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Title: Speciation of the Wandering Shrew
Author: Findley, James S.
Language: English
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  ==================================================================
                  UNIVERSITY OF KANSAS PUBLICATIONS
                     MUSEUM OF NATURAL HISTORY

               Volume 9, No. 1, pp. 1-68, figures 1-18
  --------------------    December 10, 1955    ---------------------


                    Speciation of the Wandering Shrew


                                  BY
                           JAMES S. FINDLEY


                        UNIVERSITY OF KANSAS
                               LAWRENCE
                                 1955



     UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

       Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
                Robert W. Wilson

              Volume 9, No. 1, pp. 1-68, figures 1-18
                    Published December 10, 1955

                        UNIVERSITY OF KANSAS
                           Lawrence, Kansas

                              PRINTED BY
                    FERD VOILAND, JR., STATE PRINTER
                            TOPEKA, KANSAS
                                 1955
                      [Illustration: union label]
                               25-7903



Speciation of the Wandering Shrew

BY

JAMES S. FINDLEY



CONTENTS


                                                                PAGE

 INTRODUCTION                                                      4

 MATERIALS METHODS AND ACKNOWLEDGMENTS                             4

 NON-GEOGRAPHIC VARIATION                                          7

 CHARACTERS OF TAXONOMIC WORTH                                     8

 PELAGE CHANGE                                                     9

 GEOGRAPHIC DISTRIBUTION AND VARIATION                             9
   Pacific Coastal Section                                         9
   Inland Montane Section                                         11
   Great Basin and Columbia Plateau Section                       12
   Summary of Geographic Variation                                13

 ORIGIN OF THE _Sorex vagrans_ RASSENKREIS                        16

 RELATIONSHIPS WITH OTHER SPECIES                                 26

 CONCLUSIONS                                                      60

 TABLE OF MEASUREMENTS                                            62

 LITERATURE CITED                                                 66



FIGURES


 FIGS. 1-2.--CRANIAL MEASUREMENTS                                  5

 FIG. 3.--GRAPH ILLUSTRATING WEAR OF TEETH                         8

 FIG. 4.--GRAPH ILLUSTRATING HETEROGONIC GROWTH OF ROSTRUM        10

 FIG. 5.--PRESENT GEOGRAPHIC DISTRIBUTION OF _Sorex vagrans_      15

 FIG. 6.--SKULLS OF _Sorex vagrans_                               17

 FIGS. 7-10.--PAST GEOGRAPHIC DISTRIBUTION OF SHREWS     19-20-22-27

 FIGS. 11, 12.--MEDIAL VIEW OF LOWER JAWS OF TWO SHREWS           30

 FIGS. 13, 14.--SECOND UNICUSPID TEETH OF SHREWS                  30

 FIG. 15.--DIAGRAM OF PROBABLE PHYLOGENY OF SHREWS                32

 FIGS. 16-18.--GEOGRAPHIC DISTRIBUTION OF SUBSPECIES        33-40-53



INTRODUCTION


The purpose of this report is to make clear the biological relationships
between the shrews of the _Sorex vagrans-obscurus_ "species group." This
group as defined by H. H. T. Jackson (1928:101) included the species
_Sorex vagrans_, _S. obscurus_, _S. pacificus_, _S. yaquinae_, and _S.
durangae_. The last mentioned species has been shown (Findley, 1955:617)
to belong to another species group. _Sorex milleri_, also assigned to
this group by Jackson (1947:131), seems to have its affinities with the
_cinereus_ group as will be explained beyond. The position of the
_vagrans_ group in relationship to other members of the genus will be
discussed.

Of this group, the species that was named first was _Sorex vagrans_
Baird, 1858. Subsequently many other names were based on members of
the group and these names were excellently organized by Jackson in his
1928 revision of the genus. Subsequent students of western mammals,
nevertheless, have been puzzled by such problems as the relationship of
(1) _Sorex vagrans monticola_ to _Sorex obscurus obscurus_ in the Rocky
Mountains, (2) _Sorex pacificus_, _S. yaquinae_, and _S. obscurus_ to
one another on the Pacific Coast, and (3) _S. o. obscurus_ to
_S. v. amoenus_ in California. Few studies have been made of these
relationships. Clothier (1950) studied _S. v. monticola_ and _S. o.
obscurus_ in western Montana and concluded that the two supposed kinds
actually were not separable in that area. Durrant (1952:33) was able to
separate the two kinds in Utah as was Hall (1946:119, 122) in Nevada.
Other mammalogists who worked within the range of the _vagrans-obscurus_
groups have avoided the problems in one way or another. Recently Rudd
(1953) has examined the relationships of _S. vagrans_ to _S. ornatus_.



MATERIALS METHODS AND ACKNOWLEDGMENTS


Approximately 3,465 museum study skins and skulls were studied. Most
of these were assembled at the University of Kansas Museum of Natural
History, but some were examined in other institutions.

Specimens were grouped by geographic origin, age, and sex. Studies of
the role of age and sex in variation were made. Because it was
discovered that secondary sexual variation was negligible, both males
and females, if of like age and pelage, were used in comparisons
designed to reveal geographic variation.

External measurements used were total length, length of tail, and
length of hind foot. After studying a number of cranial dimensions I
chose those listed below as the most useful in showing differences in
size and proportions of the skull. Figures 1 and 2 show the points
between which those measurements were taken.

_Condylobasal length._--From anteriormost projection of the
premaxillae to posteriormost projection of the occipital condyles
(a to a´).

_Maxillary tooth-row._--From posteriormost extension of M3 to
anteriormost extension of first unicuspid (b to b´).

_Palatal length._--From anteriormost projection of premaxillae to
posteriormost part of bony palate (c to c´).

_Cranial breadth._--Greatest lateral diameter of braincase (d to d´).

_Least interorbital breadth._--Distance between medialmost superior
edges of orbital fossae, measured between points immediately above and
behind posterior openings of infraorbital foramina (e to e´).

_Maxillary breadth._--Distance between lateral tips of maxillary
processes (f to f´).

   [Illustration: FIGS. 1 AND 2. Showing where certain cranial
      measurements were taken. × 3-1/2. (Based on _Sorex vagrans
      obscurus_, from Stonehouse Creek, 5-1/2 mi., W junction of
      Stonehouse Creek and Kelsall River, British Columbia,
      [Female], 28545 KU.)]

In descriptions of color, capitalized terms refer to those in Ridgway
(1912). In addition the numerical and alphabetical designations of
these terms are given since a knowledge of the arrangements of these
designations enables one quickly to evaluate differences between
stated colors. Color terms which are not capitalized do not refer to
any precise standard of color nomenclature.

In the accounts of subspecies, descriptions, unless otherwise noted,
are of first year animals as herein defined. Descriptions of color are
based on fresh pelages.

Unless otherwise indicated, specimens are in the University of Kansas
Museum of Natural History. Those in other collections are identified
by the following abbreviations:

 AMNH  American Museum of Natural History
 CM    Carnegie Museum
 ChM   Chicago Museum of Natural History
 CMNH  Cleveland Museum of Natural History
 FC    Collection of James S. Findley
 HC    Collection of Robert Holdenreid
 SGJ   Collection of Stanley G. Jewett
 CDS   Collection of Charles D. Snow
 AW    Collection of Alex Walker
 NMC   National Museum of Canada
 OSC   Oregon State College
 PMBC  British Columbia Provincial Museum of Natural History
 SD    San Diego Natural History Museum
 BS    United States Biological Surveys Collection
 USNM  United States National Museum
 UM    University of Michigan Museum of Zoology
 OU    University of Oregon Museum of Natural History
 UU    University of Utah Museum of Zoology
 WSC   Washington State College, Charles R. Conner Museum

In nature, the subspecies of _Sorex vagrans_ form a cline and are
distributed geographically in a chain which is bent back upon itself.
The subspecies in the following accounts are listed in order from the
southwestern end of the chain clockwise back to the zone of overlap.

The synonymy of each subspecies includes the earliest available name
and other names in chronological order. These include the first usage
of the name combination employed by me and other name combinations
that have been applied to the subspecies concerned.

In the lists of specimens examined, localities are arranged first by
state or province. These are listed in tiers from north to south and
in any given tier from west to east. Within a given state, localities
are grouped by counties, which are listed in the same geographic
sequence as were the states and provinces (N to S and W to E). Within
a given county, localities are arranged from north to south. If two or
more localities are at the same latitude the westernmost is listed
first. Marginal localities are listed in a separate paragraph at the
end of each account. The northernmost marginal locality is listed
first and the rest follow in clockwise order. Those records followed
by a citation to an authority are of specimens which I have not
personally examined. Marginal records are shown by dots on the range
maps. Marginal records which cannot be shown on the maps because of
undue crowding are listed in Italic type.

To persons in charge of the collections listed above I am deeply
indebted. Without their generous cooperation in allowing me to examine
specimens in their care this study would not have been possible.
Appreciated suggestions in the course of the work have been received
from Professors Rollin H. Baker, A. Byron Leonard, R. C. Moore, Robert
W. Wilson, and H. B. Tordoff, and many of my fellow students. Mr.
Victor Hogg gave helpful suggestions on the preparation of the
illustrations. My wife, Muriel Findley, devoted many hours to
secretarial work and typing of manuscript. Finally I am grateful to
Professor E. Raymond Hall for guidance in the study and for assistance
in preparing the manuscript. During the course of the study I received
support from the University of Kansas Endowment Association, from the
Office of Naval Research, and from the National Science Foundation.



NON-GEOGRAPHIC VARIATION


Non-geographic variation, that is to say, variation within a single
population of shrews, consists of variation owing to age and normal
individual variation. In _Sorex_ I have detected no significant
secondary sexual differences between males and females; accordingly
the two sexes are here considered together.

Variation with age must be considered in order to assemble comparable
samples of these shrews. Increased age results in wear on all teeth
and in particularly striking changes in the size and shape of the
first incisors. Skulls of older shrews develop sagittal and lambdoidal
ridges, and further differ from skulls of young animals in being
slightly broader and shorter, and in developing thicker bone,
particularly on the rostrum which thus seems to be, but is not always
in fact, more robust. Pruitt has recently (1954) noted these same
cranial differences in specimens of _Sorex cinereus_ of different
ages.

Several students of American shrews, notably Pearson (1945) on
_Blarina_, Hamilton (1940) on _Sorex fumeus_, and Conaway (1952) on
_Sorex palustris_, have shown that young are born in spring and
summer, usually reach sexual maturity the following spring, and rarely
survive through, or even to, a second winter. The result is that
collections made, as most of them are, in spring and summer, contain
two age classes, first year and second year animals. These two age
classes are readily separable on the basis of differences in the skull
as well as on the decreased pubescence of the tail and the increased
weight of second year animals. My own examination of hundreds of
museum specimens confirms this for the _Sorex vagrans_ group.
Separation of the two age classes in an August-taken series of _Sorex
vagrans_ from coastal Washington is shown in figure 3, in which two
tooth-measurements that are dependent upon wear are plotted against
one another.

First year animals are more abundant in collections than are second
year animals. Within the first year, that is to say from spring to
late fall, animals vary but little. Dental characters are best studied
in first year shews. For this reason I have used them as the basis for
the study of geographic variation, and descriptions are based on first
year animals unless otherwise noted.



CHARACTERS OF TAXONOMIC WORTH


Within the _Sorex vagrans_ complex, the only characters of taxonomic
significance that I have detected are in size and color. It is true
that cranial proportions, such as relative size of rostrum, may change
from population to population, but these proportions seem to me to be
dependent upon actual size of the individual shrew as I shall
elsewhere point out. Of the cranial measurements here employed,
palatal length and least interorbital breadth are the most significant
and useful. Color in the _S. vagrans_ group seems to be in Orange and
Cadmium Yellow, colors 15 and 17 of Ridgway (1912). No specimens
actually possess these pure colors, but most colors in these shrews
are seen to be derived from the two mentioned by admixture of black
and/or neutral gray. In color designations an increase in neutral gray
is indicated by an increased number of prime signs (´), whereas
increase in black is indicated by progressive characters of the Roman
alphabet (_i_, _k_, _m_). Thus, 17´´_k_ is grayer than 17´_k_ and
17´´_m_ is blacker than 17´´_k_. In subspecific diagnoses in this
report, color and size, and sometimes relative size, are the
characters usually mentioned.

   [Illustration: FIG. 3. Two measurements (in millimeters)
      reflecting tooth-wear plotted against one another. First year
      and second year individuals of _Sorex vagrans vagrans_, all
      taken in August at Willapa Bay, Washington, are completely
      separated. Open circles represent teeth of second year shrews;
      solid circles represent teeth of first year shrews.]



PELAGE CHANGE


In general, winter pelage is darker than summer pelage in these
shrews. Winter pelage comes in first on the rump and spreads caudad
and ventrad. The growth line of incoming hair is easily detected on
the fur side of the skin. Throughout the winter the color of the
pelage changes, often becoming somewhat browner, although no actual
molt takes place. This was noted by Dalquest (1944) who assumed that
the color change resulted from molt although he was unable to detect
actual replacement of hairs. Summer pelage usually comes in first on
the back or head and moves posteriorly and laterally. Time of molt
depends on latitude and altitude. Summer pelage may appear fairly late
in the season and may account for the anomalous midsummer molt noted
by Dalquest. Fresh pelages of summer and winter are best seen in first
year animals and are less variable than are worn pelages and hence are
used as the basis of color descriptions.



GEOGRAPHIC DISTRIBUTION AND VARIATION


Pacific Coastal Section

The largest shrews of the _vagrans_ group (large in all dimensions)
occur in the coastal forests of northern California and of Oregon.
Those shrews are reddish, large-skulled, large-toothed, and have
rostra that are large in proportion to the size of the skull as a
whole. The very largest of these shrews live along the coast of
northwestern California. To the southward they are somewhat smaller,
and at successively more northern localities, to as far as
southwestern British Columbia, they are likewise progressively smaller
and also somewhat less reddish. The relative size of the rostrum
decreases with the decrease in size of the skull; consequently smaller
shrews have relatively smaller rostra (see fig. 4). In addition the
zygomatic ridge of the squamosal decreases in relative size with
decrease in actual size of the skull. Thus, these features change in a
clinal fashion as one proceeds from, say, Humboldt County, California,
northward to Astoria, Oregon.

Turning our attention now farther inland to the Cascade Mountains of
northern Oregon, the shrews there also are smaller and less reddish
(more brownish) than in northwestern California, and the trend to
smaller and darker shrews culminates in the northern Cascades of
Washington. Shrews from there, and from the southwestern coast of
British Columbia, compared with those from northwestern California,
are much smaller and have so great a suffusion of black that they
appear brown rather than red. At places along the coast successively
farther north of southwestern British Columbia the shrews become
larger again, the largest individuals being those from near Wrangell,
Alaska. From that place northwesterly along the coast of Alaska, size
decreases again.

   [Illustration: FIG. 4. Condylobasal length (in millimeters)
      plotted against palatal index (palatal length/condylobasal
      length × 100) in several subspecies of _Sorex vagrans_ to show
      relative increase in size of rostrum with actual increase in
      size of skull.]

The shrews so far discussed inhabit forests in a region of high
rainfall and a minimum of seasonal fluctuation in temperature. Such a
habitat seems to be the optimum for shrews of the _vagrans_ group
since the largest individuals are found there. In addition, shrews
seem to be as common, or commoner, in this coastal belt, than they are
in other places.

The large shrews of the _vagrans_ group on the Pacific coast were
divided into three species by H. H. T. Jackson in his revision of the
North American _Sorex_ in 1928. The large reddish shrews of the coast
of California and southern Oregon were called _S. pacificus_. The
somewhat smaller ones from the coast of central Oregon were called _S.
yaquinae_. Still smaller shrews from northwestern Oregon and from the
rest of the Pacific coast north into Alaska were called _S. obscurus_.
I find these kinds to intergrade continuously one with the next in the
manner described and conclude that all are of a single species.


Inland Montane Section

Inland from the coasts of British Columbia and Alaska the size of the
_vagrans_ shrew decreases rapidly. Specimens from western Alaska,
central Alaska, and the interior of British Columbia are uniformly
smaller than coastal specimens. In addition the red of the hair is
masked more by neutral gray than by black with the result that the
pelage is grayish rather than brownish or reddish. Shrews of this
general appearance are found southward through the Rocky Mountain
chain to Colorado and New Mexico. On the more or less isolated
mountain ranges of Montana east of the continental divide the
_vagrans_ shrew is somewhat smaller still. On the Sacramento Mountains
of southeastern New Mexico the shrew is somewhat larger and slightly
darker. Southwestward from the Colorado Rockies this shrew becomes
smaller and slightly more reddish (less grayish).

All of these montane populations of the _vagrans_ shrew are commonest
in hydrosere communities, that is to say, streamsides and marshy areas
where the predominant vegetation is grass, sedges, willows, and
alders. Since these animals are less common within the montane
forests, hydrosere communities, rather than the actual forest, seem to
be the positive feature important for the shrews.

The shrews of the montane region just described were regarded by
Jackson as belonging to two species: _Sorex obscurus_, occupying all
the Rocky Mountains south to, and including, the Sacramento Mountains;
_S. vagrans_, made up of small individuals from various places in
Wyoming, Montana, and Colorado, and all the shrews of western New
Mexico and all of Arizona. My study of these animals has led me to
conclude that the smaller shrews of Arizona and New Mexico intergrade
in a clinal fashion with the shrews of Colorado and in fact represent
but one species. Since some individuals from Colorado are as small as
larger individuals from this southwestern population of small animals,
I conclude that such specimens are the basis for reports of _S.
vagrans_ from Colorado. The shrews of the Sacramento Mountains
resemble those of the Colorado Rockies more than they do the smaller
shrews of western New Mexico and Arizona, possibly because the climate
is similar in the Sacramento Mountains and the higher Colorado
Rockies. There is less precipitation in the more western mountain
ranges in New Mexico and in Arizona in April, May, and June than in
the Colorado Rockies. These months are critical for the reproduction
and growth of shrews.

As mentioned above, the shrews from east of the continental divide in
Montana are smaller than those of the other mountains of the state,
and it is upon such small animals that the name _Sorex vagrans_ has
been based in this area. It is clear, however, that these smaller
animals intergrade with the larger shrews of the more western
mountains. The small size might be an adaptation to the lesser
precipitation and harsher continental climate east of the continental
divide in Montana.


Great Basin and Columbia Plateau Section

The vagrant shrews of the Great Basin and adjoining Columbia Plateau
and Snake River Plains are smaller than their relatives in the Rocky
Mountains and, by virtue of less gray in their pelage, are reddish in
summer and blackish rather than grayish in winter. There is little
significant geographic variation in shrews throughout this region,
although owing to their restriction to the vicinity of water, the
populations of shrews are more or less isolated from one another and
each is somewhat different from the next. Those from nearest the
Rockies are sometimes slightly larger and those from some places in
Nevada are slightly paler than the average. This small reddish shrew
is found all the way to the Pacific coast of California, Oregon, and
Washington. In these coastal areas it is somewhat darker and sometimes
a trifle larger than elsewhere. It intergrades with a somewhat larger,
grayer shrew in the Sierra Nevada of California. Along the Wasatch
front in Utah, this Great Basin shrew intergrades with the larger,
grayer shrew of the Rockies. Owing to the abrupt change in elevation,
the zone of intergradation is rather narrow horizontally. In the
latitude of Salt Lake City, populations of intergrades occur at
between 8,700 and 9,000 feet elevation. The lowland shrew occurs in
the eastern part of the Snake River Plains, and along the valleys of
the Bear and Salt rivers into Wyoming. Along the northern edge of the
Snake River Plains and on the western edge of the mountains of central
Idaho the transition from lowland to montane habitats is abrupt and in
consequence the zone of contact between small and large shrews is
narrow. In northern Idaho and northwestern Montana the transition from
lowland to highland is more gradual. Tributaries of the Columbia River
system, especially the Clark Fork, provide a path for movement of
lowland forms into intermontane basins of western Montana. In
addition, the vegetational zones are found at lower elevations, and
there are boreal forests in the lowlands rather than only in the
mountains as is the case in Utah and Colorado. In this area,
therefore, the zone of intergradation between the smaller lowland
shrew and the larger montane shrew is more gradual and gradually
intergrading populations are found over a relatively large area. This
has been well demonstrated for northwestern Montana by Clothier
(1950). In southern British Columbia and northern Washington this
shrew in the mountains is large and in the intermontane valleys is
small. There is extensive interdigitation of valleys and mountain
ranges, and, consequently, of life-zones in this region. In a few
places, recognizably distinct populations of the vagrant shrew occur
within a few miles of one another, but in other places there are
populations of intergrades. West of the Cascades no evidence of
intergradation has been found and the two kinds occur almost side by
side and maintain their distinctness.

These Great Basin shrews dwell in hydrosere communities as do their
Rocky Mountain counterparts. In this arid region such a habitat
obviously is the only one habitable for a shrew of the _vagrans_
group. These shrews often maintain their predilection for such
habitats when they reach the Pacific coast, and are commonly found in
such places as coastal marshes, marshy meadows, and streamsides, while
the woodlands are inhabited by other species.

These small shrews of the Great Basin and the small vagrant shrews of
the Pacific Coast were called _Sorex vagrans_ by Jackson.


Summary of Geographic Variation

Large reddish shrews of the coast of California and southwestern
Oregon become smaller and darker to the north. From southwestern
British Columbia they again become larger as one proceeds northward
along the coast to Wrangell, Alaska, and north of that they again
become smaller. Moving inland from the coast the shrews become
markedly smaller in Alaska and British Columbia. The smaller inland
and montane form occurs south through the Rocky Mountains, becoming
slightly smaller in central Montana, slightly larger in southeastern
New Mexico, and slightly smaller in western New Mexico and in Arizona.
This montane form intergrades with a smaller more reddish Great Basin
shrew, the zone of intergradation roughly following the western slope
of the Rocky Mountains. The Great Basin shrew occurs westward to the
Pacific Coast; there the Great Basin shrew occurs with, although in
part it is ecologically separated from, the large reddish coastal
shrews.

There seems to be an intergrading chain of subspecies of one species,
the end members of which (the small Great Basin form and the large
coastal form) are so different in size and ecological niche that they
are able to coexist without interbreeding. In southern British
Columbia the morphological differences are not so marked as farther
south along the Pacific Coast. There, in British Columbia,
reproductive isolation is not complete and occasional populations of
intergrades occur. In Montana extensive intergradation occurs in a
broad zone of transitional habitat. Along the western edge of the
Rockies from Idaho south to Utah the zone of transition from montane
to basin habitat is sharp and the zone of intergradation, although
present, is fairly narrow, perhaps because there is little
intermediate habitat which logically might be expected to be most
suitable for intergrading populations.

The oldest name applied to a shrew of the group under consideration is
_Sorex vagrans_ Baird, 1858, the type locality of which is Willapa
Bay, Pacific County, Washington. The name applies to the small vagrant
shrew of this area, rather than to the larger forest dweller which has
been known as _Sorex obscurus_. The name _S. vagrans_, in the specific
sense, must therefore apply to all the shrews discussed which have
heretofore been known by the names _S. pacificus_, _S. yaquinae_, _S.
obscurus_, and _S. vagrans_.

A situation such as the one here described where well differentiated
end members of a chain of subspecies overlap over an extensive
geographic range throughout the year without interbreeding--thus
reacting toward one another as do full species--so far as I know has
not previously been found to exist in mammals. The overlapping
end-members of the chain of subspecies of _Sorex vagrans_ really do
coexist; specimens of the overlapping subspecies have been taken
together at the same localities from California to British Columbia.
I have taken a specimen of _S. v. vagrans_ and several of _S. v.
setosus_ in the same woodlot at Fort Lewis, Pierce County, Washington.
Two subspecies of deer, _Odocoileus hemionus_, in the Sierra Nevada of
California, occur together over a sizeable area but for only a part of
each year that does not include the breeding season (Cowan,
1936:156-157). In the deer mouse, _Peromyscus maniculatus_, the
geographic ranges of several pairs of subspecies meet at certain
places without intergradation of the two kinds. In these instances
well marked ecological differences exist between the subspecies
involved. In western Washington, for example, the geographic range of
the lowland subspecies, _P. m. austerus_, interdigitates to the
east and west with the range of the montane and coniferous
forest-inhabiting subspecies, _P. m. oreas_, and the two kinds have
not been shown to intergrade. _Peromyscus maniculatus artemesiae_ and
_P. m. osgoodi_ come together without interbreeding in Glacier
National Park, Montana. _P. m. artemesiae_ is almost entirely a
forest-dwelling subspecies, whereas _osgoodi_ is an inhabitant of open
country. The two kinds do not actually occur together ecologically
although they occur together in buildings at the edge of the woods
(A. Murie, 1933:4-5).

   [Illustration: FIG. 5. Probable present geographic distribution
      of _Sorex vagrans_. The range of _S. v. vagrans_ and its
      derivatives _S. v. vancouverensis_, _S. v. halicoetes_, and
      _S. v. paludivagus_, is shown by lines slanting in a different
      direction than those which mark the range of all the other
      subspecies of _S. vagrans_. The region in which _S. v. vagrans_
      occurs together with other subspecies of _S. vagrans_ is shown
      by the superposition of one pattern upon the other.]

Cases of sympatric existence of two subspecies of one species are
known in birds and in reptiles. Notable examples are in the gull,
_Larus argentatus_ (Mayr, 1940), in the Old World warbler,
_Phylloscopus trochiloides_ (Ticehurst, 1938), and in the great
titmouse, _Parus major_ (Rensch, 1933), of the Old World. In the first
species the two end-members, the herring gull and the lesser
black-backed gull, occur together over an extensive region from
northern Europe and the British Isles throughout Fennoscandia. Fitch
(1940) described a rassenkreis with overlapping subspecies in the
garter snake _Thamnophis ordinoides_.

The geographic distribution of the species _Sorex vagrans_ is shown in
figure 5. The geographic range of the Great Basin subspecies is shown
by a different pattern of lines than the other subspecies of _S.
vagrans_. In the region in which the geographic range of the Great
Basin subspecies overlaps those of the subspecies of the Pacific
Coast, the pattern of shading for the Great Basin subspecies is
superimposed on the patterns for the other subspecies.



ORIGIN OF THE _SOREX VAGRANS_ RASSENKREIS


The distribution of the species _Sorex vagrans_ and that of its
immediate ancestors obviously has not always been the same; during
glacial ages much of the present range of the species in Canada and in
some of the higher mountains of the United States was covered with ice
and not available to the shrew. Furthermore, large areas that are now
too hot and dry to permit the existence of_ S. vagrans_ were at one
time habitable. If we are to speculate on the manner in which the
_Sorex vagrans_ rassenkreis originated we must inquire into the nature
and extent of these climatic changes.

The most recent epoch of geological time, the Pleistocene, is known to
have been divided into a series of alternating glacial and
interglacial ages. During the glacial ages continental and montane
glaciers are judged to have covered much of Canada and the northern
United States. Concurrently the major storm tracks of the west
probably were shifted southward; in any event much of the now arid
intermontane west was much better watered than it is today.

The increased precipitation, and probably glacial meltwater, formed
large lakes in the closed basins of the Great Basin. There were boreal
forests at lower elevations than there are today in comparable
latitudes and continuous boreal habitat probably connected many of the
isolated mountain ranges of the southwest. That probability is
supported by the presence of boreal animals and plants on many of
these isolated ranges today. A boreal tree squirrel, such as
_Tamiasciurus_, could hardly be suspected of crossing a treeless,
intermontane desert valley, miles wide.

   [Illustration: FIGS. 6_a_-6_f_. Fig. 6_a_. _Sorex vagrans
      pacificus_, 1 mi. N Trinidad, Humboldt Co., California,
      FC 1442. Fig. 6_b_. _S. v. yaquinae_, Newport, Lincoln Co.,
      Oregon, AW 707. Fig. 6_c_. _S. v. yaquinae_ (near _bairdi_),
      McKenzie Bridge, Lane Co., Oregon, AW 82. Fig. 6_d_. _S. v.
      setosus_, Reflection Lake, Jefferson Co., Washington, CMNH
      4275. Fig. 6_e_. _S. v. obscurus_, 10 mi. SSW Leadore, Lemhi
      Co., Idaho, FC 1499. Fig. 6_f_. _S. v. vagrans_, Baker Creek,
      White Pine Co., Nevada, 88042 (after Hall, 1946:113).]

Interglacial ages were characterized by warmth and aridity as compared
to the glacial ages. Glaciers retreated or disappeared, boreal forests
became montane in much of the United States, and the lakes in the
Great Basin were reduced or disappeared. One can envision that during
such times boreal mammals were isolated, their geographic ranges were
restricted, and Sonoran mammals expanded their ranges.

Evidence is more extensive concerning the number and extent of glacial
ages in the eastern than in the western part of North America. This
evidence suggests a division of the Pleistocene into four glacial ages
and four interglacial ages, the fourth interglacial age corresponding
to the present time. More information is available about the
Wisconsinan, or last, glacial age, than about the earlier ones,
because the last glaciation in many montane areas destroyed evidence
of earlier glaciations. The names of currently recognized glacial and
interglacial ages of the Pleistocene are listed below. The names of
interglacial ages are in Italic type.

    Wisconsinan
    _Sangamonian_
    Illinoian
    _Yarmouthian_
    Kansan
    _Aftonian_
    Nebraskan

We may think of these ages as an alternating series of cool moist and
warm dry periods during which boreal mammals, and other organisms,
alternately moved southward (disappearing in the glaciated regions)
and northward into previously glaciated areas (while disappearing from
southern areas except on isolated mountain ranges). _Sorex vagrans_
probably followed this pattern of movement and now is restricted to
forested or well-watered places.

One possible series of events culminating in the formation of the
_Sorex vagrans_ rassenkreis may be thought of as having begun during
the Illinoian age. With much of Canada, and perhaps also many areas in
the Rockies, Cascades, and the Sierra Nevada covered with glacial ice,
the shrew-stock ancestral to _Sorex vagrans_ may well have occupied a
more or less continuous range over the Colorado Plateau, the Columbian
Plateau, the Great Basin, and in the forests of the Pacific Coast (as
well as over part of eastern United States, as will be explained
beyond; see fig. 7). At that time the species probably was a
continuously interbreeding unit.

   [Illustration: 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.

   [Illustration: 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.

   [Illustration: FIG. 9. Probable changes in the distribution of
      _Sorex vagrans_ concurrent with and following the dissipation
      of Wisconsinan ice. Dark arrows in Washington, Idaho, Oregon,
      and California, shows_ S. v. vagrans_.]

Deglaciation of the Sierra Nevada opened it up for reoccupation from
the east by _Sorex vagrans_ of the Great Basin. In response to the
montane environment the subspecies _obscuroides_, resembling the
subspecies _obscurus_ of the Rockies, developed.

Desiccation of the intermontane parts of New Mexico, Arizona, and
Chihuahua, left "marooned" populations of _Sorex vagrans_ on suitable
mountain ranges. In this way _Sorex vagrans orizabae_ may have been
isolated in southern Mexico. The isolated populations of Arizona and
New Mexico differentiated _in situ_ into the subspecies _monticola_
and _neomexicanus_.

Western Canada and Alaska were populated by shrews which originated in
the habitable parts of the Rocky Mountains and Colorado Plateau during
Wisconsinan time (as opposed to shrews originating, as subspecies, in
the Great Basin or on the Pacific Coast). These shrews differentiated
into the currently recognized subspecies of the west coast and coastal
islands of British Columbia and Alaska in response to the different
environments in these places, many of which were isolated; the
subspecies _isolatus_, _mixtus_, _setosus_, _longicauda_, _elassodon_,
_prevostensis_, _malitiosus_, and _alaskensis_ are thought to have
originated in this fashion after the areas now occupied by them were
freed of Wisconsinan ice.

This group of shrews from the Rocky Mountains probably came into
contact with the Pacific coastal segment of the species somewhere in
northwestern Oregon. The clinal decrease in size from _S. v.
pacificus_ to _S. v. setosus_ seems steepest in this area. Upon the
establishment of this contact reproductive continuity was resumed,
probably because the temporal separation of the two stocks involved
was not so great as, say, that between _S. v. vagrans_ and _S. v.
pacificus_, and in addition the morphological differentiation was not
so great.

On the eastern side of the Rockies the montane stock moved
northeastward, occupying suitable territory opened up by the
dissolution of the Laurentide ice sheet. Still later changes in the
character of the northern plains owing to desiccation divided the
range of the species and isolated _S. v. soperi_ in Manitoba and
central Saskatchewan and a population of _S. v. obscurus_, in the
Cypress Hills. A number of semi-isolated stocks in central Montana
became differentiated as a recognizable subspecies there.

A number of other boreal mammals have geographic ranges which resemble
that of _Sorex vagrans_, except that the geographic ranges of
subspecies do not overlap. Because of the general similarities of
these geographic ranges, it is pertinent to examine the reasons
suggested by students to account for the present geographic
distributions of some of these other boreal species.

The red squirrel genus, _Tamiasciurus_, has a Rocky Mountain (and
northern coniferous forest) species, _T. hudsonicus_, that occurs all
along the Rocky Mountain chain and northward into Alaska. In the
Cascade Mountains of Washington and British Columbia this species
meets the range of a well marked western species, _T. douglasii_, with
no evidence of intergradation. Dalquest (1948:86) attributes the
divergence of the two species to separation in a glacial age but feels
that the degree of difference between the two is too great to have all
taken place during the Wisconsinan. Perhaps he has overemphasized the
importance of the differences between the two, but, be that as it may,
it seems that the two kinds differentiated during a glacial age when
they were isolated, perhaps by ice on the Cascades into a coastal
population and an inland population. One difference between the
distribution of the red squirrels and vagrant shrew is that the
squirrel of the Sierra Nevada is the species of the Pacific Coast,
whereas the vagrant shrew of the Sierra Nevada was derived from the
Great Basin population, which in turn was derived from the Rocky
Mountain kind. Red squirrels do not occur on any of the boreal montane
"islands" of Nevada. During the pluvial periods when hydrosere-loving
shrews populated the Great Basin, that region may have been a treeless
grassland. Vagrant shrews, then as now, probably depended on hydrosere
communities, while red squirrels required trees. Therefore the shrews
were able to traverse the Great Basin, while the Sierran red squirrels
were of necessity derived from the coastal population.

The ecological requirements of jumping mice, genus _Zapus_, and the
subspecies of _Sorex vagrans_ that dwell in hydroseres are essentially
similar. The species _Zapus princeps_ lives in the Rocky Mountains,
the Great Basin, the Sierra Nevada, and north to Yukon (Krutzsch,
1954:395). Its geographic range is similar to that of the montane and
basin segments of _S. vagrans_. The species _Z. trinotatus_ occurs
along the Pacific coast and in the Cascades north to southwestern
British Columbia. Its distribution thus coincides in general with that
of the large red coastal subspecies of _S. vagrans_. Krutzsch
(1954:368-369) thought that these two kinds of jumping mice were first
separated by the formation of the Cascade Mountains and the Sierra
Nevada and finally by Pleistocene glaciation. The Sierran jumping
mouse (_Zapus princeps_), as is the Sierran vagrant shrew, is more
closely related to the jumping mouse of the Great Basin and of the
Rocky Mountains than it is to the jumping mouse (_Z. trinotatus_) of
the Pacific Coast, just as the Sierran vagrant shrew is related to the
shrew of the Great Basin and Rocky Mountains. The jumping mouse also
is limited in its distribution by hydrosere communities, not by
forests.

In western North America there are two species of water or marsh
shrews: _Sorex palustris_ and _S. bendiri_. They have been placed in
separate subgenera, but, as pointed out beyond, are closely related
and here are placed in the same subgenus. The species _palustris_ is
found throughout the Rocky Mountains, north into Alaska, across the
Great Basin into the Sierra Nevada, and west to the Pacific coast in
Washington. The species _bendiri_ is found from northwestern
California north along the Pacific coast to southwestern British
Columbia and east to the Cascades. Where the ranges of the two species
overlap in western Washington they do not interbreed so far as is
known, and are somewhat different in their ecology, _bendiri_ being a
lowland, and _palustris_ being a montane, species. The two species
probably were separated in a glacial period as seems to have been the
case with the wandering shrews. Also, the water shrew of the Sierra
Nevada is derived from that of the Great Basin and Rocky Mountains.
_Sorex palustris_ is tied closely in its distribution to hydrosere
communities and is not dependent upon the presence of forests.

Red-backed mice, genus _Clethrionomys_, occur throughout the Rocky
Mountains and west to the Cascades in Washington as the species _C.
gapperi_. The species _C. californicus_ is found along the Pacific
Coast from California north to the Olympic Peninsula. Where the ranges
of the two species meet in Washington they seem not to intergrade. In
some glacial interval these two species may have evolved in the same
manner as has been described for the species of _Zapus_ and those of
_Tamiasciurus_. No _Clethrionomys_ are found in the Sierra Nevada, nor
are red-backed mice found in the boreal islands of the Great Basin. It
is not known why _Clethrionomys californicus_ does not occur in the
Sierra Nevada. Some boreal birds have distributional patterns similar
to those of the mammalian examples cited above. One kind of sapsucker,
_Sphyrapicus varius nuchalis_, occurs in the Rocky Mountains north
into British Columbia and west to the Cascades and Sierra Nevada. A
related kind, _S. varius ruber_, occurs along the Pacific Coast from
California north into British Columbia. Recently Howell (1952) has
shown that some intergradation takes place between _ruber_ and
_nuchalis_ in Washington and British Columbia, although they do not
intergrade freely. Previously the two kinds were thought not to
intergrade and were regarded as two species. The two kinds intergrade
also in northeastern California, although in that state _S. v.
daggeti_, rather than _S. v. ruber_, is involved in the
intergradation. Howell considered the two kinds to be conspecific with
one another as well as with the eastern _S. varius_. He attributed a
measure of the distinctness of _nuchalis_ and _ruber_ to their
separation during a glacial period, but felt that the separation was
much older than Wisconsinan. Whatever the time of separation, the
pattern seems clear: _nuchalis_ and _ruber_ (as well as _varius_) were
separated into montane, coastal, and eastern segments respectively,
probably by glaciation (it seems to me in the Pleistocene), and have
since re-established contact with one another.

The grouse genus _Dendrogapus_ is divided into a Great Basin species,
_D. obscurus_, which extends northward into British Columbia, and a
Rocky Mountain species, _D. fuliginosus_, that is found in the Sierra
Nevada and northward along the coast and Cascades into British
Columbia. Although the two kinds have at times been considered
conspecific, they differ in voice, hooting mechanism, and characters
of the downy young, and so far no actual intergradation between the
two has been shown (Grinnell and Miller, 1944:113). These grouse thus
seem to offer additional evidence for a Pleistocene, possibly
Wisconsinan, separation of the boreal fauna into a Rocky Mountain and
a Pacific coastal segment.

A notable sidelight on these data is the frequency with which species
in the Sierra Nevada have their closest relatives in the Rocky
Mountains, rather than in the geographically nearer Cascades or
coastal areas. This similarity in fauna of the Sierra Nevada and the
Rockies was noted long ago by Merriam (1899:86).



RELATIONSHIPS WITH OTHER SPECIES


During the Sangamonian interval, isolated segments of the once
widespread ancestral _Sorex vagrans_ quite possibly persisted in such
places as the Sierra Nevada, coastal southern California, the
mountains of Arizona, New Mexico, and southern Mexico, and in the
Black Hills (see fig. 6). One might expect that by Wisconsinan time
these populations would have become reproductively isolated from their
parent stock. They would therefore have remained specifically distinct
when Wisconsinan _Sorex vagrans_, reoccupied these outlying areas, and
may still be found isolated in places peripheral to the range of the
ancestral species.

   [Illustration: FIG. 10. Probable distribution of
      _S. veraepacis_, _S. longirostris_, and the _S. ornatus_ group
      (stipple) and of their Wisconsinan ancestors (lines). Heavy
      line indicates limits of Wisconsinan glaciation.]

In fact, we do find species closely related to _Sorex vagrans_ in just
such places today (fig. 10). Probably _Sorex ornatus_, including
members of the _ornatus_ group such as _S. trigonirostris_, _S.
sinuosus_, _S. willeti_, _S. tenellus_, and _S. nanus_, and also _S.
veraepacis_, arose by separation from the ancestral _vagrans_ stock in
Sangamonian time. Probably the eastern _S. longirostris_ arose in a
like manner. The ancestor of _S. ornatus_ may have been isolated in
southwestern California during Sangamonian time, spread north and
south during the Wisconsinan age, and afterward given rise to _S.
trigonirostris_ and the modern _S. ornatus_ complex of California and
Baja California. In at least one place reproductive isolation between
_ornatus_ and the invading _S. vagrans_ has broken down (Rudd, 1953);
the place is a salt marsh along San Pablo Bay, where a hybrid
population between _S. vagrans_ and _S. sinuosus_, an _ornatus_
derivative, has formed. _Sorex tenellus_ may have been isolated in the
Sierra Nevada in the Sangamonian interval, moved into the valleys
east of the mountains during the Wisconsinan age, and become
restricted to its present range since the retreat of the last ice.
_Sorex nanus_ may have occurred in the Black Hills and isolated
mountains of Arizona and New Mexico during the Sangamonian interval
and remained in these general areas during the Wisconsinan age. Its
present range is peripheral to the main body of the Rockies and the
Colorado Plateau.

The eastern species _Sorex longirostris_ has many similarities with
shrews of the _ornatus-vagrans_ stock. _S. l. longirostris_ is close
in many ways to _S. nanus_. Indeed, the differences between the
species _S. nanus_, _S. ornatus_, and _S. longirostris_ seem to me to
be of the same magnitude and indicate a similar period of
differentiation from a common ancestor. The ancestor of _S.
longirostris_ may have gained access to the eastern United States in
the Illinoian Age _via_ the northern Great Plains south of the glacial
boundary (fig. 7). The ancestor of _Sorex veraepacis_ of southern
Mexico probably reached that area in Illinoian time as part of the
ancestral _vagrans_ stock and probably attained its differentiation
during the Sangamonian interval.

All the kinds of shrews so far discussed, including the _S. vagrans_
complex, might thus be thought of as having had a common ancestor in
the Illinoian Age. This entire group of shrews has the third unicuspid
smaller than the fourth, a pigmented ridge from the apex to the
cingulum of each upper unicuspid, and, in most individuals, lacks a
post-mandibular foramen in the lower jaw (Findley, 1953:636-637). The
pigment is not always prominent in _S. longirostris_.

Two other species of North American shrews,_ Sorex palustris_, the
water shrew, and _Sorex bendiri_, the marsh shrew, show these three
characters to a greater or lesser degree, and it seems that these two
species and the _vagrans-ornatus-veraepacis_ group had a common
ancestor, probably before Illinoian time for reasons stated beyond. I
judge, however, that far from being subgenerically distinct as they
have been considered to be, _S. palustris_ and _S. bendiri_ are
actually closely related species of the same subgenus and may have
differentiated from one another because of separation into eastern
(_palustris_) and western (_bendiri_) segments in the Sangamonian
interval, much as has been postulated concerning the eastern and
western stocks of _Sorex vagrans_. Indeed, Jackson (1928:192) has
noted that in the Pacific northwest the characters of the two kinds
approach one another and become differences of degree only.

The widespread species _Sorex cinereus_ resembles all the foregoing
species in the ridges on the unicuspid teeth and in the lack of a
post-mandibular foramen, but differs from those other species in
having the third upper unicuspid larger than the fourth. The
subspecies _S. cinereus ohionensis_, however, often has the sizes of
these teeth reversed. With _S. cinereus_ I include _S. preblei_
(eastern Oregon) and _S. lyelli_ (Sierra Nevada), both obviously
closely related to _cinereus_ as Jackson (1928:37) recognized when he
included them in the _cinereus_ group. _Sorex milleri_ (Coahuila and
central western Nuevo Leon) seems to me to resemble _S. cinereus_ more
than it does other species of North American _Sorex_, and I judge that
it also belongs to the _cinereus_ group. _Sorex cinereus_ and its
close relatives seem more closely related to the species which have
thus far been discussed than they do to such other North American
species as _S. arcticus_, _S. fumeus_, _S. trowbridgii_, _S. merriami_,
and the members of the _S. saussurei_ group; most of these five
species last mentioned possess a post-mandibular foramen, lack
pigmented unicuspid ridges, and have the third unicuspid larger than
the fourth. Because of the morphological resemblances mentioned
above, it seems likely to me that _S. cinereus_ and the
_vagrans-ornatus-veraepacis-palustris_ complex had a common ancestor
in early Pleistocene time. _Sorex cinereus_ has recently been
considered to be conspecific with the Old World_ S. caecutiens_
Laxmann (Van den Brink, 1953) which name, being the older, would apply
to the circumpolar species.

Hibbard (1944:719) recovered _S. cinereus_ and a species of _Neosorex_
(a name formerly applied to the water shrew) from the Pleistocene
(late Kansan) Cudahy Fauna. This indicates that the ancestors of the
modern _S. cinereus_ and of the water shrew had diverged from one
another before that time. Brown (1908:172) recorded _S. cinereus_ and
_S. obscurus_ from the Conard Fissure in Arkansas. These materials
were deposited probably at a later time than was the Cudahy Fauna. The
_S. obscurus_ from Conard Fissure probably represents the ancestral
_S. vagrans_ stock which I think reached eastern United States in
Illinoian time and gave rise to _S. longirostris_. The Conard Fissure
material was deposited at a time (Illinoian?) when northern faunas
extended farther south than they do today.

All of the species mentioned as having structural characters in common
with _S. vagrans_ seem to have arisen from a common ancestor which had
already differentiated from the ancestor of such species as _S.
arcticus_, _S. saussurei_, and others. Consequently all are here
included in a single subgenus. The oldest generic name applied to a
shrew of this group, other than the name _Sorex_, is _Otisorex_ DeKay,
1842, type species _Otisorex platyrhinus_ DeKay, a synonym of _Sorex
cinereus_. The subgenus can be characterized as follows.


Subgenus =Otisorex= DeKay

1842. _Otisorex_ DeKay, Zoology of New York, pt. 1, Mammalia, p. 22,
and pl. 5, fig. 1. Type, _Otisorex platyrhinus_ DeKay (= _Sorex
cinereus_ Kerr).

Third unicuspid usually smaller than fourth; upper unicuspids usually
with pigmented ridge extending from apices medially to cingula,
uninterrupted by antero-posterior groove; post-mandibular foramen
usually absent. Includes the species _S. cinereus, S. longirostris, S.
vagrans, S. ornatus, S. tenellus, S. trigonirostris, S. nanus, S.
juncensis, S. willeti, S. sinuosus, S. veraepacis, S. palustris, S.
bendiri, S. alaskanus_, and _S. pribilofensis_.

   [Illustration: FIGS. 11-14. Characters of the subgenera _Sorex_
      and _Otisorex_.

   FIG. 11. Medial view of right ramus of _Sorex (Otisorex)
      vagrans_. × 14.

   FIG. 12. Medial view of right ramus of _Sorex (Sorex)
      arcticus_. × 14.

   FIG. 13. Anterior view of left second upper unicuspid of _Sorex
      (Otisorex) vagrans_. × 45.

   FIG. 14. Anterior view of left second upper unicuspid of _Sorex
      (Sorex) arcticus_. × 45.]

Other species of _Sorex_ now occurring in North America differ from
_Otisorex_ in having the 3rd unicuspid usually larger than 4th, in
lacking a pigmented ridge from the apices to the cingula of the upper
unicuspids, and in usually possessing a well-developed post-mandibular
foramen. Exceptions to the last mentioned character are _S. fumeus_
and _S. dispar_. The subgenus _Sorex_ in North America should include
only the following species: _S. jacksoni_, _S. tundrensis_, _S.
arcticus_, _S. gaspensis_, _S. dispar_, _S. fumeus_, _S. trowbridgii_,
_S. merriami_, and all the members of the Mexican _S. saussurei_
group.

The subgenera _Otisorex_ and _Sorex_ probably separated in early
Pleistocene or late Pliocene. _Sorex_ is unknown in North America
earlier than the late Pliocene (Simpson, 1945:51).

In the genus _Microsorex_ the characters of the subgenus _Otisorex_
are carried to an extreme; the unicuspid ridges are prominent and end
in distinct cusplets, and the 3rd unicuspid is not merely smaller than
the 4th, but is reduced almost to the vanishing point. In addition,
the post-mandibular foramen is absent. Although it is closer
structurally to _Otisorex_ than to _Sorex_, the recognition of
_Microsorex_ as a distinct genus seems warranted.

Figure 15 is intended to represent graphically some of the
relationships discussed above. It must be re-emphasized that much of
it is purely speculative, especially as regards actual time when
various separations took place. It will be noted that I have indicated
most separations as having taken place in interglacial ages. They are
generally regarded as periods of warmth and aridity and, therefore,
probably are times of segmentation of the ranges of boreal mammals and
hence times exceptionally favorable to the process of speciation.
Glacial ages, characterized by extensive and continuous areas of
boreal habitat, probably were times of relatively unrestricted gene
flow between many populations of boreal mammals and hence not
favorable to rapid speciation.


=Sorex vagrans=

Wandering Shrew

The size of the wandering shrew varies from small in the subspecies
_monticola_ and _vagrans_ to large in the subspecies _pacificus_. The
tail makes up from a little more than a third to almost half of the
total length. The color pattern ranges from tricolored through
bicolored to almost monocolored. Color ranges from reddish (Sayal or
Snuff Brown) to grayish in summer pelage and from black to light gray
in winter. Diagnostic dental characters include: 3rd upper unicuspid
smaller than 4th, and unicuspids, except 5th, with a pigmented ridge
extending from near apex of each tooth medially to cingulum and
sometimes ending as internal cusplet. _S. vagrans_ differs from
members of the _ornatus_ group in less flattened skull, and in more
ventrally situated foramen magnum that encroaches more on the
basioccipital and less on the supraoccipital. The wandering shrew
differs from _S. trowbridgii_ and _S. saussurei_ in the dental
characters mentioned above. These dental characters also serve to
distinguish _S. vagrans_ readily from _S. cinereus_, _S. merriami_,
and _S. arcticus_ which may occur with _vagrans_. The large marsh
shrew and water shrew, _S. palustris_ and _S. bendiri_, can be
distinguished at a glance from _S. vagrans_ by larger size and darker
color.

   [Illustration: FIG. 15. Diagrammatic representation of the
      probable phylogeny of _Sorex vagrans_ and its near relatives.]

In the following treatment of the 29 subspecies of _Sorex vagrans_,
the subspecies are arranged in geographic sequence, beginning with the
southernmost large subspecies on the California coast and proceeding
clockwise, north, east, south, and then west back to the starting
point.


=Sorex vagrans sonomae= Jackson

    _Sorex pacificus sonomae_ Jackson, Jour. Mamm., 2:162, August
    19, 1921.

_Type._--Adult female, skin and skull; No. 19658, Mus. Vert.
Zool.; obtained on July 2, 1913, by Alfred C. Shelton, from
Gualala, on the Sonoma County side of the Gualala River, Sonoma
Co., California.

_Range._--Coastal California from Point Reyes north to Point Arena.

_Diagnosis._--Size large; average and extreme measurements of 3
topotypes are: total length, 141.7 (141-143); tail, 59 (54-63); hind
foot, 17 (17-17). Color reddish in summer, somewhat grayer in winter.

   [Illustration: FIG. 16. Probable geographic ranges of 16
      subspecies of _Sorex vagrans_.

   Guide to subspecies

     1. _S. v. shumaginensis_
     2. _S. v. obscurus_
     3. _S. v. alascensis_
     4. _S. v. soperi_
     5. _S. v. isolatus_
     6. _S. v. setosus_
     7. _S. v. bairdi_
     8. _S. v. permiliensis_
     9. _S. v. yaquinae_
    10. _S. v. pacificus_
    11. _S. v. sonomae_
    12. _S. v. longiquus_
    13. _S. v. parvidens_
    14. _S. v. monticola_
    15. _S. v. neomexicanus_
    16. _S. v. orizabae_
   ]

_Comparisons._--Differs from _S. v. pacificus_, with which it
intergrades to the north, in average smaller size and somewhat darker
color; differs from the sympatric _S. v. vagrans_ in much larger size
and more reddish color in both summer and winter.

_Remarks._--This subspecies inhabits the Transition Life-zone below
300 feet, and occurs on moist ground in forests and beneath dense
vegetation.

_Marginal records._--CALIFORNIA: Point Arena (Grinnell, 1933:82);
Monte Rio (Jackson, 1928:144); Inverness (Grinnell, 1933:82).


=Sorex vagrans pacificus= Coues

    _Sorex pacificus_ Coues, Bull. U. S. Geol. and Geog. Surv.
    Terr., 3 (3):650, May 15, 1877.

    _Sorex pacificus pacificus_, Jackson, Jour. Mamm., 2:162,
    August 19, 1921.

_Type._--Adult, sex unknown, skin and skull; No. 3266 U. S. Nat. Mus.;
date of capture unknown; received from E. P. Vollum and catalogued on
March 8, 1858; obtained at Ft. Umpqua, mouth of Umpqua River, Douglas
Co., Oregon.

_Range._--Coast of California and Oregon from Mendocino north to
Gardiner.

_Diagnosis._--Size large, largest of the species; average and extreme
measurements of 8 specimens from Orick, Humboldt Co., California, are:
total length, 143.1 (134-154); tail, 65.5 (59-72); hind foot, 17.5
(16-19). Color reddish in summer, browner or grayer in winter.

_Comparisons._--See account of _S. v. sonomae_ for comparison with
that subspecies; averaging larger in all dimensions than _S. v.
yaquinae_ with which it intergrades to the north; much larger and has
more reddish than the sympatric _S. v. vagrans_.

_Remarks._--This subspecies occurs in the Canadian and Transition
life-zones below 1500 ft. where there is found moist ground in or
adjacent to heavy forests.

_Specimens examined._--Total number, 76.

OREGON: _Douglas Co._: Umpqua, 1 BS. _Coos Co._: Marshfield, 1 BS;
Myrtle Point, 1 BS. _Josephine Co._: Bolan Lake, 1 SGJ.

CALIFORNIA: _Del Norte Co._: Smith River, 2 BS; Gasquet, 4 BS;
Crescent City, 17 BS. _Humboldt Co._: Orick, 13 BS; 1 mi. N Trinidad,
18 FC; Trinidad Head, 1 BS; Carson's Camp, Mad River, Humboldt Bay, 5
BS; Arcata, 3 BS; Cape Mendocino, 2 BS; 5 mi. S Dyerville, 1 BS.
_Mendocino Co._: Mendocino, 6 BS.

_Marginal Records._--OREGON: Marshfield; Umpqua. CALIFORNIA: Gasquet;
5 mi. S Dyerville; Mendocino, thence up coast to point of beginning.


=Sorex vagrans yaquinae= Jackson

    _Sorex yaquinae_ Jackson, Proc. Biol. Soc. Washington, 31:127,
    November 29, 1918.

    _Sorex pacificus yaquinae_, V. Bailey, N. Amer. Fauna, 55:364,
    August 29, 1936.

_Type._--Adult female, skin and skull; No. 73051 U. S. Biol. Surv.
Coll., obtained on July 18, 1895, by B. J. Bretherton, from Yaquina
Bay, Lincoln Co., Oregon.

_Diagnosis._--Size large for the species; average and extreme external
measurements of 11 specimens from Oakridge, Lane Co., Oregon, are:
total length, 125.3 (11-136); tail, 55.1 (49-61); hind foot, 14.9
(14-16). Color reddish in summer, browner or grayer in winter.

_Comparisons._--See account of _S. v. pacificus_ for comparison with
that subspecies. Larger and more reddish than _S. v. bairdi_ with
which it intergrades to the north and east. Much larger and more
reddish than the sympatric _S. v. vagrans_.

_Remarks._--The name _yaquinae_ actually applies to a population of
intergrades between _pacificus_ and _bairdi_. There is much variation
over the range of the subspecies, and individuals from the western and
southern parts are larger than those from the west slope of the
Cascades. Specimens from Vida and McKenzie Bridge are smaller than
those from Mapleton, Mercer, and the type locality but still seem
closer to _yaquinae_ than to topotypes of _bairdi_. Between Marshfield
and Umpqua on the one hand, and the Columbia River and the Cascade
Mountains on the other, the size of _Sorex vagrans_ decreases quite
rapidly from the large_ pacificus_ to the smaller _permiliensis_. Size
decreases less rapidly northward along the coast than it does eastward
toward the mountains; consequently, at any given latitude, coastal
shrews are larger than mountain shrews. In this area of rapid change
in size it is difficult to draw subspecific boundaries between
_pacificus_, _yaquinae_, and _bairdi_, and this must be done somewhat
arbitrarily.

Jackson (1928:141) remarked upon the possibility that intergradation
between _pacificus_ and _yaquinae_ took place. He noted also the close
resemblance between _yaquinae_ and _bairdi_, and stated (_loc. cit._)
that specific affinity between the two might be demonstrated with more
specimens. He had a series of eight specimens from Vida, Oregon, seven
of which he assigned to _S. o. bairdi_ and one to _yaquinae_. I have
examined these specimens and find no more variation between the
largest and the smallest than would be expected in any normally
variable series of shrews. Vernon Bailey (1936:364) arranged
_yaquinae_ as a subspecies of _pacificus_ without giving his reasons
for so doing.

_Specimens examined._--Total number, 65. OREGON: _Lincoln Co._: type
locality, 2 AW. _Benton Co._: Philomath, 2 BS. _Lane Co._: Mable, 1
OU; Vida, 4 BS, 1 OSC, 3 OU; McKenzie Bridge, 8 OSC, 3 AW, 17 OU, 2
SGJ; Mercer, 1 OSC, 1 OU; Mapleton, 3 BS; Oakridge, 11 OU. _Douglas
Co._: Gardiner, 2 BS; Elkhead, 1 BS. _Klamath Co._: Crescent Lake, 3
OU.

_Marginal Records._--OREGON: Yaquina Bay; _Philomath_; McKenzie
Bridge; Prospect (Jackson, 1928:140); Crescent Lake; Gardiner.


=Sorex vagrans bairdi= Merriam

    _Sorex bairdi_ Merriam, N. Amer. Fauna, 10:77, December 31,
    1895.

    _Sorex obscurus bairdi_, Jackson, Proc. Biol. Soc. Washington,
    31:127, November 29, 1918.

_Type._--Adult female, skin and skull; No. 17414/24318, U. S. Biol.
Surv. Coll.; obtained on August 2, 1889, by T. S. Palmer, from
Astoria, Clatsop Co., Oregon.

_Range._--Northwestern Oregon, south to Otis and east to Portland.

_Diagnosis._--Size medium for the species; average and extreme
external measurements of 6 specimens from the type locality are: total
length, 126.3 (124-130); tail, 55.0 (52-57); hind foot, 15.0 (14-15).
Color Fuscous to Sepia in summer, darker in winter, underparts buffy.

_Comparisons._--For comparisons with _yaquinae_ see account of that
subspecies. More reddish and larger than _permiliensis_ with which
_bairdi_ intergrades to the east; specimens from Portland show
evidence of such intergradation. Some specimens from southern
Tillamook County show an approach to _yaquinae_.

_Remarks._--_S. v. bairdi_ lives primarily in forests as do _yaquinae_
and _pacificus_.

_Specimens examined._--Total number, 39. OREGON: _Clatsop Co._: type
locality, 12 BS; Seaside, 3 BS. _Tillamook Co._: Netarts, 1 OU;
Tillamook, 2 OSC; Blaine, 1 AW; Hebo Lake, 1 SGJ; 5 mi. SW Cloverdale,
1 AW. _Multnomah Co._: Portland, 6 USNM. _Lincoln Co._: Otis, 7 USNM;
Delake, 1 KU. _Lane Co._: north slope Three Sisters, 6000 ft., 4 BS.

_Marginal Records._--OREGON: type locality; Portland; north slope Three
Sisters; Taft (Macnab and Dirks, 1941:178).


=Sorex vagrans permiliensis= Jackson

    _Sorex obscurus permiliensis_ Jackson, Proc. Biol. Soc.
    Washington, 31:128, November 29, 1918.

_Type._--Adult male, skin and skull; No. 91048, U. S. Biol. Surv.
Coll.; obtained on October 2, 1897, by J. A. Loring from Permilia
Lake, W base Mt. Jefferson, Cascade Range, Marion Co., Oregon.

_Range._--The Cascade Mountains of Oregon from Mt. Jefferson north to
the Columbia River.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 14 specimens from the type locality are: total length,
117.7 (110-124); tail, 51.9 (45-58); hind foot, 14.0 (14-15). Pale
reddish in summer, darker and brownish in winter.

_Comparisons._--For comparison with _S. v. bairdi_ see account of that
subspecies. Larger than _S. v. setosus_ except tail relatively
shorter. More reddish in summer pelage than _setosus_.

_Remarks._--_S. v. bairdi_ is larger in the southern part of its range
than elsewhere. Specimens from McKenzie Bridge, herein referred to
_yaquinae_, are intermediate in character between _yaquinae_ and
_bairdi_ or between _yaquinae_ and _permiliensis_. The transition
between _yaquinae_ and _bairdi_ is much more gradual than between
_yaquinae_ and _permiliensis_.

_Specimens examined._--Total number, 21. OREGON: _Hood River Co._: Mt.
Hood, 2 BS. _Wasco Co._: Camas Prairie, E base Cascade Mts., SE Mt.
Hood, 1 BS. _Marion Co._: Detroit, 1 BS; type locality, 17 BS.

_Marginal Records._--OREGON: Mt. Hood; type locality; Detroit.


=Sorex vagrans setosus= Elliott

    _Sorex setosus_ Elliott, Field Columb. Mus. Publ. 32, zool.
    ser. 1:274, May 19, 1899.

    _Sorex obscurus setosus_, Jackson, Proc. Biol. Soc.
    Washington, 31:127, November 29, 1918.

_Type._--Adult male, skin and skull; No. 6213/238, Chicago Nat. Hist.
Mus.; obtained on August 18, 1898, by D. G. Elliott from Happy Lake,
Olympic Mts., Clallam Co., Washington.

_Range._--Washington from the Cascades west; southwestern British
Columbia west of 120° W Longitude north to Lund.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 20 specimens from the Olympic Mountains, Washington,
are: total length, 117.3 (107-125); tail, 49.8 (41-54); hind foot,
13.4 (12-14). Color dark in both summer and winter.

_Comparisons._--For comparison with _permiliensis_ see account of that
subspecies. Darker, longer-tailed, and somewhat larger cranially than
_S. v. obscurus_ with which it intergrades in southwestern British
Columbia. Smaller in all dimensions, but much the same color as _S. v.
longicauda_ with which it intergrades along the British Columbian
coast north of Lund. Larger, darker, less reddish, and longer-tailed
than the sympatric _S. v. vagrans_.

_Remarks._--_S. v. setosus_ lives mostly in forests. According to
Dalquest (1948:139) it is commonest at high altitudes in western
Washington. In the Hudsonian Life-zone where shrew habitat is more
restricted and marginal than it is at lower altitudes in the humid
part of Washington, _setosus_ might be expected to compete with _S. v.
vagrans_ and to supplant it. Records of occurrence in the Olympic
Mountains suggest a degree of such separation there.

_Specimens examined._--Total number, 135.

BRITISH COLUMBIA: Lund, Malaspina Inlet, 4 BS; Gibson's Landing, 10
BS; Port Moody, 19 BS; Langley, 2 BS; Chilliwack, 1 BS; Manning Park, 2
PMBC.

WASHINGTON: _Whatcom Co._: Mt. Baker, 6 WSC; Barron, 1 BS. _Chelan
Co._: Clovay Pass, 1 WSC; Stehekin, 6 (4 WSC, 2 BS); Cascade Tunnel, 1
WSC. _King Co._: Scenic, 1 WSC. _Kittitas Co._: Lake Kachess, 1 WSC;
Easton, 10 BS. _Clallam Co._: 8 mi. W Sekin River, 1 WSC; mouth of
Sekin River, 1 WSC; Clallam Bay, 1 CMNH; 7 mi. W Port Angeles, 1 WSC;
Ozette Lake, 1 CMNH; 12 mi. S Port Angeles, 4 WSC; Forks, 1 CMNH; Deer
Lake, 7 CMNH; Hoh Lake, 1 CMNH; Bogachiel Peak, 1 CMNH; Sol Duc Hot
Springs, 3 CMNH; Sol Duc Park, 1 CMNH; Canyon Creek, 1 WSC; Sol Duc
Divide, 2 WSC; Cat Creek, 2 WSC. _Jefferson Co._: Jackson Ranger
Station, 1 CMNH; Mt. Kimta, 2 CMNH; Reflection Lake, 6 CMNH; Blue
Glacier, 3 CMNH. _Gray's Harbor Co._: Westport, 1 WSC. _Pierce Co._:
Fort Lewis, 1 FC; Mt. Rainier, 19 (16 BS, 3 WSC). _Pacific Co._:
Tokeland, 2 BS. _Yakima Co._: Gotchen Creek, 3 WSC; Mt. Adams, 1 WSC.
_Skamania Co._: Mt. St. Helens, 1.

OREGON: _Hood River Co._: 2 mi. W Parkdale, 2 BS.

_Marginal Records._--BRITISH COLUMBIA: Rivers Inlet (Anderson,
1947:20); _Agassiz_ (Jackson, 1928:136); Chilliwack Lake. WASHINGTON:
Barron; Lyman Lake (Jackson, 1928:137); Mt. Stuart (Dalquest,
1948:141); Mt. Adams. OREGON: _2 mi. W Parkdale_. WASHINGTON: Ilwaco
(Jackson, 1928:137); Lund, Malaspina Inlet.


=Sorex vagrans longicauda= Merriam

    _Sorex obscurus longicauda_ Merriam, N. Amer. Fauna, 10:74,
    December 31, 1895.

_Type._--Adult male, skin and skull; No. 74711, U. S. Biol. Surv.
Coll.; obtained on September 9, 1895 by C. P. Streator, from Wrangell,
Alaska.

_Range._--The British Columbian and Alaskan coasts from Rivers Inlet
north to near Juneau and also certain islands including Etolin,
Gravina, Revillagigedo, Sergeif, and Wrangell.

_Diagnosis._--Size medium for the species, tail relatively long;
average and extreme measurements of 17 specimens from the type
locality are: total length, 128.4 (122-138); tail, 57.8 (53-66); hind
foot, 15.1 (14-16). Color dark in summer and winter.

_Comparisons._--For comparison with _S. v. setosus_ see account of
that subspecies. Larger and darker than _S. v. obscurus_ with which it
intergrades east of the humid coastal region; larger and darker than
_S. v. alascensis_ with which it intergrades in the Lynn Canal area;
larger and darker than _S. v. calvertensis_ which occurs on Calvert
Island and Banks Island, British Columbia; differs from _S. v.
insularis_ of Smythe, Townsend, and Reginald islands in larger size
and blackish rather than brown winter pelage; larger and relatively
longer-tailed than _S. v. elassodon_ which occurs on most of the
islands west of the range of _longicauda_; larger and relatively
longer-tailed than _S. v. isolatus_.

_Specimens examined._--Total number, 151.

ALASKA: Wrangell, 54 BS; 8 AMNH; Crittenden Creek, 1 BS; Ketchikan, 2
BS; Loring, 11 BS.

BRITISH COLUMBIA: Port Simpson, 25 BS; Inverness, 15 BS; head of
Rivers Inlet, 35 BS.

_Marginal Records._--BRITISH COLUMBIA: Great Glacier, Stikine River
(Jackson, 1928:133). ALASKA: Burroughs Bay (_ibid._). BRITISH
COLUMBIA: Bella Coola region (Anderson, 1947:19); head of Rivers
Inlet; Spider Island (Cowan, 1941:101); Goose Island (Cowan, 1941:99);
Princess Royal Island (Cowan, 1941:98); Pitt Island (_ibid._);
Metlakatla (Jackson, 1928:133); Port Simpson. ALASKA: Gravina Island
(_ibid._); Helm Bay (_ibid._); Etolin Island (_ibid._); Sergeif
Island, mouth of Stikine River (_ibid._); Sumdum Village (_ibid._);
Port Snettisham (_ibid._).


=Sorex vagrans mixtus= Hall

    _Sorex obscurus mixtus_ Hall, American Nat., 72:462, September
    10, 1938.

_Type._--Adult male, skin and skull; No. 70376, Mus. Vert. Zool.;
obtained on May 4, 1936, by R. A. Cumming, from Vanada, Texada Island,
Georgia Strait, British Columbia.

_Range._--Known only from the type locality.

_Diagnosis._--Size medium; average and extreme measurements of 5
specimens from the type locality are: total length, 111 (108-117);
tail, 48 (44-49); hind foot, 12 (12-13) (Hall, 1938:463).

_Comparisons._--Color much as in _S. v. setosus_ or _S. v. isolatus_;
palate longer than that of _isolatus_ or _setosus_; hind foot shorter
than either; smaller than _S. v. longicauda_.


=Sorex vagrans isolatus= Jackson

    _Sorex obscurus isolatus_ Jackson, Jour. Washington Acad.
    Sci., 12:263, June 14, 1922.

_Type._--Adult male, skin and skull; No. 177719, U. S. Biol. Surv.
Coll.; obtained on May 21, 1911, by Alexander Wetmore from mouth of
Millstone Creek, Nanaimo, Vancouver Island, British Columbia.

_Range._--Vancouver Island.

_Diagnosis._--Size medium; measurements of two from the type locality
are: total length, 113, 118; tail, 48, 49; hind foot, 14, 14. Dark in
summer and winter, underparts brownish.

_Comparisons._--Smaller than _S. v. setosus_ but color much the same;
resembles _S. v. obscurus_ in size and cranial characters but darker
in all pelages; similar in color to _S. v. vancouverensis_ with which
_isolatus_ is sympatric but with longer tail, longer hind feet,
broader rostrum and larger teeth. For comparison with _S. v. mixtus_
see account of that subspecies.

_Remarks._--_S. v. isolatus_ and _S. v. vancouverensis_ seemingly
approach one another morphologically more closely than do any other
pair of sympatric subspecies of _Sorex vagrans_. The exceptions may be
_S. v. vagrans_ and _S. v. obscurus_ which are geographically
sympatric in a few places although they may be ecologically separated.

_Specimens examined._--Total number, 9. BRITISH COLUMBIA, Vancouver
Island: Nanaimo, 3 BS; Barclay Sound, 1 AMNH; Goldstream, 5 BS.

_Marginal Records._--BRITISH COLUMBIA, Vancouver Island. (Anderson,
1947:19): Cape Scott; Victoria.


=Sorex vagrans insularis= Cowan

    _Sorex obscurus insularis_ Cowan, Proc. Biol. Soc. Washington,
    54:103, July 31, 1941.

_Type._--Adult female, skin and skull; No. 3110, Prov. Mus. British
Columbia; obtained on August 24, 1938, by T. T. and E. B. McCabe from
Smythe Island, Bardswell Group, British Columbia.

_Range._--Smythe, Townsend, and Reginald islands, British Columbia.

_Diagnosis._--Size medium; average and extreme measurements of 50
specimens from within the range of the subspecies are: total length,
122.3 (111-134); tail 52.6 (46-58); hind foot, 14.6 (13-15) (Cowan,
1941:107).

_Comparisons._--Smaller externally and cranially than _S. v.
longicauda_ and brown instead of blackish or grayish in winter pelage.
Skull broader than that of _S. v. calvertensis_ and color brown rather
than blackish or grayish in winter pelage.

_Remarks._--_S. v. insularis_ occurs together with _S. cinereus_ on
Townsend and Smythe islands. _S. vagrans_ far outnumbered the cinereus
shrew (Cowan, 1941:96).

_Records of occurrence._--BRITISH COLUMBIA (Cowan, 1941:104): Smythe
Island, Townsend Island, Reginald Island.


=Sorex vagrans calvertensis= Cowan

    _Sorex obscurus calvertensis_ Cowan, Proc. Biol. Soc.
    Washington, 54:103, July 31, 1941.

_Type._--Adult male, skin and skull; No. 1947, Prov. Mus. British
Columbia; obtained on July 14, 1937, by T. T. and E. T. McCabe from
Safety Cove, Calvert Island, British Columbia.

_Diagnosis._--Size medium; average and extreme measurements of 13
specimens from Calvert Island are: total length, 121.6 (109-129);
tail, 54.0 (52-58); hind foot, 14.7 (13-15) (Cowan, 1941:106).
Blackish or grayish in winter pelage.

_Comparisons._--Smaller externally and cranially and paler in winter
and summer than _S. v. longicauda_; for comparisons with _S. v.
insularis_ see account of that subspecies.

_Remarks_.--_S. v. calvertensis_ seems to be the only shrew on Calvert
and Banks islands.

_Records of occurrence._--BRITISH COLUMBIA (Cowan, 1941:103): Safety
Cove, Calvert Island; Larson Harbor, Banks Island.

_Marginal Records._--BRITISH COLUMBIA: Larson Harbor, Banks Island;
type locality.


=Sorex vagrans malitiosus= Jackson

    _Sorex obscurus malitiosus_ Jackson, Proc. Biol. Soc. Washington,
    32:23, April 11, 1919.

_Type._--Adult female, skin and skull; No. 8401; Mus. Vert. Zool.;
obtained on May 21, 1909, by H. S. Swarth from east side of Warren
Island, Alaska.

_Range._--Warren and Coronation islands, Alaska.

_Diagnosis._--Size medium; average and extreme measurements of 5
topotypes are: total length, 129.8 (126-135); tail, 56.4 (53-61); hind
foot, 15.4 (15-16). Color brownish in summer, brownish rather than
blackish in winter.

_Comparisons._--Somewhat more brownish than _S. v. longicauda_ but
resembling it in size; skull slightly more flattened and rostrum
broader. Larger than _S. v. elassodon_. Larger and relatively
longer-tailed than _S. v. alascensis_.

_Records of occurrence._--ALASKA (Jackson, 1928:130): Warren Island;
Coronation Island.


=Sorex vagrans elassodon= Osgood

    _Sorex longicauda elassodon_ Osgood, N. Amer. Fauna, 21:35,
    September 26, 1901.

    _Sorex obscurus elassodon_, Elliott, Field Columb. Mus. Publ.
    105, zool. ser. 6:450, 1905.

_Type._--Adult male, skin and skull; No. 100597, U. S. Biol. Surv.
Coll.; obtained on June 13, 1900, by W. H. Osgood from Cumshewa Inlet,
near old Indian village of Clew, Moresby Island, Queen Charlotte
Islands, British Columbia.

_Range._--Alaskan and British Columbian islands from Admiralty Island
south to Moresby Island.

_Diagnosis._--Size medium; average and extreme measurements of 4 from
the type locality are: total length, 126, (119-131); tail, 53.5
(52-55); hind foot, 13.8 (13-14). Color dark.

   [Illustration: FIG. 17. Probable geographic ranges of the
      subspecies of _Sorex vagrans_ on the coast of British Columbia
      and southeastern Alaska.

    1. _Sorex vagrans malitiosus_
    2. _Sorex vagrans elassodon_
    3. _Sorex vagrans prevostensis_
    4. _Sorex vagrans calvertensis_
    5. _Sorex vagrans insularis_
    6. _Sorex vagrans longicauda_
    7. _Sorex vagrans obscurus_
   ]

_Comparisons._--Smaller with relatively smaller tail and hind feet
than _S. v. longicauda_, but resembling it in color. Smaller and paler
than _S. v. prevostensis_ with relatively narrower rostrum. Larger,
darker, and with relatively longer tail than _S. v. obscurus_.
Resembles _S. v. alascensis_ but hind foot smaller and skull
relatively narrower. Smaller than _S. v. malitiosus_.

_Remarks._--In the northern part of its range _S. v. elassodon_ occurs
with _Sorex cinereus_. In the southern part it is the only shrew
present.

_Specimens examined._--Total number 93.

ALASKA: near Killisnoo, Admiralty Island, 2 BS; Kupreanof Island, 15
BS; Petersburg, Mitkof Island, 10 BS; Woewodski Island, 4 AMNH; Kasaan
Bay, Prince of Wales Island, 18 BS.

BRITISH COLUMBIA: Cumshewa Inlet, Moresby Island, 25 BS; Massett,
Graham Island, 6 BS; Queen Charlotte Islands, 13 AMNH.

_Marginal Records._--ALASKA: Hawk Inlet, Admiralty Island (Jackson,
1928:131); Kupreanof Island; Mitkof Island; St. John Harbor, Zarembo
Island (Jackson, 1928:131); Kasaan Bay, Prince of Wales Island; Duke
Island (Jackson, 1928:131). BRITISH COLUMBIA: Massett, Graham Island,
Queen Charlotte Islands; type locality; Langara Island, Queen
Charlotte Islands (Jackson, 1928:131). ALASKA: Forrester Island
(_ibid._); Rocky Bay, Dall Island (_ibid._); Shakan (really on
Kosciusko Island) (_ibid._); Point Baker (_ibid._); Kuiu Island
(_ibid._); Port Conclusion, Baranof Island (_ibid._).


=Sorex vagrans prevostensis= Osgood

    _Sorex longicauda prevostensis_ Osgood, N. Amer. Fauna, 21:35,
    September 26, 1901.

    _Sorex obscurus prevostensis_, Elliott, Field Columb. Mus.
    Publ. 105, zool. ser. 6:450, 1905.

_Type._--Adult male, skin and skull; No. 100618, U. S. Biol. Surv.
Coll.; obtained on July 3, 1900, by W. H. Osgood from north end of
Prevost Island (Kunghit Island on some maps) on coast of Houston
Stewart Channel, Queen Charlotte Islands, British Columbia.

_Range._--Known only from the type locality.

_Diagnosis._--Size medium; measurements of two specimens from the type
locality are: total length, 132, 142; tail, 53, 59; hind foot, 14, 15.
Color dark.

_Comparisons._--Larger and darker than _S. v. elassodon_. Resembles
_S. v. longicauda_ but darker, tail relatively somewhat shorter on the
average and rostrum relatively slightly broader.

_Specimens examined._--Total number, 14. BRITISH COLUMBIA: Prevost
Island, Queen Charlotte Group, 14 BS.


=Sorex vagrans alascensis= Merriam

    _Sorex obscurus alascensis_ Merriam, N. Amer. Fauna, 10:76,
    December 31, 1895.

    _Sorex glacialis_ Merriam, Proc. Washington Acad. Sci., 2:16,
    March 14, 1900, type from Point Gustavus, east side of
    entrance to Glacier Bay, Alaska.

    _S[orex]. alascensis_, Merriam, Proc. Washington Acad. Sci.,
    2:18, March 14, 1900.

    _[Sorex glacialis] alascensis_, Elliott, Field Columb. Mus.
    Publ. 45, zool. ser. 2:372, 1901.

    _Sorex alascensis alascensis_, Miller, U. S. Nat. Mus. Bull.,
    79:16, December 31, 1912.

_Type._--Adult female, skin and skull; No. 73539, U. S. Biol. Surv.
Coll.; obtained on July 10, 1895, by C. P. Streator from Yakutat,
Alaska.

_Range._--The coast of southern Alaska from the vicinity of Juneau
west to include eastern part of the Kenai Peninsula.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 9 specimens from 9 mi. W and 4 mi. N of Haines,
Alaska, are: total length, 110 (104-128); tail, 45.4 (41-52); hind
foot, 14 (14-14). Color grayish brown.

_Comparisons._--For comparison with _S. v. longicauda_ and _S. v.
elassodon_ see accounts of those subspecies. Resembles _S. v.
obscurus_ in color but differs in larger skull, longer hind foot and
in somewhat darker color. Larger and darker than _S. v.
shumaginensis_; the two intergrade near the base of the Kenai
Peninsula.

_Remarks._--This subspecies is transitional between the large, usually
dark subspecies of the southeastern Alaskan and British Columbian
coast and islands, and the smaller, paler subspecies of western and
interior Alaska. There seem to be no sharp breaks between _alascensis_
and _shumaginesis_. North of Haines, Alaska, size of shrews decreases
in a short distance across a narrow intergradational zone between
_alascensis_ and _obscurus_. Throughout most of its range _S. v.
alascensis_ occurs with _Sorex cinereus_.

_Specimens examined._--Total number, 88.

ALASKA: Orca, 1 BS; Montague Island, Prince William Sound, 2 BS;
Yakutat, 8 BS; north shore Yakutat Bay, 2 BS; Yakutat Bay, 1 BS; E
side Chilkat River, 100 ft., 9 mi. W and 4 mi. N Haines, 12 KU; 1 mi.
S Haines, 5 ft., 10 KU; 7 mi. SSE Haines, 10 ft., 2 KU; N end Sullivan
Island, 10 ft., 6 KU; SE end Sullivan Island, 10 ft., 2 KU; Glacier
Bay, 3 BS; Mendenhall River, 1 BS; Juneau, 36 BS.

BRITISH COLUMBIA: Sheslay River, 1 AMNH; headwaters Sheslay River, 1
AMNH.

_Marginal Records._--ALASKA: Valdez Narrows, Prince William Sound
(Jackson, 1928:128); north shore Yakutat Bay; east side Chilkat River,
100 ft., 9 mi. W and 4 mi. N Haines. BRITISH COLUMBIA: Sheslay River
(Jackson, 1928:128). ALASKA: Juneau; Glacier Bay; Montague Island,
Prince William Sound (ibid.); Port Nell Juan (ibid.).


=Sorex vagrans shumaginensis= Merriam

    _Sorex alascensis shumaginensis_ Merriam, Proc. Washington
    Acad. Sci., 2:18, March 14, 1900.

    [_Sorex glacialis_] _shumaginensis_, Elliott, Field Columb.
    Mus. Publ. 45, zool. ser. 2:373, 1901.

    _Sorex obscurus shumaginensis_, Allen, Bull. American Mus.
    Nat. Hist., 16:228, July 12, 1902.

_Type._--Adult male, skin and skull; No. 97993, U. S. Biol. Surv.
Coll.; obtained on July 17, 1899, by De Alton Saunders from Popof
Island, Shumagin Islands, Alaska. (Measured by C. Hart Merriam and
numbered 2210 in A. K. Fisher's catalog.)

_Range._--Southwestern Alaska from Seward Peninsula southeasterly to
western part of Kenai Peninsula and southwesterly to the southwestern
end of the Alaskan Peninsula.

_Diagnosis._--Size medium to small for the species; average and
extreme measurements of 6 specimens from King Cove, Alaska, are: total
length, 112.7 (107-118); tail, 48.3 (45-52); hind foot, 13.8 (13-14).
Tending toward the development of a tricolor pattern, the back
darkest, the sides buffy, and the venter paler.

_Comparisons._--Paler and more definitely tricolored than _S. v.
obscurus_; also with relatively shorter palate, narrower rostrum and
smaller teeth. For comparison with _S. v. alascensis_ see account of
that subspecies.

_Remarks._--_S. v. shumaginensis_ occurs together with _Sorex
cinereus_ over much of southwestern Alaska. Part of the range of
_shumaginensis_ falls within the tundra of the Arctic Life-zone. This
may be a partial explanation of the tricolored pattern of the animal.
_Sorex tundrensis_, _S. cinereus ugyunak_, and _S. cinereus haydeni_,
shrews which dwell mostly in treeless areas, are markedly tricolored,
or bicolored. _Sorex arcticus_, however, although tricolored, is found
in forested areas.

_Specimens examined._--Total number, 340. ALASKA: Sawtooth Mts., Nome
River, 2 AMNH; Nulato, 5 BS; St. Michaels, 1 BS; Bethel, 7 BS; Aniak,
1 BS; Skeventna River, 1 BS; 6 mi. WSW Snowshoe Lake, 1 KU; 1 mi. NE
Anchorage, 1 KU; Tyonek River, 48 BS; Hope, 15 BS; Hope, Mts. near, 13
BS; Moose Camp, 3 AMNH; Kenai Peninsula, 24 AMNH; Kakwok River, 80 mi.
up, 1 BS; Kakhtul River, 5 BS; Kakwok, 3 BS; Goodnews Bay, 1 BS; Lake
Aleknagik, 6 BS; Nushagak River, 25 mi. above Nushagak, 1 BS;
Dillingham, 1 BS; Nushagak Village, 15 BS; Homer, 1 AMNH; Kenai Mts.,
37 AMNH; Seldovia, 24 AMNH; Barabor, 1 AMNH; Caribou Camp, 7 AMNH;
Ugagik River, 3 BS; Becharof Lake, 8 BS; Cold Bay, 14 BS; Kanatak,
Portgage Bay, 4 BS; Chignik, 6 BS; Moller Bay, 1 BS; Alaska Peninsula,
near Popof Island, 6 AMNH; Frosty Peak, 15 BS; Morzhovoi Bay, 7 BS;
Ungu Island, 3 BS; Sand Point, Popof Island, 45 AMNH; Popof Island, 3
BS.

_Marginal Records._--ALASKA: Nome River; Nulato; Kuskokwim River, 200
mi. above Bethel, Crooked Creek (Jackson, 1928:126); 6 mi. WSW
Snowshoe Lake; Seldovia; mts. near Hope; Morhzovoi Bay; thence along
coast to St. Michael.


=Sorex vagrans obscurus= Merriam

    _Sorex vagrans similis_ Merriam, N. Amer. Fauna, 5:34, July
    31, 1891, _nec. S. similis_ Hensel, Zeitschr. der Deutsch.
    Geolog. Gesellsch., 7:459, 1855 (= _Neomys similis_).

    _Sorex obscurus_ Merriam, N. Amer. Fauna, 10:72, December 31,
    1895, new name for _Sorex vagrans similis_ Merriam.

    _Sorex obscurus obscurus_, Miller, Bull. U. S. Nat. Mus.,
    79:15, December 31, 1912.

_Type._--Adult female, skin and skull; No. 23525/30943, U. S. Biol.
Surv. Coll.; obtained on August 26, 1890, by Vernon Bailey and B. H.
Dutcher from near Timber Creek, 8200 ft., Lemhi Mts., 10 mi. SSW
Junction (now Leadore), Lemhi Co., Idaho.

_Range._--Mountainous interior of western North America from central
Alaska east across Yukon and southwestern Northwest Territories to
northeastern Alberta, south in the mountains through north-central and
western Washington, Idaho, western Montana, Wyoming, Utah, and
Colorado, into northern New Mexico.

_Diagnosis._--Size medium to small for the species; average and
extreme measurements of 9 topotypes are: total length, 110.3
(105-117); tail, 46.4 (42-50); hind foot, 13.1 (12.5-13.5). Color
grayish or brownish gray in summer, light grayish in winter.

_Comparisons._--For comparisons with _S. v. setosus_, _S. v.
longicauda_, _S. v. alascensis_ and _S. v. shumaginensis_ see accounts
of those subspecies. Paler and slightly larger than S. v. soperi.
Larger than the subspecies from central Montana herein described as
new. Smaller than _S. v. neomexicanus_. Averaging larger in all
dimensions than _S. v. monticola_ with which _obscurus_ intergrades in
northern New Mexico and northern Arizona. Larger than _S. v. vagrans_
with more grayish rather than reddish fresh summer pelage and light
gray rather than dark grayish-black fresh winter pelage.

_Remarks._--Intergradation of _S. v. obscurus_ with _S. v. setosus_,
_S. v. longicauda_, _S. v. alascensis_, and the new subspecies from
Montana takes place in the usual way with specimens from intermediate
localities being intermediate in size and color. However the
relationship of _S. v. obscurus_ and _S. v. vagrans_ (as the latter
subspecies is defined in this study) is rather complicated. In
southern British Columbia where the two subspecies come together a
situation of remarkable complexity prevails. Series from some
localities seem to represent intergrades between _obscurus_ and
_vagrans_; from other localities some specimens seem to be referable
to one and some to the other subspecies; from other localities all
specimens seem referable to one subspecies. A similar situation is
seen in specimens from northeastern Washington, northern and central
Idaho, and extreme western Montana. The region mentioned is one of
extensive interfingering of life-zones. In southern British Columbia
the main axes of the rivers, valleys and mountain ranges are north and
south. Most of the valleys are in the Transition Life-zone; the
forests are rather dry and of pine with more or less isolated
hydrosere communities about streams and ponds. These hydrosere
situations are the habitat of _Sorex vagrans_. Shrews from these
situations are usually referable to _vagrans_. The high ridges and
mountain ranges are usually in the Canadian Life-zone or higher and
most of the shrews referable to _obscurus_ come from such places.
Marginal localities with regard to life-zone produce most of the
populations which seem to represent intergrades between the two
subspecies. Isolated areas of Canadian Life-zone, even though
surrounded with Transition Life-zone, often harbor a population of
_obscurus_, whereas the streams in the nearby dry valleys harbor
populations of _vagrans_. Farther south in the Rocky Mountain chain,
_obscurus_ seemingly intergrades regularly with _vagrans_. This
intergradation is seen in populations from several localities in Utah.
There the lower elevations west of the Wasatch and Uinta mountains
are inhabited by _S. v. vagrans_, the higher elevations by _obscurus_
and where the ranges of the two abut intergrading populations occur.
In these series of intergrades there are specimens which, using size
as a subspecific criterion, would unhesitatingly be assigned, as
individuals, to _obscurus_, and others would be assigned to _vagrans_,
but these individuals represent extremes of a normally variable
population. At Cuddy Mountain, Idaho, the two subspecies seemingly
abut without intergradation; anyhow the available specimens from this
locality are referable to one or the other subspecies and none is
intermediate. The situation just described understandably has been the
source of much anguish to students who sought to identify shrews from
the Rocky Mountains. The reason for the relationship just described
has been discussed at length in a previous section.

In the Rocky Mountains of Wyoming and Colorado the subspecies _S. v.
obscurus_ ranges almost uninterruptedly over relatively large areas,
but southward in New Mexico and southwestward into Utah and Arizona,
suitable boreal habitat becomes insular in nature and obscurus there
is confined to the higher mountains. With one exception, once the
shrew populations become 'insular' in this region they become smaller
and show intergradation with _Sorex vagrans monticola_. The exception
is the population in the Sacramento Mountains of southeastern New
Mexico which is larger than _obscurus_ and has been rightly recognized
as a distinct subspecies, _neomexicanus_.

Almost without exception the range of typical _Sorex vagrans obscurus_
is sympatric with that of _Sorex cinereus_, usually the subspecies _S.
c. cinereus_. So close is this correspondence that the presence of _S.
cinereus_ comes near to being a useful aid in identifying _S. v.
obscurus_. In areas where individuals of _obscurus_ show
intergradation with _vagrans_, _Sorex cinereus_ is absent or rare. The
implication is that as the species _S. vagrans_ approaches the size of
the species S. cinereus, competition between the two increases with
resultant displacement of _cinereus_.

_Specimens examined._--Total number, 982.

ALASKA: Wahoo Lake, 69° 08' N, 146° 58' W, 2350 ft., 2 KU; Chandler Lake,
68° 12' N, 152° 45' W, 2900 ft., 1 KU; Bettles, 1 KU, 5 BS; Alatna, 1 BS;
Yukon River, 20 mi. above Circle, 1 BS; Tanana, 1 BS; Mountains near
Eagle, 18 BS; Richardson, 8 BS; head of Toklat River, 11 BS; Savage
River, 8 BS.

YUKON: MacMillan Pass, Mile 282, Canol Road, 1 NMC; MacMillan River,
Mile 249, Canol Road, 1 NMC; S. fork MacMillan River, Mile 249, Canol
Road, 2 NMC; Sheldon Lake, Mile 222, Canol Road, 5 NMC; Rose River,
Mile 95, Canol Road, 1 NMC; McIntyre Creek, 3 mi. NW Whitehorse, 2250
ft. 1 KU; Nisutlin River, Mile 40, Canol Road, 6 NMC; SW end Dezadeash
Lake, 2 KU; 3 mi. E and 1-1/2 mi. S Dalton Post, 2500 ft., 1 KU.

MACKENZIE: Nahanni River Mtns., Mackenzie River, 1 BS; Fort Simpson, 3
BS; Fort Resolution, Mission Island, 1 BS.

BRITISH COLUMBIA: W. side Mt. Glave, 4000 ft., 14 mi. S and 2 mi. E
Kelsall Lake, 1 KU; Stonehouse Creek, 5-1/2 mi. W jct. Stonehouse Creek
and Kelsall River, 4 KU; Bennett City, 6 BS; Wilson Creek, Atlin, 1
PMBC; McDame Post, Dease River, 6 BS; McDame Creek, 3 BS; Hot Springs,
3 mi. WNW jct. Trout River and Liard River, 1 KU; NW side Muncho Lake,
1 KU; Little Tahtlan River, 1 AMNH; Junction (4 mi. N Telegraph
Creek), 7 BS; Raspberry Creek, 16 AMNH; Klappan River Valley, 1 BS;
Chapa-atan River, 4 BS; Fort Grahame, 3 BS; Kispiox Valley, 23 mi. N
Hazleton, 1 BS; Bear Lake, site of Fort Connully, 2 BS; Tetana Lake, 1
PMBC; Hudson Hope, 2 BS; Charlie Lake, 3 PMBC; Babine Mts., 6 mi. N
Babine Trail, 5200 ft., 1 BS; Big Salmon River (S branch near Canyon),
1 BS; Ootsa Lake, 2 PMBC; Indianpoint Lake, 4 PMBC; Barkerville, 7 BS;
Yellowhead Lake, 2 NMC, 1 PMBC; N. fork Moose River, 1 BS; Moose
Lake, 2 BS; Moose Pass, 1 BS; Glacier, 7 AMNH, 12 BS; Golden, 1 BS;
Field, 2 BS; Caribou Lake, near Kamloops, 2 BS; Sicamous, 1 BS;
Monashee Pass, 4 PMBC; Paradise Mine, 3 PMBC; Level Mtn., 4 AMNH; 6
mi. S Nelson, 6 BS; Morrissey, 5 NMC; Wall Lake, 1 BS.

ALBERTA: Hays Camp, Slave River, Wood Buffalo Park, 1 NMC; Kinuso,
Assineau River, 1920 ft., 2 KU; Athabaska River, 30 mi. above
Athabaska Landing, 8 BS; Smokey Valley, 50 mi. N Jasper House, 1 BS;
Sulfur Prairie, Grande Cache River, 3 BS; Stoney River, 35 mi. N
Jasper House, 1 BS; Moose Mtn., 1 NMC; Rodent Valley, 25 mi. W Henry
House, 1 BS; Henry House, 3 BS; Jasper, 2 NMC; Shovel Pass, 4 NMC;
mouth of Cavell Creek, Jasper Park, 1 NMC; 11 mi. S Henry House, 2 BS;
15 mi. S Henry House, 1 BS; Red Deer River, 1 AMNH; 27 mi. W Banff, 3
NMC; 12 mi. WNW Banff, 4500 ft., 1 NMC; N. Fork Saskatchewan River,
5000 ft., 1 NMC; Cypress Hills, 1 NMC; Waterton Lakes Park, 53 NMC.

SASKATCHEWAN: Cypress Hills, 21 NMC.

WASHINGTON: _Okanogan Co._: Pasayten River, 1 BS; Bauerman Ridge, 1
BS; Conconully, 2 BS. _Pend Oreille Co._: 2 mi. N Gypsy Meadows, 2
WSC; Round Top Mtn., 1 WSC; head Pass Creek, 1. _Chelan Co._:
Stehekin, 4 BS; head Lake Chelan, 4 BS; Wenatchee, 1 BS. _Kittitas
Co._: Easton, 10 BS.

IDAHO: _Boundary Co._: Cabinet Mtns., E Priest Lake, 2 BS. _Adams
Co._: 1/2 mi. E Black Lake, 1 KU; 1 mi. N Bear Creek R. S., SW slope
Smith Mtn., 2 KU. _Washington Co._: 1 mi. NE Heath, SW slope Cuddy
Mtn., 4000 ft., 4 KU. _Lemhi Co._: 10 mi. SSW Leadore (type locality),
4 BS; 5 FC. _Fremont Co._: 7 mi. W West Yellowstone, 4 KU. _Custer
Co._: head Pahsimeroi River, Pahsimeroi Mtns., 1 BS. _Blaine Co._:
Perkins Lake, 1 KU. _Bear Lake Co._:--_Caribou Co._ line: Preuss Mts.,
1 BS.

MONTANA: _Glacier Co._: Sherburne Lake, 3 UM; 2-1/2 mi. W and 1-1/2 mi. S
Babb, 1 KU; St. Mary's, 6 UM; St. Mary Lakes, 9 BS; Fish Creek, 2 BS;
Gunsight Lake, 2 BS. _Flathead Co._: Nyack, 3 UM, 1 BS; 1 mi. W and
2 mi. S Summit, 1 KU. _Ravalli Co._: 8 mi. NE Stevensville, 3 BS;
Sula, 1 BS. _Meagher Co._: Big Belt Mtns., Camas Creek, 4 mi. S Fort
Logan, 7 BS. _Gallatin Co._: West Gallatin River, 4 BS. _Park Co._:
Emmigrant Gulch, 3 mi. SE Chico, 2 BS; Beartooth Mtns., 2 BS; _Carbon
Co._: Pryor Mtns., 2 BS.

WYOMING: _Yellowstone Nat'l Park_: Mammoth Hot Springs, 11 BS; Tower
Falls, 1 BS; Astringent Creek, 1 BS; Flat Mtn., 1 BS; Yellowstone
Park, 1 UM. _Park Co._: Beartooth Lake, 15 BS; SW slope Whirlwind
Peak, 1 KU; Pahaska Tepee, 6300 ft., 8 BS; Pahaska, mouth Grinnell
Creek, 15 BS; Pahaska, Grinnell Creek, 7000-7500 ft., 18 BS; 25 mi. S
and 28 mi. W Cody, 1 KU; Valley, Absaroka Mts., 14 BS; Needle Mtn., 2
BS. _Big Horn Co._: 28 mi. E Lovell, 9000 ft., 12 KU; head Trapper's
Creek, W slope Bighorn Mtns., 7 BS; 17-1/2 mi. E and 4-1/2 mi. S Shell, 1
KU. _Teton Co._: Two Ocean Lake, 6 FC; Emma Matilda Lake, 2 BS; 1 mi.
N Moran, 1 FC; 2-1/2 mi. E and 1/4 mi. N Moran, 6230 ft., 2 KU; Moran, 7
FC, 1 KU; 2-1/2 mi. E Moran, 6220 ft., 1 KU; 1 mi. S Moran, 1 FC; 3-3/4 mi.
E and 1 mi. S Moran, 9 KU; 7 mi. S Moran, 3 FC; Timbered Island, 14
mi. N Moose, 6750 ft., 3 KU; Bar BC Ranch, 2-1/2 mi. NE Moose, 6500 ft.,
1 KU; Beaver Dick Lake, 1 UM; Teton Mtns., Moose Creek, 6800 ft., 9
BS; Teton Mtns., S Moose Creek, 10,000 ft., 3 BS; Teton Pass, above
Fish Creek, 7200 ft., 15 BS; Whetstone Creek, 4 UM; Flat Creek-Gravel
Creek Divide, 2 UM; Flat Creek-Granite Creek Divide, 1 UM; Jackson, 3
KU, 2 UM. _Fremont Co._: Togwotee Pass, 5 FC; Jackey's Creek, 3 mi. S
Dubois, 1 BS; Milford, 5400 ft., 2 KU; Mosquito Park R. S. 17-1/2 mi. W
and 2-1/2 mi. N Lander, 1 KU; 17 mi. S and 6-1/2 mi. W Lander, 9300 ft., 1
KU; Mocassin Lake, 19 mi. W and 4 mi. N Lander, 10,000 ft., 1 KU; 23-1/2
mi. S and 5 mi. W Lander, 8600 ft., 1 KU; Green Mts., 8 mi. E Rongis,
8000 ft., 4 BS. _Washakie Co._: 9 mi. E and 5 mi. N Tensleep, 7400
ft., 2 KU; 9 mi. E and 4 mi. N Tensleep, 7000 ft., 2 KU. _Lincoln
Co._: Salt River Mtns., 10 mi. SE Afton, 5 BS; Labarge Creek, 9000
ft., 1 BS. _Sublette Co._: 31 mi. N Pinedale, 8025 ft., 3 KU;
Surveyor's Park, 12 mi. NE Pinedale, 8000 ft., 2 BS; N. side Half Moon
Lake, 7900 ft., 1 KU; 2-1/2 mi. NE Pinedale, 7500 ft., 2 KU. _Natrona
Co._: Rattlesnake Mts., 7000-7500 ft., 18 BS; Casper Mts., 7 mi. S
Casper, 6 BS. _Converse Co._: 21-1/2 mi. S and 24-1/2 mi. W Douglas, 7600
ft., 7 KU; 22 mi. S and 24-1/2 mi. W Douglas, 7600 ft., 4 KU; 22-1/2 mi. S
and 24-1/2 mi. W Douglas, 7600 ft., 2 KU. _Uinta Co._: 1 mi. N Fort
Bridger, 6650 ft., 1 KU; Fort Bridger, 3 KU; Evanston, 1 BS; 9 mi. S
Robertson, 8000 ft., 6 KU; 9 mi. S and 2-1/2 mi. E Robertson, 8600 ft., 1
KU; 10 mi. S and 1 mi. W Robertson, 8700 ft., 3 KU; 10-1/2 mi. S and 2
mi. E Robertson, 8900 ft., 1 KU; 13 mi. S and 1 mi. E Robertson, 9000
ft., 1 KU; 13 mi. S and 2 mi E Robertson, 9200 ft., 1 KU. _Carbon
Co._: Ferris Mts., 7800 to 8500 ft., 13 BS; Shirley Mts., 7600 ft., 7
BS; Bridget's Pass, 18 mi. SW Rawlins, 7500 ft., 2 KU; 10 mi. N and 12
mi. E Encampment, 7200 ft., 1 KU; 10 mi. N and 14 mi. E Encampment,
8000 ft., 6 KU; 9-1/2 mi. N and 11-1/2 mi. E Encampment, 7200 ft., 2 KU; 9
mi. N and 3 mi. E Encampment, 6500 ft., 1 KU; 9 mi. N and 8 mi. E
Encampment, 7000 ft., 1 KU; 8 mi. N and 14 mi. E Encampment, 8400 ft.,
3 KU; 8 mi. N and 14-1/2 mi. E Encampment, 8100 ft., 2 KU; 8 mi. N and 16
mi. E Encampment, 4 KU; 8 mi. N and 21-1/2 mi. E Encampment, 9400 ft., 2
KU; S. base Bridger's Peak, 8800 ft., Sierra Madre Mts., 3 BS; 8 mi. N
and 19-1/2 mi. E Savery, 8800 ft., 2 KU; 7 mi. N and 17 mi. E Savery,
8300 ft., 1 KU; 6-1/2 mi. N and 16 mi. E Savery, 8300 ft., 1 KU; 6 mi. N
and 15 mi. E Savery, 8500 ft., 1 KU; 5 mi. N and 10-1/2 mi. E Savery,
8000 ft., 2 KU; 14 mi. E and 6 mi. S Saratoga, 8800 ft., 1 KU. _Albany
Co._: Springhill, 12 mi. N Laramie Peak, 6300 ft., 10 BS; Laramie
Peak, N. slope, 8000 to 8800 ft., 7 BS; Bear Creek, 3 mi. SW Laramie
Peak, 7500 ft., 6 BS; 2-1/2 mi. ESE Brown's Peak, 10,500 ft., 2 KU; 3 mi.
ESE Brown's Peak, 10,000 ft., 1 KU; 27 mi. N and 5 mi. E Laramie, 6960
ft., 2 KU; 1 mi. SSE Pole Mtn., 8350 ft., 3 KU; 2 mi. SW Pole Mtn., 3
KU; 3 mi. S Pole Mtn., 8100 ft., 2 KU; 8-3/4, mi. E and 6-1/2 mi.
S Laramie, 8200 ft., 2 KU; Woods P. O., 1 BS. _Laramie Co._: 5 mi.
W and 1 mi. N Horse Creek P. O., 7200 ft., 2 KU.

UTAH: _Weber Co._: Mt. Willard, Weber-Box Elder Co. line, 9768 ft., 2
UU. _Salt Lake Co._: Butterfield Canyon, 7000 ft., 1 UU; Brighton,
Silver Lake P. O., 8700 ft., 2 UU; Brighton, Silver Lake P. O., 8750
ft., 8 UU; Brighton, Silver Lake P. O., 9000 ft., 2 UU; Brighton,
Silver Lake P. O., 9500 ft., 1 UU. _Summit Co._: Jct. Bear River and
East Fork, 2 CM; Smith and Morehouse Canyon, 7000 ft., 1 UU; Mirror
Lake, 10,000 ft., 1 UU. _Daggett Co._: Jct. Deep and Carter creeks,
7900 ft., 1 UU. _Utah Co._: Nebo Mtn., 1 mi. E Payson Lake, 8300 ft.,
1 UU; Nebo Mts., 12 mi. SE Payson Lake, 1 UU. _Wasatch Co._: Current
Creek, Uinta Mts., 1 BS; Wasatch Mts., 1 BS. _Uintah Co._: Paradise
Park, 21 mi. W and 15 mi. N Vernal, Uinta Mts., 10,050 ft., 2 CM, 3
KU; Paradise Park, Uinta Mts., 10,100 ft., 6 UU. _Sanpete Co._: Manti,
3 BS. _Sevier Co._: 7 mi. Creek, 20 mi. SE Salina, 5 CM; Fish Lake
Plateau, 2 BS. _Emery Co._: Lake Creek, 11 mi. E Mt. Pleasant, 4 CM.
_Grand Co._: Warner R. S., La Sal Mts., 9750 ft., 2 UU; La Sal Mts.,
11,000 ft., 1 BS. _Beaver Co._: Puffer Lake, Beaver Mts., 2 BS. _Wayne
Co._: Elkhorn G. S., Fish Lake Plateau, 14 mi. N Torrey, 9400 ft., 3
UU. _Garfield Co._: Wildcat R. S., Boulder Mtn., 8700 ft., 6 UU; 18
mi. N Escalante, 9500 ft., 1 UU. _Washington Co._: Pine Valley Mts., 7
BS. _San Juan Co._: Geyer Pass, 18 mi. SSE Moab, 3 CM; Cooley, 8 mi. W
Monticello, 3 CM.

COLORADO: _Larimer Co._: Poudre River, 1 KU. _Rio Blanco Co._: 9-1/2 mi.
SW Pagoda Peak, 2 KU. _Grand Co._: Arapaho Pass, Rabbit Ears Mts., 2
BS. _Boulder Co._: Willow Park, Rocky Mtn. Nat'l Park, 8 UM; Longs
Peak, 1 BS; 3/4 mi. N and 2 mi. W Allenspark, 8400 ft., 5 KU; Ward, 9500
ft., 1 BS; Buchanan Pass, 1 BS; 3 mi. S Ward, 1 KU; 7 mi. NW
Nederland, 1 KU; 5 mi. W Boulder, 3 BS; Boulder, 3 BS, 1 ChM;
Nederland, 6 BS, 4 ChM; Eldora, 1 BS. _Garfield Co._: Baxter Pass,
8500 ft., 2 BS. _Eagle Co._: Gores Range, 1 BS. _Gilpen Co._: Black
Hawk, 1 BS. _Lake Co._: 3 mi. W Twin Lakes, 2 KU; 12 mi. S and 1 mi. W
Leadville, 1 KU. _Gunnison Co._: 2 mi. W Gothic, 2 FC; Copper Lake, 2
FC; Gothic, 1 FC. _Chaffee Co._: St. Elmo, 10,100 ft., 2 BS; E side
Monarch Pass, 7 mi. W Salida, 2 ChM. _Teller Co._: Glen Core, Pikes
Peak, 2 UM. _El Paso Co._: Hunters Creek, a tributary of Bear Creek,
7250-7400 ft., 1 AMNH. _Montrose Co._: Uncomphagre Plateau, 8500 ft.,
3 BS. _Saguache Co._: 3 mi. N and 16 mi. W Saguache, 8500 ft., 2 KU;
Cochetopa Pass, 10,000 ft., 4 KU; Monshower Meadows, 27 mi. W
Saguache, 2 BS. _San Juan Co._: Silverton, 4 BS. _Mineral Co._: 23 mi.
S and 11 mi. E Creede, 1 KU. _Costilla Co._: Fort Garland, 2 BS.
_Huerfano Co._: 5 mi. S and 1 mi. W Cuchara Camps, 8 KU.

NEW MEXICO: _Taos Co._: 3 mi. N Red River, 2 BS; Taos, 1 BS. _Colfax
Co._: 1 mi. S and 2 mi. E Eagle Nest, 8100 ft., 2 KU. _Sandoval Co._:
Jemez Mts., 3 BS. _Santa Fe Co._: Hyde Park, 5 mi. NE Santa Fe, 2 HC;
Santa Fe Field Station, 1 HC; Santa Fe Ski Basin, 1 KU; Pecos Baldy, 4
BS. _Torrance Co._: Manzano Mts., 2 BS.

_Marginal Records._--ALASKA: Chandler Lake, 68° 12' N, 152° 45' W; Yukon
River, 20 mi. above Circle; Mountains near Eagle. MACKENZIE: Nahanni
River Mts.; Fort Simpson; Fort Resolution, Mission Island. ALBERTA:
Wood Buffalo Park; Athabaska River, 30 mi. above Athabaska Landing.
SASKATCHEWAN: Cypress Hills. MONTANA: St. Mary; 4 mi. S Fort Logan;
Pryor Mts. WYOMING: 1 mi. W and 1 mi. S Buffalo, 27424 KU; Springhill,
12 mi. N Laramie Peak; 5 mi. W and 1 mi. N Horse Creek PO. COLORADO:
Boulder; Hunters Creek; 5 mi. S and 1 mi. W Cuchara Camps. NEW MEXICO:
3 mi. N Red River, 10,700 ft.; Pecos Baldy; Manzano Mts.; Jemez Mts.
COLORADO: Navajo River (Jackson, 1928:120); Silverton. UTAH: La Sal
Mts., 11,000 ft. COLORADO: Baxter Pass. UTAH: junction Trout and
Ashley Creeks, 9700 ft. (Durrant, 1952:35); Mirror Lake, 10,000 ft.;
Mt. Baldy R. S. (Durrant, 1952:53); Wildcat R. S.; Pine Valley Mts.;
Puffer Lake; Butterfield Canyon. IDAHO: Preuss Mts.; 4 mi. S Trude
(Davis, 1939:104); head Pahsimeroi River, Pahsimeroi Mts.; Perkins
Lake; 1 mi. NE Heath; _1/2 mi. E Black Lake_. MONTANA: Sula; 8 mi. NE
Stevensville. WASHINGTON: head Pass Creek; Conconully; Wenatchee;
Easton; Stehekin; Pasayten River. BRITISH COLUMBIA: Second Summit,
Skagit River, 5000 ft., (Jackson, 1928:120); Babine Mts., 6 mi. N
Babine Trail, 5200 ft.; Hazleton (Jackson, 1928:120); 23 mi. N
Hazleton; Flood Glacier, Stikine River (Jackson, 1928:120); Cheonee
Mts. (_ibid._); Level Mtn.; west side Mt. Glave, 4000 ft., 14 mi. S
and 2 mi. E Kelsall Lake. ALASKA: head Toklat River; Tanana; Alatna;
Bettles.


=Sorex vagrans soperi= Anderson and Rand

    _Sorex obscurus soperi_ Anderson and Rand, Canadian
    Field-Nat., 59:47, October 16, 1945.

_Type._--Adult male, skin and skull; No. 18249, Nat. Mus. Canada;
obtained on September 21, 1940, by J. Dewey Soper, from 2-1/2 mi. NW Lake
Audy, Riding Mtn. Nat'l Park, Manitoba.

_Range._--Southwestern Manitoba to central Saskatchewan.

_Diagnosis._--Size medium to small for the species; measurements of
type and two topotypes are: total length, 107, 108, 117; tail, 45, 45,
45; hind foot, 12.1, 12.3, 12.5. Color dark brownish or fuscous in
summer pelage; winter pelage unknown.

_Comparison._--Resembles _S. v. obscurus_ in size; darker than
_obscurus_ in summer pelage; cranium slightly higher and top more
nearly flat; larger and darker in summer pelage than the new
subspecies from central Montana.

_Remarks._--In their description of this subspecies Anderson and Rand
pointed out that specimens from the type locality and from central
Saskatchewan represent the dark extreme in a color cline which begins
in south-central British Columbia with "pale, brownish-tinged
animals." These authors referred shrews from Cypress Hills,
southwestern Saskatchewan and southeastern Alberta to _S. o. soperi_,
although they noted that these specimens, taken by themselves, are not
strikingly different from _S. o. obscurus_ from the Rocky Mountains.
The specimens from the Cypress Hills were included in _soperi_ because
the authors felt that the subspecific boundary should be drawn "where
specimens average about half way between the extremes (of the cline)
in characters."

It is true, as Anderson and Rand say, that the shrews from Cypress
Hills are hardly separable from those from, say, Waterton Lakes Park.
The specimens from the Cypress Hills are noticeably different from
specimens from the Okanagan area, but some of the latter, in my
opinion may represent intergrades between _S. v. obscurus_ and the
more reddish _S. v. vagrans_ and are not, at any rate, typical
_obscurus_. In view of the similarity of shrews from Cypress Hills to
typical _S. v. obscurus_ and since the Cypress Hills are much nearer
to the range of _S. v. obscurus_ than to the record-stations of
occurrence in central Saskatchewan and Manitoba, I have chosen to
restrict the name _soperi_ to shrews from these latter two localities.
Seemingly _S. vagrans_ is absent from the plains separating the
Cypress Hills from the Rocky Mountains and from Riding Mountain
National Park.

_Specimens examined._--none.

_Marginal records._--SASKATCHEWAN: Prince Albert National Park, 1700
ft. (Anderson and Rand, 1945:48). MANITOBA: Riding Mountain National
Park, 2-1/2 mi. NW Audy Lake (ibid.).


=Sorex vagrans longiquus= new subspecies

_Type._--First year male, skin and skull; No. 87332, Univ. Michigan
Mus. Zool.; obtained on July 21, 1942, by Emmet T. Hooper from 25 mi.
ESE Big Sandy, Eagle Creek, Chouteau Co., Montana, original no. 2184.

_Range._--Central Montana; marginal localities are: Bearpaw Mts.,
Zortman, Big Snowy Mts., Buffalo, Little Belt Mts.

_Diagnosis._--Size small for the species; measurements of three
topotypes are: total length, 101, 105, 108; tail, 39, 40, 42; hind
foot, 11.5, 11.5, 12. Color pale; summer pelage: back near (17´´´_k_)
Olive Brown but hairs of dorsum with a pale, buffy band proximal from
the tips which imparts a pale over-all appearance; flanks near Wood
Brown; underparts Pale Smoke Gray, usually not with a buffy wash;
color of underparts often extending along margin of upper lip. Skull
small for species; rostrum relatively broad and heavy; relatively
broad interorbitally.

_Comparisons._--From _S. v. obscurus_, _S. v. longiquus_ differs as
follows: size smaller; skull smaller in all dimensions although
similar in proportion. From _S. v. soperi_, _S. v. longiquus_ differs
in: size smaller; color paler in summer pelage. From _S. v. vagrans_,
_S. v. longiquus_ differs in: color paler in summer pelage, less
brownish; color of venter extending higher on flanks; venter Pale
Smoke Gray, rarely tinged with buffy rather than usually tinged with
buffy. From _S. v. monticola_, _S. v. longiquus_ differs in: summer
pelage slightly paler, venter Pale Smoke Gray rather than suffused
with buffy.

_Remarks._--The subspecies _longiquus_ is obviously derived from the
neighboring _S. v. obscurus_ and differs from it mainly in size. Some
specimens of obscurus from western Montana show evidences of
intergradation with _S. v. vagrans_ in possessing a somewhat buffy
belly and these are thus more strikingly different from _longiquus_
than are other specimens of _obscurus_. Many specimens of _obscurus_
from the eastern slope of the Lewis and Clark Range in Montana show
the tricolored pattern seen in many specimens of _longiquus_. The
smallest individuals of longiquus are found on the Big Snowy
Mountains. Intergradation with _obscurus_ is seen in specimens here
referred to _S. v. obscurus_ from the Big Belt Mountains.

_Specimens examined._--Total number, 45. MONTANA: _Hill Co._: Bearpaw
Mts., 5 UM, 2 BS. _Phillips Co._: Zortman, 1 BS. _Chouteau Co._: type
locality, 3 UM; Highwood Mts., 13 BS. _Cascade Co._: Neihart, Little
Belt Mts., 1 BS. _Judith Basin Co._: 3 mi. W Geyser, 4100 ft., 1 KU;
Otter Creek, 10 mi. SW Geyser, 1 BS; Dry Wolf Creek, 20 mi. SW
Stanford, 1 BS. Buffalo, 13 mi. W Buffalo Canyon, 2 BS. _Fergus Co._:
Moccasin Mts., 15 mi. NW Hilger, 3 BS; Judith Mts., 17 mi. NE
Lewiston, 1 BS; 15 mi. S Heath, N. fork Flat Willow Creek, Big Snowy
Mts., 1 BS; Timber Creek, Big Snowy Mts., 1 BS; Crystal Lake, 6000
ft., Big Snowy Mts., 2 UM; Rocky Creek, 5600 ft., Big Snowy Mts., 3
UM; Big Snowy Mts., 3 BS. _Meagher Co._: Sheep Creek, 16 mi. N White
Sulphur Springs, Little Belt Mts., 1 BS.

_Marginal records._--MONTANA: Bearpaw Mts.; Zortman; Big Snowy Mts.;
16 mi. N White Sulphur Springs; Highwood Mts.


=Sorex vagrans neomexicanus= Bailey

    _Sorex obscurus neomexicanus_ Bailey, Proc. Biol. Soc.
    Washington, 26:133, May 21, 1913.

_Type._--Adult male, skin and skull; No. 100440, U. S. Biol. Surv.
Coll.; obtained on May 29, 1900, by Vernon Bailey, from Cloudcroft,
9000 ft., Otero Co., New Mexico.

_Range._--Sacramento and Capitan Mountains of New Mexico.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 4 topotypes are: total length, 105.2 (103-107); tail,
41.0 (39-42); hind foot, 13.1 (12.5-14). Color near Olive Brown in
summer; winter pelage unknown; skull large and relatively broad; teeth
relatively large.

_Comparisons._--Skull larger than that of _S. v. obscurus_ and
relatively somewhat broader; much larger in all cranial dimensions
than _S. v. monticola_.

_Remarks._--_S. v. neomexicanus_ is a well-marked subspecies seemingly
limited to the mountains of southeastern New Mexico. It is the only
species of _Sorex_ thus far recorded from that area.

_Specimens examined._--Total number, 12. NEW MEXICO: _Otero Co._: SW
slope Capitan Mts., 2 BS; 10 mi. NE Cloudcroft, 2 BS; Cloudcroft, 7
BS, 1 UM.

_Marginal records._--NEW MEXICO: SW slope Capitan Mts.; 10 mi. NE
Cloudcroft; type locality.


=Sorex vagrans monticola= Merriam

    _Sorex monticolus_ Merriam, N. Amer. Fauna, 3:43, September
    11, 1890.

    _Sorex vagrans monticola_, Merriam, N. Amer. Fauna, 10:69,
    December 31, 1895.

    _Sorex melanogenys_ Hall, Jour. Mamm., 13:260, August 9, 1932,
    type from Marijilda Canyon, 8600 ft., Graham Mts. [= Pinaleno
    Mts.] Graham Co., Arizona.

_Type._--Adult male, skin and skull; No. 17599/24535, U. S. Biol.
Surv. Coll.; obtained on August 28, 1899, by C. Hart Merriam and
Vernon Bailey from San Francisco Mtn., 11,500 ft., Coconino Co.,
Arizona.

_Range._--Mountains of western New Mexico, eastern Arizona, and the
northern Sierra Madre Occidental of Mexico.

_Diagnosis._--Size small for the species; average and extreme
measurements of 12 specimens from the White Mountains, Arizona, are:
total length, 104.3 (98-112); tail, 41.2 (37-45); hind foot, 12.0
(11-13). Summer pelage between (15´_m_) Proutts Brown and (15´´_m_)
Bister, venter tinged with (15´_f_) Pale Ochraceous Buff; winter
pelage near (17´´´_k_) Olive Brown; skull relatively broad.

_Comparisons._--For comparisons with _S. v. obscurus_ and _S. v.
neomexicanus_ see accounts of those subspecies. Skull slightly larger
and relatively broader than that of _S. v. orizabae_, and color
slightly paler. Differs from _S. v. vagrans_ in: winter pelage
grayish (near 17´´´_k_ Olive Brown) rather than blackish (17´´´´_k_ or
17´´´´_m_ Chaetura Drab or Chaetura Black); summer pelage slightly
grayer; skull relatively slightly broader rostrally and
interorbitally.

_Remarks._--_S. v. monticola_ intergrades gradually with _S. v.
obscurus_ to the north and east; indeed the type locality is actually
in this area of intergradation. So far as I know, _monticola_ is not
in reproductive continuity with any other subspecies of _Sorex
vagrans_. Specimens from southeastern Arizona are the smallest and
seem to be the most "typical" in the sense that they are most
different from _S. v. obscurus_. Some specimens from the whole length
of the Rocky Mountain chain in the United States have for years been
referred to _monticola_. Some of these, as I have pointed out, belong
to _S. v. longiquus_, and others are intergrades between _S. v.
obscurus_ and _S. v. vagrans_. Since _vagrans_ and _monticola_
resemble one another somewhat, and since topotypes of _S. v.
monticola_ actually show the influence of intergradation with
_obscurus_, it is easy to understand how intergrades between
_obscurus_ and _vagrans_ could have been assigned to _monticola_.

Throughout most of its range, _S. v. monticola_ is the only _Sorex_
present. In some places _monticola_ may occur with _S. nanus_ or _S.
merriami_. _S. v. monticola_ occurs with the water shrew in
southeastern Arizona.

_Specimens examined._--Total number, 80.

ARIZONA: _Coconino Co._: San Francisco Mtn., 3 BS, 6 CMNH. _Apache
Co._: Spruce Creek, Tunitcha Mts., 7 BS; Springerville, 1 BS; North
Fork White River, White Mts., 12 SD; White River, Horseshoe Cienega,
8300 ft., White Mts., 5 BS; Mt. Thomas, 9500 to 11,000 ft., White
Mts., 12 BS; Little Colorado River, White Mts., 4 BS; near head Burro
Creek, 9000 ft., White Mts., 1 BS. _Graham Co._: Graham Mts., 9200
ft., 2 BS. _Greenlee Co._: Prieto Plateau, 9000 ft., S. end Blue
Range, 1 BS. _Pima Co._: Summerhaven, 7500 ft., Santa Catalina Mts., 3
BS, 1 SD. _Cochise Co._: Fly Park, Chiricahua Mts., 4 BS; Rustler
Park, Chiricahua Mts., 1 SD; Long Park, Chiricahua Mts., 1 UM;
Huachuca Mts., 1 BS. _Santa Cruz Co._: Stone Cabin Canyon, 8500 ft.,
Santa Rita Mts., 1 BS.

NEW MEXICO: _San Juan Co._: Chusca Mts., 1 BS. _Catron Co._: Mogollon
Mts., 3 BS; 10 mi. E Mogollon, 1 KU. _Socorro Co._: Copper Canyon,
Magdalena Mts., 3 BS. _Sierra Co._: Mimbres Mts., near Kingston, 1 BS.

CHIHUAHUA: Sierra Madre, near Guadalupe y Calvo, 5 BS.

_Marginal records._--ARIZONA: Tunitcha Mts. NEW MEXICO: Chusca Mts.;
Copper Canyon, Magdalena Mts.; Mimbres Mts., near Kingston. CHIHUAHUA:
Guadalupe y Calvo. ARIZONA: Huachuca Mts.; Santa Catalina Mts.; White
River, Horseshoe Cienega, 8300 ft., White Mts.; San Francisco Mtn.


=Sorex vagrans orizabae= Merriam

    _Sorex orizabae_ Merriam, N. Amer. Fauna, 10:71, December 31,
    1895.

    _Sorex vagrans orizabae_, Jackson, N. Amer. Fauna, 51:113,
    July 24, 1928.

_Type._--Adult female, skin and skull; No. 53633, U. S. Biol. Surv.
Coll.; obtained on April 24, 1893, by E. W. Nelson from W slope of Mt.
Orizaba, 9,500 ft., Puebla.

_Range._--Transverse volcanic belt of mountains at the southern end of
the Mexican Plateau.

_Diagnosis._--Size small for the species; measurements of 3 specimens
from Volcan Toluca, Mexico, are: total length, 98, 100, 108; tail, 35,
39, 40; hind foot, 13, 13, 14. Summer pelage Mummy Brown tending
toward Olive Brown; Fuscous to Fuscous-Black in winter; skull and
teeth relatively narrow.

_Comparisons._--For comparison with _S. v. monticola_ see account of
that subspecies.

_Remarks._--The range of _S. v. orizabae_ probably is not now in
contact with that of any other subspecies of _S. vagrans_, although
judging by the slight degree of difference between _orizabae_ and
_monticola_ the separation between the two has not been of great
duration.

_Sorex vagrans orizabae_ occurs with _S. saussurei saussurei_
throughout the transverse volcanic belt.

_Specimens examined._--Total number, 23.

MICHOACÁN: Patambán, 1 BS; Nahuatzín, 3 BS; Mt. Tancítaro, 4 BS.

MEXICO: Salazar, 2 BS, 1 KU; Volcan de Toluca, 3 BS.

TLAXCALA: Mt. Malinche, 2 BS.

PUEBLA: Mt. Orizaba, 6 BS.

VERACRUZ: Cofre de Perote, 1 BS.

_Marginal records._--MICHOACÁN: _Patambán_. VERACRUZ: Cofre de Perote.
PUEBLA: _Mt. Orizaba_. MICHOACÁN: Mt. Tancítaro.


=Sorex vagrans vagrans= Baird

    _Sorex vagrans_ Baird, Rep't Pacific R. R. Survey 8: pt. 1,
    Mammals, p. 15, July 14, 1858.

    _Sorex suckleyi_ Baird, Rep't Pacific R. R. Survey 8: pt. 1,
    Mammals, p. 18, July 14, 1858, type from Steilacoom, Pierce
    Co., Washington.

    _Sorex dobsoni_ Merriam, N. Amer. Fauna, 5:33, July 30, 1891,
    type from Alturas or Sawtooth Lake, altitude about 7200 ft., E
    base Sawtooth Mts., Blaine Co., Idaho.

    _Sorex amoenus_ Merriam, N. Amer. Fauna, 10:69, December 31,
    1895, type from near Mammoth, 8000 ft., head Owens River, E
    slope Sierra Nevada, Mono Co., California.

    _Sorex nevadensis_ Merriam, N. Amer. Fauna, 10:71, December
    31, 1895, type from Reese River, 6000 ft., Nye-Lander Co.
    line, Nevada.

    _Sorex shastensis_ Merriam, N. Amer. Fauna, 16:87, October 28,
    1899, type from Wagon Camp, Mt. Shasta, 5700 ft., Siskiyou
    Co., California.

_Type._--Adult male, alcoholic; No. 1675, U. S. Nat. Mus.; obtained at
Shoalwater (Willapa) Bay, Pacific Co., Washington; received from J. G.
Cooper, and entered in Museum catalog on October 23, 1856.

_Range._--The Great Basin, and Columbia Plateau west across the
mountains to the Pacific coast of northern California, Oregon,
Washington and southwestern British Columbia.

_Diagnosis._--Size small for the species; average and extreme
measurements of 8 topotypes are: total length, 104.1 (99-109); tail,
43.3 (42-45); hind foot, 12.9 (12-14). Summer pelage ranging from
(15´_k_) Cinnamon Brown through (15´_m_) Proutt's Brown to (17´_m_)
Mummy Brown. Winter pelage (13´´´´_m_) Fuscous Black to (17´´´´_m_)
Chaetura Black.

_Comparisons._--For comparison with _S. v. monticola_ see account of
that subspecies. Differs from _S. v. halicoetes_ in relatively
narrower and more attenuate rostrum and in less brownish underparts in
winter pelage; smaller and more brownish (less grayish) than _Sorex
vagrans_ from the southern Sierra Nevada.

   [Illustration: FIG. 18. Probable geographic ranges of _Sorex
      vagrans vagrans_, its derivative subspecies, and _S. v.
      mixtus_.

   1. _S. v. vancouverensis_
   2. _S. v. vagrans_
   3. _S. v. halicoetes_
   4. _S. v. paludivagus_
   5. _S. v. obscuroides_
   6. _S. v. mixtus_
   ]

_Remarks._--Restriction of the range of _S. v. monticola_ to Arizona
and New Mexico leaves shrews that were formerly assigned to this
subspecies from Utah, Idaho, Washington and southern British Columbia
unassigned. Several names are available for consideration. The name
_Sorex vagrans dobsoni_ Merriam, 1891, type locality Alturas Lake,
Blaine Co., Idaho, was once applied to small shrews from Idaho,
Montana, Wyoming, and Utah, but was considered by Jackson to be
synonymous with _S. v. monticola_. The name _Sorex vagrans amoenus_
Merriam, 1895, type locality near Mammoth, Mono Co., California, has
been applied to wandering shrews from western Nevada, northeastern
California and southern Oregon. _Sorex vagrans nevadensis_ Merriam,
1895, type locality Reese River on Nye-Lander Co. line, Nevada was
considered by Hall (1946:119) to be synonymous with _S. v. amoenus_.
Specimens of _Sorex vagrans_ west of the Cascade Mountains have long
been referred to the nominate subspecies which has its type locality
at Willapa Bay, Pacific Co., Washington. Over so wide an area it is
only to be expected that some geographic variation is to be found.
Thus specimens from central Nevada average slightly paler in summer
pelage than those from the Pacific Coast or from the foothills of the
Rocky Mountains. In addition there are slight average differences in
size from place to place. Topotypes of _S. v. vagrans_, however, show
a fair degree of variability and some are nearly as pale as the paler
Great Basin stocks. Furthermore topotypical individuals of _vagrans_
can be lost in series of _S. v. amoenus_, although _amoenus_ is
shorter-tailed on the average. Specimens from the western foothills
of the Rocky Mountains show an amazing series of relationships with
the montane _S. v. obscurus_. In Utah, as previously pointed out,
complete intergradation occurs. At 1 mi. N Heath, Washington Co.,
Idaho, the lowland and the highland forms approach each other within a
short distance and still maintain a degree of distinctness, especially
in size. In northwestern Montana intergradation is extensive
(Clothier, 1950). In northeastern Washington distinctly separable
populations occur within a few miles of one another. In southern
British Columbia some populations are clearly intergrades while at 6
mi. S Yahk intergradation seemingly has not taken place. Where some
intergradation has occurred the result often has been increased size
of the lowland shrews, although they usually retain the reddish summer
pelage rather than acquiring the more grayish pelage of _obscurus_.
The name _dobsoni_ was based upon shrews from a place where lowland
and highland forms occur almost together with only a slight amount of
intergradation. Examples of "_dobsoni_" may not with certainty be
distinguished from typical _vagrans_ except that they are, as Merriam
(1895:68-69) points out, somewhat larger. Merriam (_loc. cit._)
further notes that _dobsoni_ is "intermediate in size and cranial
characters between _S. vagrans_ and _obscurus_;" a statement which
hits very close to the heart of the matter. I consider the name
_dobsoni_ to apply to intergrades. To attempt to apply the name to the
highly variable populations of intergrades from British Columbia to
southern Idaho seems inadvisable. I have examined the possibility of
using the name _amoenus_ for the animals from this region. The
characters which set _amoenus_ apart from _vagrans_, slightly shorter
tail and slightly darker summer pelage, however, are not universally
found in shrews from the Columbian Plateau and eastern Great Basin and
furthermore these differences between _amoenus_ and _vagrans_ do not
seem to me to be of great enough magnitude to warrant subspecific
recognition of the former. Thus the name _S. v. vagrans_ may apply to
shrews in the region under consideration. The subspecies, as thus
thought of, embraces several incipient subspecies, namely (1) the
populations on the isolated mountain ranges of Nevada, (2) the coastal
rain forest population and possibly (3) the population on the
Columbian Plateau.

In western British Columbia, Washington, and Oregon no evidences of
intergradation between _S. v. vagrans_ and the races _setosus_,
_permiliensis_, _bairdi_, _yaquinae_, or _pacificus_ are seen. In this
region _S. v. vagrans_ occurs sympatrically with one or the other of
these subspecies. Different degrees of differentiation thus obtain
between the subspecies _vagrans_ as here defined and the surrounding
subspecies of _Sorex vagrans_ to wit: complete intergradation and
allopatry in Utah with _S. v. obscurus_; partial intergradation and
partial sympatry with _S. v. obscurus_ in the foothill region from
Idaho to British Columbia; no intergradation and complete sympatry
with all the other races of _Sorex vagrans_ from the Cascades to the
coast and south to San Francisco Bay. The relationship of _S. v.
vagrans_ to the wandering shrews of the high Sierra is discussed on
page 58.

Throughout most of the Great Basin and Columbian Plateau _Sorex
vagrans_ is, with the exception of the rare _S. merriami_ and _S.
preblei_, the only small shrew. In the Cascades and in the coastal
lowlands it is the only small shrew except for _S. cinereus_ and _S.
trigonirostris_, both extremely rare and local in this region. _S.
vagrans_ seemingly competes to a certain extent with the larger _S.
trowbridgii_ in western Washington and seems to be partially dominant
to _trowbridgii_, at least in marshy habitats (Dalquest, 1941:171).

_Specimens examined._--Total number, 1197.

BRITISH COLUMBIA: _Osoyoos District_: Okanagan, 20 PMBC; Okanagan
Landing, 2 PMBC; Nahun Plateau, 2 PMBC. _Vancouver District_:
Vancouver, 2 PMBC. _New Westminister District_: Port Moody, 16 BS;
Westminster Jct., 4 AMNH; Langley, 1 BS; Vedder Crossing, 1 PMBC;
Huntingdon, 69 NMC; Sumas, 16 BS; Cultus Lake, 1 NMC. _Similkameen
District_: Princeton, 6 Mile Creek, 1 NMC. Hedley, Stirling Creek,
7 NMC; Fairview-Keremeos Summit, 5 NMC; Oliver, 1 NMC; Westbridge,
6 NMC; Osoyoos, 1 PMBC; Osoyoos-Bridesville Summit, 4 NMC; Cascade,
7 NMC. _Nelson District_: Kuskonook, 1 PMBC; Rossland, 14 NMC; Trail,
2 NMC; Creston, 4 PMBC, 4 NMC; near Creston, 7 NMC. _Cranbrook
District_: Cranbrook, 5 BS; Yahk, 2 NMC; Yahk Camp 6, 2 NMC; Goatfell,
2 NMC. _Fernie District_: Newgate, 3 NMC.

WASHINGTON: _Whatcom Co._: Blaine, 1 BS; Beaver Creek, 5 WSC; Glacier,
1 BS; Mt. Baker Lodge, 1 WSC; Lake Whatcom, 1 BS; Barron, 2 BS.
_Okanogan Co._: Sheep Mtn., 3 BS; E. end Bauerman Ridge, 1 BS;
Oroville, 1 BS; Hidden Lakes, 1 BS; Loomis, 1 BS; Conconully, 1 BS;
Twisp, 1 BS. _Ferry Co._: 5 mi. W Curlew, 2 BS. _Stevens Co._: Marcus,
1 BS. _Pend Oreille Co._: Canyon, 1 WSC; Metaline, 2 BS; Sullivan
Lake, 1 BS. _San Juan Co._: East Sound, Orcas Island, 3 BS; Friday
Harbor, San Juan Island, 1 BS; San Juan Park, 2 WSC; Blakely Island,
1 KU; Richardson, 6 BS. _Skagit Co._: Cypress Island, 1 KU; Hamilton,
1 BS; Sauk, 1 BS; Avon, 3 BS; Mt. Vernon, 2 BS; La Conner, 5 BS.
_Island Co._: San de Fuca, Whidby Island, 3 BS; Greenbank, Whidby
Island, 2 BS; 3 mi. N Clinton, Whidby Island, 1 BS. _Snohomish Co._:
Oso, 2 BS; Hermosa Point, Tulalip Indian Reservation, 7 mi. W and 1/2
mi. N Marysville, 3 KU. _Chelan Co._: Entiat, 2 BS. _Lincoln Co._:
6 mi. E Odessa, 4 BS. _Spokane Co._: Marshall, 7 BS. _Clallam Co._:
Neah Bay, 29 BS; 8 mi. W Sekin River, 1 WSC; mouth Sekin River, 1 WSC;
Dungeness, 1 BS; Port Townsend, 3 BS; Ozette Indian Reservation,
1 CMNH; Sequim, 4 BS; Tivoli Island, Ozette Lake, 1 CMNH; Garden Island,
Ozette Lake, 3 CMNH; Elwah, 1 WSC; Blyn, 1 BS; Soleduck River, 1 BS;
12 mi. S Port Angeles, 1 WSC; Forks, 9 CMNH, 1 BS; Cat Creek, 1 WSC;
Lapush, 5 BS. _Jefferson Co._: Jefferson Ranger Station, N Fork Hoh
River, 5 CMNH; Duckabush, 6 BS. _Kitsap Co._: Vashon Island, 2 BS.
_King Co._: Redmont, 2 BS; Kirkland, 20 BS; Seattle, 1 WSC, 3 KU;
Northbend, 2 BS: Lake Washington, near Renton, 2 BS; Kent, 1 BS;
Enumclaw, 1 BS. _Grays Harbor Co._: Lake Quinault, 9 BS; Aberdeen,
20 BS; Westport, 5 BS, 2 WSC; Oakville, 1 BS. _Mason Co._: Lake Cushman,
11 BS; Hoodsport, 1 BS; North Fork Skokomish River, 1 BS; Shelton,
2 BS. _Pierce Co._: Puyallup, 6 BS; Steilacoom, 1 BS; 6 mi. S Tacoma,
2 BS; Roy, 3 BS; Bear Prairie, Mt. Rainier, 1 BS; Reflection Lake, Mt.
Rainier, 1 WSC. _Kittitas Co._: Blewett Pass, 3 BS; Easton, 3 BS;
2 mi. E Cle Elum, 4 FC; Ellensburg, 2 BS. _Grant Co._: Moses Lake, 1 BS;
9 mi. S and 1 mi. W Neppel, 1 UM. _Whitman Co._: Hangman Creek, Tekoa,
1 WSC; 4 mi. ENE Pullman, 1 KU; 2 mi. N Pullman, 2 WSC; 2 mi. NW
Pullman, 1 WSC; 2 mi. W Pullman, 1 WSC; Pullman, 5 WSC; Armstrong,
1 WSC; 5 mi. NE Wawawai, 1 BS; Wawawai, 5 WSC. _Thurston Co._: Nisqually
Flats, 2 BS; Nisqually, 1 BS; 4 mi. S Olympia, 1 BS; Tenino, 4 BS.
_Pacific Co._: Tokeland, 4 BS; 1 mi. S Nemah, 2 FC; 1 mi. N Bear
River, Willapa Bay, 8 FC; 1/4 mi. N Bear River, 3 FC; 3-1/2 mi. E Seaview,
6 FC; Ilwaco, 1 BS. _Lewis Co._: 8 mi. W Chehallis, 2 BS; Chehallis,
2 BS; Toledo, 1 BS. _Yakima Co._: Selah, 7 KU; Wiley City, 4 BS.
_Wahkiakum Co._: Cathlamet, 1 BS. _Skamania Co._: 45 mi. SE Toledo,
2 BS; Carson, 1 BS; Stevenson, 1 BS; 15 mi. NW White Salmon, 1 BS.
_Klickitat Co._: Trout Lake, 15 mi. S Mt. Adams, 2 BS; 15 mi. N
Goldendale, 1 WSC; Goldendale, 1 BS. _Walla Walla Co._: College Place,
1 KU. _Columbia Co._: Starbuck, 3 BS. _Garfield Co._: 1 mi. E Pomeroy,
1 SGJ. _Asotin Co._: 21 mi. SE Dayton, 1 BS; Rogersburg, 1 BS.

IDAHO: _Bonner Co._: 4 mi. S Sandpoint, 1 UM. _Kootenai Co._: Coeur
d'Alene, 2 BS. _Shoshone Co._: Osburry, 1 BS; Mullan, 2 BS. _Latah
Co._: Felton's Mills, 1 WSC; Cedar Mtn., 5 WSC. _Lewis Co._: Nezperce,
2 BS. _Idaho Co._: Seven Devils Mtn., 1 BS. _Adams Co._: Summit of
Smith Mtn., 7500 ft., 5 KU; New Meadows, 1 BS; Tamarack, 1 BS.
_Washington Co._: 1 mi. NE Heath, SW slope Cuddy Mtn., 4000 ft., 7 KU.
_Boise Co._: Bald Mtn. R. S., 10 mi. S Idaho City, 1 BS. _Elmore Co._:
Cayuse Creek, 10 mi. N Featherville, 1 BS. _Canyon Co._: Nampa, 5 BS.
_Blaine Co._: Sawtooth City, 5 BS; Alturas Lake, 1 BS. _Bonneville
Co._: 10 mi. SE Irwin, 5 BS. _Bannock Co._: Pocatello, 1 BS, 1 KU;
1 mi. W Bancroft, 1 KU; Swan Lake, 1 BS. _Owyhee Co._: Grasmere, 1 SGJ.
_Cassia Co._: 10 mi. S Albion, Mt. Harrison, 1 BS.

MONTANA: _Sanders Co._: Prospect Creek, near Thomson Falls, 4 BS.
_Lake Co._: Flathead Lake, 5 BS. _Ravalli Co._: Bass Creek, NW
Stevensville, 2 BS; 2 mi. NE Stevensville, 1 UM; Corvallis, 4 BS;
6 mi. E Hamilton, 1 KU.

OREGON: _Clatsop Co._: Seaside, 1 BS. _Washington Co._: 5 mi. SE
Hillsboro, 1 BS; Beaverton, 1 BS. _Multnomah Co._: Portland, 20 BS;
Portland, Switzler Lake, 5 BS. _Hood River Co._: 2 mi. W Parkdale,
1 BS; north slope Mt. Hood, 2 BS. _Umatilla Co._: 10 mi. W Meacham,
2 BS; Meacham, 3 BS. _Union Co._: Elgin, 2 BS; Kamela, 2 BS; Hot Lake,
2 BS. _Wallowa Co._: 25 mi. N. Enterprise, 4 BS; Wallowa Lake, 23 BS;
S Wallowa Lake, 1 BS. _Clackamas Co._: Estacada, 1 KU. _Marion Co._:
Salem, 8 BS; Permilia Lake, 2 BS. _Benton Co._: Corvallis, 2 BS; 5 mi.
SW Philomath, 5 BS. _Linn Co._: Shelburn, 1 BS. _Jefferson Co._: 20
mi. W Warm Springs, 2 BS. _Grant Co._: Beech Creek, 6 BS; Austin,
1 BS; Strawberry Butte, 1 BS; Strawberry Mts., 12 BS. _Baker Co._:
Homestead, 1 BS; Cornucopia, 11 BS; Rock Creek, 1 BS; Bourne, 7 BS;
McEwen, 1 BS; Huntington, 1 BS; Anthony, 42 AMNH. _Lane Co._: north
slope Three Sisters, 3 BS; Vida, 1 BS; Mapleton, 1 BS; Eugene, 2 BS;
10 mi. S McKenzie Bridge, 1 BS; Florence, 1 BS. _Deschutes Co._:
Paulina Lake, 7 BS; Lapine, 8 BS. _Crook Co._: 1 SGJ. _Douglas Co._:
Winchester Bay, 1 SGJ; Scottsburg, 3 BS; Drain, 5 BS; Lookingglass,
1 BS; Diamond Lake, 6 BS. _Coos Co._: Empire, 5 BS. _Curry Co._: Port
Orford, 1 BS; Gold Beach, 4 BS. _Klamath Co._: Anna Creek, Mt. Mazama,
1 BS; Crater Lake, 14 BS; Upper Klamath Marsh, 2 BS; Ft. Klamath,
35 BS; Klamath Falls, 6 BS. _Lake Co._: 10 mi. SW Silver Lake, 3 BS;
west fork Silver Creek, Yamsay Mts., 4 BS; Plush, 1 BS; Warner Creek,
Warner Mts., 1 BS; Warner Mts., 3 BS; Gearhart Mts., 17 SGJ; _Harney
Co._: Diamond, 2 BS; Keiger Gorge, Steens Mts., 3 BS. _Malheur Co._:
8 mi. W Jordon Valley, 1 BS.

WYOMING: _Lincoln Co._: 13 mi. N and 2 mi. W Afton, 6 KU; 10 mi. N
Afton, Salt River, 2 BS; 9 mi. N and 2 mi. W Afton, 6 KU; 7 mi. N and
1 mi. W Afton, 4 KU; Cokeville, 1 BS; 12 mi. N and 2 mi. E Sage, 2 KU;
6 mi. N and 2 mi. E Sage, 1 KU.

CALIFORNIA: _Del Norte Co._: Smith River, 2 BS; Crescent City, 20 BS.
_Siskiyou Co._: Beswick, 1 BS; Hornbrook, 3 BS; Brownell, Klamath
Lake, 1 BS; Mayten, 2 BS; Squaw Creek, Mt. Shasta, 5 BS; Upper Ash
Creek, Mt. Shasta, 1 BS; upper Mud Creek, Mt. Shasta, 8 BS; Wagon
Camp, Mt. Shasta, 5 BS; Warmcastle Soda Springs, Squaw Creek Valley, 2
BS; Sisson, 7 BS. _Modoc Co._: Davis Creek, Goose Lake, 1 BS.
_Humboldt Co._: _Humboldt Bay_, 10 BS. _Trinity Co._: Canyon Creek, 2
BS. _Shasta Co._: Fort Crook, 11 BS; Dana, 17 BS; Fall Lake, Fall
River Valley, 3 BS; Cassel, 2 BS; 12 mi. E Burney, 1 BS; Lassen Peak,
13 BS; Kellys, Warner Creek, 1 KU; Drakes Hot Springs, Warner Creek, 2
BS. _Mendocino Co._: Russian Gulch State Park, 2 FC. _Plumas Co._: 12
mi. NE Prattville, 2 BS; Spring Garden Ranch, Grizzly Mts., 3 BS;
Sierra Valley, 1 BS. _Sierra Co._: Lincoln Creek, 1 BS. _Sonoma Co._:
Petaluma, 3 BS; Point Reyes, 7 BS. _Placer Co._: Donner, 3 BS. _El
Dorado Co._: Tallac, 3 BS. _Mono Co._: Mt. Conness, 1 BS; Mono Lake, 1
BS; near Mammoth, 8000 ft., head of Owens River, 2 BS. _Inyo Co._:
Alvord, 1 BS.

NEVADA: _Elko Co._: Mountain City, 1 BS; Three Lakes, 1 KU; west side
Ruby Lake, 3 mi. N White Pine Co. line, 8 KU; Ruby Mts., 9 BS; W side
Ruby Lake, 3 BS. _White Pine Co._: W side Ruby Lake, 3 mi. S Elko Co.
line, 1 KU. _Nye Co._: Cloverdale, Reese River, 3 BS.

UTAH: _Weber Co._: Beaver Creek, S Fork Ogden River, 2 UU; Huntsville,
10 mi. E Ogden, 1 UU; Hooper Bay Refuge, 4200 ft., 1 UU; Riverdale,
4200 ft., 3 UU; Riverdale, 4250 ft., 1 UU; 3 mi. SE Ogden, 2 UU; Snow
Basin, 2 UU; Snow Basin, S part Wheeler Canyon, 1 UU; Uinta, 2 mi. W
Weber Canyon entrance, 4 UU; 2 mi. W Uinta, 1 UU. _Salt Lake Co._:
City Creek Canyon, 6 mi. NE Salt Lake City, 4700 ft., 2 UU; 1 mi. up
City Creek Canyon, 4600 ft., 1 UU; 3/4 mi. above Forks, City Creek
Canyon, 1 UU; The Firs, Millcreek Canyon, 1 UU; Olympus Water Box, 1
UU; Salamander Lake, Lamb's Canyon, 9000 ft., 3 UU (near _obscurus_);
Salt Lake City, 7500 ft., 1 UU; 1 mi. W Draper, 4500 ft., 6 UU;
Draper, 4500 ft., 5 UU; 1-1/2 mi. SW Draper, 4500 ft., 1 UU; 3 mi. SW
Draper, 4400 ft., 2 UU; 3 mi. S Draper, 4400 ft., 2 UU; 1 mi. S
Draper, 4500 ft., 1 UU. _Juab Co._: W side Deep Creek Mts., Queen of
Sheba Canyon, 8000 ft., 3 UU. _Wasatch Co._: Midway Fish Hatchery,
5450 ft., 1 UU.

_Marginal records._--BRITISH COLUMBIA: Okanagan; Westbridge;
Kuskonook; Cranbrook. MONTANA: Flathead Lake; 6 mi. E Hamilton;
Prospect Creek. IDAHO: Cedar Mtn.; New Meadows; Alturas Lake; 10 mi.
SE Irwin. WYOMING: 13 mi. N and 2 mi. W Afton; 6 mi. N and 2 mi. E
Sage. IDAHO: 1 mi. W Bancroft; Swan Lake. UTAH: Beaver Creek, South
Fork, Ogden River; Midway Fish Hatchery; west side Deep Creek Mts.,
Queen of Sheba Canyon, 8000 ft. NEVADA: Baker Creek (Hall, 1946:120);
Reese River (_ibid._); 2 mi. S Hinds Hot Springs (_ibid._).
CALIFORNIA: Mono Lake (Jackson, 1928:110); near Mammoth; Alvord; Mount
Conness; Donner; Buck Ranch (Jackson, 1928:110); Warner Creek, Drake
Hot Springs (_ibid._); Canyon Creek; Cuddeback (Jackson, 1928:105);
Novato Point (_ibid._), thence northward along the coast to
WASHINGTON: Friday Harbor, San Juan Island. BRITISH COLUMBIA: Port
Moody.


=Sorex vagrans obscuroides= new subspecies

_Type._--First year female, skin and skull; No. 30064/42074, U. S.
Biol. Surv. Coll.; obtained on August 9, 1891, by Frank Stephens from
Bishop Creek, 6600 ft., Inyo Co., California, original no. 811.

_Range._--The Sierra Nevada of California, north at least to El Dorado
County, intergrading northerly with _S. v. vagrans_.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 5 topotypes are: total length, 107 (103-112); tail, 47
(45-50); hind foot, 12.8 (12-13.5). Skull relatively broad
interorbitally; color of dorsum in summer pelage nearest (17´´´_k_)
Olive Brown.

_Comparisons._--Differs from _S. v. vagrans_, with which it
intergrades to the north, in: longer tail and total length; skull
larger and relatively broader interorbitally; color in summer grayer
(less reddish), the lighter subterminal color bands of the hair often
showing through the darker tips and imparting a grizzled appearance to
the dorsum. Differs from _S. v. parvidens_ to the south in: skull
relatively broader interorbitally and less flattened; teeth slightly
larger.

_Remarks._--_S. v. obscuroides_ has long been called _S. v. obscurus_.
In fact, _obscuroides_ is separated from the range of _obscurus_ by
the intervening, smaller subspecies _S. v. vagrans_. _S. v.
obscuroides_ resembles _S. v. obscurus_ in color and size but the
skull is smaller, although relatively slightly broader. The
resemblance in color is possibly due to the fact that _obscuroides_,
like _obscurus_, is a high mountain form. _S. v. obscuroides_
intergrades with _S. v. vagrans_ along the crest of the Sierra between
Yosemite National Park and Lassen Peak and on the eastern slope of the
Sierra from approximately Mammoth northward. Specimens from Donner are
intergrades but are closest to _S. v. vagrans_. Although all specimens
from Lassen Peak are referable to _S. v. vagrans_, some show cranial
characters of _obscuroides_.

_Specimens examined._--Total number, 76. CALIFORNIA: _Mono Co._: Mt.
Dana, 6 BS; Mt. Lyell, 11 BS. _Mariposa Co._: Tuolumne Meadows, Muir
Meadow, 9300 ft., 1 BS; Tuolumne Meadows, Mt. Unicorn, 1 BS; Tuolumne
Meadows, N base Mt. Lyell, 8 BS; Tuolumne Meadows, Soda Springs, 4 BS;
Lake Tenaya, 5 BS. _Madera Co._: San Joaquin River, 8000 ft., 4 BS.
_Fresno Co._: Horse Corral Meadows, 3 BS. _Mono Co._: head of Owens
River near Mammoth, 2 BS. _Inyo Co._: Bishop Creek, 5 BS; Round
Valley, 1 BS. _Tulare Co._: E. Fork Kaweah River, 7 BS; Mt. Whitney, 5
BS; Whitney Creek, Mt. Whitney, 4 ChM; Whitney Meadows, 9700 ft., 1
BS; Mineralking, 2 BS; N. Fork Kern River, 9600 ft., 1 BS; S. Fork
Kern River, 4 BS; Kern Lakes, 1 BS.

_Marginal records._--CALIFORNIA: Pyramid Peak; near Mammoth; _Round
Valley_; Bishop Creek; Mt. Whitney; Kern Lakes; Halstead Meadows;
Horse Corral Meadows; east fork Indian Canyon (Jackson, 1928:121).


=Sorex vagrans parvidens= Jackson

   _Sorex obscurus parvidens_ Jackson, Jour. Mamm., 2:161, August
   19, 1921.

_Type._--Adult male, skin and skull; No. 56561, U. S. Biol. Surv.
Coll.; obtained on October 3, 1893, by J. E. McLellan from Thurmans
Camp, Bluff Lake, 7500 ft., San Bernardino Mts., California.

_Range._--Confined, so far as known, to the San Bernardino and San
Gabriel mountains, San Bernardino Co., California.

_Diagnosis._--Size medium for the species; measurements of two
specimens from the San Bernardino Mountains are: total length, 105,
106; tail, 41, 48; hind foot, 12, 14. Upper parts in summer
Olive-Brown to Buffy-Brown; cranium flattened and relatively narrow;
unicuspids and incisors relatively small.

_Comparisons._--For comparison with _S. v. obscuroides_, the only
adjacent subspecies, see the account of that subspecies.

_Remarks._--_S. v. parvidens_ is seemingly an uncommon mammal. I have
been informed by Terry Vaughan that repeated attempts by him to obtain
it in suitable habitat in the San Gabriel Mountains failed. This shrew
is probably no longer in reproductive continuity with _Sorex vagrans_
of the Sierra Nevada.

_Specimens examined._--Total number, 4. CALIFORNIA: _San Bernardino
Co._: type locality, 4 BS.

_Marginal records._--CALIFORNIA: Camp Baldy, San Antonio Canyon
(Jackson, 1928:124); type locality.


=Sorex vagrans halicoetes= Grinnell

    _Sorex halicoetes_ Grinnell, Univ. California Publ. Zool.,
    10:183, March 20, 1913.

    _Sorex vagrans halicoetes_, Jackson, N. Amer. Fauna, 51:108,
    July 24, 1928.

_Type._--Young adult male, skin and skull; No. 3638, Mus. Vert. Zool.;
obtained on May 6, 1908, by Joseph Dixon from salt marsh near Palo
Alto, Santa Clara Co., California.

_Range._--Marshes in the southern part of San Francisco Bay,
California.

_Diagnosis._--Size small for the species; measurements of two
topotypes are: total length, 105, 106; tail, 39, 40; hind foot, 12,
13. Upper parts in winter Chaetura Black or near Fuscous-Black;
underparts brownish; upper parts in summer near (17´_m_) Mummy Brown;
underparts with a decided buffy wash, near (15´_d_) Light Ochraceous
Buff; rostrum relatively large; maxillary tooth-row relatively long;
teeth relatively large.

_Comparisons._--Darker ventrally, both summer and winter, than _S. v.
vagrans_; slightly more reddish dorsally in summer pelage than _S. v.
vagrans_, rostrum and teeth relatively larger; smaller externally than
_S. v. paludivagus_, paler; skull longer, narrower cranially and
broader rostrally.

_Remarks._--This subspecies seems to be restricted to salt marshes
where it occurs with _Sorex ornatus_.

_Specimens examined._--Total number, 12. CALIFORNIA: _San Francisco
Co._: San Francisco, 4 BS. _Alameda Co._: West Berkeley, 1 BS;
Berkeley, 1 BS; Dumbarton Point, 1 KU. _San Mateo Co._: San Mateo, 2
BS. _Santa Clara Co._: Palo Alto, 3 BS.

_Marginal records._--CALIFORNIA: Berkley, _Elmhurst_; _Palo Alto_; San
Mateo.


=Sorex vagrans paludivagus= von Bloeker

    _Sorex vagrans paludivagus_ von Bloeker, Proc. Biol. Soc.
    Washington, 52:93, June 5, 1939.

_Type._--Adult male, skin and skull; No. 5053, Los Angeles Museum of
History, Science and Art, obtained on November 3, 1938, by Jack C. von
Bloeker, Jr., from salt marsh at mouth of Elkhorn Slough, Moss
Landing, Monterey Co., California, original no. 9456.

_Diagnosis._--Size medium for the species; average and extreme
measurements of 6 topotypes are: total length, 115 (113-118); tail,
46.5 (42-48); hind foot, 14.5 (14-15) (von Bloeker, 1939:94). In
winter nearly black dorsally, deep mouse gray ventrally; in summer
nearly as dark dorsally as in winter, hairs of venter tipped with
Clove Brown; skull short, relatively broad cranially and relatively
narrow rostrally.

_Comparisons._--For comparison with _S. v. halicoetes_ see account of
that subspecies.

_Remarks._--This subspecies, occurring at the limits of the range of
the species, is uncommon in most collections. Seven specimens were
available for the original description. The summer pelage is not
completely described in the original description, but is stated to be
darker than the winter pelage of _S. v. vagrans_, and must thus be
considerably darker than the summer pelage of _S. v. halicoetes_. Two
specimens in the Museum of Vertebrate Zoology, from San Gregario,
referred by Jackson to _S. v. halicoetes_, were included in the
present subspecies by von Bloeker.

_Specimens examined._--None.

_Records of occurrence_ (von Bloeker, 1939:94).--CALIFORNIA: _San
Mateo Co._: San Gregario. _Monterey Co._: Seaside; mouth of Salinas
River; Moss Landing.

_Marginal records._--CALIFORNIA: San Gregario; Seaside.


=Sorex vagrans vancouverensis= Merriam

   _Sorex vancouverensis_ Merriam, N. Amer. Fauna, 10:70,
   December 31, 1895.

   _Sorex vagrans vancouverensis_, Jackson, N. Amer. Fauna,
   51:106, July, 1928.

_Type._--Adult male, skin and skull; No. 71913, U. S. Biol. Surv.
Coll.; obtained on May 10, 1895, by Clark P. Streator, from
Goldstream, Vancouver Island, British Columbia.

_Range._--Vancouver Island from Sayward south, and Bowen Island.

_Diagnosis._--Size small for the species; average and extreme
measurements of 6 specimens from Alberni Valley, Vancouver Island,
are: total length, 106.5 (97-115); tail, 41.7 (40-43); hind foot, 12
(11-13) (Jackson, 1928:107). Ventral parts brownish, winter pelage
reddish brown rather than grayish.

_Comparisons._--Differs from _S. v. vagrans_ in more brownish ventral
parts and more brownish, rather than grayish, winter pelage; differs
from the sympatric _S. v. isolatus_ in shorter tail, shorter hind
foot, more narrow skull, and smaller teeth.

_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


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    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
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  BAILEY, V.
    1936.    The mammals and life zones of Oregon. N. Amer. Fauna,
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  BROWN, B.
    1908.    The Conard Fissure, a Pleistocene bone deposit in northern
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    1950.    Contribution to the taxonomy and life history of _Sorex
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  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
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    1941.    Insularity in the genus Sorex on the north coast of British
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  DALQUEST, W. W.
    1941.    Ecologic relationships of four small mammals in western
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    1944.    The molting of the wandering shrew. Jour. Mamm., 25:146-148,
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    1948.    Mammals of Washington. Univ. Kansas Publ., Mus. Nat. Hist.,
             2:1-444, 140 figs. in text, April 9.

  DAVIS, W. B.
    1939.    The Recent Mammals of Idaho. The Caxton Printers, Ltd.,
             Caldwell, Idaho. Pp. 1-400, 33 figs. in text, 2 pls., April 5.

  DURRANT, S. D.
    1952.    Mammals of Utah, taxonomy and distribution. Univ. Kansas
             Publ., Mus. Nat. Hist., 6:1-549, 91 figs. in text, 30 tables,
             August 10.

  FINDLEY, J. S.
    1953.    Pleistocene Soricidae from San Josecito Cave, Nuevo Leon,
             Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 5:633-639,
             December 1.

    1955.    Taxonomy and distribution of some American shrews. Univ.
             Kansas Publ., Mus. Nat. Hist., 7:613-618, June 10.

  FITCH, H. S.
    1940.    A biogeographical study of the ordinoides artenkreis of garter
             snakes (genus Thamnophis). Univ. California Publ. Zool.,
             44:1-150, October 31.

  GRINNELL, J.
    1933.    Review of the Recent mammal fauna of California. Univ.
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  GRINNELL, J., and A. H. MILLER.
    1944.    The distribution of the birds of California. Pacific Coast
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  HALL, E. R.
    1938.    Variation among insular mammals of Georgia Strait, British
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    1946.    Mammals of Nevada. University of California Press, Berkeley
             and Los Angeles, pp. i-xi + 1-710, 11 pls., 485 figs. in text,
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  HAMILTON, W. J., JR.
    1940.    The biology of the smoky shrew (_Sorex fumeus fumeus_ Miller).
             Zoologica, 25:473-492, 4 pls., 1 fig. in text, 2 tables.

  HIBBARD, C.
    1944.    Stratigraphy and vertebrate paleontology of Pleistocene
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  HOWELL, T. R.
    1952.    Natural history and differentiation in the yellow-bellied
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  JACKSON, H. H. T.
    1928.    A taxonomic review of the American long-tailed shrews
             (genera Sorex and Microsorex). N. Amer. Fauna, 51:i-vi +
             1-238, 13 pls., 24 figs. in text, July 24.

    1947.    A new shrew (genus Sorex) from Coahuila. Proc. Biol. Soc.
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  KRUTZSCH, P. H.
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             Mus. Nat. Hist., 7:349-472, 47 figs. in text, 4 tables,
             April 21.

  MACNAB, J. A., and J. C. DIRKS.
    1941.    The California red-backed mouse in the Oregon Coast Range.
             Jour. Mamm., 22:174-180, May 14.

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    1940.    Speciation phenomena in birds. Amer. Nat., 74:249-278.

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    1895.    Synopsis of the American shrews of the genus Sorex. in
             N. Amer. Fauna, 10:57-100, December 31.

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             October 28.

  MURIE, A.
    1933.    The ecological relationship of two species of _Peromyscus_
             in the Glacier Park region, Montana. Occ. Pap. Mus. Zool.,
             Univ. Michigan, 270:1-17, 2 figs., 3 tables, July 1.

  PEARSON, O. P.
    1945.    Longevity of the short-tailed shrew. Amer. Midl. Nat.,
             34:531-546, 2 tables, 4 figs. in text, September.

  PRUITT, W. O., JR.
    1954.    Aging in the masked shrew, _Sorex cinereus cinereus_ Kerr.
             Jour. Mamm., 35:35-39, February 10.

  RENSCH, B.
    1933.    Zoologische systematik und artbildungsproblem. Ver. deutsch.
             zool. Gesellschaft, 1933:19-83.

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    1912.    Color standards and color nomenclature. Washington, D. C.,
             privately printed, i-iv + 1-44, 53 pls.

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    1953.    Differentiation in shrews of the tidal marshes of the San
             Francisco Bay region. Summary of the dissertation for the
             degree of Doctor of Philosophy, University of California
             Graduate Division, 4 pages, unnumbered, June.

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             mammals. Bull. American Mus. Nat. Hist., 85:i-xvi + 1-350,
             October 5.

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    1938.    A systematic review of the genus Phylloscopus. British Mus.,
             London, i-viii + 1-193, 8 maps, 2 pls., November 26.

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             17:96-125, 1 map, June.



UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY


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  * An asterisk designates those numbers of which the Museum's
  supply (not the Library's supply) is exhausted. Numbers published
  to date, in this series, are as follows:

  Vol. 1, Nos. 1-26 and index. Pp. 1-638, 1946-1950.

 *Vol  2. (Complete) Mammals of Washington. By Walter W. Dalquest.
          Pp. 1-444, 140 figures in text. April 9, 1948.

  Vol. 3. *1. The avifauna of Micronesia, its origin, evolution, and
              distribution. By Rollin H. Baker. Pp. 1-359, 16 figures
              in text. June 12, 1951.

          *2. A quantitative study of the nocturnal migration of birds.
              By George H. Lowery, Jr. Pp. 361-472, 47 figures in text.
              June 29, 1951.

           3. Phylogeny of the waxwings and allied birds. By M. Dale
              Arvey. Pp. 473-530, 49 figures in text, 13 tables.
              October 10, 1951.

           4. Birds from the state of Veracruz, Mexico. By George H.
              Lowery, Jr. and Walter W. Dalquest. Pp. 531-649,
              7 figures in text, 2 tables. October 10, 1951.

          Index. Pp. 651-681.

 *Vol. 4. (Complete) American weasels. By E. Raymond Hall. Pp. 1-466,
          41 plates, 31 figures in text. December 27, 1951.

  Vol. 5.  1. Preliminary survey of a Paleocene faunule from the
              Angels Peak area, New Mexico. By Robert W. Wilson.
              Pp. 1-11, 1 figure in text. February 24, 1951.

           2. Two new moles (Genus Scalopus) from Mexico and Texas.
              By Rollin H. Baker. Pp. 17-24. February 28, 1951.

           3. Two new pocket gophers from Wyoming and Colorado.
              By E. Raymond Hall and H. Gordon Montague. Pp. 25-32.
              February 28, 1951.

           4. Mammals obtained by Dr. Curt von Wedel from the barrier
              beach of Tamaulipas, Mexico. By E. Raymond Hall.
              Pp. 33-47, 1 figure in text. October 1, 1951.

           5. Comments on the taxonomy and geographic distribution of
              some North American rabbits. By E. Raymond Hall and
              Keith R. Kelson. Pp. 49-58. October 1, 1951.

           6. Two new subspecies of Thomomys bottae from New Mexico
              and Colorado. By Keith R. Kelson. Pp. 59-71, 1 figure in
              text. October 1, 1951.

           7. A new subspecies of Microtus montanus from Montana and
              comments on Microtus canicaudus Miller. By E. Raymond
              Hall and Keith R. Kelson. Pp. 73-79. October 1, 1951.

           8. A new pocket gopher (Genus Thomomys) from eastern
              Colorado. By E. Raymond Hall. Pp. 81-85. October 1, 1951.

           9. Mammals taken along the Alaskan Highway. By Rollin H.
              Baker. Pp. 87-117, 1 figure in text. November 28, 1951.

         *10. A synopsis of the North American Lagomorpha. By E.
              Raymond Hall. Pp. 119-202, 68 figures in text.
              December 15, 1951.

          11. A new pocket mouse (Genus Perognathus) from Kansas.
              By E. Lendell Cockrum. Pp. 203-206. December 15, 1951.

          12. Mammals from Tamaulipas, Mexico. By Rollin H. Baker.
              Pp. 207-218. December 15, 1951.

          13. A new pocket gopher (Genus Thomomys) from Wyoming and
              Colorado. By E. Raymond Hall. Pp. 219-222.
              December 15, 1951.

          14. A new name for the Mexican red bat. By E. Raymond Hall.
              Pp. 223-226. December 15, 1951.

          15. Taxonomic notes on Mexican bats of the Genus Rhogeëssa.
              By E. Raymond Hall. Pp. 227-232. April 10, 1952.

          16. Comments on the taxonomy and geographic distribution of
              some North American woodrats (Genus Neotoma). By Keith R.
              Kelson. Pp. 233-242. April 10, 1952.

          17. The subspecies of the Mexican red-bellied squirrel,
              Sciurus aureogaster. By Keith R. Kelson. Pp. 243-250,
              1 figure in text. April 10, 1952.

          18. Geographic range of Peromyscus melanophrys, with
              description of new subspecies. By Rollin H. Baker.
              Pp. 251-258, 1 figure in text. May 10, 1952.

          19. A new chipmunk (Genus Eutamias) from the Black Hills.
              By John A. White. Pp. 259-262. April 10, 1952.

          20. A new piñon mouse (Peromyscus truei) from Durango,
              Mexico. By Robert B. Finley, Jr. Pp. 263-267.
              May 23, 1952.

          21. An annotated checklist of Nebraskan bats. By Olin L.
              Webb and J. Knox Jones, Jr. Pp. 269-279. May 31, 1952.

          22. Geographic variation in red-backed mice (Genus
              Clethrionomys) of the southern Rocky Mountain region.
              By E. Lendell Cockrum and Kenneth L. Fitch. Pp. 281-292,
              1 figure in text. November 15, 1952.

          23. Comments on the taxonomy and geographic distribution of
              North American microtines. By E. Raymond Hall and
              E. Lendell Cockrum. Pp. 293-312. November 17, 1952.

          24. The subspecific status of two Central American sloths.
              By E. Raymond Hall and Keith R. Kelson. Pp. 313-337.
              November 21, 1952.

          25. Comments on the taxonomy and geographic distribution of
              some North American marsupials, insectivores, and
              carnivores. By E. Raymond Hall and Keith R. Kelson.
              Pp. 319-341. December 5, 1952.

          26. Comments on the taxonomy and geographic distribution of
              some North American rodents. By E. Raymond Hall and
              Keith R. Kelson. Pp. 343-371. December 15, 1952.

          27. A synopsis of the North American microtine rodents.
              By E. Raymond Hall and E. Lendell Cockrum. Pp. 373-498,
              149 figures in text. January 15, 1953.

          28. The pocket gophers (Genus Thomomys) of Coahuila, Mexico.
              By Rollin H. Baker. Pp. 499-514, 1 figure in text.
              June 1, 1953.

          29. Geographic distribution of the pocket mouse, Perognathus
              fasciatus. By J. Knox Jones, Jr. Pp. 515-526, 7 figures
              in text. August 1, 1953.

          30. A new subspecies of wood rat (Neotoma mexicana) from
              Colorado. By Robert B. Finley, Jr. Pp. 527-534, 2 figures
              in text. August 15, 1953.

          31. Four new pocket gophers of the genus Cratogeomys from
              Jalisco, Mexico. By Robert J. Russell. Pp. 535-542.
              October 15, 1953.

          32. Genera and subgenera of chipmunks. By John A. White.
              Pp. 543-561, 12 figures in text. December 1, 1953.

          33. Taxonomy of the chipmunks, Eutamias quadrivittatus and
              Eutamias umbrinus. By John A. White. Pp. 563-582,
              6 figures in text. December 1, 1953.

          34. Geographic distribution and taxonomy of the chipmunks of
              Wyoming. By John A. White. Pp. 584-610, 3 figures in text.
              December 1, 1953.

          35. The baculum of the chipmunks of western North America.
              By John A. White. Pp. 611-631, 19 figures in text.
              December 1, 1953.

          36. Pleistocene Soricidae from San Josecito Cave, Nuevo Leon,
              Mexico. By James S. Findley. Pp. 633-639. December 1, 1953.

          37. Seventeen species of bats recorded from Barro Colorado
              Island, Panama Canal Zone. By E. Raymond Hall and
              William B. Jackson. Pp. 641-646. December 1, 1953.

           Index. Pp. 647-676.

 *Vol. 6.  (Complete) Mammals of Utah, _taxonomy and distribution_.
           By Stephen D. Durrant. Pp. 1-549, 91 figures in text,
           30 tables. August 10, 1952.

  Vol. 7. *1. Mammals of Kansas.  By E. Lendell Cockrum. Pp. 1-303,
              73 figures in text, 37 tables. August 25, 1952.

           2. Ecology of the opossum on a natural area in northeastern
              Kansas. By Henry S. Fitch and Lewis L. Sandidge.
              Pp. 305-338, 5 figures in text. August 24, 1953.

           3. The silky pocket mice (Perognathus flavus) of Mexico.
              By Rollin H. Baker. Pp. 339-347, 1 figure in text.
              February 15, 1954.

           4. North American jumping mice (Genus Zapus). By Philip H.
              Krutzsch. Pp. 349-472, 47 figures in text, 4 tables.
              April 21, 1954.

           5. Mammals from Southeastern Alaska. By Rollin H. Baker and
              James S. Findley. Pp. 473-477. April 21, 1954.

           6. Distribution of Some Nebraskan Mammals. By J. Knox Jones,
              Jr. Pp. 479-487. April 21, 1954.

           7. Subspeciation in the montane meadow mouse, Microtus
              montanus, in Wyoming and Colorado. By Sydney Anderson.
              Pp. 489-506, 2 figures in text. July 23, 1954.

           8. A new subspecies of bat (Myotis velifer) from
              southeastern California and Arizona. By Terry A. Vaughan.
              Pp. 507-512. July 23, 1954.

           9. Mammals of the San Gabriel mountains of California.
              By Terry A. Vaughan. Pp. 513-582, 1 figure in text,
              12 tables. November 15, 1954.

          10. A new bat (Genus Pipistrellus) from northeastern Mexico.
              By Rollin H. Baker. Pp. 583-586. November 15, 1954.

          11. A new subspecies of pocket mouse from Kansas. By
              E. Raymond Hall. Pp. 587-590. November 15, 1954.

          12. Geographic variation in the pocket gopher, Cratogeomys
              castanops, in Coahuila, Mexico. By Robert J. Russell and
              Rollin H. Baker. Pp. 591-608. March 15, 1955.

          13. A new cottontail (Sylvilagus floridanus) from
              northeastern Mexico. By Rollin H. Baker. Pp. 609-612.
              April 8, 1955.

          14. Taxonomy and distribution of some American shrews.
              By James S. Findley. Pp. 613-618. June 10, 1955.

          15. The pigmy woodrat, Neotoma goldmani, its distribution
              and systematic position. By Dennis G. Rainey and Rollin
              H. Baker. Pp. 619-624, 2 figs. in text. June 10, 1955.

           Index. Pp. 625-651.

  Vol. 8.  1. Life history and ecology of the five-lined skink,
              Eumeces fasciatus. By Henry S. Fitch. Pp. 1-156, 26 figs.
              in text. September 1, 1954.

           2. Myology and serology of the Avian Family Fringillidae, a
              taxonomic study. By William B. Stallcup. Pp. 157-211,
              23 figures in text, 4 tables. November 15, 1954.

           More numbers will appear in volume 8.

  Vol. 9.  1. Speciation of the wandering shrew. By James S. Findley.
              Pp. 1-68, 18 figures in text. December 10, 1955.

           More numbers will appear in volume 9.



TRANSCRIBER'S NOTES


Except for the movement of the list of publications to the end, the
typographical corrections noted below and a number of minor corrections
not detailed, the text is the same as the original printed version.

Whole and fractional parts of numbers are displayed as follows:
8-3/4 = eight and three quarters; 10-1/2 = ten and one half; etc.


Typographical Corrections

  Page          Correction
  ====   =============================
  13      predeliction => predilection
  36      Clallum => Clallam
  37      Mt. Ranier => Mt. Rainier
  39      Towsend => Townsend
  41      Admiraltry => Admiralty
  49      Okanagon => Okanagan
  57      Lookinglass => Lookingglass
  64      Popoff Is. => Popof Is.
  ii      Vaughn => Vaughan


Emphasis Notation

  _Text_ - Italics

  =Text= - Bold





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