned to distinguish S. vagrans from S. ornatus. Acknowledgments: For loan of or access to specimens in their care, I thank M. A. Bogan (U.S. Fish and Wildlife Service, Albuquerque, New Mexico), M. S. Boyce (Univ. Wyoming), J. R. Choate (Fort Hays State Univ., FHSU). J. A. Cook (Univ. Alaska Museum, UAM), L. S. Ellis (Northeast Missouri State Univ.), M. D. Engstrom (Royal Ontario Mu- seum, ROM), R. D. Fisher (National Museum of Natural History, USNM), K. R. Foresman (Univ. Montana). E. K. Fritzell (formerly at Univ. Mis- souri-Columbia), J. E. Heyning (Los Angeles County Museum of Natural History, LACM), A. Gluesenkamp (Univ. California-Davis, UCD), A. H. Harris (Univ. Tex.as-El Paso), E. J. Heske (Univ. Illinois at Urbana- Champaign, UIMNH), M. L. Kennedy (Univ. Memphis, MSUMZ), G. L. Kirkland, Jr. (Shippensburg State Univ.), H. G. McCartney (Univ. Arkan- sas-Fayetteville), V. R. McDaniel (Arkansas State Univ.), S. B. McLaren (Carnegie Museum of Natural History), D. Paulson and G. Shugart (Univ. Puget Sound, PSM), C. L. Pritchett (Brigham Young Univ.), B. Stein (Museum of Vertebrate Zoology, Univ. California-Berkeley), R. M. Timm (Univ. Kansas, KU), P. Unitt (San Diego Museum of Natural History, SDMNH), C. G. van Zyll de Jong (formerly at Canadian Museum of Nature), B. J. Verts (Oregon State Univ., OSUFW), M. R. Voorhies (Univ. Nebraska State Museum), and T. L. Yates (Univ. New Mexico, MSB). R. S. Hoffmann, C. G. van Zyll de Jong, and B. J. Verts commented on an earlier draft of this manuscript. This publication received partial support from the Thomas G. Scott Achievement Fund for publications. Department of Fish- eries and Wildlife, Oregon State University. This is Technical paper No. 10,668, Oregon Agricultural Experiment Station. LITERATURE CITED Butler. P. M., R. S. Thorpe, and M. Greenwood. 1989. Interspecific relations of African crocidurine shrews (Mammalia: Soricidae) based on multivariate analy- sis of mandibular data. Zool. J. Linnean Soc. 96:373-412. Carraway, L. N. 1987. Analysis of characters for distinguishing Sorex trowbridgii from sympatric S. vagrans. Murrelet 68:29-30. Carraway. L. N. 1990. A moiphologic and morphometric analysis of the "Sorex vagrans species complex" in the Pacific Coast region. Spec. Publ., The Museum, Texas Tech Univ. Press, 32:1-76. Carraway. L. N., and B. J. Verts. 1994. Relationship of mandibular morphology to relative bite force in some Sorex from western North America. Pp. 201-210, in Merritt, J. F., G. L. Kirkland, Jr., and R. K. Rose (eds.). Advances in rhe Biology of Shrews. Carnegie Mus. Nat. Hist. Spec. Publ. 18:1^58. 36 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 Cho.ate, J. R. 1970. Systematics and zoogeography of Middle American shrews of the genus Cryptotis. Univ. Kansas Publ. Mus. Nat. Hist. 19:195-317. Dannelid, E. 1989. Medial tines on the upper incisors and other dental features used as identification characters in European shrews of the genus Sorex (Mammalia, Soricidae). Z. Saugetier. 54:205-214. DiERSiNG, V. E. 1980. Systematics and evolution of the pygmy shrews (subgenus Micwsorex). J. Mammal. 61:76-101. FoRESMAN, K. R., AND R. E. Jensen, Jr. 1992. A morphometric comparison of the masked shrew {Sorex cinereus) and Hayden's shrew (Soir.x haydeni) in Montana and bordering states. Northwest. Nat. 73:15-21. George, S. B., H. H. Genoways, J. R. Choate, and R. J. Baker 1982. Karyotypic relationships within the short-tailed shrews, genus Blariiui. J. Mammal. 63:639- 645. GuREEV, A. A. 1971. [Shrew (Soricidae) Fauna of the Worhl]. Akad. nauk. SSSR, Lenningrad. [In Russian] Hall, E. R. 1 98 1 . The Mammals of North America. 2nd Ed. Vol. 1 . New York: John Wiley & Sons. Handwerk, J. 1987. Neue Daten zur Morphologic, Verbreitung und Okologie der Spitzmiiuse Sorex araneus und S. coronatus im Rheinland. Bonn. Zool. Beit. 38:273-297. Harris, A. H., and L. N. Carraway. 1993. Sorex prehlei from the Late Pleistocene of New Mexico. Southwest. Nat. 38:56-58. Hausser, J. 1984. Genetic drift and selection: their respective weights in the morphological and genetic differentiation of four species of shrews in southern Europe (Insectivora, Soricidae). Z. Zool. Syst. Evol.-forsch. 22:302-320. Hausser, J., and D. Jammot. 1974. Etude biometrique des machoires chez les Sorex du groupe araneus en Europe continentale (Mammalia, Insectivora). Mammalia 38:324-343. Hennings, D.. and R. S. Hoffmann. 1977. A review of the taxonomy of the Sorex vagrans species complex from western North America. Univ. Kansas Mus. Nat. Hist. Occas. Papers 68:1-35. Hershkovitz, p. 1971. Basic crown patterns and cusp homologies of mammalian teeth. Pp. 95-150 in Dahlberg, A. A. (ed.). Dental Morphology and Evolution. Chicago: Univ. Chicago Press. Jackson, H. H. T. 1928. A review of the American long-tailed shrews: (genus Sorex and Microsorex). N. Amer. Fauna 51:1-238. Jammot, D. 1972^. Principes et methodes d'une etude moderne des insectivores. Application aux Soricidae fossiles. Mammalia 36:435^48. Jammot, D. 1972/?. Relationships between the new species Sorex scottensis and the fossil shrews Sorex cinereus Ken-. Mammalia 36:449^58. Junge, J. A., AND R. S. Hoffmann. 1981. An annotated key to the long-tailed shrews (genus Sorex) of the United States and Canada, with notes on Middle American Sorex. Univ. Kansas Mus. Nat. Hist. Occas. Papers 94:1^8. Junge, J. A., R. S. Hoffmann, and R. W. Debry. 1983. Relationships within the Holarctic Sorex arcticus — Sorex tundrensis species complex. Acta Theriol. 28:339-350. A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 37 MuLLiCAN, T. R., AND L. N. Carraway. 1990. Shrew remains from Moonshiner and Middle Butte caves, Idaho. J. Mammal. 71:351-356. Ognev, S. I. 1962. Mammals of Eastern Europe and Northern Asia: Insectivora and Chiroptera. Translated from the 1928 Russian edition for the National Science Foundation by the Israel Program for Scientific Translations, Jerusalem, Vol. 1. Repenning, C. a. 1967. Subfamilies and genera of the Soricidae. Geol. Surv. Prof. Paper 565:1-74. RuPRECHT, A. L. 1971. Taxonomic value of mandible measurements in Soricidae (Insectivora). Acta Theriol. 16:341-357. ScHAEFER, H. 1975. The shrew mice (Soricidae) from the High Tatra Mountains since 30 000 years (mandibular study). Zool. Anz. 195:89-1 1 1. VAN Zyll de Jong, C. G. 1980. Systematic relationships of woodland and prairie forms of the common shrew, Sore.x cinereus cinereus Kerr and S. c. haydeni Baird, in the Canadian prairie provinces. J. Mammal. 61:66-75. van Zyll de Jong, C. G. 1982. Relationships of amphiberingian shrews of the Sorex cinereus group. Canadian J. Zool. 60:1580-1587. Vesmanis, I. E., V. Sans-Coma, R. Pons, A. Vesmanis, and J. A. Alcover. 1980. tjber die Coronar-hohe des Unterkiefers als trennendes Merkmal (?) verschiedener Wimperspitzmaus-taxa im Mittelmeerraum (Mammalia, Insectivora). Misc. Zool. 6:135-139. Williams, D. F. 1984. Habitat associations of some rare shrews (Sorex) from California. J. Mammal. 65:325-328. Wilson, R. W. 1985. The dentition of the Paleocene "insectivore" genus Acmeodon Matthew and Granger (?Palaeoryctidae, Mammalia). J. Paleontol. 59:713-720. Woodman, N., and R. M. Timm. 1992. A new species of small-eared shrew, genus Cryptotis (Insectivora: Soricidae), from Honduras. Proc. Biol. Soc. Washington 105:1-12. Zar, J. H. 1984. Biostatistical Analysis. Englewood Cliffs, New Jersey: Prentice- Hall, Inc. 38 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 "^ s^ ITi r<-i rr, ri f ^ CJ ~^ 0 p 0 p c ^ s d p 0 Tt; d r~; 0 '-r r" r«^ (N nC ri oc ^^ rj —^ — ^ 'b 2 ■5 5 +1 1 ^- +1 1 f*^ +1 1 ^-' +1 1 — li rt H. c 0 r^ ^^ — r-- ,.—.* ^M y-i^ ON d C -C OJ J3 r~~ ^—^ ir, ^"^ -^ ^^ -* _aj U Q E 13 X3 ~™ — ' "~ rj ~ 0 O (N cs ^— ^-H ' o ^ ^ "2 13 d rn (in de; 4°W(i a: E op 'S. _o "i +1 ? ^ '^ +1 ^ 0 +1 ? CnI +1 7 ON 0 O a. 0 -* d d d ON 10 d d d d -* ON d d a On 3 d d d — ' ^ C Oil ^ r- y — V ^ 'd- C-l r-i n of a States 0 0 0 p d +1 ON T m rj d +1 T ^ d +1 T r<-, +1 7 ON. 0 alio ted '53 X 0 0 00 d d p d p d d ON ON d d > C ,— < ^ I— 1 »— 1 ^ 15 VO ^ in C«^ ca > u-^ 0 0 r<-, 0 0 ar mean 5sippi Ri aj O rv-) d r<", d 00 d ON d — ; X) .5 oi) S +1 1 0 00 +1 1 rrj 0 +1 t' NO 0 +1 ml 1 s ■~. 00 d iri 0 0 NO d r- ^ d 00 d 3 '3; •o \6 ^ ^ ^ 1> a S c :3 00 \o ^ ^ ■o (u C '..^ 1 "O C/5 0 0 NO 0 0 r3 "Z: r3 o -a '0 t/5 d ON d NO d in d On .^ ^^— 1 ?J ^ "o c« nD r^ r<". in NO r<~l NO f<~i NO P O c c '>^ aj +1 1 0 +1 1 0 +1 1 0 +1 1 ^.^ a ■a c 0 0 0 q d '^ 00 0 NO d "* r~; d r^ p d X t2 o o 0 0 in ON ^' rj r^i r^i 'J: c >i C P OJ S -i: ^ ON VO ^ r~ 'o ."2 0 0 d ON 0 d NO in 0 d ^ 0 d r~~ 1- -^ C/5 ^>- ^c .2 0 c s 0 0 +1 1 00 0 p +1 ^\ m 0 +1 T >n 0 +1 ■nI- ON 0 (U CQ X 03 s d uS 0 On 00 in d r) (^j d 0 p d e! E On 00 in vo u g. O U-i 0 r^i 0 rj 0 0 -a '- ^ 0 >^ d d d d — ■_ d r-) C (11 Ch ^ cd ^" >r-, 00 -^ r~^ '^ r~-' in ges, ai d thre( Q. -J 4— • C aj 13 +1 0 T 00 06 0 d +1 in T 00 00 0 d +1 NO in NO 0 d +1 NO 00 1 NO 0 d c c ON On NO NO 2 « o w 5 c/5 ^ C 0 -- (^1 0 +1 s 3 "■ — ^ -i^ X3 UO *- = -5 I— > ^ c o o ^ s ^ *— o 2 :S ■2 Q F^ ^ • ^ T3 53 2 "p •2^ a >• 5 s g ^ c 0 e2 5 "^ 0 53 NJ t2 'Z (U 5 U :^ A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 39 J ° (L) ^ •5 S ■t: ^ 13 O o c < u o c "^^ rt '"^ •t— t ^+- y: O 5 T3 U a c o U e2 c o X +1 7 o VO (N d ^_ ^™ — (^ o d \q ,__, •* +1 1 — 00 ON d d o d >— ; ^^ ^ +1 1 o rj oc d ON d o d ^ C^i r<-, +1 1 d ON r^ 00 +1 I On — o CO s 2 E t^ — __ f"! Z o 00 d (N d .Co S5 5 §2a +1 I X ^ ^ ^ ON O d +1 ON q fNl O d +1 00 o vO o d f^ +1 I q O d o ' 00 +1 1 o o On 00 d d d CM o d ^ o +1 j_ d vq ON +1 00 00 d r^i >/-i ^^ o +1 1 q d 00 o d +1 ^ ^—■^ CN U-, »— 1 00 ^ d ^^ 1-H o o d "^ d , ', ^ +1 1 o +1 o On d 1/-J n I r<-i O en O d +1 m m ^ o oc O o d +1 in ON o ON d o d +1 ON o §^ d o d +1 00 ON ON o GO d XI O d +1 in _ ^ "*. d o o d +1 o '^- ° d o I in ON o o d +1 rn +1 ON ON d I o ON 00 ri I ^ 4 U 00 00 ON C3 N c X) 03 3 O L. CJ « c s: o ~ c "O r^ cd o 40 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 t4_ OJ o ^ yj ■a 4J o rt ^ , — V « tA D. c r" rt ." u _2 ■ — ' 2 o c ^ p3 a\ r- (U r" 3 ^ oi v; T3 OJ OJ c E D (U .^ ■ 3 r3 > U r- s C .— U— D, o .s- y. 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ON 3; +1 I p ON o 00 o Co o d -^i — ON +1 I P O o o o ^ d S +1 I P in ■— ' d o d .c ^ ri ^- Ci 00 1 o +1 1 ir, — ' +1 d r^ d (^1 ,__ d vO o d ON ori 00 o d o d +1 o 00 -^ On T o o d +1 I O d r- vo o d +1 ri — ■ o o d °o d o +1 I — r^ ^ o O d ^. +1 I -: ON OO d o m o d t--: +1 I P in in o d +1 o 00 o oo o orj o d +1 ON ri ri o d ^ d f^J +1 I — . t^ "^i o d o d +1 On 00 d d CO o [^ r-4 +1 I NO ON . <^ ^ ^ "^ 53 rv -c On o d in o d '^' in ''^ +1 I P P in in VO n ri O d +1 ON in m o O d ^ +1 I q r-i O d ^ rr, 3- +1 I q ^ ^ O rr-, cn o d "^ +1 I q CNl o d +1 ::; r. ri I ON O Si in o r«-, r<"i +1 7 q o 00 o '^ 00' ON o d ^ r^ in +1 I q On _; 1^ nO vd o d q +1 I q ^ ^i ^ ^ 0 d — ' vd 3; +1 1 0 rr, rn d NO in o d r^ (^ rq +1 I q _ ^ o C '^ o d q I^ rq +1 I q ^ <"' o ON ■ ^^ in ^ o d +1 m, nO" I in o __ "^i d sd u 3 c o U f2 On C/3 'o , ts 0 C<1 CO CO o s S o o Co o o Cj CO .§ So s; o Co 42 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 u^ _U ~, oo (N o ri n O X3 2 o O '^f o o ON o ^ -5 c c3 d o d d m d /~j d rj c d d — ' d — ' — ' -o (N 1— ( Q __ _^ <4— o c O d O d t^ d >n o d o d t^ 00 « E **.. +1 ON o +1 d 1 oc c +1 ^ 8 +1 ? o +1 in 'S _W) o o . , ^ d rn in d o mj d o in d i^ X c 3 ir^i d VO o in O r^ d m. d d d d d d (U ^1=! JL "O '^ x: o -— CO ™ O ^ O S t == o .^ r- •- T3 aj 3 '4—' C O U e2 c o X f2 oo o d +1 00 On O O d +1 00 "^i (N in o d +1 +1 TS 00 CN o r<^ ^ I o "~^ o o ~: o OO d "^ +1 I p r^ ^ o ^ o d +1 in 7 ir, d oc m o d — : +1 I p r^ '^' O \0 ^ o oo r- ^'i ^ rn o +1 n o o d ^! +i7 00 q <^ in o d ^ +1 I P ^ ^. o 00 ^ o i>^ cri 00 o d +1 in 00 ON 00 o oo d m d +1 I O r- d +1 +1 oo <~") •n en in 1 <^ 00 <^^ O o d <^ rj O rn o cnI in 2 ^ ^ +1 I P oo "^ in «N 3 1>3 Co in o d +1 in in d ON d ri O o d ^ in "^ +1 I 00 o d in in O in d ^ +i I P O o — mj ^ 00 rn rn O oo d "^ v^ '^ O 3^' o ^ o <^ oo ^ +1 i p r. ?! O o Co s ^ o ON o d o d +1 in O d +1 On ^ in d fN o d ^ +1 I P ,-. ^ o O in q 00 3! +1 I P r^ 00 O On • ^ o d (N +1 7 in r^i ON r«^ t^ o CO <>1 •>* s .Si u 'o ex ■♦*« *-* c/5 s: o s: X) o ^ o 3 <>:) s: O CO A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 43 (N (S m n _H f<^ rri O o o o o o O d •n ON d r<^ d r^ d \D d ON d q d °q (N ^« ON ,__! r^: _^ ^— d 00 ri -— ■ _i rj +1 9 ^-' +1 1 o +1 1 — +1 J^ — ' 4-1 1 o +1 1 ^- +1 1 — ' ^ 1 d ON On d in '-^ d NO d O r~~ d '^ n"; o -* d d t^ d ON d r<-) ^"^ n '"^ 00 o NO ""^ "*. ° d d ^^ — ' d — ' ' — ' , , T-H , 1 ,__ (N »-H o d o o d 1^ o d 00 o d NO d O d q o d ^ 9 CO d I-*", d ON On ^— ^^ ^^ (^ ON +1 1 >n ^- +1 1 — ' +1 1 o +1 9 O +1 O 1 — +1 1 — ' +1 1 o oc d ON r<-; d NO d oo 1 d ON 1 d ^ r~; d •^ ^ d ^ d m d r- d in d -* d 00 d t^ o d d d d d d d (N r<-i c^ r<-i (N ^ m p q O o O ■*. O o o d d (N d ITj d d q d r-^ o ^ '"■' >n , \ ■* ^ r-j ^^ ^ ,_« in ._ in >n +1 1 ■ — 1 +1 1 -^ +1 1 — ' +1 1 in ' — ' +1 1 ' — ' +1 1 — ' +1 7 ' — ' ON d n >n On rn rj >n VO "^ ^ ^ in >n r<-i NO >n -st- m (^ ^ r^ "^ in p O o o o o o d q d NO d r-; d r4 d r-1 d r- d 9 +1 1 00 o +1 1 o +1 ""■ o +1 T mj O +1 rn o +1 ^" o +1 in 1 in o +1 •^ O +1 r<-i 1 O +1 NO 1 o in +1 1 in o 00 vc d i^ q d r<-i d q d ON On d ON '"* d o 2 d 00 (N VO rn »n VO VO r- in '^ ON r- q q o o o o o d r~; d r^ d ri d q d \D d d <^ ^ I/-, r^ in On -xi- ON n-i NO r^ ""^ >n ON •* +1 1 o +1 o +1 1 o +1 1 O +1 1 O +1 1 o +1 1 o (N in d ■"^ n-. d ^ q d o q d NO NO d On 1 00 d d ini ON NO NO 00 in 00 in in 00 r- oo vO <5 00 od NO o< 00 o o o ^^ r<-) o o -o m m m m m ro m 3 C '■t^ c tn o ;2 U l>3 o S 1>5 3 -5 5 1 5 >5 omae lelliod (U c s o ?" s: c J? 2 c « "a o o ^ 1 o Cl, c^ s. ^ ^ CO Co CO ^ CO Co CO 44 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 u- _aj ■-~, -^ <^ r^ , 1 ...H (N o ■3 s o o o o p O ^ ^ o — ; d rn d n-) d rj d p d r^, Q. ^^ 00 ^^' 00 ^ On __' )0 , r- ' o c3 _o +1 1 — +1 1 o +1 1 O +1 1 o +1 1 o +1 1 Q E 1j o r~; d ^ p d ON On d oo p d NO 00 d ON d 00 d ^™ p d p '""' OO d p d d — ' — — ' d — ■ ■o ^^ »-H ri ^^^ , 1 ^4-H CJ p o o o o o o ^ E o ir, d VD d r^ d NO d ^ d NO E '■5 s +1 d 00 o +1 d 1 ON o +1 ^ 00 +1 ? NO o +1 d 1 00 o +1 d On P 3j X (=0 'B. c 3 o o 00 d o d d in d d d On d m o d d d O in d d in d NO in d d in d O -t O ■2 '^^ d ' — 1 d rr-j d r<-) d (N d p d oi s ^^ r~- ^^ r) „^ o ^ -+ ^ ^^ ^ o +1 1 — +1 1 — ' +1 1 ri +1 1 — +1 1 — 1 +1 1 — ' X o ON d d ON o oc d d ITi On d d o d d ON OO d d ON ON d d dJ NJ ^ _i ^ '^ _ •— O c/i -C O — V3 S fc; c o n 00 O d +1 I ON OC o 00 o d <^! +1 I P r^ ^ O in "^ XI vo O d ^ in 2 +1 I P ^ s ° in n in I OO o CO O ° ^ ^ +1 I P r. ^! o r-) o d +1 ri T O Z -- o d '^! +1 I P fN "^ ° o CO :~. O OD c/5 r^i o d P •^ ri +1 I P o ^i o O d +1 NO Sd ^u •2 73 '0 ^ (U i^ -S t. D. ;^ OJ -§ NO I ON ri ri o d +1 NO ON o rn O d ri T o o d ^. +1 I P r. °^ o ^. "^ x> ON d +1 ON rj in T rJ o — : d X) r^^ o d P r^ >n +1 I P r- ri ri XI r-) o o +1 en I o o '^ o d — : +1 I NO o O CO s Co CO o d P +1 I p _ ^: o \0 '^ o d '^' n-; in +1 I P ^ ^ o S: ri CO O n-i T in 0 +1 0 d On r^ d in r<-i ro in o d 1^ +1 I P ^ "^^ o On ,^ ON ^ O CO A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 45 ^^ (N ^« r-i ^^ 0 P 0 0 0 -C <^i d -* m, d rn rr~i d mi in d r-; 1^ d NO ^ M ^ .— 1 0 ^— 'O — ^ 0 ^^ 0 — 0 C '^ +1 1 d +1 ri d +1 1 r| d +1 1 d +1 1 d -J o »n n n ^-^ m ri ^^ ri ^^ r<-i ^^ in '"^ ^ ^^ ^ " --^ '^ ^ ^ M (N m ^^ 0 0 0 0 0 ^ ~~< XI d VD Tt- d ^. •^ d oo_ NO d p ON d On m, W) 5 Cm , 00 0 NO ^ 0 ^^ r^ __ NO ■o p +1 1 0 +1 1 0 +1 1 +1 1 p +1 1 0 ■^ o ^ t^ r^ d r- 00 d m. 00 d m, p ^^ NO p d 00 d 00 d ON d ^^ p '"^ d d d ^^ -^ ^^ CN ^^ CNl ^ 0 0 0 0 0 _c: d 00 d 00 d p d ^ d p ■4— > "3; XI 0 ^ d NO ,^ ON ,__, 00 ,__, t^ ■o Ci. r- +1 1 ^— s +1 1 0 +1 1 0 +1 1 0 +1 1 0 ^ o d t^ t^ d CJ r-; d 0 r^ d r^ On d ON 00 d r- d r- d 00 d p d 00 d d d d -^ d ^^ (N ^H ra ^ p 0 0 0 0 f-; d 00 d ^ d ON d -^ d p *^ ^ d r~~ d ON d ON ^^ ON ,_« r~- "S ^ (^ +1 1 0 +1 1 0 +1 1 0 +1 1 0 +1 1 0 ^ c-t— 1 o d ^ d in in d 1— H NO d 00 00 d ^ r~; d r- d in d r- d ON d 00 d d d d d d ^ , oi , r^ ^^ 0 0 0 0 0 \ d >o r<^ d -* rn d —^ in d ■"""^ ^ c s _« 0 •— 1 0 0 1— < 0 •— 0 1— — 1 0 +1 1 0 d +1 1 d .5 d +1 1 q d +1 1 q d +1 1 ON d 0 00 (N ^o >n VO ri • q ^^ q d q -^ q ^^ q ^^ q 0 ^ ^ ^ ^ ' — ' ' — ' (N n d CN q d d d d d d — 1 ^^ ^^ _ ,_ — q 0 0 0 0 0 '4— t 0 q d ON d ^ d 00 d r- d f^j , 1 ^^ 0 ^^ 0 t^ 0 00 ^^ in ^« r<-) •0 +1 1 -^ +1 1 ' — ' +1 1 0 +1 1 0 +1 1 0 +1 1 0 ■^ 0 ON d 0 d d 00 d q d d 01 0.70 0.6 d d 0 d d d 0 0 d d ON 0 d d r~; 0 00 d ^ d vo d in d ^ ^ 0 ^^ 0 r-i 0 ON 0 00 0 0 d 00 -a =x +1 1 — +1 1 — ' +1 1 0 +1 1 0 +1 1 0 +1 1 0 ^ 0 d X) 0 d d n d 0 d d (N d in d d 00 d 0 d d <1> 0 0 't T^ r<-i ^ d — : ^ d m 00 d ^ 00 d r<-i ON C > 0 0 oi d ,__, 0 r<-) d 0 d (N d 03 ~ .i2 0 Q +1 m ri d d d +1 CnI d 1 d d 0.09 ± 0.07- d +1 d 1 d d +1 ON d 1 (N d +1 CN d 1 d d X ^ On vq ON NO 0) . Q 5 s ^ X e2 G -« •« -c — = Co CO CO CO CO CO A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 47 r<-, __ r<~i fN ._ _. ^^ O' 0 0 0 0 0 0 o r] r- c-j vO d ON '^ d ^_ ^ d r<-) >r-, d ' — ; ^ d r«". NO _^ o ^« 0 ? >/-) 0 1— > 0 ^^ 0 ^— 0 ^^ 0 +1 1 o d +1 1 q d +1 d +1 1 q d +1 _^ d +1 1 q d +1 1 q d 00 r~r, 00 00 ^^ ■* Ol 00 q ^^ CA '""' 00 d c-j "~ r-j ~ q "^ ' — ' ^ ^^ ^ d ^^ ^^ ""^ ^^ (N , «"- r<~, _j ,__, n O O 0 0 r^ 0 0 0 d ^ ^ d ^ >n d — _ t^ d ON d NO ITi d r«-, ITi d ir, oc ^« s^ 1— q ^— 0 1 •— c — 0 ^— 0 +1 1 d +1 1 +1 ci d +1 q d +1 1 d +1 J^ d +1 1 ON d r<~i 00 ir, ON NO vO "* d r«^i ^^ '^ "^ q '"^ »n — I ^ ~~ — ; "^ rn — < ^ ^ ^ ^^ — ' — ' (N , m r<-) , ^^ .« O o 0 0 0 0 0 d 00 >sO d q ON d 00 d n-j ri d q 00 d 00 ON d ON r-~ d o 1-^ 0 d d 0 — 1—1 0 d 0 d 0 +1 i d +1 1 d +1 d +1 1 00 d +1 1 d +1 ^ d +1 1 d -* oc d -* ON d 0 00 d d NO d d d d d d d d m ^^ ■ 0 r<^ __ ^« , — , o q 0 ^D 0 0 0 d 00 d q d ir, d "1 d q d 00 d ON o S 1 ON d 0 „_ r<-, _^ ON d 00 d 00 +1 1 +1 1 0 +1 1 0 +1 1 ' — ' +1 1 0 +1 1 0 +1 1 0 o d d OC d ^ 0 d d q ON d d 00 1^ d d NO NO d d t^ r^ d d d d d --^ d d d m _j 0 __ ^ , 1 ^^ q o 0 0 0 0 0 0 d t^ d 00 d vO d 00 d ON d NO d r~- d ir^ d t— d 0 d S d 0 d ON d ON +1 1 — ' +1 1 ^- +1 1 c +1 1 0 +i 1 ^" +1 0 +1 1 0 d d d d c nC d 1^ 1^ 0 d 1^ NO NO d d 0 d d s d d d d d d d d d M 1 m ^^ ,,^ ,_^ ^ q o 0 0 r^i 0 0 0 d u-j d MD d d 00 d t~~; d in d r-- o GO 0 ON d 1 ^^ d 3 d 00 d "* d 0 +1 1 o +1 0 +1 — ' +1 1 0 +1 1 0 +1 1 — ' +1 1 ' — ' 00 '^ d r*". "v|- d rr-, 1 d ON d 00 in d r<-j ro d ^ >n d '^ d >/", d m d VO d m d ^ d m 0 d d d d d d d r<-j ^ 0 _ _ , ^.^ O o q 0 0 0 0 d ■* (N d m (N 0 ri 0 d >n NO d n^, ■^ d 00 d r<-, 0 d CN d <^1 ^ 0 d d *— 1 d r^) +1 1 d +1 1 d +1 1 d +1 1 d +1 1 d +1 1 d +1 1 d c, d o d 0 d d ON d ri '^— ' NO d d d d d d d d CM 00 1/-J vq On 00 +1 1 +1 0 ri 1 d +1 1 +1 00 +1 1 +1 1 +1 1 r^i r«-, OS n ir-, ^ ri "" rn 1 -H u-i 00 00 ■^ u-i ^ u^ 00 d d ^ O 0 5 5 0 .2- &n ts .0 .^ ;_ D. ^ a "S 0 ^-.* C 0 4— ' X3 3 5 j,^ e2 ^ s i s S 0 tin 0 00 03 to '^ to CO CO 48 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 175 g)S ^ o -a 5 T3 1= '^ ^ o .22 o 2 c ri +1 7 ON en ■■—A Co c d ^! r- — o +1 I ~ 00 ° o o o d f^; o — o ^~ ^ ° o d °o _ d -- +1 I Q Iq o d o d =» d 2^ +1 I q t- ^ o d o d +1 o d +1 ^ ON c d ^ d NO o d I +1 I —. NO 9 22 >n o (N o d +1 7 oo ""' NO CO § !5 "^ +1 7 § ^ ^ ° o d +1 ON +1 I m 00 '- o d o ir, 8o o d '^' I 9 q o CNl o d +1 ON o — (N OO o d O d +1 00 ON ON o d NO q d -^ oo o — +' rl, O d ON I^ +1 1 o oo '^^ ""^ o ro 5 5b S o to o * ly^j O ' u~i -^ O +1 I ~ r-\ ^ 00 o d +1 I oo 00 ON 1^ o 00 NO o d NO + 1 I c ON ON o d — : +1 7 __ ^ o d o d +1 I o oo o 00 d o d +1 I — d CO +1 7 O !>1 i ^ o d o d "^ — +i°d ^. d d o d ^i r-, +1 I -7 O _ O c d +1 o (N O d '^' +1 d o d ^' +1 1 ^ o d ir-j ri .„ +1 in d Si ■§ CO I oo "n o d o d "^^ vo O O +1 I ~ 00 _; d o d r^ o S +1 I q 00 ^ ° d o d ^ O 00 +1 I P d in o O 00 I 9 -^ o d n I A KEY TO RECENT SHREWS OF THE FAMILY SORICIDAE 49 lU 3 C O U O d +1 ^ O ri o d +1 oc O d ^ +1 I ON d (N O d +1 ON 00 d (N O d ^ +1 "T' ^ O 00 +1 7 o6 NO o 00 d o 00 o O d °° d o +1 I — . r, ^ O d (^) o ^; d NO o d ON. .to I I CO , — 1 ^^ ^^ CM 0 0 0 0 ^ "* V~i d ^i ^, d ^ >r^ d ^ ON ^— C , , V 0 ^ 0 +1 1 d +' i d +1 1 ON d +1 1 0 d 00 • 00 _; >^' ^ Tf ^ r-l "" ^^ ^— ^ r-- o d ■ . -^ 00 +1 o d "^ I o +1 00 o d r^ d o d r^ o 00 +1 I P '^' d ° NO O d o d ^ o o +1 I -: ^ ^ o NO • — in o d o +1 ^ o r<-i d :^ O c« C<] +i7S ON O d °9 +1 7 o d "^ IT) NO ^* d o d +1 T ON d o d +1 00 (N O o ir, d d I d NO r~; r^; — < r<-, r^i 0 +1 T +1 1 NO •n NO 06 NO 0 0 m c^ yi '& - • 2 ^ 'u to OJ i^ -5 D. o d '^r uo — o +1 ^ d q 0 0 d ON ON d 00 00 d r- ;^ 0 ^ s.^ ^.^ +1 r-- d +1 1 nC d +1 1 0 r<~i 00 d r^ d NO d d d d ■n d S o d +1 NO 03 o a 00 o p d ^ +1 o ^ O Tt o •^i d o p d ^ +1 o )S p ^ o d O O d ^i ^, 04 >o => d o •a +1 I fN O d 10 IT) o +1 I rr; NO • o d P _ +1 "i" d ^ NO => nD _; CNl o d +1 nO o O d r^ o »< +1 I -: "n o 00 _• '— ^ IT) O d o d ^ o )0 +1 I P „ "^i o 10 o d o d ^. o +1 I ^, d d o +1 7 06 ON o CO l>3 00 00 ON N c 3 Xl C3 3 0 L. 0 rt c 0 ■"^ "^ ■0 C/5 5 03 ■^ ^-* X> 1/5 1/3 _0J CJ — *^— » w 0 c 0 ^ (U 1) c/^ _N N ^ 'i^ '^ 03