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OCCASIONAL PAPERS

MUSEUM OF NATURAL HISTORY
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NUMBER 68, PAGES 1-35 JULY 15, 1977

A REVIEW OF THE TAXONOMY OF THE

SOREX VAGRANS SPECIES COMPLEX

FROM WESTERN NORTH AMERICA

By
Darwen Hennings^ and Robert S. Hoffmann"

Before the publication of Findley's monograph, Speciation of
the Wandering Shreiv (1955a), the Sorex vagrans complex was
thought to consist of at least three closely related species: Sorex
vagrans, S. obscurus, and S. pacificus. Jackson (1928) separated
these species on the basis of size and pelage color; throughout much
of the Pacific Northwest at least two of the taxa can be collected at
the same localities. However, in the northern Rocky Mountains,
especially in western Montana, vagrans and ohscurus are so similar
in size and pelage color that mammalogists found them impossible
to distinguish (Clothier, 1950; R. S. Peterson, pers. comm.; P. L.
Wright, pers. comm. ) . Findley ( 1955a ) interpreted these similari-
ties as an indication that barriers to interbreeding between vagrans
and obscurus had broken down along much of their zone of contact.
He speculated that the two forms had a connnon origin prior to the
Wisconsin glaciation and had diverged while isolated from one an-
other during the final glacial period. Sorex obscurus appeared to
intergrade also with Sorex pacificus (including S. yaquinae) in
western Oregon. Findley concluded that the three species were in
fact one — a complex ring of overlapping subspecies to which he
applied the name of the earliest described form, Sorex vagrans. He

1 Department of Zoology, University of Montana, Missoula, Montana 59801
(present address, 15623 Jiin Creek Rd., Arlington, Washington 98223).

~ Museum of Natural History and Department of Systematics and Ecology,
The University of Kansas, Lawrence, Kansas 66045.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

compared this "Rassenkreis" to those reported for the Herring Gull,
Larus argentatus (see Mayr, 1963), the Old World titmouse, Pariis
major (Rensch, 1933), and the garter snake, Thamnophis ordinoides
(Fitch, 1940) among others.

Findley distinguished obscuriis from vagrans as subspecies, but
his descriptions do not pemiit easy identification. Mammalogists
working with shrews from the Rocky Mountains and other areas
where the two fonns differ subtly, tended to lump vagrans and
ohscurus together as "Sorex vagrans'^ with the unfortunate result
that important differences between the two were obscured (e.g.,
Brown 1967; Hoffmann and Taber, 1960; Ingles, 1961; Spencer and
Pettus, 1966; in all cases the shrew identified as vagrans was actu-
ally ohscurus ( Fig. 1 ) . Findley 's revision was greeted cautiously by
some. Johnson and Ostenson (1959) suggested that the biological
relationships among the subspecies of Findley 's Sorex vagrans were
too poorly known to justify his sweeping revision of the nomen-
clature, and proposed that the basic framework of Jackson's 1928
classification be upheld until further studies could be made.

Findley's conclusions were based on his interpretation of the
nature of character convergence between vagrans and ohscurus in
the northern Rocky Mountains (1955a: 12-13, 44-45, and 54-55).
We attempted to locate "pure" populations of the two supposed
subspecies, in order to define their differences and clarify their
morphological and ecogeographic relationships (Hennings, 1970).
We found differences in the structure and pigmentation of the up-
per incisor teeth and in cranial dimensions which clearly separated
vagrans and ohscurus, and confirm their specific integrity, as will be
shown.

Sorex monticolus, Merriam 1890, long considered a synonym of
Sorex vagrans, is the oldest available name for the species previ-
ously known as S. ohscurus Merriam, 1895. The familiar name is
applicable only to the most widely distributed subspecies, S. m.
ohscurus (Fig. 2). Sorex pacificus is morphologically distinct from

Fig. 1. — Sorex monticolus ohscurus, Sierra Nevada, Fresno Co., Calif.
(Adapted from a photo in Ingles, 1961, where the shrew was identified as Sorex
vagrans.) X 4/5.

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 3

S. monticohis (=S. ohsctinis), but as Findley observed, there ap-
pears to be a narrow zone of hybridization between the two taxa.
We discuss certain taxonomic problems which may arise as the
nature of their relationship becomes more fully understood. In the
rest of this paper we use the names Sorex monticohis (=zol)scurus),
S. pacificus and S. vo grans.

Materials and Methods

Hennings collected, at key localities in western Montana, 94
Sorex V. vagrans and S. nionticolus obscnrus, using snap-traps and
sunken cans (Hennings, 1970). Subsequently, we also examined
2,502 additional specimens, including all subspecies of Sorex va-
grans, S. monticohis and S. pacificus (including S. yaqiiinae), as
well as S. durangae, which Jackson (192S) included in the Sorex
vagrans group, and S. trigonirostris, which he allied with S. ornatus
(Fig. 2). Specimens were borrowed from the following collections
(see also Specimens Examined, Appendix): Brigham Young Uni-
versity Life Sciences Museum ( 13YU ) ; California Academy of Sci-
ences (CAS); California State University, Long Beach (CSLB);
Carol Terry (CT); Dartmouth College Museum"(DCM); Harvard
University, Museum of Comparative Zoology (MCZ); Michigan
State University, The Museum (MSU); Montana State University,
Bozeman, Department of Zoology (MSUDZ); National Museum of
Natrual History (USNM); National Museum of Natural Science,
Ottawa (NMC); San Diego Natural History Musevmi (SDSNH);
Southern Oregon College (SOMNH); Stanford University Collec-
tion, at the California Academy of Sciences (CAS-SU); University
of Alberta, Museum of Zoology (UAMZ); University of British Co-
lumbia Vertebrate Museum (UBC); University of California, Berke-
ley, Museum of Vertebrate Zoology (MVZ); University of Kansas,
Museum of Natural History (KU); University of Montana, Zoolog-
ical Museum, Missoula (UM); Universidad National Autonoma de
Mexico (UNAM); University of Utah (UU); Washington State
University, Charles R. Connor Museum (CRCM).

In the initial phase of the study 40S post-juvenile S. v. vagrans
and 115 S. monticohis ohscnriis from the northern Rocky Mountains
and Columbia Plateau regions were selected for statistical analysis.
Specimens were identified by body measurements and by tooth
characteristics described below. Six cranial dimensions described
by Findley ( 1955a : 5 ) , plus cranial height, were measured with
Helios metric dial calipers. Age was determined by Rudd's (1955a)
subjective toothwear method.

Data from the largest homogeneous population of each species
were grouped by age and month of capture and analyzed statisti-
cally. ( Sex differences in Sorex have no significant effect on cranial

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 5

morphology; see Rudd, 1955a.) Of the six measurements used by
Findley only one, maxillary tooth-row length, varied significantly
with age. Cranial height varied according to the patterns noted by
Dehnel (1949), Pucek (1955, 1957), and Dapson (1968). (See dis-
cussion in Hennings, 1970).

Hennings ( 1970 ) grouped specimens into several geographical
populations, the cranial dimensions of which were compared sta-
tistically. For this report we have regrouped some of the smaller
samples (Fig. 3), and will compare data from only the three most
variable measurements; condylobasal length, palatal length, and
least interorbital breadth ( Fig. 7, A ) . Pelage color and body meas-
urements were not analyzed in detail (but see Hennings, 1970).

Acknowledgements

We thank the Department of Zoology, and the Biological Station
of the University of Montana for use of facilities and equipment
which made this study possible. Hennings' graduate studies were
financed in part by an NDEA Title IV Fellowship. Phillip Wright,
Donald Jenni, Lee Metzgar, Oliver Pearson, Seth Benson, Robert
Rudd, R. James Brown, James Patton, Bill Lidicker, Robert Orr,
James Findley, Murray Johnson, Chris Maser, Don Wilson, Myrnal
and David Hawes, Matt Greenstone, J. Maiy Taylor, Clyde Pritch-
ett, Victor Diersing, Donald Hoffmeister, and the late Richard S.
Peterson, at various times and in various ways gave us advice, en-
couragement and technical assistance. We are grateful to the cura-
tors of institutions listed previously, who have graciously loaned us
material. From the senior author, a final word of thanks to my wife
Vally. We dedicate this paper to Darwin and Adriele, children of
Myrnal and David Hawes.

Analysis of Morphological Characters

Jackson (192S) distinguished Sorex ohscurus from Sorex vagrans
on the basis of total length - ohscurus being 110 mm or greater and
vagrans less than 110 mm. Figure 4 is a review of total length
measurements from shrews collected in a number of regions in the
Pacific Northwest. Sorex monticolus ohscurus tends to be larger

Fig. 2. — Approximate distribution of tlie races of Sorex vagrans, S. monti-
colus, and S. pacificus (modified from Findley, 1955a). The range of S. monti-
colus ohscurus is stippled darkly; tliose of other monticolus subspecies, and of
pacificus are lightly stippled. (The boundary between ohscurus and coastal
subspecies in Canada and Alaska is based in part on vegetation patterns and is
somewhat speculative. ) The range of S. vagrans is cross-hatched. Ranges of
S. V. vagrans and S. m. ohscurus are considered in more detail in Fig. 3 (in-
set area ) .

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

pad

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 7

than S. V. vagrans, but the overlap is so broad that the 110 mm
separation point is unworkal)le.

Heptner and Dolgov (1967) and Yudin (1969), described the
medial tines on the first upper incisors of the Asian shrew Sorex
mirabilis, and of S. v. vagrans from Montana. Sorex pacificus, with
which mirabilis had been synonymized (Brobinskii et ah, 1944),
appears typically to lack this stiiicture ( Heptner and Dolgov, 1967;
Hoffmann, 1971; see Fig. 5C). Sorex vagrans and S. nionficolus can
be distinguished easily by differences in the position of the medial
tines and their relationship to the red pigment on the first upper
incisors ( Fig. 5A, B ) . In vagrans the tines are typically small, and
located near the upper limit of the pigment; they usually are set off
distinctly by a break in the color. In monticolus the tines are typi-
cally large, and situated well below the top of the pigment, which
is darker and more extensive than that of vagrans. In live-trapping
S. V. vagrans and S. monticolus setosus in British Columbia, Myrnal
Hawes has noticed that the teeth of monticolus are more robust,
harder and more brittle than those of vagrans. Thus, they appear
to wear more slowly but break more easily than do those of vagrans
(pers. comm.).

These differences in tooth pattern remain unchanged with age,
although they may be obscured in old shrews as the teeth break and
the pigment wears away ( Fig. 5A, B ) . The two patterns do not
grade into each other. Measurements with optical micrometer of
tine-pigment relationships in 83 S. v. vagrans and 57 S. monticolus
obscurus from two homogeneous allopatric populations showed no
overlap (Hennings, 1970). We have distinguished vagrans and
monticolus with a hand lens while live-trapping in Montana, al-
though the technique is difficult to apply to old adults. Mymal and
Dave Hawes (pers. comm.) have had similar success in British
Columbia, with a population that Cowan and Guiguet (1956:52)
thought "demonstrates complete intergradation between S. m.
setosus . . . and S. v. vagrans. suggesting conspecificity."

Sorex monticolus generally is larger than S. vagrans, even in
western Montana where size differences between the subspecies
S. V. vagrans and S. m. obscurus are slight. There is as much over-

FiG. 3. — Approximate distribution of Sorex vagrans (cross-hatched), S.
monticolus and S. pacificus (stippled), with details of tlie synipatric ranges of
S. V. vagrans and S. m. obscurus in the intermountain West. Selected speci-
mens of vagrans examined are represented by hollow triangles; monticolus, by
intact circles; pacificus, by broken circles; not all specimens examined are
plotted. Ranges are based on literature and collection records combined with
vegetation patterns from U.S. and Canadian National Atlases. Symbols outside
shaded areas are records from insular montane populations adjacent to areas of
continuous distribution. Localities enclosed by dashed lines comprise the popu-
lations used in statistical analysis of cranial morphology (Figs. 6 and 7).

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

lap in condylobasal length, which appears to be a function of body
size, as there is in total length ( Fig. 6A ) . However, differences in
some skull proportions are conspicuous. The skull of S. m. obscurus
is significantly broader through the orbits than that of S. v. vagrans
(Fig. 6B), and the palate is significantly longer (Fig. 6C). A tri-
variate analysis, plotting the three proportions as percentages of
their total, shows virtually no overlap between the two species
(Fig. 7B).

Systematic Accounts

The following synonymies include only those references relevant
to the scope of this report. Fonnal treatment centers on the two
most widely distributed taxa in the Sorex vagrans complex, S. v.
vagrans and S. monticolus obscurus, which were studied intensively
in western Montana. Other subspecies are discussed briefly, as is
S. pacificus.

Sorex vagrans Baird
Vagrant or Wandering Shrew

Sorex vagrans Baird, 1858:15. (Type locality, Shoalwater Bay, Pa-
cific Co., Washington ) .

Sorex suckleyi Baird, 1858:18. (Type locahty, Steilacoom, Pierce
Co., Washington).

Sorex amoenus Merriam, 1895:69. (Type localit)% near Mammoth,
8000 ft. E. slope Sierra Nevada, Mono Co., California).

Sorex nevadensis Merriam, 1895:71. (Type locality, Reese R., 6000
ft., Nye-Lander Co. Line, Nevada).

Sorex shastensis Merriam, 1899:87. (Type locality, Mt. Shasta, 5700
ft., Siskiyou Co., California).

Sorex trigonirostris Jackson, 1922:264. (Type locality, Ashland, 1975
ft., Jackson Co., Oregon).

Holotype. — Adult male, alcoholic; USNM 1675, from Shoalwater
(Willapa) Bay, Pacific Co., Washington.

Range. — The subspecies Sorex vagrans vagrans is found in ripar-
ian and montane habitats throughout the northern Great Basin and
the Columbia Plateau, north to southern British Columbia and east
almost to the Continental Divide in Montana, the Wasatch Moun-
tains in Utah, and barely entering Wyoming; coastal mountains of
Washington, Oregon and California, as far south as the San Fran-
cisco Bay and the east-central Sierra Nevada (Fig. 3). One sub-
species, vancouverensis, is found on \'ancouver Island and its
nearby offshore islands. Sorex vagrans Jialicoefcs and S. v. paludi-
vagus are restricted to the southern San Francisco Bay and Monte-
rey Bay Region. Sorex vagrans orizabae appears to be an isolated

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 9

80

90

TOTAL LENGTH
100 110 120

V. vagrans

TOPOTYPES (W. WASH.)

SE. WASHINGTON

E. OREGON

S. IDAHO (SNAKE R.)
SW. MONTANA
NW. MONTANA

SW. MONTANA

NW. MONTANA

CENT. IDAHO

m. obscurus TOPOTYPES
(CENT. IDAHO)

NW. WYOMING
(BEARTOOTH PLATEAU)

m. longiquusTOPOJYPES
(CENT. MONTANA)

S. CENT. MONTANA

m. obscurus

E. OREGON, S. IDAHO,
N. NEVADA

(INSULAR MT. RANGES)

130

80

90

100

110

120

130

140

1 I
VAGRANS

1 1 1

1 8

33

-

-

24

1

' 18

54

-

89

MONTICOLUS

8

-

20

-

1 19

-

9

-

66

-

3
8

13

-

-

1 1 1

1 1

140

Fig. 4. — Range and mean of total length (mm) from intermountain popu-
lations of Sorex vagrans vagrans and S. 7notiticolus obscurus. Sample size is
indicated along tlie right margin. Samples are grouped by region, and are
somewhat broader than the statistical populations identified in Figure 3.

relict restricted to coniferous forests on the Transverse Volcanic Belt
along the southern edge of the Mexican Plateau (Goldman, 1951)

(Fig. 2).

Diagnosis. — Small to medium-sized shrews of the subgenus Oti-
sorex; third unicuspid tooth smaller than fourth, and foramen mag-
num situated more or less ventrally ( Findley, 1955a ) . Medial tines
on first upper incisors situated at approximately the upper edge of
the pigment; set off distinctly by a gap in color (Fig. 5, A). This
character is fairly uniform throughout the range of Sorex vagrans,
and in S. v. vancouverensis and S. t;. haUcoetes, although upper in-

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

VAGRANS

MONTICOLUS

\ c

H

■' Fig. 5. — Frontal views of first upper incisors from Sorex vagrans, S. monti-
cohis, S. pacificus, and related shrews in the subgenus Otisorex: ( A. and B. )
S. V. vagrans (Hamilton, Mont.) and S. m. obscitrus ( Beartooth Plateau),
compared by Rudd age classes 1-4. Note differences in structure of medial
tines (circled). (Specimens, A: DCM 3030, 3076, 2922, 2993; B: UM, 7186,
8954, 7185, 8188); (C.) S. pacificus pacificus, Requa, Calif. (CAS .3810);
(D.) S. p. yaquinae x? S. m. hairdi, "hybrid", Taft, Ore. ( MVZ 71219); (E.)
S. monticolus longicauda, Wrangel, Alaska. (CAS 9196); (F.) S. v. vagrans,
Cassel, Calif. (USNM, BS 49127), hybrid-like pattern; (G.) S. cinereus
cinereus, Atlin, B.C. (CAS 7436); (H.) S. ornatus calif ornicus. Piedmont,
Alameda Co., Calif. (CAS 3941); (I.) S. pribilofensls, St. Paul Is., Alaska
(CAS 2869); and (J.) S. "durangae", (=m. monticolus). El Salto, Durango,
Mex. (USNM, 94539).

cisors of some S. v. vagrans in northeastern California, southern
Idaho and northern Utah are more heavily pigmented and have

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 11

larger tines (Fig. 5F). This is also true of the one specimen of S. v.
paludivagus examined. This deviation occurs sporadically in other
populations, and will be discussed below. The geographically iso-
lated subspecies, S. v. orizabae, has a typical vagrans pattern; al-
though Findley (1955a) felt it was little differentiated from S. mon-
ticolus, Jackson (192S) correctly allied it with Sorex vagrans.

Among populations of S. vagrans vagrans, total length is quite
variable, ranging from 86 mm to at least 120 mm, with means gen-
erally from 98-108 mm. (compared with means of 100-112 mm in
S. monticolns obscurus. Condylobasal length varies similarly, rang-
ing from 15.5 to 17.5 mm in the populations examined, with means
16.2-16.9 ( Hennings, 1970). Larger skulls are found among mon-
tane populations. Least interorbital breadth provides a real measure
of distinction between S. v. vagrans and S. m. obscurus. Although it
ranges from 2.7 to 3.5 mm, the least interorbital breadth of at least
80 percent of all specimens of vagrans is below 3.2 mm; in contrast,
about the same percentage of obscurus have least interorbital
breadths greater than 3.2 mm. (See Fig. 6B; estimate based on over-
lap of standard deviations.) Palatal length in S. v. vagrans ranges
from 6.1 to 7.3 mm, but is less than 7.0 mm in at least 80 percent
of the specimens; S. m. obscurus is generally greater than 7.0 mm
(Fig. 6C). See Findley ( 1955a) for size and color variation in other
subspecies.

Remarks. — Findley (1955a) was the first to recognize that the
small shrew of the northern intermountain region, S. v. vagrans,
was not the same as Sorex "vagrans" nionticola from Arizona. He
also united the several populations which Jackson (1928) had
treated as subspecies with vagrans. West of the Cascades, south
along the Pacific Coast, and into the Sierra Nevada and Great Basin,
S. V. vagrans is found in a variety of habitats. On the Columbia
Plateau it occupies riparian habitats within grassland, sagebiTish
prairie and ponderosa pine forest. Where it enters the mountains in
northern Nevada, Oregon, ^Vashington, Idaho, and western Mon-
tana and Wyoming, the subspecies is usually found in forests of
Douglas fir, lodgepole pine, western larch, grand fir, western hem-
lock, and red cedar, at elevations below 5,000 ft. Sorex monticolus
obscurus, S. c. cinereus, S. palustris navigator, and Microsorex hoiji
toashingtoni may live in geographic sympatry in this area.

Sorex V. vagrans from steppe and arid forest in Washington,
Oregon, and Idaho are smaller than the typical fonii from the
Washington coast (Fig. 4). The so-called "Oregon dwarf shrew,"
S. trigonirostris, is a small S. v. vagrans. Comparison of the t\'pe,
and a near-topotype on which Jackson ( 1928 ) based the name, with
series of vagrans to the east and west, as well as other specimens
from Jackson County indicates that although the type specimen is
at the lower end of the size range for S. v. vagrans (condylobasal

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

length, 15.6 mm), other specimens from the area show nonnal size
variation. Characteristics of the incisors, unicuspids, and foramen
magnum position also agree with those of S. vagrans, and distin-
guish " trigonhostris" from the ornatus group, in which it was placed
by Jackson (1928).

The largest S. v. vagrans are found in moist grassland and forest
in western Montana, where winters in the shrew's microhabitat are
less severe than on the more barren Columbia Plateau. Mezhzherin
(1964) demonstrated that size variation in shrews does not follow
Bergmann's Rule; shrews from colder, more severe, climates are
smaller rather than larger than those from regions of milder winters.
This appears to be an adaptation to decreased food supply and
increased metabolic demands in regions of colder weather. Gebc-
zynski (1965) suggested that seasonal size reduction in individual
shrews (Dehnel, 1949; Pucek, 1963, 1965) has a similar adaptive
function.

Sorex monticolus Merriam
Montane or Duskv Shrew

Sorex monticolus Merriam, 1890:43. (Type locality, San Francisco
Mtn., 11,500 ft., Coconino Co., Arizona).

Sorex dohsoni Merriam, 1891:33. (Type locality, Altin-as Lake, E
base Sawtooth Mts., 7200 ft., Blaine Co., Idaho; Nomen ohlitum,
see below).

Sorex vagrans similus Merriam, 1891:34. (Type locality, near Tim-
ber Cr., 8200 ft., Lemhi Mts., Lemhi Co., Idaho). Name pre-
occupied by Sorex similus Hensel, 1855:459 (=Neomijs similus).

Sorex ohscurus Merriam, 1895:72. (New name for S. vagrans similus
Merriam, 1891; see below for validity of ohscurus as a sub-
species name).

Sorex vagrans ohscurus Findley, 1955:54. (New combination).

Sorex vagrans longiquus Findley, 1955:49. (Type locality, 25 mi
ESE Big Sandy, Choteau Co., Montana).

Sorex vagrans ohscuroicles Findley, 1955:58. (Type locality, Bishop
Cr., 6600 ft., Inyo Co., California).

Ho/o/ype.— Adult male, skin and skull, USNM 17599/24535, Biol.
Surv. Coll., from San Francisco Mtn., 11,500 ft., Coconino Co.,
Arizona.

Range. — The nominate race S. m. monticolus is found in the
isolated mountain ranges of eastern Arizona, western New Mexico,

Fig. 6. — Geographical variation in three skull dimensions (mm, see Fig.
7A). The three S. v. vagrans populations from "SW Montana" are from 1)
Missoula and the lower Bitteroot Valley, 2) the Sapphire Range, and 3) Ham-
ilton and vicinity. The "NW Wyoming" S. rn. ohscurus population is from the
Beartooth Plateau ( see map, Fig. 3 ) .

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 13

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and northwestern Mexico. The subspecies S. m. ohscurus occupies
the interior of western North America from central Alaska south to
New Mexico, primarily in the Rocky Mountains. Two geograph-
ically isolated subspecies, S. m. neomexicanus and S. m. soperi, are
probably derived from these Rocky Mountain populations (Fig. 2).
Around the northern and southern edges of the Columbia Plateau
S. m. ohscurus grades into S. m. setosus in the Washington Cas-
cades and approaches S. m. permiUiensis and S. m. hairdii in west-
ern Oregon (Fig. 3). Populations of S. m. ohscurus extend through
the mountains of Utah and northern Nevada, to the Sierra Nevada,
with an isolated montane race, S. m. parvidens, in southern Cali-
fornia (Fig. 2). A series of coastal subspecies, some of which are
characterized by a variant tooth pattern, extends northward from
Vancouver Island to Alaska - isohitus, mixtus, ccdvertensis, insularis,
prevostensis, longicauda, malitiostis, elassodon, aJascensis, and shu-
maginensis - the last also extending into interior Alaska (Fig. 2).
Sorex monticolus is appropriately named; its habitat is mainly mon-
tane, boreal and coastal coniferous forest and alpine barrens.

Diagnosis. — Small to medium size shrews of the subgenus Oti-
sorex, similar to S. vagrans, but with medial tines on the first upper
incisors typically situated well below the top edge of the tooth pig-
ment. The tines are larger, the pigment darker, and the incisors
more robust than those of vagrans (Fig. 5B). The tooth pattern
sometimes approaches that of vagrans among the northwest coastal
races ( Fig. 5E ) ; this confusing anomaly is discussed below.

The races of Sorex monticolus are exceedingly variable. Clines
of increasing and decreasing size, and changes in intensity of pelage
pigmentation proceed in many directions, especially among the
coastal mountain ranges (Findley, 1955a). S. monticohis ohscurus
were as small as 86 mm long, and as large as 128 mm; mean total
length varied from 100-112 mm. Condylobasal length ranged from
16.1 to 17.7 mm, averaging 16.7-17.1 mm. Least interorbital breadth
averaged about 3.3 mm, ranging from 3.0-3.6 mm among the four
populations analyzed statistically. Palatal length ranged from 6.7-
7.7 mm, averaging 7.1-7.3. (See diagnosis of S. v. vagrans for com-
parisons; also Fig. 6A-C.)

Remarks. — The usual habitat of Sorex monticohis ohscurus on
the Columbia Plateau and in the northern Rocky Mountains is high
altitude spruce-fir forest and alpine timdra. ( See U.S. National At-
las, vegetation map: A. W. Kuchler, ed., 1967.) Sorex monticolus
confronts S. vagrans as low as 3,000 ft. in mid-altitude forests of
Douglas fir, lodgepole pine, western larch, and grand fir ( Fig. 3 ) .
It may descend to pinyon-juniper woodland on the northern end of
the Great Basin, but except for the Ruby Range has not been col-
lected from the mountains of central Nevada (see Brown, 1971). It
occurs along streams and rivers in the high prairie east of the Con-

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 15

tinental Divide (where va^rans does not occur) and on isolated
mountain ranges of central Montana and ^^''yoming, whose forests
are of the same t\'pe as those of the Rocky Mountains. No monti-
colus have been collected from the pine forests of eastern Montana
and Wyoming, and western South Dakota.

The incisor tine/pigment configurations we discovered in S.
monficohis ohsciirus is characteristic also of the subspecies ala-
sccnsis, hairdi, inixtus, monticolus, parvidcns, permiUiensis, setosiis,
shiimaginensis, and soperi. Except for monticolus, these forms were
previously described as sub.species of "Sorex ohscurus." S. m. mon-
ticolus was confused by Merriam (1895) and Jackson (1928) with
Sorex vagrans from the northern Rockies. As Findley recognized
(1955a), populations of S.m. monticolus from New Mexico, Arizona,
Chihuahua and Durango appear to represent a southern offshoot of
S. m. ohscurus from the central Rockies. Sorex durcingae, which
Jackson (1928) viewed as ". . . in many respects similar to S. oh-
scurus . . ." is also a synonym of Sorex m. monticohis; its skull and
teeth are virtually indistinguishable (see Fig. 5J). It is not assign-
able to Sorex saussurei as Findley thought (1955b).

The few specimens of the race neomexicanus that we examined
showed a somewhat aberrant tooth pattern, the accessory tines be-
ing very small to nearly absent; however, the general pattern is that
of monticohis rather than vagrons. James Findley (pers. comm.)
checked all specimens of neomexicanus at the University of New
Mexico and found none in which accessory tines were entirely
lacking.

Findley's .subspecies longiquus and ohscuroides do not, in our
opinion, deserve recognition apart from S. monticohis ohscurus. The
three fonns are identical in tooth structiu-e. Findley (1955a: 49)
distinguished longiquus from ohscurus mainly because of its small
size. However, Figure 4 shows that specimens of S. m. ohscurus
from south and west of the Snake River and from central Montana
outside the range of longiquus are, on the average, even shorter
than the three Findley topotypes of longiquus. This is consistent
with Mezhzherin's hypothesis (1964) discussed earlier. On the other
hand, cranial measurements of several longiquus in Findley's report
(1755a: 64) fell below minimum values we noted in populations of
typical ohscurus. Further data are needed, but ^^Sorex vagrans
longiquus" probably represents no more than a cluster of ecotypic
variants of S. monticolus ohscurus, similar to isolated montane pop-
ulations from the northern edge of the Great Basin.

Similarly, Findley distinguished "S. vagrans ohscuroides" as
smaller, and ". . . separated from the range of ohscurus by the in-
tervening smaller subspecies S. v. vagrans." (1955a: 58). However,
Sierra Nevada populations of monticolus are no smaller in size than
small S. m. ohscurus (see above), and are not as isolated geograph-

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ically as Findley indicated (Fig. 3). Sorex monticolus obscurtis ap-
pears to have reached the Sierra Nevada somtime during the Wis-
consin glacial period. Martin (1958) noted that during the height
of the Wisconsin the southwestern spiiice-fir forest zone was dis-
placed downward 4,000-5,000 ft. in southern New Mexico (and
probably most of the southwest). Spmce pollens from this period
have been found as far south as the Valley of Mexico. During this
time monticolus and vagrans probably moved through continuous
forest far into Mexico, giving rise to S. m. monticolus in the moun-
tains of Chihuahua and Durango. As noted above, S. vagrans
orizahae appears to be a relict of similar origin.

A similar corridor of boreal forest must have spanned the south-
em rim of the Great Basin, allowing monticolus access to the Sierra
Nevada from southern Utah; Major and Bamberg (1967) presented
Horistic evidence for such a dispersal corridor. Sorex hjelli, a rela-
tive of S. cinereus, and subspecies of Eutamias umbrinus and Phe-
nacomijs intermeclius, may have reached California by this pathway,
and Sorex nanus, in the ornatus group, seems to have moved into
the Rocky Mountains from west to east. Findley believed "ohscu-
roides" to be an offshoot of S. vagrans vagrans, but eveiy specimen
of California S. m. obscurus examined bears the distinctive incisor
pattern of Sorex monticolus. Sorex monticolus parvidens, a rare
shrew from the San Bernardino Mountains, California, also appears
to be an isolated offshoot of S. m. obscurus. It may also occur in
the Piute and Tehachapi mountains, between the known range of
S. m. parvidens and the southernmost records of S. m. obscurus in
the Sierra Nevada (Fig. 3, question marks). The few specimens
from these isolated intennediate mountain ranges available to us
(USNM 135947-48; 159416-17) appear to be closer to S. monticolus
than to S. ornatus, to which they are currently assigned. Additional
specimens are needed before a decision can be reached.

Populations of Sorex monticolus from southeastern Alaska and
coastal British Columbia i.e., calvertensis, elassodon, insularis, iso-
Jatus, longicauda, and prevostensis) have a variable incisor pattern
which in some specimens is intennediate between those observed
for vagrans and monticolus (Fig. 5E). The color is less intense and
the tines are somewhat higher on the face of the tooth than in typ-
ical monticolus. We have observed somewhat similar patterns in
occasional specimens of Sorex vagrans (Fig. 5F), and in the related
S. ornatus (Fig. 5H). They may recall a primitive condition from
which the distinctive patterns of vagrans, monticolus, cinereus (Fig.
5G), pribilofensis (Fig. 51), and other Otisorex shrews diverged.
At present these island races assigned to monticolus probably main-
tain a genetic link with interior populations through S. m. longi-
cauda, S. m. obscurus, and S. m. setosus (Findley, 1955a). Foster
(1965) suggested that ancestors of the insular races may have been

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 17

isolated from the main body of monticohis during the most recent
glacial period, on a Queen Charlotte Refugium. The discontinuity
in gene flow may have produced or perpetuated the abberant tooth
pattern found among Sorex monticohis from this region; in some
ways this situation parallels the relationship of Sorex monticohis
and S. pacifictis.

The vahditij of the name "obscums." — Our decision to recognize
"Sorex ohscuriis" as a valid species led to the discoveiy that the
names of the two shrews that had been previously considered sub-
species of Sorex vagrans (monticohis, dobsoni) had priority over the
name obscums. Merriam (1890) named S. monticohis from speci-
mens collected in the San Francisco Mts., Arizona, and subse-
quently QSQl) described S. dobsoni from Altin-as (^Sawtooth)
Lake, Idaho. By 1895 he had come to regard both as subspecies of
Sorex vagrans. Jackson (1928) followed Merriam in his definition
of S. vagrans monticola, but reduced S. v. dobsoni to the status of
a junior synonym of monticola, which he believed to range from
northern Mexico to southern British Columbia. However, Findley
(1955a), restricted the name S. v. monticola to shrews from Ari-
zona, New Mexico, and Chihuahua. He included the northern
shrews in the subspecies S. v. vagrans, along with the races of S. v.
amoenus and S. v. nevadensis, fomierly recognized by Jackson
(1928), and considered that the name dobsoni applied to inter-
grades between S. v. vagrans and S. v. obscums in the northern
Rockies.

We have examined the holotypes of Sorex monticolus and S.
dobsoni. They are clearly members of the species "S. obsctirus"
(sensu Jackson, 1928). The earliest valid description is that of S.
monticohis Merriam, 1890; Sorex dobsoni Merriam, 1891, and S.
obscums Merriam, 1895, are junior synonyms of S. monticohis.

The name dobsoni has priority over obscums as the name for
the Rocky Mountains subspecies of the dusky shrew; i.e. "S. oh-
scurus obscurus" (sensu Jackson, 1928). However, we choose to
consider dobsoni a nomen obhtum ( cf . Art. 23b, International Code
of Zoological nomenclature, 1964: 23). The code provides that a
name may be considered "forgotten" if it has remained unused as a
senior synonym in the primaiy taxonomic literature for more than
fifty years. The name dobsoni has been applied incorrectly to Sorex
vagrans since 1895; Jackson (1928) made it a junior synonym. The
name Sorex obscurus on the other hand was applied correctly and
consistent with its type from 1895 to 1966, when it was made a
synonym of Sorex vagrans (Findley, 1955a). It is still widely known
and used. Obscurus is no longer a valid species name, but to dis-
card it as a name for the subspecies in favor of dobsoni would cause
a great deal of unnecessary taxonomic confusion. We therefore em-
ploy the name-combination Sorex monticohis obscurus.

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Sorex pacificus Coues, 1877
Pacific Giant Shrew

Sorex pacificm Cones, 1877:650. (Type locality. Ft. Umpqua, Doug-
las Co., Oregon).

Sorex yaqiiinae Jackson, 1918:127. (Type locality, Yaquina Bay,
Lincoln Co., Oregon).

Sorex pacificus yaqiiinae V. Bailey, 1936:364. (New Combination).

Sorex vagrans pacificv^ Findley, 1955:34. (New Combination).

Sorex vagrans yaquinae Findley, 1955:34. (New Combination).

Holotype. — Adult, sex unknown, skin and skull; USNM 3266,
from Ft. Umpqua, mouth of Umpqua R., Douglas Co., Oregon.

Range. — Three subspecies occupy humid to dry coniferous and
mixed evergreen forests in coastal Oregon and California, often to
high altitude on the western flanks of the southern Cascades (Fig.
2). Sorex pacificus yaquinae ranges south from about the Siletz and
McKenzie Rivers (near Albany and Eugene), where it may hy-
bridize with S. monticolus hairdii (and S. m. permiUiensis?), to the
Umpqua River on the coast and inland to the Siskiyou Mountains
on the California border. Sorex pacificus pacificus is confined more
closely to coastal forests, from the mouth of the Umpqua River in
Oregon, inland to the western Siskiyous, and south as far as Men-
docino, California. Sorex pacificus sonomae continues south in the
redwood forests nearly to the Golden Gate.

Diagnosis. — Medium to large sized shrews of the subgenus Oti-
sorex, more similar to S. monticolus than to S. vagrans. As in vagrans
and monticolus the third unicuspid tooth is smaller than the fovnth,
and the foramen magnum is situated more or less ventrally (Find-
ley, 1955a). Pigment on the first upper incisors is extensive and
dark, but there are typically no medial tines (Fig. 5C). Some
specimens of S. p. yaquinae, in the zone of contact with S. monti-
colus hairdii have tiny tines suggesting the typical monticolus pat-
tern (Fig. 5D).

Size varies north to south and from the coast inland; the largest
pacificus are on the coast in southwest Oregon and northwestern
California. For example, three S. p. pacificus from near Gold Beach,
Oregon, had total lengths of 155-163 mm, and condylobasal lengths
of 22.3-22.8. Eight S. p. pacificus from near Happy Camp, inland
from the coast (Siskiyou Co., California), were 134-149 mm long,
with condylobasal length 21.2-21.9 mm. Five specimens of S. p.
yaquinae from Newport, Oregon (near the type locality), were 128-
141 mm long, with condylobasal length 19.3-20.5; specimens in the
zone of contact with S. monticolus hairdii are even smaller. See
Findley (1955a) for details of size and color variation.

Remarks. — The three subspecies of Sorex pacificus are larger
than any of the races of S. monticolus. Nonnally there are no tines

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 19

oil tlie upper incisors, although the pigment is high and very dark
(Fig. 5C'). Specimens assigned to S. p. yaqtiinae from Taft and
Philomath, Oregon (near localities from which S. monticolus hairdi
have been collected) have traces of medial tines which suggest that
pacificus may hybridize with Sorex monticolus (Fig. 5D). Incisor
structure varies in populations of S. m. ])airdi and S. m. permilliensis
which approach yaquinae in central Oregon, but father north (e.g.,
Astoria, Portland, Tillamook) S. m. hairdi consistently shows large,
low tines and heav)' pigment characteristic of S. monticolus. From
his studies of the biology of hairdi and yaquinae, Chris Maser (pers.
comm.) concurs with Findley (1955a) that the cline in body size
between Sorex monticohis and S. pacificus represents genie intro-
gression, the nature of which is poorly understood. The ancestors
of S. pacificus were probabh' isolated from typical monticohis dur-
ing the Wisconsin glaciation, and limited gene flow may have been
reestablished with subsequent post-glacial contact.

Brown (1970, 1974) has done preliminary work on the chromo-
somes of Sorex vagrans, S. monticohis, S. pacificus, S. ornatus, and
S. sinuosus from the coastal region of California, Oregon, and
Washington. The normal diploid number in all species is 54, with
three small pairs of metacentric autosomes. \^ariation occurs in the
number of submetacentric and acrocentric autosomes. Sorex orna-
tus and S. sinuosus consistently have 18 (9 pairs) of submetacen-
trics (47 specimens from nine localities in central California). Sorex
vagrans varies, from 6 submetacentrics (3 pairs) in the San Fran-
cisco Bay region (3 specimens of S. v. vagrans; 1 of S. v. hahcoetes)
to 4 ( 2 pairs ) in northern California and Oregon ( 7 specimens from
four localities). Karyotypes from Sorex monticohis and S. pacificus
show a north to south decline in numbers of submetacentric auto-
somes, this cline paralleling the morphological intergradation de-
scribed by Findley (1955a). Sorex monticohis setosus has 4 to 5
submetacentrics (5 specimens from two localities), and S. m. hairdi
has 2 or 4 (4 specimens from two localities). S. pacificus yaquinae
has 0 or 1 submetacentrics ( 11 specimens from 3 localities) as does
S. m. permihensis (6 specimens from 1 locality) and S. p. pacificus
appears to have no submetacentrics (but only one specimen was
examined ) .

More intensive work needs to be done to substantiate these
trends, and to detennine the extent of gene exchange between Sorex
p. yaijuinae and S. m. hairdi and between hairdi and permiUiensis
south of, and setosus north of the Columbia River. If there are no
substantial reproductive barriers between the taxa another nomen-
clatoral change will be necessary as Sorer pacificus Coues, 1877, has
priority over S. monticohis Merriam, 1890. For the present the evi-
dence is not sufficient to justify further revision.

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

% PALATAL LENGTH

Fig. 7 — (A.) Dorsal (left) and ventral (right) views of skull of Sorex
monticolus obscunts ( KU 28545), from northern British Columbia, showing
the three measurements in millimeters, used in Figs. 6 and 7. CBL, condylo-
basal length; LIB, least interorbital breadth; PL, palatal length. (Redrawn
from Findley, 1955a. ) Approx. X 3/4; and ( B. ) Trivariate analysis of cranial
proportions in 50 S. v. vagrans from eastern Washington and western Mon-
tana, and 50 S. ?n. obscurus from western Montana and the Beartooth Plateau
(NW Wyoming). The two specimens darkly circled have "hybrid" skull char-
acteristics. Both are from areas of parapatry or sympatry, the vagrans from
Camas Cr., Glacier National Park, Mont., and the obscurus from Three-Mile
Cr., Sapphire Range, Ravalli Co., Mont.

Discussion

Although Sorex vap,rans and Sorex monticolus are sympatric
over major portions of their ranges, close study reveals that they
tend to occupy different microhabitats, and only rarely are they col-
lected at the same trap site. In western Montana S. v. vagrans is
largely a shrew of riparian habitat, ranging widely along water-
courses throughout the intemiountain region. Its ability to survive at

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 21

high altitudes, on dn- soils, and in boreal forests, appears limited.
In Montana the Continental Divide may limit the eastward ex-
pansion of its range. Sorex monticoJus ohscurus ranges more widely
through montane forests, is less dependent on water, and is found at
high altitudes and on dry acidic soils. Where vagrans and monti-
colus occur together in the lowlands of western Montana (3,000-
5,000 ft.), we found monticoJtis much less common than vagrans.
In British Columbia, Hawes (1975) reported biological and eco-
logical differences, including odor (Hawes, 1976), between S. v.
vagrans and S. m. setostis which are consistent with our observa-
tions, as has Terry (1974) in western Washington.

Of all the shrews examined from western Montana, only one or
two specimens has the blend of vagrans and ohscurus characters
one would expect of a hybrid (see Fig. 7B). Hybridization per se
is not uncommon in mammals (Mayr, 1963); it does not follow that
hybrids are fertile, or that backcrossing into the parent populations
is also occurring. We see no morphological evidence in Montana of
broad intergradation cited by Findley when he synonymized Sorex
ohscurus (^=monticohrs) with S. vagrans. Moreover, S. v. vagrans
differs from S. monticolus and S. pacificus in that the two largest
metacentric pairs of chromosomes are subequal in size in the latter
whereas in vagrans one is considerably larger than the other; neither
is the morphology of the submetacentric chromosomes completely
comparable (Brown, 1974).

Findley also inferred that intergradation between vagrans and
monticohis occurred along the Wasatch Front in Utah and the Don-
ner Pass region in the Sierra Nevada of California. Almost all of
the specimens we examined from these areas are clearly either S.
vagrans or S. monticolus, even when the two occur at the same
localities. However, a few "hybrid" tooth patterns (Fig. 5F) were
found in specimens from Nevada Co. (Boca Springs; Donner; In-
dependence Lake), and Plumas Co. (Spring Garden Ranch, Grizzly
Mts.) in central California, while in Utah, possible hybrids were
found in Davis Co. (Clearfield), Salt Lake Co. (Parley's Canyon,
5000'; Spring Run, 4400'), and Weber Co. (2 mi. W. ^Roy P. O.;
Snow Basin). We also identified possible hybrid specimens from
Anthony, Oregon, and southeast of Pocatello, Idaho.

In California Sorex monticohis is limited to the continuously
mountainous central and southern Sierra Nevada. Of all the po-
tential "hybrid" individuals, a single specimen ( from Spring Garden
Ranch, Plumas Co., California) more closely resembles monticohis
than vagrans in the shape and placement of the accessoiy tines, and
is so assigned. This place is separated from montane habitats
around Mt. Lassen and Mt. Shasta by deep canyons and broad
stretches of lowland forest. Nonetheless, a few shrews from the
lowlands between Mt. Shasta and Mt. Lassen (Shasta Co.: Cassel;

22 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Siskiyou Co.: Medicine Lake, and Wagon Camp), where vagrans
is common but no monticoJus have been collected, also have incisors
conforming to the "hybrid" pattern (Fig. 5F), and Jackson (1928)
thought that ohscurus (^monticolus) inhabited the northern Sierra
Nevada. (His northernmost record, at McCloud River, Siskiyou Co.,
was a misidentified Sorex troivhridgu. )

The "hybrid" pattern, wherever it occurs, is reminiscent of mon-
ticohis only because the tines are larger and the pigment higher
than is characteristic for ?)Orex vagrcins. But the pattern is in other
ways essentially vagrans (compare Figs. 5A1 and F); the shape and
placement of tines in monticohis is markedly different (see Fig.
5B1). We conclude that the hybrid patterns are actually character
convergence - the product of genetic drift or selective factors oper-
ating within local populations of Sorex vagrans. The available evi-
dence demonstrates no intergradation between vagrans and mon-
ticohis.

Rudd ( 1955b ) documented a range of gradation in color and
morphology of salt marsh shrews in the northern part of San Fran-
cisco Bay, which he believed to represent introgression between
Sorex vagrans and S. [ornatus] sinuosus. However, Brown (1970)
examined the karyot)^pes of these shrews and found that all 13 in-
dividuals of the so-called hybrid population at Tolay Creek were
karyotypically S. sinuosus (see above) as were 10 shrews from the
vicinity of Petaluma Creek which Rudd had regarded as S. v.
vagrans. Character convergence in this case, too, must be accounted
for by something other than genie introgression. Pizzimenti (1976)
discovered a similar situation among prairie dogs (Cynomys).

Findley (1955a) regarded shrews of the S. ornatus group (orna-
tus, nanus, tenellus, etc.), S. longirostris and S. veraepacis as closely
related to the vagrans group, from which they diverged during the
late Pleistocene. Presently, S. ornatus is sympatric with S. vagrans
only in the area around San Francisco and Monterey bays, and with
S. monticohis only in the San Bernardino Mountains; its incisor tine
pattern ( Fig. 5H ) is closer to that of monticohis than vagrans, as is
that of S. nanus, which is broadly sympatric with S. monticohis in
the central and southern Rocky Mountains. In contrast, S. longi-
rostris of the southeastern United Stated, regarded by Findley as
"close in many ways to S. nanus ^ ( 1955a: 28), has incisor tines which
resemble those of S. vagrans; its distribution is wholly allopatric
and disjunct from its above-mentioned relatives. These new data
mandate re-interpretation of the biogeography and evolution of the
vagrans-ornattis-longirostris-veraepacis groups, but will require
still fuller understanding of the relationships within and between
these groups.

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 23

Summary

Morpliological similarities between Sorex vagrans vagrans and
S. monticolus (=obscurus) ohscurus were re-examined. Findley
(1955a) interpreted convergence in size and color as evidence of
introgression, and redefined the species Sorex vagrans to include
also S. ohscurus, S. pacificus, and S. yaquinae (sensu Jackson,
1928). In this paper we describe differences in tooth morphology
and cranial proportions, which demonstrate more clearly than past
criteria the integrity of S. vagrans and S. monticolus, and to a cer-
tain extent, that of S. pacificus. The name Sorex monticolus Mer-
riam, 1890, takes priorit)^ over S. ohscurus Merriam, 1895. The latter
is conserved as a subspecies name; Sorex dohsoni Merriam, 1891, is
considered a nomen oblitum.

Sorex vagrans and S. monticohis are biologically and ecologically
distinct, although their ranges overlap extensively. There is no evi-
dence that character convergence in some localities represents genie
introgression, although occasional hybrids may occur.

Secondary introgression may occur between Sorex monticolus
and S. pacificus, but they should be recognized as distinct species
until further research clarifies their relationship to each other.

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Dapson, R. W. 1968. Growth patterns in a post-juvenile population of short-
tailed shrews (Blarina brevicauda). Amer. Midi. Nat. 79:118-129.

24 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Dehnel, a. 1949. Badania nad rodzajem Sorex L. [Studies on the genus Sorex

L.] Ann. Univ. M. Curie-Sklodowska, Lublin Sect. C, 4:14:17-102 (in

Polish, English Summary, 75-97).
FiNDLEY, J. S. 1955a. Speciation of the wandering shrew. Univ. Kansas Publ.,

Mus. Nat. Hist., 9:1-68.
FiNDLEY, J. S. 1955b. Taxonomy and distribution of some American .shrews.

Univ. Kansas Publ., Mus. Nat. Hist., 7:613-618.
Fitch, H. S. 1940. A biogeographical study of the ordinoides artenkreis of

garter .snakes (genus Tliaimwphis). Univ. California Publ. Zool., 44:1-

150.
Foster, J. B. 1965. The evolution of the mammals of the Queen Charlotte

Islands, British Columbia. Occ. Papers, B. C. Prov. Museum, 14:1-130.
Gebczynski, M. 1965. Seasonal and age changes in the metabolism and activ-
ity of Sorex araneus Linnaeus 1958. Acta Theriol., 10:303-331.
Goldman, E. A. 1951. Biological investigations in Mexico. Smithson. Mis.

Coll., 1 15 :xiii-f 1-476.
Hawes, M. L. 1975. Ecological adaptations in two species of shrews. Unpubl.

Ph.D. dissertation, Univ. British Columbia, Vancouver, xiii+211 pp.
Hawes, M. L. 1976. Odor as a possible isolating mechanism in sympatric

species of shrews {Sorex vagrans and Sorex ohscurus). J. Mamm., 57:

404-406.
Hennings, D. N. 1970. Systematics of the Sorex vagrans-obscurus complex,

revisited. Unpubl. M. A. thesis, Univ. of Montana, Missoula. 118 pp.
Hensel, R. 1855. Beitriige zur Kenntniss fossiler Siiugethiere. Insektenfressen

und Nagethiere der DiluvialfoiTiiation. Zeitsch. der Deutsch. Geolog.

Gesellsch., 7(3):458-462.
Heptner, V. C, DoLGOv, V. A. 1967. O sistematicheskom polozhenni Sorex

mirabilis Ognev, 1937 [Systematic position of Sorex mirabilis Ognev,

1937. (Mammalia, Soricidae)]. Zool. Zhur., 56:1419-1422. (In Russian,

English summary. )
Hoffmann, R. S. 1971. Relationships of certain Holarctic shrews, genus Sorex.

Zeitsch. f. Saugetierk., 36:193-200.
Hoffmann, R. S., Taber, R. D. 1960. Notes on Sorex in the northern Rocky

Mountain alpine zone. J. Mamm., 41:230-234.
Ingles, L. G. 1961. Home range and habitats of the wandering shrew. T-

Mamm., 42:455-462.
Jackson, H. H. T. 1918. Two new shrews from Oregon. Proc. Biol. Soc.

Washington, 31:127-130.
Jackson, H. H. T. 1922. New species and subspecies of Sorex from western

America. J. Washington Acad. Sci., 12:262-264.
Jackson, H. H. T. 1928. A taxonomic review of the American longtailed

shrews (genera Sorex and Microsorex). N. Amer. Fauna, 51:1-238.
Johnson, M. L., Ostenson, B. T. 1959. Comments on the nomenclatiue of

some mammals of the Pacific Northwest. J. Mamm., 40:571-577.
Kuchler, a. W. 1967. Vegetation [map], in The National Atlas of the United

States of America. U.S. Geol. Survey, Washington, D.C.
Major, J., Bamberg, S. A. 1967. Some cordilleran plants disjunct in the Sierra

Nevada of California, and tlieir bearing on Pleistocene ecological condi-
tions, in Arctic and alpine environments (H. E. Wright, Jr. and W. H.

Osbum, eds.) Indiana Univ. Press, Bloomington: 171-188.
Martlv, p. S. 1958. Pleistocene ecology and biogeography of North America.

Pp. 372-420 in Zoogeography, ( C. L. Hubbs, ed. ) AAAS, Washington,

D. C, Publ. 51:x+509pp.
Mayr, E. 1963. Animal species and evolution. Har\'ard Univ. (Belknap)

Press, Cambridge, Mass. xv+797 pp.

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 25

Merriam, C. H. 1890. Results of a biological sui-vey of the San Francisco
Mountain region and desert of the Little Colorado, Arizona. N. Anier.
Fauna, 3:vii+136 pp., 13 pi., 5 maps.

Merriam, C. H. 1891. Results of a biological recoimoissance of south-central
Idaho. N. Anier. Fauna, 5-113 pp., 4 pi.

Merriam, C. H. 1895. Synopsis of the American shrews of the genus Sorcx.
N. Amer. Fauna, 10:57-98.

Merriam, C. II. 1899. Results of a biological survey of Mount Shasta, north-
ern California. N. Amer. Fauna, 16:179 pp., 5 pi.

Mezhzherin, V. A. 1964. Yavlenie Dehelya i evo vozmozhnoe ob'y^snenie.
[Dehnel's phenomenon and its possible explanation.] Acta Theriol.,
8:95-114 (in Russian, English summary).

Pizzimenti, J. J. 1976. Genetic divergence and morphological convergence in
the prairie dogs, Cijnomys gunnisoni and Cijnoinijs leucunis. I. Morpho-
logical and ecological analyses. II. Genetic analyses. Evolution, 30:
345-379.

PucEK, Z. 1955. Untersuchungen iiber die Veranderlichkeit der Schadels im
Lebenszyklus von Sorex aninciis arancus, L. Ann. Univ. M. Currie-
Sklodowska, C10:399-428.

PucEK, Z. 1957. Histomorphologische Untersuchungen iiber die Winterdepres-
sion des Schadels bei Sorex L. und Neomtjs Kaup. Ann. Univ M. Currie-
Sklodowska, C9: 399-428.

PucEK, Z. 1963. Seasonal changes in the braincase of some representatives of
the genus Sorer from tlie Palearctic. J. Mamm. 44:523-536.

PucEK, Z. 1965. Seasonal and age changes in the weight of internal organs of
shrews. Acta Theriol., 10:369-438.

Rensch, B. 1933. Zoologische systematik imd artbildingsproblem. Ver.
deutsch. zool. Gesellschaft, 1933:19-83.

RuDD, R. L. 1955a. Age, sex and weight comparisons in three species of shrews.
J. Mamm., 36:323-339.

RuDD, R. L. 1955b. Population variation and hybridization in some California
shrews. Syst. Zool., 4:21-34.

Spencer, A. W., Pettus, D. 1966. Habitat preferences in five sympatric spe-
cies of long-tailed shrews. Ecology, 47:677-683.

Terry, C. J. 1974. Ecological differentiation of three species of Sorex and
Neurotrichus gibbsi in western Washington. Unpubl. M. S. thesis, Univ.
of Washington, Seattle. 101 pp.

YuDiN, B. S. 1969. Novye dannye po sistematike nekotorykh vidov zemleroek
(Soricidae) Palearktiki i Nearktiki. [Taxonomy of some species of shrews
(Soricidae) from Palearctic and Nearctic] Acta. Theriol., 14:21-.34 (in
Russian, English summary).

Appendix: Specimens Examined

Locality data for each specimen examined are arranged as follows: alpha-
betically by species and subspecies; by state or pnnince, county and locality;
alphabetically by museum abbreviation. Asterisks indicate specimens included
in statistical analysis of cranial measurements.

Sorex monticohts

S. monticoJus alascensis. Total number, 65. — AL.4SKA: head of Cordova
Inlet, 1 MVZ; Disc Is., 1 MVZ; Douglas Is., 0.5 mi N of bridge (near Juneau),
1 MVZ; Eleanor Is., 1 MVZ; Glacier Bay Nat. Monument (58°22'-26'N,
136°52'-54'W), 14 CT; Glacier Bay Nat. Monument, Bartlett Cove, 2 USNM;
Glacier Bay Nat. Monument, Pt. Gustavus, 1 USNM (holotype S. glacialis);

26 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

4 mi N 9 mi W Haines, Chilkat R., 100', 9 KU; 1 mi S Haines, 5', 10 KU; 1
mi SSE Haines, 10', 2 KU; Jmieau, Sheep Cr., 2 UM; Knight Is., Drier Bay, 1
MVZ; Muir Inlet, Nunatak Camp, 1 USNM; Prince WilHam Sound, Orca, 1
USNM; Seward, 4 USNM; NE end SulHvan Is., 8 KU; Valdez, Sawanill Bay,
1 USNM; Yakutat Bay, 3 USNM (incl. holotype); no e.xact locahty, 2 USNM.

S. monticolus bainlii. Total number, 86. — OREGON: Clackamas Co.: Arra
Wanna, near Mt. Hood, 1 MVZ; 1.75 mi SW Colton, 1 MVZ; Estacada, 1 KU,
1 MVZ; Lake Grove, 10 mi SW Portland, 1 UM; Clatsop Co.: Astoria, 5 MVZ,
12 USNM (incl. holotype); Old Fort Clatsop, 100', 5 MVZ; Seaside, 3 USNM;
Columbia Co.: 3 mi SW Rainier, 500', 1 MVZ; 7 mi SE Rainier, 100', 12 MVZ;
Desdiutcs Co., McKenzie Pass, 1 USNM; Deschutes or Lane Co.: N slope.
Three Sisters, 4 USNM; Lane Co.: Blue R., 1 USNM; McKenzie Bridge, 1
USNM; Vida, 6 USNM; Lincoln Co.: Delake ( =Lincoln City), 1 KU; Lincoln
Beach, 2 USNM; Otis, 10 USNM; Rose Lodge, 1 USNM; Siletz R., 1 USNM;
Multnomah Co.: Larch Mt., 1 USNM; Portland, 3 USNM; Tillamook Co.,
Tillamook, 2 KU, 2 MVZ, 2 USNM; 5 mi WNW Timber (in W^ashington Co.),
2200', 1 UM; 7 mi WNW Timber (in Washington Co.), 2100', 1 UM; W«.s/i-
in^ton Co.: 18.5 mi WNW Portland, 1300', 3 MVZ.

S. rnonticolus calvertensis. Total number, 4.— BRITISH COLUMBIA:
Safety Cove, Calvert Is., 2 USNM (topotypes); Calvert Is., 2 UBC.

S. monticolus elassodon. Total number, 21. — ALASKA: Admiralty Is. (3
localities), 8 MVZ; Duke Is., 1 MVZ; Forester Is., 2 MVZ, 7 CAS-SU; Mitkof
Is., 2 USNM; BRITISH COLUMBIA: Queen Charlotte Islands, Moresby Is.,
Cumshewa Inlet, 1 USNM (holotype).

S. monticolus instilaris. Total number, 2. — BRITISH COLUMBIA: Smytlie
Is., 2 USNM (topotypes).

S. monticolus isolatus. Total number, 21.— BRITISH COLUMBIA (Van-
couver Island): Albemi, 1 MVZ; Comox, 1 USNM; Errington, 1 MVZ; French
Cr., 1 MVZ; Golden Eagle (Mine), 1 MVZ; Coldstream, 1 MVZ, 5 USNM;
Little Qualicum R., 1 MVZ; Nanaimo, 4 USNM (incl. holotype); Parksville,
1 MVZ; Quatsino, 1 USNM; Satuma Is., 2 MCZ; Mt. Washington, 1 KU.

S. monticolus longicaucla. Total number, 18. — ALASKA: Bocade Quadra,
1 MVZ; Bradfield Canal, 1 MVZ; Chickamin R. ( Behm Canal), 1 MVZ; Etolin
Is., 1 MVZ: Gravina Is. (opp. Ketchikan), 1 MVZ; Hehm Bay, 1 MVZ;
Wrangell Is., Polk Point, 1 CAS; Wrangell Is., Wrangell, 6 CAS, I USNM
(holotype); BRITISH COLUMBIA: B Is., Hunter group, 1 UBC; Port Simp-
son, 2 USNM; Ruth Is., 1 UBC.

S. tiu^nticolus malitiosus. Total number, 5. — ALASKA: Coronation Is., Egg
Harbor, 3 MVZ; E side Warren Is., 2 MVZ (incl. holotype).

S. monticolus mixtus. Total number, 3.— BRITISH COLUMBIA: Texada
Is., Vanada, 3 MVZ (incl. holotype).

S. monticolus monticolus. Total number, 59. — ARIZONA: Apache Co.:
Springei-ville, 1 USNM; Tunitcha Mts., Spruce Cr., 8 USNM; White Mts., 10
USNM; White Mts., Mt. Thomas, 9 USNM; Cochise Co.: Chiricahua Mts.,
Fly Park, 4 USNM; Coconino Co.: San Francisco Mts., 3 USNM (incl. holo-
type); Graham Co.: Graham Mts., 9200', 2 USNM; Greenlee Co.: Prieto Pla-
teau, 9000', S end Blue Range, 1 USNM; Pitna Co.: Santa Catalina Mts.,
7500', Summediaven, 3 USNM; CHIHUAHUA: Sierra Madre, near Gauda-
lupe y Calvo, 5 USNM; DURANGO: El Salto, 2 USNM (incl. holotype S.
clurangae); 5 mi S El Salto, 1 MCZ; Cueva, 8 mi S El Salto, 1 MSU; NEW
MEXICO: Catron Co.: Mogollon Mts., 3 USNM; 10 mi E Mogollon, 1 KU;
San Juan Co.: Chusca Mts., 1 USNM; Sierra Co. Mts. 4 mi W Kingston, 1
USNM; Soccoro Co.: Magdalena Mts., Copper Canyon, 3 USNM.

S. monticolus neomexicanus. Total number, 19. — NEW MEXICO: Otero
Co.: SW slope Capitan Mts., 2 USNM; Cloudcroft, 10 MCZ, 7 USNM (incl.
holotype ) .

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 27

S. monticolus obscurus. Total number, 903. — ALASKA: Anaktmuk Pass,

4 USNM; Bettles, 66°04'N, I5P34'W, 671', 1 KU; Chandler Lake, 68°12'N,
152°45'W, 2900', 1 KU; Chilkat Valley, Wells, 1 USNM; Mts. iiear Eagle, 1
USNM; Kilikniak \'alley, head, Kilikniak Cr., 1 USNM; Noatak Valley, Anuk
Lake, 1 USNM; Tanana, 3 USNM; Head, Toklat R., 3 USNM; Wahoo Lake,
69°08'N, 146°58'W, 2350', 1 KU; Yukon R., 20 mi above Circle, 2 USNM;
ALBERTA: Assineau R., 4 mi N 10 mi E Kimise, 2 KU; Blindmans Red R.,
1 MCZ; BRITISH COLUMBIA: Atlin, 6 CAS; Barkerville, 7 USNM; Bennett
(City), 6 USNM; S branch, Big Salmon R., 1 USNM; Caraboo L., near Kam-
loops (^Cariboo L. 40 mi N Kamloops), 2 USNM; Chapa-atan R., 4 USNM;
Cranbrook, 1 MVZ, 1 USNM; Field, 2 USNM; Ft. Grahame, 3 USNM; Glacier
(Nat. Park), 6 USNM; Glenora, 1 USNM; Golden, 1 USNM; Hope, 2 MCZ;
Hudson's Hope, 1 USNM; Hot Springs, 3 mi WNW jet. Trout and Liard Rs.,
1 KU; 4 mi S 2 mi E Kelsall Lake, W side Mt. Clave, 1 KU; N end Kerry
Lake, E bank Crooked R., 3 KU; Klappan R., 2 USNM; Kootenay Nat. Park,
Veniiillion Crossing, 1 USNM; Laurier Pass, 4 USNM; McDame Post, Dease
R., 5 USNM; Mt. Revelstoke Nat. Park, 3 USNM; NW side Muncho Lake, 1
KU; 9 mi S 44 mi W Muskwa, 2 KU; Nelson, 8 USNM; 70 mi E Prince
Rupert, 5 USNM; 30 mi N Radium Hot Springs, 2 USNM; Redfern Lake, 1
USNM; Sicamous, 1 USNM; Sikanni Chief R., 6 USNM; Stonehouse Cr., 5.5
mi W jet. with Kelsall R., 3 KU; Tweedsmuir Prov. Park, Fenton Lake, 1
UM**; Tweedsmuir Prov. Park, NE end Goodrich Lake, 3 UM"; Wall Lake,
1 USNM; Wind Summit Lake, Mile 393 Alaska Huy., 1 KU; Yale District, 7
mi W Bridesville, Anarchist Mtn., 400', 1 MVZ; CALIFORNIA: Eldorado Co.:
China Flat, Roekbound Valley, 7800', 1 M\'Z; Echo (Lake), 4 MCZ; Gilmore
Lake, Mt. Tallac, 1 MVZ; 1 mi E Phillips, 1 U\7y, Fresno Co.: Horse Corral
Meadows, 3 USNM; Humphreys Basin, 10800', 1 MVZ; Kea(r)sarge Ledges,
Kea(r)sarge Basin, 11500', 2 KU; Inyo Co.: 3 mi S 8 mi W Big Pine, 7700',
1 MVZ; 7 mi W Big Pine, 7000', 2 CSLB; 10 mi W Big Pine, Big Pine Cr.,
8000', 1 MVZ; 11 mi SW Big Pine, 9000', 1 CSLB; S Fork Big Pine Cr., ca.
10800'. 1 CAS; Bishop Cr., 6600', 5 USNM (inch holotype S. ohscuroidcs)- 5
mi W 1.25 mi S Independence, 6000', 1 MVZ; Inyo Nat. For., Sage Flat
Camp, 7700', 1 CSLB; Madera Co.: San Joachin R., 8000', 3 USNM; Mariposa
Co.: Lake Tenaya, 3 USNM; Tuolomne Meadows, 5 mi N base Mt. Lyell, 8
USNM; Tuohmine Meadows, Muir Meadow, 9300', 1 USNM; Tuolomne Mea-
dows, Soda Springs, 3 USNM; Tuolumne Meadows, Mt. Unicorn, 1 USNM;
Tuolomne Meadows, 2 USNM; Mono Co.: Mt. Conness, 1 USNM; Mt. Dana,
6 USNM; Lake Marv, 9000', 7 CAS; Lee Vining, 1 MVZ; Mt. Lyell, 12
USNM; Mammoth, 2 MCZ, 1 USNM; Pine City, near Mammoth, 8700', 1
MVZ; Pine City, 9000', 1 USNM; Nevada Co.: Independence Lake, 1 MVZ;
Placer Co.: Donner, 1 USNM: Plumas Co.: Spring Garden Ranch, Grizzly
Mts., 1 USNM; Tulare Co.: Kem Lakes, 1 USNM; N fork, Kern R., 9600', 2
USNM; S fork, Kem R., 4 USNM; Mineral King, 2 USNM; Moltkes Meadows,
9000', 1 USNM; W slope Olancha Peak, Little Brush Meadow, 9750', 1 MVZ;
Round Valley, 12 mi S Mt. Whitney, 1 USNM; Sequoia Nat. Park, Gaboon
Meadow, 1 USNM; Sequoia Nat. Park, Halstead Meadows, 4 USNM; Sequoia
Nat. Park, E fork Kaweah R., 7 USNM; Sequoia Nat. Park, Round Meadow,
1 USNM; 3 mi SE Three Rivers, 2 CSLB; Mt. Whitney, 2 MCZ, 6 USNM;
Tuolumne Co.: 0.5 mi NW Sonora Pass, 9400', 1 MVZ; COLORADO: Boulder
Co.: 0.75 mi N 2 mi W Allenspark, 8400', 5 KU; 3 mi S Ward, 9000', 1 KU;
Chaffee Co.: Poncho Cr., 10 mi SW Salida, 8500', 5 KU; Conejos Co.: 5 mi

5 24 mi W Antonita, 1 KU; Delta Co.: 12 mi S 5.5 mi E Collbran, Grand
Mesa, 10400', 4 KU; 0.5 mi S 8 mi E Skyway, Grand Mesa, 10000', 2 KU; 1
mi S 8 mi E Skyway, Grand Mesa, 10200', 1 KU; 1.5 mi S 8 mi E Sky\vay,
Grand Mesa, 9500', 3 KU; 2 mi S 8 mi E Skyway, Grand Mesa, 9500', 2 KU;
Dolores Co.: T40N R13W Sec 13, 8100', 1 KU; Gaifeld Co.: Deep Lake, 16

28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

mi N Glenwood Springs, 3 KU; Gunnison Co.: Gothic, 1 UM; 1.5 mi N
Gothic, 7 KU; 9 mi WNW Sapinero, Black Mesa, 9500', 1 KU; Huerfano Co.:
4 mi S Cucharas Camps, 3 KU; 5 mi S 1 mi W Cuchaias Camps, 10000', 10
KU; Jackson Co.: 2 mi N 2 mi E Gould, 8600', 1 KU; Lake Co.: Halfmoon
Cr., 8 mi SW Leadville, 10000', 3 KU; 12 mi S 1 mi W Leadville, 1 KU; 3
mi W Twin Lakes, 2 KU; Larimer Co.: 3.5 mi S 4 mi W Estes Park Village,
2 KU; Poudre R., 1 KU; Mineral Co.: 23 mi S 11 mi E Creede, 9300', 1 KU;

4 mi S 6 mi E Wagon Wlieel Gap, 8500', 1 KU; Montrose Co.: 13 mi N 7
mi E Nonvood, 8400', 1 KU; T48N R14W Sec. 11 (SE K), 9000', 2 KU;
Montezuma Co.: 1 mi W Mancos, 7000', 1 KU; Mesa Verde Nat. Park, upper
well. Prater Canyon, 7575', 2 KU; Mesa Verde Nat. Park, .25 mi. N middle
well, Prater Canyon, 7575', 2 KU; Mesa Verde Nat. Park, Morefield Canyon,
7600', 2 KU; Rio Blanco Co.: 9.5 mi SW Pagoda Peak, 7700', 2 KU; Saguache
Co.: Cochetopa Pass, 33 mi W Saguache, 10000', 3 KU; 3 mi N 16 mi W
Saguache, 8500', 2 KU; IDAHO: Adams Co.: 0.5 mi E Black Lake, 6800', 1
KU; 3 mi W Payette Lake, 5400', 1 MVZ; SW slope. Smith Mt., 1 mi N Bear
Cr. R. S., 5400', 1 KU; summit, Smith Mt., 7500', 2 KU; Bannock Co.: 4 mi

5 Pocatello, Indian Cr., 6500', 1 MVZ; Blaine Co.: Alturas (=Sawtooth)
Lake, 7000', 1 MVZ, 1 USNM (holotype S. dohsoni)- Perkins Lake, 7000', 1
KU; Sawtootli City, 5 USNM; Bonneville Co.: 10 mi SE Ir^vin, 2 USNM;
Boundanj Co.: Cabinet Mts., E of Priest Lake, 1 USNM; Boise Co.: Bald Mt.
R. S., 10 mi S Idaho City, 7400', 2 USNM; Bear Lake— Caribou Co. Line:
Preuss Mts., head of Crow Cr., 1 USNM: Cassia Co.: 10 mi S Albion, Mt.
Harrison, 1 MVZ; Custer Co.: Head of Pahsimeroi R., Pahsimeroi Mts., 1
MVZ, 1 USNM; Fremont Co.: 17 mi E 4 mi N Ashton, 6275', 9 MVZ; 7 mi
W West Yellowstone, 7000', 4 KU; Idaho Co.: 6 mi SW Selway Falls, 5890',

1 CAS; Seven Devils Mt., 1 USNM; Lemhi Co.: "Salmon R. Mts." (=Lemhi
Mts.), 6 USNM; Timber Cr., Lemhi Mts., 8200', 1 USNM (holotype S. ob-
scurus); Shoshone Co.: Enaville, 1 USNM; Teton Co.: 3 mi SW Victor; 2 MVZ;
Valley Co.: Landmark R. S., 10 mi E Wann Lake, 6000', 1 USNM; Washing-
ton Co.: SW slope Cuddy Mt., 1 mi NE Heath, 4000', 8 KU; MONTANA:
Beaverhead Co.: 18 mi NW Dillon, Birch Cr., 7100', 6 MVZ; S of Wise River
(towTi), on Wise R., 1 UM; Broadwater Co.: 5 mi W Townsend, 2 UM**;
Carbon Co.: Beartoodi Mts., 2 USNM; Pryor Mts., 5 USNM; 2 mi E Shriver,
6500', 3 MVZ; Chotcau Co.: Highwood Mts., 13 USNM; 0.5 mi N Arrow Cr.
Divide, 5400', Highwood Mts., 1 KU; Fergus Co.: Big Snowy Mts., 1 UM, 4
USNM; Big Snowy Mts., 13 mi W Buffalo Canyon, 2 USNM; Big Snowy Mts.,
6400', 1 mi S Crystal Lake, 1 KU; Big Snowy Mts., N fork Flatwillow Cr., 15
mi S Heath, 1 USNM; Judith Mts., 4600', 7 mi N 9 mi E Lewistown, 3 KU;
Juditli Mts., Lime Kiln Gulch, 7 mi NE Lewistown, 1 USNM; Moccasin Mts.,
5 mi NW Hilger, 2 USNM; Mocassin Mts., 4500', Kendall, 1 KU; Flathead
Co.: Aeneas Peak, 1 UM', Big Mountain Ski Area (near Whitefish), 1 DCM*;
Glacier Nat. Park, Arrow Lake snowshoe cabin, 4000', 1 MVZ; Glacier Nat.
Park, Logan Pass, 7000', 4 UM"; Moran R. S. (N fork Flathead R.), 2 UM";

2 mi S 1 mi W Summit, 5000', 1 KU; Upper Stillwater Lake, 1 USNM;
Gallatin Co.: 5 mi N Bozeman, 1 MSUDZ'; 5 mi NE Bozeman, 1 MSUDZ";
Bridger Range, Fairy Lake Camp, 1 UM'; W fork, W Gallatin R., 4 USNM;
Upper Hyalite Cr., Palace Butte Camp, 3 UM"; Glacier Co.: 1.5 mi S 2.5 mi
W Babb, 4700', 1 KU; 3 mi SW Babb, N bank Swiftcurrent Cr., 4700', 2
UM"; 1 mi W Cut Bank, 3650', 1 KU; Glacier Nat. Park, Crossley Lake, 1
USNM; Glacier Nat. Park, Cut Bank (Campground), 1 UM"; Glacier Nat.
Park, Gunsight Lake, 2 USNM; Glacier Nat. Park, Many Glacier, 4900', 2
UM; Glacier Nat. Park, McDermitt Lake, 1 USNM; Glacier Nat. Park, Rising
Sun (Campground), 2 UM"; Glacier Nat. Park, 2.5 mi N 2.5 mi W Two
Medicine Chalet, 7000', 1 MVZ; St. Marys Lake, 9 USNM; Granite Co.: 1.2
mi W Beannouth, 1 UM"; Hill Co.: Bear Paw Mts., 20 mi. SE Ft. Assini-

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 29

boine, 2 USNM, Jefferson Co.: 0.5 mi E Whitehall, 1 UM; Judith Basin Co.:
3 mi W Ge>ser, 4100', 1 KU; Little Belt Mts., Dry Wolf Cr., 20 mi SW
Stanford, 1 USNM; Little Belt Mtns., Hoo\er Spring, 1 UM; Little Belt Mts.,
Neiliart, 1 USNM; Little Belt Mts., Otter Cr., 10 mi SW Geyser, 1 USNM;
Little Belt Mts., Sheep Cr., 16 mi N White Sulphur Springs, 1 USNM; Little
Belt Mts., Trask Gulch, 2 UM; Lake Co.: Flathead Lake, Yellow Bay, UM
Biol. Sta., 2900', 3 UM*; Yellow Bay Cr.. .3000', 1 UM*; 3.5 mi E UM Biol.
Sta., 5600', 4 UM; 4 mi W Ronan, 1 UM**; Lewis and Clark Co.: Rogers
Pass, 5600', 1 MSUDZ; Steeker's Camp, Sun R., 2 UM; Madison Co.: Ruby
Mts., Hinch Cr., 12 mi SW Alder, 4 USNM; Ward Peak, Washington Cr.,
6000', 1 USNM; Meagher Co.: Camas Cr., 4 mi S Ft. Logan, 7 USNM; Little
Belt Mts., N fork Smith R., lower Sawmill Cr., 1 UM; Missoula Co.: Lu-
brecht Exp. Forest, 1 UM*; Missoula, Miller Cr., 1 UM*; Mi.ssoula, Pattee
Canyon, 1 UM*; Park Co.: 2 mi NE Cooke, 8500', 6 MVZ; Emigrant Gulch,
3 mi SE Chico, 6500', 2 USNM; 12 mi S Livingston, Pine Cr., Luccock Park
Camp, 1 UM; Phillips Co.: Zortman, 1 USNM; Ravalli Co.: Bass Cr., NW
Stevensville, 4000', 1 USNM; 7 mi W Hamilton, Blodgett Canyon, 1 DCM*;
7 mi N 9 mi E Stevensville, Three-mile Cr., 4500', 2 UM*; 8 mi S 9 mi E
Stevensville, Burnt Fork, 5050', 2 UM*; 8 mi NE Stevensville, 4000', 3 USNM;
Sula, 1 USNM; Siherhow Co.: Fleecer Mt., head Indian Cr., 1 UM; Sweet-
grass Co.: Crazy Mts., head Big Timber Cr., 4 UM*; Teton Co.: 6.5 mi N
17.13 mi W Augusta, 5100', 1 KU; No exact locality, 1 UM; Toole Co.:
Sweetgrass Hills, 1 UM; NEVADA: Elko Co.: 22 mi N Deeth, Marys R.,
5800', 1 MVZ; Granite Range, headwaters Trout Cr., 1 MVZ; Ruby Mts., S
fork Long Cr., 3 KU; Ruby Mts., summit. Secret Pass, 6200', 1 KU; Ruby Mts.,
Three Lakes, 4 KU; Rubv Mts., 2 USNM; Humboldt Co.: 13 mi N Paradise
Valley, 6700', 1 MVZ; Ormshy Co.: Marlette Lake, 8000', 1 MVZ; 0.5 mi S
Marlette Lake, 8150', 1 M\'Z; Washoe Co.: Marlette Lake, 8000', 2 MVZ; 3
mi S Mt. Rose, 2 MVZ; NEW MEXICO: Colfax Co.: 1 mi S 2 mi E Eagle
Nest, 8100', 2 KU; Sandoval Co.: Jemez Mts., 3 USNM; San Miguel Co.:
Hoover's Ranch, 20 mi NW Las Vegas, 1 KU; Santa Fe Co.: Pecos Baldy,
1100-11700', 4 USNM; Santa Fe Ski Basin, 1 KU; 5 mi NE Santa Fe, 1 CAS;
Taos Co.: 3 mi N Red River, 10700', 1 USNM; Taos, 1 USNM; Twining,
9800-12500', 6 USNM; Torrance Co.: Manzano Mts., 2 USNM; NORTH-
WEST TERRITORY: Ft. Resolution, Mission Is., 1 USNM; Ft. Simpson, 3
USNM; Nahanni R. Mts., Mackenzie R., 1 USNM; OREGON: Baker Co.:
Anthony, 4 MVZ, 1 USNM; Bourne, 3 USNM; Grant Co.: Beech Cr., 5
USNM; Strawberry Butte, 1 USNM; Harney Co.: Diamond, 1 USNM; Steens
Mts., 2 MVZ, 1 USNM; Steens Mts., Keiger Gorge, 7500-8000', 2 USNM;
Hood River Co.: 2 mi W Parkdale, 1500', 2 USNM; Wallowa Co.: Wallowa
Lake, 4800', 2 MVZ, 3 USNM; Wallowa Mts., S Wallowa Lake, 8100', 2
USNM; SASKATCHEWAN: Cypress Hills, 10 NMC; Cypress Hills, near
Maple Cr., 1 MVZ, 2 NMC; Middle Cr. Res. (SW Cypress Lake), 1 UAMZ;
UTAH: Beaver Co.: Beaver Mts., Puffer Lake, 3 USNM; Davis Co.: Bounti-
ful Canyon, Mueller Park, 1 UU; Kaysville, 1 mi W L.D.S. Will [sp?], 1 UU;
Steed's Canyon, Centerville, 1 UU; Grand Co.: La Sal Mts., 11000', 1 USNM;
Iron Co.: Brian Head (Mammoth Summit), 11000', 2 MVZ; Parowan Mts.,
2 USNM; Millard Co.: Richfield, 1 USNM; Salt Lake Co.: Big Cottonwood
Canyon, Spruces, Camp 67, 1 UU; Brighton, 8700', 2 UU; Brighton, 8750', 1
UU; Brighton, 9500', 1 UU; 1 mi W Brighton, 8600', 1 UU; 2 mi from
Brighton, 8600', 1 UU; Butterfield Canyon, 1.5 mi above tunnel, 1 UU;
Butterfield Canvon, 3 mi SW of tunnel, 7000', 1 UU; Little Cottonwood Can-
yon, 1 mi E of mouth, 5500', 1 UU; Mill Cr. Canyon, The Firs, 7000', 1
UU; Mill Cr. Canyon, 2 mi E Mill Cr. G. S., 7000', 1 UU; Mill Cr. Canyon,
Wilson's Fork, 8000', 2 UU; Salt Lake City. 1 USNM; Sanpete Co.: Ephraim,
1 USNM; Manti, 3 USNM; Sevier Co.: Fish Lake Plateau, 2 USNM; Summit

30 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Co.: Holiday Park, 1 MCZ; Umtah Co.: Paradise Park, 15 mi N 21 mi W
Vernal, 1050!)', 3 KU; Utah Co.: Aspen Grove, Mt. Timpanogos, 2 BYU; Basal
American Fork Cirque, Mt. Timpanogos, 1 BYU; Hidden Lake, Mt. Timpa-
nogos, 1 BYU; Payson Lake, Nebo Mts., 8300', 1 UU; 1 mi E Payson Lake,
Nebo Mts., 8300', 1 UU; Spanish Fork, 1 mi E BYU Farm, 1 BYU; Spanish
Fork Bench, 1 BYU; Timpooneke Basin, Mt. Timpanogos, 4 BYU; Wasatch
Co.: Lost Lake, 1 BYU; Uinta Forest, Current Cr., 8000', 1 USNM; Washing-
ton Co.: Pine Valley Mts., 3-5 mi E Pine Valley; 10 USNM; 1.5 mi E Pine
Valley, 6 USNM; WASHINGTON: Chelan Co.: Entiat, 1 USNM; Wenatchee,
1 USNM; Ferry Co.: 5 mi W Curlew, 2800', 2 USNM; Kittitas Co.: Easton,
10 USNM; 4.5 mi N 6.5 mi W Easton, 1 KU; Okanogan Co.: W end of
Bauerman Ridge, at tungsten mine, 6800', 1 USNM; Head, Lake Chelan, 4
USNM; E fork, Pasayten R., 3900', 1 USNM; Stehekin, 4 USNM; Pend
'Oreille Co.: 2 mi N Gypsy Meadows, 2 CRCM*; Yakima Co.: Yakima Indian
Reservation, Signal Peak, 4000', 5 USNM; WYOMING: Albany Co.: Woods
P. O., 1 USNM; Carbon Co.: Bridger Pass, 2 USNM; Sierra Madre Mts., S
base Bridger Pk., 8900', 1 USNM; Johihson-Washaki Co. Line: Bighorn Mts.,
8400-9000', 3 USNM; Lincohi Co.: 10 mi SE Afton, 7500-9000', 4 USNM; 3
mi W Stanley, 3 USNM; Park Co.: Beartooth Plateau, 26 UM*; Beartooth
Pass, 0.5 mi N ski to\\% 10325', 1 UM'; Beartooth Plateau, Mont.-Wyo. State
Line, 2 UM*; 25 mi S 28 mi W Cody, 6350', 1 KU; Cooke City Hwy., S Red
Lodge, 2 UM"; Cooke City Hwy., Rennet Cr. divide, 10150-10931', 4 UM*;
Cooke City Hwy., Boundary Lake(s), 7 UM*; Cooke City Hwy., opposite
Gardner Lake viewpoint, 10500', 1 UM*; Cooke City Hwy., 0.5 mi N Gard-
ner Lake viewpoint, 10325', 16 UM"; Cooke City Hwy., 0.75 mi N Gardner
Lake viewpoint, 6 UM*; Cooke City Hwy., 5 mi N Gardner Lake view^joint,
1 UM*; Cooke City Hwy., 6 mi N Gardner Lake viewpoint, 1 UM*; Cooke
City Hwy., 7 mi N Gardner Lake viewpoint, 1 UM*; Cooke City Hwy., 0.33
mi S Frozen Lake, 1 UM"; Ishawopa Cr., 19 mi S 19 mi W Cody, 2 KU;
Pahaska, mouth, Grinnell Cr., 6300', 7 USNM; 0.5 mi W Twin Lake Basin, 1
UM; Yellowstone Nat. Park, Blacktail Deer Cr., 6700', 1 MVZ; Yellowstone
Nat. Park, Mammotli Hot Springs, 1 USNM; Yellowstone Nat. Park, Slough
Cr., 6300', 2 M\^Z; SW slope Whirlwind Peak, 9000', 1 KU; Sublette Co.: N
side Half Moon Lake, 1 KU; 2.5 mi NE Pinedale, 1 KU; 2.25 mi N Pinedale,
1 KU; 31 mi N Pinedale, 8025', 3 KU; Sui-veyors Park, 12 mi NE Pinedale,
8000', 2 USNM; Teton Co.: Jackson Hole Wildlife Park, 7 USNM; 2.5 mi NE
Moose, 6500', Bar BC Ranch, 1 KU; 4 mi N Moose, 6750', Timberland Is., 3
KU; Moran, 6244', 1 KU; Moran, Lake Emma Matilda, 2 USNM; 1 mi N
Moran, 3 USNM; Upper Arizona Cr., N Moran, 2 USNM; 0.25 mi N 2.5 mi
E Moran, 6230', 3 KU; 1 mi E Moran, 4 KU; 2.5 mi E Moran, 1 KU; 3 mi
E Moran, 6210', 1 KU; 3.75 mi E Moran, 6300', 1 KU; 1 mi S 3.75 mi E
Moran, 6200', 6 KU; Teton Mts., Moose Cr., 6800', 7 USNM; Teton Pass,
above Fish Cr., 7200', 2 USNM; Whetstone Cr., 3 USNM; Head, Wolverine
Cr., 1 USNM; Yellowstone Nat. Park, Old Faithful, 3 USNM; Uinta Co.:
Evanston, 1 USNM; Ft. Bridger, 6650'. 3 KU; 1 mi N Ft. Bridger, 6650', 1
KU; 9 mi S Robertson, 8000', 3 KU; 9 mi S 2.5 mi E Robertson, 8600', 1 KU;
10 mi S 1 mi W Robertson, 8700', 3 KU; 10.5 mi S 2 mi E Robertson, 8900',
1 KU; 13 mi S 1 mi E Robertson, 9000', 1 KU; 13 mi S 2 mi E. Robertson,
9200', I KU; Washakie Co.: 4 mi N 9 mi E Tensleep, 7000', 2 KU; 5 mi N 9
mi E Tensleep, 7400', 1 KU; YUKON: 1.5 mi S 3 mi E Dalton Post, 2500',
1 KU; SW end Dezadeash Lake, 1 KU; Mclntyre Cr., 3 mi NW Whitehorse,
2250', 1 KU.

S. monticohis parvidens. Total number, 7. — CALIFORNIA: San Bernar-
dino Co.: San Bernardino Mts., Bluff Lake, 7500', 4 USNM (inch holotype);
Deep Cr., near Lake Arrowhead, 1 CSLB; Sunnnit, 1 USNM; Los Angeles
Co.: San Gabriel Mts., 0.4 mi W Wrightwood, 6000', 1 CSLB.

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 31

S. monticohis pcrwilicnsis. Total number, 24. — OREGON: Hood River
Co.: Brooks Meadow, 9 mi ENE Mt. Hood, 4300', 1 MVZ; Near timberline,
Mt. Hood, 2 USNM; Marion Co.: Detroit, 1 USNM; Permilia Lake, W base
Mt. Jefferson, 19 USNM (incl. holotype); Wasco Co.: Camas Prairie, 12 mi
S Mt. Hood, 1 USNM.

S. tiiouticohis prevosten.sis. Total number, 13.— BRITISH COLUMBIA:
Queen Charlotte Islands, Prevost Is., 13 USNM (incl. holotype).

S. monticolus scfosiis. Total number 43.— BRITISH COLUMBIA: Howe
Sound, Gibson's Landing, 1 USNM; WASHINGTON: Chilian, Co.: Sol Due
Hot Springs, 1 KU; Snow Cr., 1200', 2 USNM; Clark Co.: 3.5 mi E 5 mi N
Yacolt, 500', 1 M\'Z; CotcJitz Co.: 4 mi E mouth Kalama R., 1 MVZ; 6 mi E
mouth Kalama R., 2 MVZ; Gratj's Harbor Co., Quinault Lake, 7 USNM;
Okanogan Co.: W fork, Pasayten R., 4700', 2 USNM; Pacific Co.: 2.5 mi SE
Chinook, 10', 13 MVZ; Pierce Co.: E gate McChord Air Force Base, 1 UM;
Puyallup, 1 USNM; Skagit Co., Mount Vernon, 2 USNM; Skamania Co., Mt.
St. Helens, Spirit Lake, 3200', 4 USNM; Snohomish Co.: Silverton, Dickennan
Pk., 5000', 1 USNM; Wahkiakum Co., Cathlamet, 2 USNM; Whatcom Co.:
Barron, 5000', 2 USNM.

S. monticohis shumaginensis. Total number, 19. — ALASKA: 1 mi NE
Anchorage. 100', 1 KU; Bethel, 1 USNM; Chevak, 61M0'N, 165°18'W, 1
USNM; Chignik, 1 USNM; Izembek Bay, 3 USNM; Kenai Peninsula, Indian
Cr., Lower Kenai Lake, 1 CAS; Nagai Is., 1 USNM; Petra Cr., 20 mi NE
Anchorage, 300', 1 KU; Popof Is., Shumagin Islands, 3 USNM (incl. holo-
type); Sanak Is., 3 USNM; 6 mi WSW Snowshoe Lake, Glenn Hwv., 2 KU;
Takotna, 1 USNM.

S. monticohis sopcri. Total number, 3. — MANITOBA: Riding Mountain
Nat. Park, 2.5 mi NW Lake Audy, 1 UAMZ (topotype); Riding Mountain
Nat. Park, Swanson Cr., 1 UAMZ; SASKATCHEWAN: Prince Albert Nat.
Park, Spruce R. Twnshp., 1 UAMZ.

Sorex pacificus

S. pacificus pacificus. Total number, 24. — CALIFORNIA: Del Norte Co.:
Crescent City, 1 KU; 8 mi E Crescent Citv, 1 MVZ; Klamath, 1 CAS; 6 mi
N Klamath, 2 MVZ; Requa, 2 CAS; Humbohh Co.: Eureka, 1 KU; Siskiyou
Co.: 12 mi WNW Happy Camp, Poker Flat, 5000', 9 MVZ; 3 mi W Klamatli
R., Clear Cr., 1400', 1 MVZ; OREGON: Coos Co.: MarshReld (=:Coos Bay),
1 USNM; Myrde Point, 1 USNM; Currij Co.: 3 mi above Gold Beach, S side
Rogue R., 3 MVZ; Douglas Co.: Ft. Umpqua, 1 USNM (holotype).

S. pacificus sonomae. Total number, 2. — CALIFORNIA: Marin Co.: In-
verness, 1 USNM; Sonoma Co.: Gualala, 1 MVZ (holotype).

S. pacificus yaquinae. Total number, 40. — CALIFORNIA: Siskiyou Co.:
15 mi W Hilt, Donomore Meadow, 5800', 1 MVZ; OREGON: Bei^ton Co.:
Philomadi, 2 USNM; Douglas Co.: Gardiner, 7 MVZ, 2 USNM; 1.3 mi E Gardi-
ner, 5 MVZ; Elkhead, 1 USNM; Jackson Co.: E. Fork, Ashland Cr., 0.6 mi N
junct. F.S. rd. 3903, 3 SOMNH; Klamath Co., Crater Lake Nat. Park, Munson
Valley, 1 USNM; S end Lake of the Woods, 1 M\'Z; Lane Co.: Mapleton, 3
USNM; Mercer, 1 MVZ; Vida, 2 USNM; Lincoln Co.: Newport, Yaquina Bay,
1 MCZ, 5 MVZ, 1 USNM (holotype); Taft (^Lincoln City), 4 MVZ.

Sorex vagrans

S. vagrans halicoetes. Total number, 39.— CALIFORNIA: .Mameda Co.:
Berkeley, 1 USNM; Dumbarton Point, 1 Kll; West Berkeley, 1 USNM; San
Francisco Co., San Francisco, 6 CAS, 4 USNM; San Mateo Co.: Colma, 20

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

CAS; San Mateo, 2 USNM; Santa Clara Co.: Palo Alto, 3 USNM, 1 MVZ
( holotype ) .

S. vagrans orizabae. Total number, 29. — MEXICO: Salazar, 2 USNM; 1
mi W Salazar, 9850', 1 KU; Nevada de Toluca, 13700', 1 KU; Nevada de To-
liica, 5 mi S Raicer, 1 USNM; Nevada de Toluca, 16 mi SSW Toluca, 3
USNM; N slope Volcan de Toluca, 3 USNM; MICHOACAN: Nahuatzin, 3
USNM; Patamban, 1 USNM; Mt. Tancitaro, 4 USNM; MORELOS: Cerro
Ci-uz del Marquez, 2440 m, 1 UNAM; PUEBLA: Mt. Orizaba, 6 USNM (incl.
holotype); TLAXCALA: Mt. Malinche, 2 USNM; VERACRUZ: Cofre de
Perote, 1 USNM.

S. vagrans pahidivagus. Total number, 1. — CALIFORNIA: Moiiteretj Co.:
4 mi. W Pacific Grove, Bird Rocks, 1 KU. (Identification cjuestionable; may be
S. ornatus. )

S. vagrans vagrans. Total number, 1130.— BRITISH COLUMBIA: Cran-
brook, 10 USNM; Creston, Kootcnav Valley, 4 CAS; Enderby, 1 KU; Glacier
Nat. Park, 4 USNM; Hope, 2 MCZ; Okanagan, 1 USNM; o'kanagan Lake, 3
USNM; Mt. Revelstoke Nat. Park, 1 USNM; Mt. Richter, 3 mi E Similkameen
R., 1 USNM; Sumas, 1 MCZ; \'ancouver. Point Grey, 2 UM; CALIFORNIA:
Del Norte Co.: Requa, 1 CAS; El Dorado Co.: Talks, 3 USNM; Humboldt
Co.: Capetown, 2 CAS; Samoa Pen., 0.5 mi N Coast Guard, 1 CSLB; Inyo
Co.: Alvord, Owens Valley, 1 USNM; Lassen Co.: Bogard R. S., 2 CAS;
Marin Co.: 5 mi W Inverness, 1 KU; Head, Limantour Bay, 2 CAS; Wliite
Gulch, 5', 1 UM; Mendocino Co.: 6 mi NE Cummings, 2500', 1 CAS; Hearst
P. O., 1 CAS; Laytonville, 1 CAS; Modoc Co.: Davis Creek, Goose Lake, 1
USNM; Mono Co.: 1 mi W June Lake, 1 CSLB; Mammotli, 1 USNM; Mam-
motli Pass, 1 USNM; Mono Lake, 1 USNM; Head Owens R., near Mammoth,

1 USNM (holotype, S. amoenus); Nevada Co.: Boca Springs, 6000', 1 MVZ;

2 mi N 5 mi W Hobart Mills, 6900', 3 MVZ; Independence Lake, 9 M\^Z;
Sagehen Cr., 6500', 23 MVZ; Sagehen Cr., 3 mi NW Hobart Mills, 1 CSLB;
Placer Co.: Donner, 3 USNM; 5 mi S Sagehen Cr., west bank Tnickee R., 4
MVZ; Plumas Co.: 8 mi NW Greenville, 1 USNM; Spring Garden Ranch,
Grizzly Mts., 2 USNM; 12 mi NE Prattville, 2 USNM; Sierra Valley, 1
USNM; Shasta Co.: Carbenys Ranch, 4 USNM; Casell, 2 USNM; Fall R.
Valley, Fall Lake, 2 USNM; Ft. Crook, 10 USNM; Goose Valley, 2 CAS;
Kellys, Warner Cr., 5200', 1 KU; SE side, Lassen Pk., 2 USNM; Sierra Co.:
3.2 mi W 1 mi N Calpine, McNair Meadow, 5300', 3 MVZ; 3 mi N Inde-
pendence Lake, 1 CAS; Lincoln Cr., 1 USNM; Siskiyou Co.: Beswick, 1
USNM; Big Springs, 1 CAS; Brownell, 1 USNM; Goose Nest Mt, 2 USNM;
Gordon Ranch, 1 CAS; Hornbrook, 3 USNM; Indian Tom Lake, 4 mi N Dor-
ris, 1 CSLB; May ten, Shasta Valley, 2 USNM; Medicine Lake, 6671', 4 MVZ;
Mt. Shasta, upper Ash Cr., 1 USNM; Mt. Shasta, Mud Cr., 1 USNM; Mt.
Shasta, upper Mud Cr., 2 USNM; Mt. Shasta, timberline. Mud Cr., 6 USNM;
Mt. Shasta, Squaw Cr., 7800'-8100', 3 USNM; Mt. Shasta, head. Squaw Cr.,

2 USNM; Mt. Shasta, Wagon Camp, 3 MVZ, 5 USNM (incl. holotype, S.
shastensis); 0.5 mi SW Mt. Shasta City, 3400', 4 MVZ; Sisson, 1 MVZ; Squaw
Cr. Valley, Warmcastle Soda Springs, 2 LTSNM; Tehama Co.: Battle Cr.
Meadow, 4800', 3 MVZ; Lvonsxille P. O., Turner's, 3500', 2 MVZ; Mineral,

3 MVZ; IDAHO: Adams Co.: New Meadows, 1 USNM; 3 mi W Payette
Lake, 5400', 2 MVZ; summit. Smith Mt., 7500', 4 KU; Tamarack, 1 USNM;
Bannock Co.: 1 mi W Bancroft, 5300', 1 KU; Pocatello, 2 KU, 1 USNM; 9.5
mi E Pocatello, W fork Rapid Cr., Barrett Ranch, 1 MVZ; 6.5 mi E Pocatello,
N fork Pocatello Cr., 1 M\^Z; 2.5 mi NW Pocatello, 2 UM; E Pocatello,
Schutt's Mine, 6500', 4 MVZ; 5 mi E Pocatello, S fork Pocatello Cr., 1 MVZ;
Swan Lake, 1 USNM; Blaine Co.: Alturas (=:Sawtooth) Lake, 7000', 1 MVZ,
1 USNM; Boise Co.: Bald Mt. R. S., 10 mi S Idaho City, 1 USNM; Boise
Nat. For., Deer Park Guard Station, 4800', 1 USNM; Boise Nat. For., Hunter

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 33

Cr., 4900', 1 USNM; Bonner Co.: Cabinet MLs., Priest Lake, 4 USNM; 5 mi
W Cocolalla, 3500', 7 M\'Z; Bonneville Co.: Irwin, 1 USNM; 10 mi SE Irwin,
2 USNM; Boundary Co.: Cabinet Mts., E of Priest Lake, 1 USNM; Butte Co.:

1 mi W Arco, 1 DCM; Canyon Co.: Nampa, 1 MVZ, 5 USNM; Cassia Co.:
Alljion, 1 USNM; Clearwater Co.: 2 mi NE Weippe, 3000', 1 MVZ; Elmore
Co.: Cayuse Cr., 10 mi N Featherville, 1 USNM; Trail Cr., Boise Nat. For.,
4850', 1 USNM; Franklin Co.: 20 mi NE Preston, 1 MVZ; Goodinfi Co.: 2
mi E Hagerman, 1 M\'Z; Idaho Co.: 4 mi SSW Lolo Pass, Bmshy Cr., 4000',
7 CAS; 3 mi SW Selway Falls, 1 CAS; 4 mi SW Selway Falls, 2 CAS;
Kootenai Co.: Coeur d'Alene, 3 USNM; Coeur d'Alene, Deception Cr. Exp.
Forest, 1 UM"; Coeur d'Alene, near mouth, Montford Cr., 2 USNM; Hoodoo
\'alley, 2 CAS-SU; Latah Co.: Cedar Mts., 1 CRCM**; Cedar Mts., 1 mi S Cedar
Peak, 1 CRCM"; Cedar Mts., WSC Camp, 2 CRCM"; Leivis Co.: Nez Perce,

2 USNM; Poicer Co.: (near?) Pocatello (in Bannock Co.), 1 KU; 4 mi S
Portnouf R., Bannock Cr., 1 MVZ; Shoshone Co.: Glidden Lakes, 5700', 1
MVZ; Mullan, 2 USNM; Osburn, 1 USNM; Tivin Falls Co.: 8 mi W Roger-
son, Salmon Cr., 1 MVZ; Valley Co.: Payette Nat. For., Landmark Ranger
Station, 10 mi E Warm Lake, 6000', 1 USNM; 5 mi E Warm Lake, 7000'^, 2
MVZ; Washington Co.: SW slope Cuddy Mt., 1 mi NE Heath, 4000', 5 KU;
MONTANA: Flathead Co.: Glacier Nat. Park, Camas Cr. (cabin) 2.5 mi ENE
Kinda Lake Rd., 3800', 8 UM*; Glacier Nat. Park, 2 mi N Camas Cr. cabin, 3700',
1 UM"; Glacier Nat. Park, Fern Cr., on Kintla Lake Rd., 3500', 3 UM°; Glacier
Nat. Park, Fish Cr., on Kinda Lake Rd., 3300', 2 UM"; Glacier Nat. Park,
Fish Cr., 4 USNM; Nyack, 1 USNM; Flathead-Glacier Co. Line: Summit, 2
USNM; Granite Co.: Rock Cr. drainage. Cougar Cr., 1 UM; Frog Pond Basin,
1 UM; Lake Co.: Flathead Lake, 1 UM*", 7 USNM; Flathead Lake, base
Finley point, 1 UM"; Flathead Lake, E shore (Yellow Bay), MSU [=UM]
Biol. Sta., 2950', 4 UM°; 0.25 mi NW UM Biol. Sta., 2950', 2 UM*; 0.5 mi
E MSU [=UM] Biol. Sta., 3200', 1 UM*; 1.25 mi S UM Biol Sta., 1 UM*;
4.5 mi N UM Biol. Sta., 3100', 6 UM*; 5 mi N UM Biol. Sta., 3200', 10
UM*; Yellow Bay Cr., 3000', 11 UM*; Yellow Bay State Park, 2950-3000', 5
UM*; Moiese, Nat. Bison Range, 3000', 4 UM*; 0.2 mi N Moiese, 2600', 2
KU; Ninepipe Reservoir, 1 UM*; 2 mi W Ronan, 1 UM*; Lincoln Co.: Leigh
Cr., 3600', 1 UM; Missoida Co.: Lolo, 1 UM*; 6.5 mi W Lolo, Lolo Cr.,
3470', 2 MVZ; Missoula, Buckhouse Bridge, 3 UM*; Missoula, Deer Cr. (.sev-
eral localities), 11 UM*; Missoula, Greenough Park, 12 UM*; Missoula, Pat-
tee Canyon (several localities), 11 UM*; Missoula, Missoula Country Club, 1
UM*; Missoula, Rattlesnake Cr. (several localities), 6 UM*; Mis.soula, UM
Arboretum, 3 UM*; 15 mi WNW Missoula, 1 UM*; Ravalli Co.: Bass Cr.,
1 UM*; Bass Cr., NW Stevensville, 4600', 3 USNM; Conner, Medicine Hot
Springs, 1 DCM*; Corvallis, 12 DCM*, 7 USNM; 3 mi S Darby, 2 DCM*;
Florence, 6 MSUDZ*, 3 USNM; Florence, Sweeney Cr., 1 MSUDZ*; 1 mi E
Florence, 1 UM*; 2 mi W Florence, 3 USNM; 3 mi SW Florence, 1 USNM;
Hamilton, 3600', 10 DCM*; Hamilton, S city park, 3 UM*; 0.5 mi W Hannl-
ton, 1 DCM*; 1 mi E Hamilton, 6 DCM*; 2 mi E Hamilton, 3700', 35 DCM*;

3 mi E Hamilton, 3700', 129 DCM*; 4 mi E Hamilton, 3800', 8 DCM*; 5 mi
E Hamilton, 8 DCM*; 6 mi E Hamilton, 3700', 1 KU; 10 mi S Hamilton, 2
DCM*; Ross Hole (Sula), 3 DCM*; Steven.sville, 6 USNM; 7 mi N 9 mi E
Stevensville, Three-mile Cr., 4500', 35 UM*; 8 mi S 9 mi E Stevens\ille,
Burnt Fork, 5050', 10 UM*; Sanders Co.: Prospect Cr., 5 USNM; Thompson
Pass, 1 USNM; NEVADA: Elko Co.: Jarbidge Mts., summit between heads.
Copper and Coon Crs., 4 KU; Mountain City, 1 USNM; Ruby Lake, 3 USNM;
Ruby Mts., 7 USNM; Ruby Mts., Harrison R. S., Green Mt. Canyon, 6 KU;
Ruby Mts., S fork Long Cr., 4 KU; Ruby Mts., W side Ruby Lake, 3 mi N
White Pine Co. Line, 8 KU; Ruby Mts., summit. Secret Pass, 6200', 1 KU;
Ruby Mts., Thomas Cr., 7600', 1 MVZ; Eureka Co.: Evans, 1 MVZ; Lander

34 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Co.: Kingston R. S., 7500', 3 MVZ; Lander-Nye Co. line: Reese R., 6000-
6890', 3 USNM (incl. holotype S. nevadensis); Nye Co.: Bell's Ranch, Reese
R., 6890', 2 MVZ; Cloverdale, 1 USNM; Ormshy Co.: Marlette Lake, 8000',
1 MVZ; White Pine Co.: Baker Cr., 6600', 1 MVZ; Baker Cr., 8000', 2 MVZ;
Baker Cr., 11100', 4 MVZ; Ruby Mts., W side Ruby Lake, 3 mi S Elko Co.
Line, 6100', 1 KU; OREGON: Baker Co.: Anthony, 5 MCZ, 3 USNM;
Bourne, 2 USNM; Cornucopia, 8 USNM; Homestead, 1800', 1 USNM; 3 mi
NE Huntington, 2100', 1 USNM; E Pine Cr., 2.5 mi NE Cornucopia, 1 USNM;
Rock Cr., 1 USNM; McEwen, 1 USNM; Benton Co.: Coivallis, 1 KU; 0.5 mi
W Corvallis, 1 KU; 1 mi S Corvallis, 1 KU; Clackamas Co.: Estacada, 1 KU;

10 mi S Skinner Ranch, 1 UM; Crook Co.: Maury Mts., 3 USNM; Curry Co.:
Gold Beach, 3 USNM; Port Orford, 1 USNM; Grant Co.: Austin, 1 USNM; 15
mi NW Austin, 5400', 1 USNM; 21 mi SE Prairie City, N fork Malheur R.,
5000', 1 MVZ; Strawberry Mts., 12 USNM; Harney Co.: Burns, 4 USNM;
Diamond, 1 USNM; Steens Mts. (T32S R33E Sec 32), 1 MVZ; Hood River
Co.: W slope Mt. Hood, near timberline, 1 USNM; N slope Mt. Hood, 2800',
1 USNM; Jackson Co.: Ashland, 1975', 1 USNM (holotype S. tri^onirostris);
Ashland, 5 SOMNH; Ashland, Scenic Hills Mem. Park, 6 SOMNH; Ashland,
Hy. W., 1 mi, 1 SOMNH; Field N of Ashland airport, 1 SOMNH; Denman
Game Mgt. Area (T36S RIW Sec 30), 2 SOMNH; E fork, Ashland Cr., 0.6
mi N junct. F.S. rd. 3909, 1 SOMNH; Emigrant Cr., below Dead Indian Bridge,
1 SOMNH; W slope. Grizzly Peak, 3500', 1 USNM; 6 mi S Medford, 1 MVZ;
No exact locality, 1 SOMNH; Klamath Co.: Diamond Lake, 1 USNM; Ft.
Klamath, 2 USNM; Klamath Falls, 5 USNM; W Silver Cr., 10 mi SW Silver
Lake, 3 USNM; Yamsay Mts., Yamsay R., 4800', 1 USNM; Lake Co.: 10 mi
SE Lakeview, 1 USNM; Malheur Co.: Jordan R., 8 mi W Jordan Valley, 1
USNM; Multnomah Co.: Portland, 1 KU; Tillamook Co.: Tillamook, 1 USNM;
No exact locality, 3 KU; Umatilla Co.: Meacham, 3 USNM; 10 mi W
Meacham, 2 USNM; 15 mi E Weston, Langdon Lake (=Tollgate), 3 MVZ;

11 mi E Weston, 4 mi W Langdon Lake, 2 MVZ; Blue Mts., Meacham L.,
3800', 2 MVZ; Union Co.: Elgin, 2 USNM; Hot Lake, 2 USNM; Kamela, 2
USNM; Wallowa Co.: Enterprise, 25 mi N Sled Springs, 4600', 4 USNM; 16
mi S 3 mi E Lostine, 5500', 1 MVZ; Wallowa Lake, 4000-4800', 6 MVZ; 21
USNM; Wheeler Co.: 11 mi W 7 mi S Mitchell, 4850', 3 MVZ; UTAH: Box
Elder Co.: Mouth of Bear R., 2 USNM; Bear R. Refuge, 1 USNM; Brigham
City, 4500', 1 USNM; Cache Co.: Tony Grove, Logan Canyon, 6400', 1 MVZ;
Davis Co.: Bountifvil, 0.5 mi SW of oil refinery, 4300', 1 UU; W Clearfield,
W of Smithfield cannery, 1 UU; Farmington Bay area, 4 UU; Farmington Bay,
jet. Tuq:)in and Main ditches, 4250', 1 UU; Farmington Lake, 1 UU; W Lay-
ton, 1 UU; Steeds Canyon, 1 UU; Sah Lake Co.: City Cr. Canyon, 4600', 1
UU; 1 mi W Draper, 4500', 2 UU; Rocky Glen at mouth of Emigration
Canyon, 4880', 1 UU; Lamb's Canvon, Salamander Lake, 9000', 1 UU^ Mid-
vale, 1 USNM; Midvale, "9000 S" [block] and Jordan R., 4200', 1 UU; Mount
Dell Canyon, 5400', 1 UU; Parley's Canyon, 5000', 2 UU; Salt Lake City, 1
USNM; 1 mi W Salt Lake City aiiport, 1 UU; Spring Run, 9tli E and 56 S,
4400', 1 UU; Utah Co.: Palmyra, 1 BYU; Provo, 1 BYU, 1 USNM; Provo City
Park, Provo Canyon, 1 BYU; Provo, Rock Canyon, 1 BYU; 4 mi NE Provo,
1 UU; Spanish Fork Bench, 1 BYU; Weber Co.: Farr West, 1 UU; Hooper
Bay Refuge, 1 UU; Ogden, 4 USNM; Ogden, Melan's Basin, Mt. Ogden,
9000', 1 UU; N side Pine View Lake, 1 UU; Riverdale, 4200', 1 UU; River-
dale, 4250', 1 UU; 2 mi W Roy P. O., 5 UU; South Ogden, 1 UU; Birch Cr.,
1 mi N South Ogden, 4500', 1 UU; Snow Basin, 2 UU; Snow Basin, S part
of Wheeler Canyon, 1 UU; Uintah, 1 UU; WASHINGTON: Adams Co.:
Twelve-mile Slough, 1 CRCM"; Asotin Co.: 5 mi W Anatone, 1 MVZ; 9 mi
W Asotin, 1 CRCM^ Bly, 1000', 1 USNM; Rogersburg, 1 USNM; Chellan
Co.: Entiat R., 20 mi from moutli, 1680', 2 USNM; Clallam or Jefferson Co.:

REVIEW OF THE TAXONOMY OF SOREX VAGRANS SPECIES 35

Olympic Mts., 1 USNM; ColumhUi Co.: Blue Mts., 21 mi SE Dayton, 1
USNM; Godinan Spring, 2 MVZ; Ilumfrcy C'llunpeg"?) Falls, 1 MVZ; Star-
buck, 645', 1 CRCM", 1 USNM; Grant Co.: Moses Lake, 1000', 1 USNM; 10
mi S Moses Lake, 2 USNM; Kin^ Co.: Seattle, 3 KU; Kiftitas Co.: 21 mi S
Blewett Pass, 3000', 3 USNM; Easton, 2 USNM; Ellensburg, 1500', 2 USNM;
Leivis Co.: 7 mi E Centralia, 1 USNM; Lincoln Co.: 4.5 mi SW Davenport,
1 CRCM*; 11 mi N Odessa, 1 CRCM"*; Sylvan Lake, 6 mi E Odessa, 4
USNM; Sprague Lake, 1 USNM; Okanogan Co.: E end, Bauerman Ridge,
near head, Haig Cr., 6500', 1 USNM; Conconullv, 2 USNM; Hidden Lake,
4100', 1 USNM; Loomis, 1300', 1 USNM; Oroville, 1 USNM; Oroville,
Osoyoos Lake, 1000', 1 USNM; Sheep Mt. (Park Mt.), 6500', 3 USNM; Twisp,
1600', 1 USNM; Pacijic Co.: Shoalwater (Willapa) Bay, 2 USNM (incl. holo-
type); Shoalwater Bay, Tokeland, 3 USNM; Pcnd 'Oreille Co.: 9 mi N
Metaline, 2600', 2 USNM; Newport, 1 USNM; Salmo Mt., Salmo Pass (sum-
mit), 5600', 1 CRCM**; Sullivan Lake, 3000', 1 CRCM", 1 USNM; Fierce Co.:
Mt. Rainer, Bear Prairie, 1 USNM; Puyallup, 2 USNM; Steilacoom, 3 USNM
(incl. holotype, S. suckleiji); Tacoma, 2 UM; San Juan Co.: San Juan Is., 4.5
mi NW Friday Harbor, 4 KU; 1.5 mi W Friday Harbor, 1 KU; Skafiit Co.:
Cypress Is., 1 KU, 2 MVZ; Mt. Vernon, 1 USNM; Skamania Co.: 5 mi N
Willard, 1 USNM; Snohomi^ih Co.: 0.5 mi N 7 mi W Marysville, 3 KU;
Spokane Co.: Marshall, 7 USNM; Stevens Co.: Chewelah, 2 KU; 5 mi S Col-
ville, 1 KU; 15 mi E Colville, 1 USNM; Marcus, 1 USNM; ThurMon Co.:
Olympia, 5 USNM; Wahkiakum Co.: Cathlamet, 1 USNM; Walla Walla Co.:
College Place, 1 KU; Walla Walla, 1 KU; Whatcom Co.: Barron, 5000', 1
USNM; Whitman Co.: Armstrong, 1 CRCM'; Pullman, 1 CRCM*, 2 USNM;
1 mi E Pullman, 1 CRCM"; 2 mi N Pullman, 2 CRCM*; 4 mi ENE Pullman,
2400', 1 KU; 13 mi S Pullman, 1 CRCM^ Tekoa, Hangman Cr., 1 CRCM*;
Wawawai, 4 CRCM*; 5 mi NE Wawavvai, 1 USNM; Yakima Co.: Mt. Adams,
Bird Lake, 3 USNM; Selah, 5 KU; Wiley City, 10 mi W Yakima, 2000', 4
USNM; Yakima Indian Resen'ation, Signal Peak, 4000', 1 USNM; WYOMING:
Lincoln Co.: 10 mi N Afton, 6200', 3 USNM; 13 mi N 2 mi W Alton, 6100',
6 KU; 9 mi N 2 mi W Afton, 6 KU; 7 mi N 1 mi W Afton, 3 KU; Cokeville,
6400', 1 USNM; 12 mi N 2 mi E Sage, 2 KU; 6 mi N 2 mi E Sage, 1 KU.

S. vagrans vancoucerensis. Total number, 16. — BRITISH COLUMBIA
(Vancouver Island): Cowichan Lake, 1 USNM; Colwood, 1 USNM; Departure
Bav, 1 USNM; Errington, 1 MVZ; French Cr., 1 MVZ; Golden Eagle Mine,
1 MVZ; Coldstream, 1 M\'Z, 1 USNM (holotype); Nanaimo, 3 USNM; Parks-
ville, 1 MVZ; Satuma Is., 4 MCZ.