Remarks.--This is a poorly differentiated subspecies which is closely related to S. v. vagrans. The differences in color noted are average ones. Some individuals of this shrew might be difficult to separate from S. v. isolatus. The slight degree of morphological divergence is such that intergrades might be expected to occur. Possibly some habitat separation occurs, but such has not been reported.
Specimens examined.--Total number, 3. BRITISH COLUMBIA: Vancouver Island: Mt. Washington, 1 KU; Nanaimo, 1 BS; type locality, 1 BS.
Marginal records.--BRITISH COLUMBIA: Sayward (Anderson, 1947:18); Bowen Island (Hall, 1938:463); Alberni (Jackson, 1928:107).
CONCLUSIONS
1. Sorex vagrans, S. obscurus, S. pacificus, and S. yaquinae are conspecific with one another. Each is a valid subspecies but all should bear the specific name Sorex vagrans Baird, 1858.
2. The subspecies of Sorex vagrans form a cline from large (pacificus) to small (vagrans). The cline is bent in such a manner that the terminal subspecies occur together. Where the two subspecies occur together, individuals of one subspecies do not crossbreed with individuals of the other subspecies and therefore react toward one another as do full species. Sorex vagrans vagrans occurs sympatrically with S. v. sonomae, S. v. pacificus, S. v. yaquinae, S. v. bairdi, S. v. permiliensis, and S. v. setosus. S. v. vancouverensis occurs sympatrically with S. v. isolatus.
3. The sympatric existence of the terminal subspecies of the Sorex vagrans rassenkreis is made possible by marked differences between them in size and in ecological preference.
4. The west-coast subspecies, sonomae, pacificus, yaquinae, bairdi, and permiliensis probably differentiated from the Great Basin and Rocky Mountain subspecies, vagrans, obscurus and monticola, during a separation caused first by aridity in the Great Basin, and secondly by glaciation of the Cascade Mountains and the Sierra Nevada, possibly in the Sangamonian and Wisconsinan ages respectively.
5. Sorex v. vagrans originated in the Great Basin and arrived on the Pacific Coast after the last deglaciation of the Cascades and Sierra Nevada.
6. In S. vagrans, heterogonic growth is illustrated; the larger the skull, the larger the rostrum in proportion to the skull as a whole.
7. In the species S. vagrans, size and color vary geographically more than do other features.
8. The S. ornatus group, S. longirostris, and S. veraepacis had a common ancestor with S. vagrans, possibly in the Illinoian Age.
9. S. vagrans, the S. ornatus group, S. veraepacis, S. longirostris, S. palustris, S. bendiri, and the S. cinereus group, because of structural resemblances, should be placed in a single subgenus, Otisorex. S. trowbridgii, the S. arcticus group, the S. saussurei group, S. merriami, S. fumeus, and S. dispar, should be included in the subgenus Sorex.
10. Sorex cinereus occurs with the medium-sized and large-sized S. vagrans in the Rocky Mountains and in Canada, but does not occur with the smaller subspecies of S. vagrans, probably because competition between two shrews of like size excludes S. cinereus.
TABLE 1--CRANIAL MEASUREMENTS OF SOREX VAGRANS
========================================================================== Catalog | | | | | | number or | | | | | Least | number of |Condylobasal|Palatal|Maxillary|Cranial|interorbital|Maxillary individuals| length |length |tooth-row|breadth| breadth | breadth averaged | | | | | | ------------+------------+-------+---------+-------+------------+--------- | Sorex vagrans pacificus, Orick, California. 8 av | 21.8 | 9.6 | 8.6 | 10.4 | 4.1 | 6.6 Max | 22.8 | 10.2 | 9.0 | 11.1 | 4.3 | 6.8 Min | 21.3 | 9.3 | 8.4 | 10.2 | 4.1 | 6.4 +------------+-------+---------+-------+------------+--------- | Sorex vagrans yaquinae, Newport, Oregon. 707 AW | 20.1 | 8.9 | 7.6 | 9.3 | 3.7 | 5.7 706 AW | 19.3 | 8.8 | 7.3 | 9.3 | 4.0 | 5.8 +------------+-------+---------+-------+------------+--------- | Mapleton, Oregon. 205273 USBS| 20.6 | 9.0 | 8.2 | 9.9 | 4.2 | 6.0 205270 USBS| 20.4 | 8.9 | 7.9 | 9.3 | 3.7 | 6.0 205272 USBS| | 9.2 | 8.3 | | 4.0 | 6.1 +------------+-------+---------+-------+------------+--------- | Vida, Oregon. 4 av | 19.5 | 8.4 | 7.5 | 9.2 | 3.6 | 5.5 Max | 20.3 | 8.8 | 7.9 | 9.5 | 3.7 | 5.7 Min | 19.3 | 8.2 | 7.3 | 8.9 | 3.5 | 5.3 +------------+-------+---------+-------+------------+--------- | McKenzie Bridge, Oregon. 6 av | 18.9 | 8.2 | 7.1 | 9.1(5)| 3.7 | 5.6 Max | 19.5 | 8.6 | 7.6 | 9.4 | 3.8 | 5.7 Min | 18.7 | 8.0 | 6.8 | 8.4 | 3.6 | 5.4 +------------+-------+---------+-------+------------+--------- | Sorex vagrans bairdi, Astoria, Oregon. 6 av | 18.5 | 7.8 | 7.0 | 8.9(4)| 3.4 | 5.3 Max | 19.2 | 8.1 | 7.4 | 9.0 | 3.5 | 5.5 Min | 18.0 | 7.6 | 6.9 | 8.9 | 3.2 | 5.2 +------------+-------+---------+-------+------------+--------- | Sorex vagrans permiliensis, Mt. Jefferson, Oregon. 14 av | 18.0 | 7.5 | 6.8 | 9.0 | 3.5 | 5.2 Max | 18.9 | 7.9 | 7.2 | 9.4 | 3.7 | 5.3 Min | 17.2 | 7.2 | 6.5 | 8.6 | 3.3 | 4.8 +------------+-------+---------+-------+------------+--------- | Sorex vagrans setosus, Olympic Mts., Washington. 12 av | 17.2(9) | 7.1 | 6.5 | 8.5(8)| 3.3 | 5.0 Max | 17.9 | 7.4 | 6.8 | 8.7 | 3.4 | 5.3 Min | 16.7 | 6.9 | 6.2 | 8.3 | 3.0 | 4.8 +------------+-------+---------+-------+------------+--------- | Mt. Rainier, Washington. 16 av | 17.2 | 7.1 | 6.5 |8.4(14)| 3.3 | 5.0 Max | 17.6 | 7.3 | 6.7 |8.7 | 3.5 | 5.2 Min | 16.4 | 6.5 | 6.1 |8.1 | 3.2 | 4.7 +------------+-------+---------+-------+------------+--------- | Sorex vagrans longicauda, head Rivers Inlet, B.C. 15 av | 18.0 | 7.4 | 6.7 | 8.7 | 3.2 | 5.0 Max | 18.4 | 7.6 | 6.9 | 8.9 | 3.3 | 5.2 Min | 17.6 | 7.2 | 6.4 | 8.4 | 3.1 | 4.8 +------------+-------+---------+-------+------------+--------- | Port Simpson, British Columbia. 10 av | 18.1(9) | 7.6 | 7.0 | 8.9 | 3.4 | 5.1 Max | 18.8 | 7.8 | 7.2 | 9.2 | 3.6 | 5.4 Min | 17.2 | 7.2 | 6.6 | 8.5 | 3.3 | 4.9 +------------+-------+---------+-------+------------+--------- | Fort Wrangell, Alaska. 18 av | 18.5 |7.8(15)| 7.1 |9.0(15)| 3.3 | 5.1 Max | 18.9 |8.0 | 7.3 |9.2 | 3.5 | 5.3 Min | 17.8 |7.5 | 6.7 |8.6 | 3.2 | 5.0 +------------+-------+---------+-------+------------+--------- | Sorex vagrans elassodon, Woewodsky Is., Alaska. 20550 AMNH | 18.0 | 7.5 | 6.7 | 8.7 | 3.3 | 5.0 20553 AMNH | 17.5 | 7.1 | 6.3 | 8.3 | 3.2 | 4.7 +------------+-------+---------+-------+------------+--------- | Sorex vagrans alascensis, | 9 mi. W and 4 mi. N Haines, Alaska. 10 av | 17.2 | 7.2 | 6.7 | 8.5 | 3.1 | 4.9 Max | 17.6 | 7.4 | 6.9 | 8.8 | 3.3 | 5.0 Min | 16.9 | 6.9 | 6.5 | 8.2 | 3.0 | 4.7 +------------+-------+---------+-------+------------+--------- | Yakutat Bay, Alaska. 73543 USBS | 18.0 | 7.5 | 6.8 |... | 3.2 | 5.0 73536 USBS | 18.0 | 7.6 | 6.8 |8.8 | 3.4 | 5.3 73541 USBS | 17.9 | 7.4 | 6.7 |8.8 | 3.1 | 5.2 +------------+-------+---------+-------+------------+--------- | Sorex vagrans shumaginensis, Sandpoint, Popof Is., Alaska. 9 av | 17.2(5) | 7.0 | 6.3 |8.3(7) | 3.1 | 4.8 Max | 17.6 | 7.2 | 6.6 |8.5 | 3.2 | 5.1 Min | 16.8 | 6.8 | 6.1 |8.0 | 3.0 | 4.7 +------------+-------+---------+-------+------------+--------- | Sorex vagrans obscurus, Barkerville, British Columbia. 5 av | 17.1 | 7.1 | 6.5 |8.5(4) | 3.3 | 4.8 Max | 17.3 | 7.3 | 6.6 |8.6 | 3.3 | 5.0 Min | 16.6 | 6.7 | 6.4 |8.2 | 3.2 | 4.6 +------------+-------+---------+-------+------------+--------- | 10 mi. SSW Leadore, Idaho. 7 av | 17.2(4) | 7.3(9)| 6.6 |8.6(4) | 3.3 | 5.0 Max | 17.3 | 7.5 | 6.8 |8.9 | 3.4 | 5.1 Min | 17.0 | 7.1 | 6.4 |8.3 | 3.2 | 4.7 +------------+-------+---------+-------+------------+--------- | Albany Co., Wyoming (several localities). 20 av | 17.3 | 7.3 | 6.8 |8.7(19)| 3.2 | 5.2 Max | 17.9 | 7.6 | 6.9 |9.0 | 3.4 | 5.5 Min | 16.7 | 6.9 | 6.5 |8.4 | 3.1 | 5.0 +------------+-------+---------+-------+------------+--------- | Sorex vagrans longiquus, 25 mi. ESE Big Sandy, Montana. 87332 UM | 16.4 | 6.8 | 6.2 |8.2 | 3.0 | 4.8 87334 UM | 16.8 | 7.1 | 6.3 |8.1 | 3.2 | 4.7 87335 UM | 15.8 | 6.7 | 6.0 |8.4 | 3.1 | 4.9 +------------+-------+---------+-------+------------+--------- | Highwood Mts., Montana. 10 av | 16.3(9) | 6.7 | 6.2 |8.0(9) | 3.1 | 4.7 Max | 16.9 | 6.9 | 6.4 |8.3 | 3.3 | 5.0 Min | 15.6 | 6.5 | 6.0 |7.8 | 3.0 | 4.5 +------------+-------+---------+-------+------------+--------- | Sorex vagrans neomexicanus, Cloudcroft, New Mexico. 4 av | 17.6(3) | 7.6 | 7.0 | 8.7 | 3.3 | 5.2 Max | 17.7 | 7.7 | 7.1 | 8.8 | 3.4 | 5.4 Min | 17.4 | 7.4 | 7.0 | 8.5 | 3.2 | 5.1 +------------+-------+---------+-------+------------+--------- | Sorex vagrans monticola, White Mts., Arizona. 12 av | 16.1(9) | 6.6 | 5.9(9) | 8.2 | 3.1(11) | 4.7 Max | 16.6 | 7.0 | 6.1 | 8.5 | 3.3 | 4.9 Min | 15.5 | 6.5 | 5.6 | 8.1 | 3.0 | 4.6 +------------+-------+---------+-------+------------+--------- | Sorex vagrans orizabae, Volcan Toluca, Mexico. 55900 USBS | 17.1 | 6.9 | 6.2 | 7.8 | 2.9 | 4.5 55898 USBS | 17.1 | 6.8 | 6.1 | 8.0 | 3.0 | 4.8 55897 USBS | 16.8 | 6.9 | 6.1 | 7.9 | 2.9 | 4.6 +------------+-------+---------+-------+------------+--------- | Sorex vagrans vagrans, Lincoln Co., Wyoming. 7 av | 16.5(6) | 6.6 | 6.1 | 8.2 | 2.9 | 4.7 Max | 17.1 | 7.0 | 6.4 | 8.5 | 3.1 | 4.9 Min | 16.0 | 6.4 | 5.9 | 7.9 | 2.9 | 4.5 +------------+-------+---------+-------+------------+--------- | Gearhart Mtn., Lake Co., Oregon. 17 av | 16.5(15) | 6.6 | 5.9 | 8.1 | 2.9 | 4.6 Max | 17.1 | 7.0 | 6.6 | 8.5 | 3.1 | 4.9 Min | 16.1 | 6.2 | 5.7 | 7.8 | 2.8 | 4.4 +------------+-------+---------+-------+------------+--------- | Willapa Bay, Washington. 9 av | 16.6 | 6.8 | 6.2 | 8.1 | 2.8 | 4.7 Max | 17.2 | 7.1 | 6.4 | 8.3 | 3.1 | 5.0 Min | 16.2 | 6.6 | 5.9 | 7.9 | 2.7 | 4.6 +------------+-------+---------+-------+------------+--------- | Sorex vagrans obscuroides, Bishop Creek, California. 4 av | 16.7 | 7.0 | 6.3 | 8.2 | 3.2 | 4.8 Max | 16.8 | 7.1 | 6.4 | 8.3 | 3.4 | 4.9 Min | 16.6 | 6.9 | 6.2 | 8.1 | 3.1 | 4.7 +------------+-------+---------+-------+------------+--------- | Mt. Whitney, California. 4 av | 16.7(3) | 6.9 | 6.3 | 8.4 | 3.3 | 4.8 Max | 16.7 | 7.0 | 6.4 | 8.5 | 3.4 | 5.0 Min | 16.7 | 6.7 | 6.1 | 8.4 | 3.1 | 4.7 +------------+-------+---------+-------+------------+--------- | Sorex vagrans parvidens, San Bernardino Peak, California. 56559 USBS | 17.1 | 7.0 | 6.1 | 8.0 | 2.9 | 4.8 56558 USBS | 16.4 | 6.8 | 6.1 | 8.0 | 3.0 | 4.8 +------------+-------+---------+-------+------------+---------
LITERATURE CITED
ANDERSON, R. M. 1947. Catalogue of Canadian Recent mammals. Nat. Mus. Canada, Bull. 102, Biol. ser. 31:i-v + 1-238, January 24.
ANDERSON, R. M. and A. L. RAND 1945. A new form of dusky shrew from the prairie provinces of Canada. Canadian Field-Nat., 59:47-48, March-April.
BAILEY, V. 1936. The mammals and life zones of Oregon. N. Amer. Fauna, 55:1-416, 52 pls., 102 figs. in text, August 29.
BROWN, B. 1908. The Conard Fissure, a Pleistocene bone deposit in northern Arkansas: with descriptions of two new genera and twenty new species of mammals. Mem. Amer. Mus. Nat. Hist., 9:157-208, pls. 14-25, February.
CLOTHIER, R. R. 1950. Contribution to the taxonomy and life history of Sorex vagrans monticola and Sorex obscurus obscurus. Master's thesis, Montana State University, Missoula, Montana--a manuscript.
CONAWAY, C. H. 1952. Life history of the water shrew (Sorex palustris navigator). Am. Midl. Nat., 48:219-248, 6 tables, 9 figs. in text, July.
COWAN, I. MCT. 1936. Distribution and variation in deer (Genus Odocoileus) of the Pacific coastal region of North America. California Fish and Game, 22(3):155-246, 13 figs., 3 graphs, 8 tables, July.
1941. Insularity in the genus Sorex on the north coast of British Columbia. Proc. Biol. Soc. Washington, 54:95-108, July 31.
DALQUEST, W. W. 1941. Ecologic relationships of four small mammals in western Washington. Jour. Mamm., 22:170-173, May 14.
1944. The molting of the wandering shrew. Jour. Mamm., 25:146-148, one fig. in text, May 25.
1948. Mammals of Washington. Univ. Kansas Publ., Mus. Nat. Hist., 2:1-444, 140 figs. in text, April 9.
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