Fig. 7. Possible distribution in Illinoian (inset) and Sangamonian times of the ancestor of the Sorex vagrans-ornatus-longirostris-veraepacis complex. Approximate southern boundary of Illinoian glaciation marked by heavy line.

In the ensuing Sangamonian interglacial age all glaciers retreated or disappeared thereby opening up extensive areas in the north and in the higher mountains which were occupied by a boreal fauna, including S. vagrans. Concurrently the Great Basin, and probably also much of the Columbian Plateau, became dry, and desert conditions developed, perhaps much as they are today. Increasing aridity eliminated shrew habitat in most places between the Rocky Mountains and the Sierra Nevada-Cascade mountain chain with the result that the geographic range of the species resembled an inverted "U", one arm lying along the Rocky Mountains and the other along the Cascade-Sierra Nevada axis; the connection between the two arms was in British Columbia (see [fig. 7]). At present Sorex vagrans does occur in isolated places in the Great Basin, but its existence there is tenuous and seemingly dependent upon the occurrence of permanent water such as Ruby Lake and Reese River. With such an arrangement as this it can readily be seen that gene flow between the eastern and western arms of the "U" would be greatly reduced by distance; consequently differentiation between the two might be expected.

Fig. 8. Possible distribution of Sorex vagrans at two different times in the Wisconsinan Age. Left, early Wisconsinan; right, mid-Wisconsinan.

Wisconsinan glaciation again rendered Canada uninhabitable, and it is quite possible that extensive areas in the Rocky Mountains, the Cascades and the Sierra Nevada were heavily glaciated. With the elimination of the northern part of the "U", the eastern and western arms became isolated, if not by the width of the Columbian Plateau at least by the glaciated Cascade Mountains. At the same time extensive areas on the Colorado Plateau and much of the area south to the Mexican highlands were again occupied by the species. Finally the Great Basin, again being well-watered, provided suitable habitat for, and was reoccupied by, Sorex vagrans (see [fig. 8]). This reoccupation of the Great Basin took place probably from the Colorado Plateau and mountains of Arizona and Utah, since the present day shrews of the species S. vagrans in the Great Basin closely resemble Rocky Mountain shrews but differ markedly from the large endemic subspecies of the Pacific Coast.

Finally, with the waning of Wisconsinan ice, the species again was able to occupy northern and montane areas as it had during Sangamonian times. Again dessication of the Great Basin caused drastic restriction of shrew habitat. The small, marsh-dwelling kind of wandering shrew which had developed there around the lakes of Wisconsinan time occupied suitable habitat all the way to the Pacific coast where its range came into contact with that of the western arm of the Sangamonian "U."-pattern of shrew distribution (see [fig. 9]). The animals of this western segment and the new arrivals from the east were by this time so different from one another that the two kinds lived in the same areas without interbreeding. The descendants of the original western arm now are known as Sorex vagrans sonomae, S. v. pacificus, S. v. yaquinae, and S. v. bairdi. The newcomers from the east are known as S. v. vagrans, S. v. halicoetes, S. v. paludivagus and S. v. vancouverensis.

In addition to occupying the Pacific Coast from San Francisco Bay north to the Fraser Delta, the Great Basin subspecies populated the Columbia Plateau and the western foothills of the central and northern Rockies. By so doing that subspecies came into secondary contact with its own parent stock with which it was still in reproductive continuity in Utah. In some places in British Columbia differentiation between the two kinds had proceeded to such an extent that some reproductive isolation was effected, but in many other places the two interbred. The Rocky Mountain form spread north and west and occupied the Cascades and coastal lowlands in southwestern British Columbia and in Washington. Here the differentiation between the Rocky Mountain subspecies and the Great Basin subspecies was great enough to cause complete reproductive isolation.