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Ss. “8 DEPARTMENT ¢ OF ‘AGRICULTURE
: BUREAU OF BIOLOGICAL SURVEY
ag) ENR W. HENSHAW, Chief

NORTH AMERICAN FAUNA,

oe No. 36.

[Actual date of publicati on, June 5, 1914]

“REVISION OF THE AMERICAN HARVEST MICE

(Genus we Een es

BY
“ARTHUR EE. HOWELL=

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY —

- c
z WASHINGION ~ es & :
ae GOVERNMENT PRINTING OFFJOB © > Loe
- - 1984

Peo DEPARTMENT OP AGRICULTURE
BUREAU OF BIOLOGICAL SURVEY

HENRY W. HENSHAW, Chief

NORTH AMERICAN FAUNA

No. 386

[Actual date of publication, June 5, 1914]

REVISION OF THE AMERICAN HARVEST MICE

(Genus REITHRODONTOMYS)

BY

me Ee Es HOW EEL

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY

\ =

WASHINGTON
GOVERNMENT PRINTING OFFICE
1914

Pehrek OF PRANSMITTAL.

UniTED States DEPARTMENT OF AGRICULTURE,
BuREAU OF BIOLOGICAL SURVEY,
Washington, D. C., January 3, 1914.

Srr: I have the honor to transmit herewith, for publication as
North American Fauna No. 36, the results of a study of the American
harvest mice by Arthur H. Howell, assistant biologist, Biological
Survey.

The American harvest mice occur abundantly in or near meadows
and cultivated lands over a large part of the United States. Their
exact economic relations are still little known, but they belong to a
group of mammals many species of which are injurious to agriculture,
and there is no doubt that they consume large quantities of forage
and some grain.

Up to the present time the relationships and distribution of the
many species have been imperfectly understood. The present report
furnishes for the first time a complete systematic synopsis of the
group, with maps showing the ranges of the species of most economic
importance.

Respectfully, H.W. HEnsHaw,
Chief, Biological Survey.
Hon. Davin F. Hovuston,
Secretary of Agriculture.

ea ie i

Hive

CONTENTS.

UpmRe OUR MeaRIPI oe A eo Nk cee wich ewe vate 2 ob we dioucieiene bance
Peete aie material. 2. 22 Se ~ es - 55-5: - ieteitaye yas ce staat os oe
Se DIS USENET ly 99 yey oo) oe had Sccaetera o ce bite nie oad

ee SMT IANS 52 so 5 on ane evAE Sones He nieie mon be es Sec sh
Sipe SPE EIN Seles oP is Ly ate ies seeks peice a
List of species and subspecies, with type localities..............-..-------
Meme spesies ord subspecies. ..... ..- 2.22242 2h se seit whee eek ede
SE RIOD OUI Seer seo A887 oo a he Salk 2 Diaries ice ein © seek
SPE ERIE ELIOIL ES Sel aera Says Gre Alia ara cyanyne aya OL cee migew obo cies we
PEM pe ALTMCASUTCIMONGS occa = - a ae sft et we Dans Se ena slew eetionse

Pruate I.

lf

Til.

WEUe

ILLUSTRATIONS.

PLATES.

Skulls (upper views) of Reithrodontomys h. humulis, R. h. merriami,
R. a. albescens, R. a. griseus, R. montanus, R. m. megalotis, R. m.
dychei, R. m. saturatus, R. m. zacatece, R. m. alticolus, R. r.
raviveniris, R. 71. haliewtes. .. . 22223 na- ee eee

Skulls (upper views) of Reithrodontomys amenus, R. otus, R. a.
australis, R. dorsalis, R. c. colime, R.r. luteolus, R. f. tenuis, R. f.
dificilis, R. f. helvolus, K.f_.doliecus.._.0-< = oes ee eee

Skulls (upper views) of Reithrodontomys hirsutus, R. levipes, R. c.
chrysopsis, R. m. mexicanus, R. m. cherru, R. m. goldmani, R.
gracilis, R. t. aureus, R. m. microdon

errr ecececesezreeceeree ee sceeree ee ecec ce

. Skulls (lower views) of Reithrodontomys h. humulis, R. h. mer-

riami, R. a. albescens, R. a. griseus, R. montanus, R. m. mega-
lotis, R. m. dychei, R. m. saturatus, R. m. zacatece, R. m. alti-
colus, R. r. raviventris, R. r. halicetes

ere eee eee ee eee eee eee see eee ese

. Skulls (lower views) of Reithrodontomys amenus, R. otus, R. a.

australis, R. dorsalis, R. c. colime, R. r. luteolus, R. f. tenuis,
RK. f. dificls, R. f. helootus; hf. oli

. Skulls (lower views) of Reithrodontomys hirsutus, R. levipes, R. c.

chrysopsis, R. m. mexicanus, R. m. cherrii, R. m. goldmani,
R- gracilis, R. t. aureus, R.m. amicrodon..- ==
Molar teeth of Reithrodontomys m. megalotis, R. t. aureus, R. levipes,
R. m. mexicanus, R. c. chrysopsis, R. h. merriami; skull of type
of R. m. mexicanus

TEXT FIGURES.

Fie. 1. Distribution of Reithrodontomys humulis, R. albescens and subspecies...
2. Distribution of Reithrodontomys montanus, R. megalotis and subspecies.
3. Distribution of Reithrodontomys fulvescens and subspecies. .--.-.--------
4. Distribution of Reithrodontomys rufescens, R. dorsalis, R. australis,

R. colimz, R. allent and subspecies .:....._.. 12222 aeeeeeeeee

5. Distribution of Reithrodontomys chrysopsis, R. perotensis, R. ips,

R. hirsutus and subspecies

6. Distribution of Reithrodontomys mexicanus, R. gracilis and subspecies. -

6

Page.

82

84

86

88

90

92

94

20
27
45

58

67
73

No, 36, NORTH AMERICAN FAUNA, June, 1914,

REVISION OF THE AMERICAN HARVEST
MICE.

(Genus REITHRODONTOMYS.)
By Artaur H. HoweEtt.

INTRODUCTION.
HISTORY AND MATERIAL.

The mice of this genus have been known to naturalists since the
days of Audubon and Bachman. At that time only a single species,
the eastern harvest mouse, was known, it having been described by
Bachman in 1841 under the name of Mus humulis from specimens
collected near Charleston, 8. C. The following year (1842) the same
authors redescribed this species as Mus leconti from a specimen
collected by Maj. John Leconte in Liberty County, Ga.

Although the species was known from only a few localities (Geor-
gia, South Carolina, and Virginia), and considered rare, its habits
were quite fully described by Bachman in ‘‘Quadrupeds of North
America,’’ published in 1851. The spelling of the name was there
changed to “‘humilis.”” Even at that time Bachman appreciated
the fact that this species was not closely related to true Mus, as is
indicated by the following statement:

In examining the teeth of this species, we have found that the tuberculous sum-
mits on the molars were less distinct than in those which legitimately belong to the
genus Mus, and that there are angular ridges on the enamel by which it approaches
the genus Arvicola; it is in fact an intermediate species, but in the aggregate of its
characteristics perhaps approaches nearest to Mus, where we for the present have
concluded to leave it.!

His keenness in noting the distinction between the molars of the
harvest mouse and of Mus makes his failure to mention the groov-
ing of the incisors all the more remarkable, but as pointed out by
Osgood ? the rest of the description fits the species so well that there

1 Aud. & Bach., Quad. N. Am., II, 1851, p. 106.
£ Proc. Biol. Soc. Wash., XX, 1907, pp. 49-50.

8 NORTH AMERICAN FAUNA. [No. 36.

can be no doubt of the applicability of the name humulis to the
harvest mouse of the Atlantic coast.

In 1853 Leconte pointed out that ‘‘ Mus Lecontet of Bachman is a
Reithrodon, and neither a Mus nor a Hesperomys.”’ }

In 1855 Baird described a second species in the genus, Reithrodon
montanus, based on a single specimen collected on one of the Govern-
ment exploring expeditions in the Rocky Mountains, the exact local-
ity being unknown. This specimen remained unique in collections
for over 50 years, and not until 1907 was a series of topotypes secured,
making possible accurate characterization of the species. In his
great work on the ‘‘Mammals of North America,” published in 1857,
Prof. Baird recognized four valid species—humilis and montanus,
already described, and megalotis and longicauda, proposed as new,
the range of the genus being extended to the Mexican border and to
the coast of California. <A fifth species, R. carolinensis (Aud. &
Bach.), was provisionally recognized.

Baird realized that considerable differences existed between true
Reithrodon of South America and the North American mice placed
in the same genus, but as he had never seen either skins or skulls of
the former, he found it impossible to indicate the discrepancies.

In 1860 De Saussure described a species from Vera Cruz, Mexico,
as R. mexicanus and the next year another, R. swmichrasti, also from
Mexico but without definite locality.

In 1874 Coues published a synopsis of the genus,” pointing out the
characters which distinguish it from true Reithrodon and proposing
the name Ochetodon for the North American species. Five species
were recognized in this paper—humulis, longicauda, mexicanus, mon-
tanus, and sumichrasti, the last two provisionally. &. megalotis was
placed in synonymy under humilis. Using practically the same
material, he published in 1877 a more detailed revision of the genus *
in which he recognized only four species, having dropped O. sumi-
chrastt even from synonymy.

The name Ochetodon quickly became current, American naturalists
having overlooked the name Reithrodontomys proposed by Giglioli
some monthsearlier than Ochetodon, and not until Merriam called atten-
tion to this fact in 1892,* was the earlier name given precedence.

In 1893 Allen revived the names megalotis and montanus, placing
them formally in the genus Reithrodontomys and naming a new form,
R. aztecus, from New Mexico.°

In 1895 Allen published an extended revision of the genus ® in
which he recognized 15 forms, 8 of which were there first described,

1 Proc. Acad. Nat. Sci. Phila., 1853, p. 410. .
2 Proc. Acad. Nat. Sci. Phila., 1874, pp. 184-186.

8 Mon. N. Am. Rodentia, Rept. U. 8. Geol. Surv. Terr., XI, 1877, pp. 120-130.

¢ Proc. Biol. Soc. Wash., VII, 1892, p. 26.

5 Bull. Am. Mus. Nat. Hist., V, 1893, p. 79.
6 Bull. Am. Mus. Nat. Hist., VII, 1895, pp. 107-143.

1914.] INTRODUCTION. 9

and the range of the genus was shown to extend practically across
the continent from the Carolinas to California and south to Costa
Rica. Of the 15 forms recognized, three (nebrascensis, deserti, and
pallidus) are considered synonyms by the present writer. Dr.
Allen’s material consisted of 920 specimens, chiefly from the United
States. The paucity of material from southern Mexico, where the
genus reaches its highest development, of course accounts for the
incompleteness of this revision in so far as the southern forms are
concerned. Only 2 specimens from southern Mexico and 17 from
Costa Rica were at that time available—only 19 specimens from a
region where over 35 subspecies are now known.

As a result of increased activity in the collection and study of
North American mammals, new species were discovered almost
every year. In 1898 Thomas described RK. séderstrémi from Ecuador,
thus extending the range of the genus into South America. In 1901
Merriam published a paper?’ giving descriptions of 23 new forms,
based chiefly on the unexampled collections of Nelson and Goldman
in Mexico and Guatemala. Since that date 9 additional species
have been described, bringing the total number of named forms up
to 67, of which 16 are at present regarded assynonyms. Fifty-eight
. forms are recognized in the present revision, of which 7 are here
described as new.

The material on which the present revision is based consists of
2,280 specimens contained in the collections of the United States
National Museum, including that of the Biological Survey, and the
Merriam Collection. This material has been supplemented by a con-
siderable number of specimens borrowed from other American muse-
ums, thus bringing the total number of specimens examined up to
2,583. All the existing types have been examined with the excep-
tion of R. tenuis, BR. séderstrémi, and R. modestus, which are in the
British Museum, and R. mexicanus and R. sumichrasti, which are in
the Geneva Museum. Of the last two I have seen photographs of
the type skulls, while of tenuis and modestus I have examined prac-
tical topotypes. The National Museum collections contain repre-
sentatives of all but 5 or 6 of the recognized forms and in most cases
considerable series showing all ages and pelages.

In this connection I wish to acknowledge the kindness of the fol-
lowing gentlemen, who have generously loaned from the collections
under their charge such material as was needed to supplement the
collections in the National Museum: Dr. J. A. Allen, of the Ameri-
can Museum of Natural History; Mr. Outram Bangs, of the Museum
of Comparative Zoology; Mr. F. J. V. Skiff, of the Field Museum of
Natural History; Mr. Witmer Stone, of the Academy of Natural Sci-

1 Proc. Wash. Acad. Sci., IIT, 1901, pp. 547-558.

10 NORTH AMERICAN FAUNA. [No. 36.

ences of Philadelphia; and Mr. Joseph Grinnell, of the Museum of
Vertebrate Zoology, Berkeley, Cal. I am indebted, also, to Mr. Ger-
rit S. Miller, jr., and Mr. Ned Hollister for valuable criticism.

DISTRIBUTION,

The genus Reithrodontomys is clearly of Austral origin, being most
abundant and highly developed in southern Mexico.

In western North America the typical subgenus ranges practi-
cally throughout the Upper and Lower Austral Zones, the northern
limits of its range being indicated by the following localities where
specimens have been taken: Prescott, Wash.; Custer, Mont.; and
Fort Clark, N. Dak. ast of the Mississippi River its range is more
restricted, and so far as known the genus does not occur north of the
Ohio and Potomac Valleys, the most northerly records being at
Ceredo, W. Va., and Falls Church, Va. Southward the subgenus
Reithrodontomys ranges through Mexico and Central America to
Panama, but it is not known from South America.

The sabe genus Aporodon’ ranges from central Mexico (Jalisco and
Vera Cruz) south throughout Central America and thence to Ecua-
dor, its exact limits being very imperfectly known.

th the United States the genus is confined to the Austral Zones,
but in Mexico and Central America ranges from the Tropical at or
near sea level through all the zones to and including the Canadian
at timber line.

HABITS AND ECONOMIC STATUS.

The harvest mice are preéminently field mice. Practically all the
known species live in more or less open grassy situations and are
partial to neglected fields overgrown with grasses or sedges and to
weedy and grassy borders of cultivated tracts. Many species seem
to prefer moist situations, and some (as R. ravwentris) live exclu-
sively in wet marshes, either salt or fresh; others (notably #. albescens
and 2. montanus) are found only in dry, sandy uplands. In the arid
West in general harvest mice are most often taken along the grassy
borders of sloughs, small streams, or irrigation ditches; but they may
be found in almost any situation where there is sufficient cover of
vegetation to hide them from their enemies.

Certain Mexican and Central American species ascend the moun-
tains to timber line and live in grassy openings or among brush in
the more open parts of the forest. R. séderstrémi, of South America,
is described as living in gardens among climbing plants.

Some species, and perhaps all, construct substantial nests of
grasses, often lined with soft materials and placed either on the

1 See page 63.

1914.] INTRODUCTION. Ti

ground or in vines, bushes, or low trees some distance above the
ground. Deserted birds’ nests are sometimes used as a base for the
nest of the mouse. Mr. H. P. Attwater has found their nests in old
woodpecker holes in fence posts, and Mr. Howard Lacey, at Kerr-
ville, Tex., has found them on cornstalks and made of cornsilk. The
breeding season extends in northern latitudes from April to October
and in tropical regions may cover the entire year. The number of
young produced at a birth varies from three to seven. ;
All the species live chiefly above ground, but burrows are also
used, and cracks and openings in the ground are often occupied by
the nests. ‘The mice sometimes travel in narrow beaten paths of their
own, and also are caught in the runways of other mammals, particu-
larly those of cotton rats (Sigmodon) and meadow mice (Microtus).
They are both nocturnal and diurnal in habit and remain active

during the entire year.
FOOD.

The food of harvest mice consists largely of seeds and grain, with
considerable green vegetation and occasionally fruit. Our knowl-
edge of their preferred habitat indicates that most of their food
moust be obtained from wild plants of little or no value to man.

Bachman, who had studied the habits of the eastern harvest mouse
rather closely, writes of it as follows:

We doubt whether this species is of much injury to the farmer. It consumes but
little grain, is more fond of residing near grass fields, on the seeds of which it subsists,
than among the wheat fields. We have observed in its nest small stores of grass
seeds—the outer husks and other remains of the broom grass (Andropogon dissiti-
florum)—also that of the crab grass (Digitaria sanguinalis), and small heaps of the
seeds of several species of paspalum, poa, and panicum, especially those of panicum
Ttalicum.*

Mr. H. P. Attwater, writing of Reithrodontomys intermedius, as
observed in Bexar County, Tex., says:

These mice seem to be fond of peaches, eating the peach and leaving the stone
hanging on the tree.”

Mr. E. A. Goldman, at Metlaltoyuca, Mexico, captured a specimen
of R. mexicanus goldmani on a bunch of bananas hanging about 8
feet above the ground.

Only rarely is any damage to crops reported. Mr. C. W. Seeg-
miller, of St. George, Utah, states that he has known harvest
mice to do some damage by climbing grain stalks and cutting off
the heads. He has found their nests built several feet from the
ground in close-growing clusters of grain stalks. Prof. D. E. Lantz
reports that in eastern Kansas Reithrodontomys dychei is often found
in the fall under shocks of wheat and corn, and in such situations it
may be expected to glean some of the grain.

1 Aud. & Bach., Quad. N. Am., IT, 1851, p. 105.
2 Allen, Bull. Am. Mus. Nat. Hist., VIII, 1896, p. 236.

12 NORTH AMERICAN FAUNA. [No. 36.

Although in most places harvest mice are only moderately numer-
ous, in a few regions they occur in great abundance.

PELAGES.

The mice of this genus usually molt but once a year, normally in
the late fall (at least in northern latitudes), but the changes produced
in the pelage by wear and fading are so great that the summer pelage
of adults differs markedly from their winter pelage. The young are
usually less ochraceous and in some species darker than adults, but
the difference between young and adults is less marked than in
Peromyscus and other related genera. Of the forms examined,
R. humulis merriami presents the greatest differences between young
and adults.

Immature or subadult individuals taken in summer and fall always
present a fresh, unworn appearance, the color usually being decidedly
paler than in fresh winter pelage. Adults taken in midsummer or
early fall are almost always in a more or less worn condition—often so
much so as to appear ‘‘ragged’’—but occasionally one may be found
in summer in fresh pelage. The fresh, full pelage is retained during
most of the winter, and little evidence of wear is seen before March
or April. As the tips of the hairs wear off, the pelage in most species
becomes redder, and by midsummer the evidences of fading are often
pronounced, individuals and local colonies, however, showing marked
differences in the amount of wear and the date of its appearance.
Excessively worn individuals of certain species QUE AI R. mega-
lotis) often become very pale and gray.

Specimens showing clearly the process of molting are comparatively
rare, and in the majority of cases the molt seems to progress insidi-
ously, new hairs coming in simultaneously over all parts of the body.
In a few cases, however, the line of demarcation is plainly evident,
new pelage appearing first on the hinder back and along the sides,
the fore part of the back being the last portion to be invested. ‘This
is equally true of young and adults.

While the fall molt is doubtless normal in northern latitudes, occa-
sional exceptions have been noted, indicating the occurrence (prob-
ably abnormal, at least rare) of a spring or midsummer molt. A
young adult male specimen of R. megalotis dyche, from Meadow,
Wyo., taken June 28, 1909, has nearly completed a full molt, the
new pelage covering the whole of the anterior portion of the body to
the rump, where the old pelage, decidedly redder than the new,
remains. A specimen of R. fulvescens aurantius (also a young adult
male), from Sour Lake, Tex., taken July 18, 1902, shows a fresh, long

1 Specimens showing this molt are as follows:
R. humulis merriami, 9 juv., Lexington, Ky., November 19.
R. megalotis longicauda, f ad., Stanford University, Cal., November 7.
R. megalotis aztecus, 9 subadult, Grand Junction, Colo., November 2.
R. montanus, 9 ad., Medano Ranch, Colo., November 4.

1914.] GENUS REITHRODONTOMYS. 13

pelage covering the anterior half of the body. As in the previous
instance, the new pelage is darker and more ochraceous (less reddish)
than the old. In both examples it will be noticed that the progress
of the molt is directly the reverse of the normal autumn molt, the
rump being the last part of the body to be invested, instead of the
first, as in the usual method.

The molting period of the species inhabiting Mexico and Guatemala
is less strictly confined to the autumn, but as a general rule (with
many exceptions) individuals taken from December to June show a
fuller and fresher pelage than those taken in the other half of the
year. In the entire collection from Mexico I have been able to find
only two specimens showing a clearly marked molting lme. These
are both adult males of R. megalotis alticolus, taken at La Parada,
Oaxaca (in the lower edge of the Transition Zone), August 19, 1894.
In these the molt is nearly completed, the new pelage covering the
anterior portion of the body to the rump.

Genus REITHRODONTOMYS Giglioli.

Reithrodon Leconte, Proc. Acad. Nat. Sci. Phila., 1853, pp. 410, 413; Baird, Mamm.
N. A., 1857, p. 447. (Not Retvthrodon Waterhouse. )

Reithrodontomys Giglioli, Ricerche intorno alla Distrib. Geog. Gener. <Boll. Soc.
Geog. Ital., Roma, XI, May—July, 1874, p. 326. Author’s separates dated 1873
(probably dated from first part of paper in Vol. X), repaged, p. 160. (North
American members of the genus Reithrodon; no species mentioned.) !

Ochetodon Coues, Proc. Acad. Nat. Sci. Phila., December 15, 1874, p. 184 (no type
selected; species included: O. humilis, O. longicauda, O. mexicanus, O. montanus,
and O. sumichrastt). |

Aporodon Howell, postea, p. 63. Type, Retthrodontomys tenwirostris Merriam.

Remarks.—Since no species were mentioned by Giglioli when he
named the genus feithrodontomys, the type must be fixed by subse-
quent designation. Miller and Rehn? have selected Mus lecontu
Aud. and Bach. [=R. humulis] as the type, but this is in violation of
Opinion 46 of the International Commission on Zoological Nomencla-
ture, which advises that the species first associated with the name be
considered the type.* Allen, in 1893, was the first to use the generic
name in connection with a specific name, his paper including three
species—R. megalotis, R. aztecus, and R. montanus.* Since both
megalotis and montanus were published prior to the naming of the
genus, either may be selected as the type. I therefore designate
Reithrodon megalotis Baird as the type of the genus Retthrodontomys.

GENERIC CHARACTERS.

Form murine; tail long, always more than one-third, often more
than one-half total length, slender, scaly, thinly haired; ears promi-

1 Elliot in Pub. Field Columb. Mus., Zool. Ser. III, 1903, p. 145, and several times subsequently, spolls
this name ‘‘ Rhithrodontomys’’.

2 Proc. Boston Soc. Nat. Hist., XXX, 1901, p. 95.

3 Smiths. Inst., Pub. 2060, February, 1912, p. 104. 4Bull. Am. Mus. Nat. Hist., V, 1893, pp. 79-80.

14 NORTH AMERICAN FAUNA. [No. 36. |

nent, often large,.more or less hairy; soles of hind feet 6-tubercu-
late; mamme, 6—P+, 12; no cheek pouches. Skull with smoothly
rounded braincase, more or less inflated, without prominent ridges;
zygomata slender; outer wall of anteorbital foramen a broad thin plate;
anterior palatine foramina relatively large, forming long narrow slits
separated by a thin septum, slightly narrower anteriorly, terminating
about at the plane of anterior border of tooth row; posterior border
of palate square, often with a slight median spine, terminating at
plane of posterior border of tooth row; pterygoids nearly parallel;
audital bull more or less inflated, longer than broad, and obliquely
situated. Descending process of mandible a broad flattened plate,
strongly deflected inward, the lower portion twisted into a nearly
horizontal position and the inner margin raised, leaving a distinct
depression in the ramus;* coronoid process short. Upper incisors
with a deep longitudinal groove near the middle of the tooth. Molars
brachyodont, tuberculate, the tubercles arranged in two longitudinal
series; first upper molar with five principal tubercles, an anterior
median one, and two pairs of lateral ones; m+ and m? with or without
accessory tubercles or enamel loops in the principal angles; upper
molars normally three-rooted, but in some species four-rooted; lower
molars two-rooted.

List of Species and Subspecies, with Type Localities.

Subgenus REITHRODONTOMYS.

R. humulis group:
Reithrodontomys humulis humulis (Bachman). .Charleston, S. ©.

ins pig pan Cee ne ee ee eee ee White Sulphur Springs, W. Va.

humauls merriqii Alena ee) eee Alvin, Tex.

albescens albescens Cary .....-.----------- 18 miles northwest of Kennedy,

Nebr.
albescens, griscusiealleyere pass eee ere San Antonio, Tex.
R. megalotis group:
Reithrodontomys montanus (Baird)......--..-- ‘“Rocky Mountains, latitude 38°”

[=San Luis Valley, Colo., near
San Luis Lakes].

megalotis megalotis (Baird).............--- ‘‘Between Janos, Sonora [=Chi-
huahua] and San Luis Spring”
[New Mexico].

AEGUIOLIS AZLECUS AMVC neta ne ie eee La Plata, N. Mex.

megaotisdyche: Allen wu is 2002.25. 8 Lawrence, Kans.

megalotis nigrescens nobis.......--------- Payette, Idaho.

megalotis longicaudus (Baird). ..--..------ Petaluma, Cal.

megalotis peninsule Elliot..........----- San Quintin, Lower California.

megalotis cinereus Merriam.........------- Chalchicomula, Puebla.

1 Less importance is to be ascribed to this character in distinguishing Reithrodontomys from Peromyscus
than the remarks of Baird (Mamm. N. Am., p. 447) and Coues (Mon. N. Am. Rodentia, p. 122) would
indicate, for although typical Peromyscus differs noticeably on the average from Reithrodontomys in the
shape of the descending process, yet certain species of the subgenus Haplomylomys show practically the
Same condition as Reithrodoniomys.

1914.] LIST OF SPECIES AND SUBSPECIES. 15

R. megalotis group—Continued.
Reithrodontomys megalotis saturatus Allen &

Chapman: 22.2.2... biadeases 2-2 Las Vigas, Vera Cruz.
megalotis alticolus lpetliane. Rwae ee weet ae Cerro San Felipe, Oaxaca.
megalotis arizonensis Allen...............-Chiricahua Mountains, Ariz.
megalotis zacatece Merriam......------.--- Valparaiso Mountains, Zacatecas.
TILES ASIC ELS Sa SMR SS Pinal de Amoles, Queretaro.
Caran WNWObs 2 son ee. os os 2 1 eHanta, Catalina Island, Cal.
raviveniris raviventris Decca cc Saban ee ee Redwood City, Cal.
raviventris halictees Dixon...........-.--- Petaluma, Cal.

R. fulvescens group:
Reithrodontomys fulvescens fulvescens Allen. ...Oposura, Sonora.

Wmineseens tenuis Allen... 222 2.22.-5.-- ++. Rosario, Sinaloa.
JSulvescens intermedius Allen..........----- Brownsville, Tex.
JSulvescens aurantius Allen..............-- Lafayette, La.
JSulvescens difficilis Merriam ..........-.-.-- Orizaba, Vera Cruz.
fulvescens ioltecus Merriam .............-- Tlalpam, D. F., Mexico.
Sulvescens helvolus Merriam.........---.--- Oaxaca, Oaxaca.
Jfulvescens chiapensis nobis........--------- Canjob, Chiapas.
lnelmescens Nelsons NObIS- 2.222525. 22 25 2 - Colima, Colima.
fulvescens mustelinus nobis......-....----- Llano Grande, Oaxaca.
MMPS ENOL. --.-...--.--.-----+--.----Relorma, Oaxaca.
MPENIOCEMAMT 0622.5. 52 c2 2. Le Sierra Nevada de Colima, Jalisco.

R. rufescens group:
Reithrodontomys rufescens rufescens Allen &

DinepiitWers = 22 e5 5525.2. - 282 2S Yo.) dalapa, Vera Cruz.

mpescens LULcOlUS NODIS: 2.0... fe Juquila, Oaxaca.

arta sees 6 he 2 ee NS RS Mountains near Ozolotepec, Oa-
xaca.

conmeiconme Merriam. --.2 22.2 222. Sierra Nevada de Colima, Jalisco;
altitude, 12,000 feet.

Calne merterus Merriam: 222s... Sierra Nevada de Colima, Jalisco;
altitude, 6,500 feet.

ManreeMerriam)- oo) oe) obs cas Calel, Guatemala.

ausizains austrans Allen........2.2...--.2 Volcan de Irazu, Costa Rica.

australis modesius Thomas. .-....-...:.-- Jinotega, Nicaragua.

Subgenus APORODON.
R. levipes group:
Reithrodontomys levipes Merriam........---.-..- San Sebastian, Jalisco.
isis Merriam. 22222222. 3 28 2: . | Ameca. Jalisce:
R. chrysopsis group:
Reithrodontomys chrysopsis chrysopsis Merriam .. Mount Popocatepetl, Mexico.

chrysopsis toluce Merriam........-.--.---- Volcan Toluca, Mexico.
chrysopsis orizabe Merriam......-.-....-- Mount Orizaba, Puebla.
perotensis Merriam..........-...........-Cofre de Perote, Vera Cruz.

R. mexicanus group:
Reithrodontomys mexicanus mexicanus (De Saus-

SOS ai OREN eae a ee Mountains of Vera Cruz.
mexicanus goldmani Merriam.....-...---- Metilaltoyuca, Puebla.
mexicanus cherria (Allen).....-....-...----San Jose, Costa Rica.
ice licnh! 2) fey ys 2 oot Rees Munchique, Colombia.
soderstromt Thomas.............-.2...+..-Quito, Ecuador.

gracilis Allen & Chapman.........-.....-- Chichen Itza, Yucatan.

16 NORTH AMERICAN FAUNA. [No. 36.

R. tenutrostris group:
Reithrodontomys tenwirostris tenuirostris Mer-

PLAID. 2 c fie AI Re PE Nii Oe Todos Santos, Guatemala.
tenwirostris aureus Merriam.........---.-- Calel, Guatemala.
ene per ame wees aie Mato hie Uae Volcan de Chiriqui, Panama.
microdon microdon Merriam...........-..- Todos Santos, Guatemala.
mucrodon albilabris Merriam............-- Cerro San Felipe, Oaxaca.

Key to Species and Subspecies.!
[Based on adults.]

a. Outer wall of anteorbital foramen decidedly broader than width of interpterygoid
fossa; upper molars usually without subsidiary enamel loops.”
6. Tail length, 120 mm. or more.

c. Greatest length of skull more than 24mm................-.. hirsutus (p. 66).
cc. Greatest length of skull less than 24 mm.
d. Dorsal.area blackish... 0 -.'..-..... 2. Ae toltecus (p. 51).
dd, Dorsaliarea not blackish... ..:..-.. 2. 2322222 eee otus (p. 55).

bb. Tail length less than 120 mm.
c. Tail length more than 65 mm.
d. Ears blackish brown, without ochraceous hairs.
e. Underparts whitish.
j. Tail length 100 mm. or more.

gi Hind foot;more than’ 20 mm: .2...|. 12. 23 see eee toluce (p. 68).

gg. Hind foot less than 20 mm. 2.2305 oe ee allent (p. 59).
ff. Tail length less than 100 mm.

g. Upperparts mainly ochraceous........ ...-.---chiapensis (p. 53).

gg. Upperparts mainly blackish or proce
h. Paler; sides light ochraceous-buff.

1. Larger; hind foot more than 18 mm-.-/_..-._.7.2 cinereus (p. 35).
u. Smaller; hind foot less than 18 mm.
7. Sides of face pure buff (Arizona) .......---- arizonensis (p. 38).

4j. Sides of face ochraceous-buff or grayish.
k. Tail length more than 80mm. (Lower California)
peninsulz (p. 35).
kk. Tail length less than 80 mm.
l. Weet eray (Calitornia) ..2...-eeeeeeee longicaudus (p. 33).
ll. Feet white :.:.....:02.22-22 4.3 nigrescens (p. 32).
hh. Darker; sides dark ochraceous-buff or pinkish cinnamon.
2. Skull length less than 21.5 mm.
j. Smaller; hind foot less than 17 mm............ amoles (p. 40).
4). Larger; hind foot more than 17 mm _.7 252/228 halicetes (p. 42).
w. Skull length more than 21.5 mm,
4. Tail length more than 80 mm.

k. Breadth of braincase more than 11 mm...... toluce (p. 68).
kk. Breadth of braincase less than 11 mm.
l. Skull larger, with wider zygomata.-.-...- dorsalis (p. 61).

il. Skull smaller, with narrower zygomata. .saturatus (p. 36).
4j. Tail length less than 80 mm.
k, Ear from notch more than 13 mm.....-...-. -alticolus (p. 37).
kk, Ear from notch less than 13 mm........-.-. modestus (p. 63).
ee. Underparts ochraceous-buff or pinkish cinnamon.
f. Tail unicolor or nearly so.

gi Daily length more than 60 mm. ....- 2405.5 465 eee rufescens (p. 56).

gg. Tai lepeth lessithan,80)mm.,.,......2:)-1.5seeer eee raviventris (p. 41).

1 Color comparisons made with Ridgway’s “‘Color standards and color nomenclature” (1912),
2 Present in levipes, hirsutus, chrysopsis, tolucz, orizabz, aud perotensis.

1914.] KEY TO SPECIES AND SUBSPECIES. 17

ff. Tail distinctly bicolor.
g. Pelage long; upper molars always with subsidiary enamel loops
(except in colimx); habitat above 9,000 feet altitude.
hk. bramease flattened: moe tric sda cl tbed ee yt |... perotensis (p. 69).
hh. Braincase globular.
1. Larger; tail length more than 95 mm.

4. Breadth of braincase more than 11 mm...... chrysopsis (p. 66).
4j. Breadth of braincase less than 11 mm.......-.- orizabx (p. 69).
u#. Smaller; tail length less than 95 mm...........-. colime (p. 59)

gg. Pelage short; upper molars usually without subsidiary enamel loops;
habitat below 9,000 feet altitude.
he Skull length less thanc22,;mmy5.dicool eR ki. - zacatece (p. 39).
hh. Skull length more than 22 mm.
i. Larger; hind foot 20-21 mm.
4: Braincase flattened. (2c es SA ees anol lt luteolus (p. 57).
qj Brainease rounded! 9345) so. (elsiee y halons ctv nerterus (p. 60).
uw. Smaller; hind foot 18-20 mm.
j. Nasals longer (8-9.7 mm.); black dorsal area well defined.
k. Skull larger, with wider zygomata.......... dorsalis (p. 61).
kk. Skull smaller, with narrower zygomata. . . .saturatus (p. 36).
4j. Nasals shorter (7.3-8.8 mm.); black dorsal area not well defined.
k. ‘Tail longer (88-101 mm.); (Mexico).-...... difficulis (p. 50).
kk. Tail shorter (82-92 mm.); (Costa Rica) ...... australvs (p. 62).
dd. Ears light brown or drab, with ochraceous hairs.
e. Underparts buffy or ochraceous.
f. Skull length more than 22.5 mm.

g- Length of nasals more than 8.5 mm........-....:..-- toltecus (p. 51).

gg- Length of nasals less than 8.5 mm................-.- levipes (p. 64).
ff. Skull length less than 22.5 mm.

ae Ma length OS mm. or moresai. wie Lodidy oaks mustelinus (p. 54).

gg. Tail length less than 105 mm.
h. Upperparts strongly mixed with blackish.

1. Underparts strongly washed with buff ............ difficilis (p. 50).

w. Underparts faintly washed with buff.......... aurantius (p. 48).
hh. Upperparts mainly ochraceous-buff.

Molarcer and paler... erp) ) rarer ert to baae tenuis (p. 45).

Hoesomaier and, brighter 204.2. ny €o wed) aeslidits nelsoni (p. 53).

ee. Underparts white.
f. Tail length more than 104 mm.
g. Larger; skull length 23 mm. or more..--.-......-..-. toltecus (p. 51).

gg. Smaller; skull length less than 23 mm.............. helvolus (p. 52).
ff. Tail length less than 104 mm.
Gaskill leneth less than.20 mm . 2 2. fide oe & amenus (p. 55).

gg- Skull length more than 20 mm.
h. Tail length more than 80 mm.
1. Upperparts tawny-ochraceous, or cinnamon mixed with black.
7. Braincase flatter; nasals shorter.............. chiapensis (p. 53).
4). Braincase higher; nasals longer .............. aurantius (p. 48).
uw. Upperparts ochraceous-buff or mixed with blackish.
4. Upperparts mainly ochraceous-buff.
k. Sides deep ochraceous-buff............-.....-.tenwis (p. 45).
kk. Sides pale ochraceous-buff,
28657°— 14-2

18 NORTH AMERICAN FAUNA. [No. 36.

l. Paler; sides of face grayish. ............- fulvescens (p. 43).
li. Darker; sides of face ochraceous-buff. .intermedius (p. 47).
jj. Upperparts mainly blackish or brownish.
k. Skull heavy; interpterygoid fossa broad.
i. Paler (less ochraceous) 201i 2. intermedius (p. 47).
ll. Darker (more ochraceous).............-.- aurantius (p. 48).
kk. Skull light; interpterygoid fossa narrow... .catalinz (p. 40).
hh. Tail length less than 80 mm.
1. Upperparts mainly deep ochraceous-buff. . ..longicaudus (p. 33).
vw. Upperparts mainly light ochraceous-buff or brownish.

j. Skull length more than 21.5 mm............... aztecus (p. 30).
4). Skull length less than 21.5 mm.
k. ‘Tail Jonger;. ear 12-13 ‘mimi, 322 megalotis (p. 26).
kk. ‘Tail shorter; ear 10-11 mim: O)7 62 ee dychet (p. 30).

cc. Tail length less than 65 mm.
d. Upperparts mainly grayish or light buff, mixed with black.
e. Black dorsal stripe distinct; ears usually with blackish patches.

J. Paler; skull heawiersis2 0. 6600.22) ea albescens (p. 22).
ff. Darker;. skull lighters. 370200 ole. Jase Sa See oe griseus (p. 23).
ee. No distinct dorsal stripe; ears without blackish patches.
jf. Darkeriand more ochraceous 2.225... 72 ee ee dychet (p. 30).
ff. Paler and less ochraceous.
g: Larger; skull ‘broader. 03.02.52 TSG eee ee anegaionis (uaa).
gg. Smaller; skullnarroweri).f0 22 .o Sak Ws eee montanus (p. 24).

dd. Upperparts mainly blackish or dark brown.
e. Upperparts Prout’s brown.

yo Har from, notch. 8-9 mmtiid.. S05 Le impiger (p. 20).
if. War from notch 9-10 mini. [22 30.(.0. 220) 2 ee humulis (p. 19).
ee. Upperparts sooty or with blackish stripe. .-.........---- merriamt (p. 21).

aa. Outer wall of anteorbital foramen usually narrower than width of interpterygoid
fossa;! upper molars always with subsidiary enamel loops.
6. Skull length more than 25 mm.
c. Upperparts tawny or ochraceous-tawny.

ad. Upperparts tawny. 210 $009 Jada oo l2 i eee tenuirostris (p. 78).
dd. Upperparts ochraceous-tawmy 20100. 2 Uae ei eee aureus (p. 78).
cc. Upperparts mummy-brown (Panama) .. - 22 Bee eee creper (p. 79).

bd. Skull length less than 25 mm.
c. Rostrum long and slender; braincase inflated.
d. Darker; underparts (in adult) pinkish cinnamon........-- microdon (p. 80).
dd. Paler; underparts white.........-- 2.2 -se 2 salbilabris (p. 80).
cc. Rostrum short and broad; braincase net inflated
d. Upperparts tawny or Siemon pea

e. Colors darker; skull smaller. 2:22. 60-0220. 5. 20 2 eee mexicanus (p. 70).
ee. Colors paler; skull larger.
j.. Upperparts bright tawny dee. 100 100.) oie 0a cherrit (p. 73).
Jf. Upperparts dull tawny or brownish.
g. Smaller; underparts white.) 2- 2/5 66 3 millert (p. 74).
gg. Larger; underparte bufty.\..2..020ei4 4) 24: séderstr6mi (p. 75).
dd. Upperparts pinkish cinnamon.
é. Skull length less than-23\mmmi. 23.9.2 14 sce ee gracilis (p. 76).
eb) Skull length more than 23-mamw 21/1. Gyo!) og ae eee goldmant (p. 72).

1In gracilis of about equal width or slightly broader.

1914.] SUBGENUS REITHRODONTOMYS. 19

Subgenus REITHRODONTOMYS Giglioli.

Subgeneric characters.—Enamel pattern of upper molars simple,
the first and second each with two outer reéntrant angles, usually
without accessory tubercles.

REITHRODONTOMYS HUMULIS GROUP.
REITHRODONTOMYS HUMULIS HUMULIS (Bachman).

EASTERN Harvest Mouse.
(Pl. I, fig. 1; Pl. IV, fig. 1.)
Mus humulis Aud. & Bach., Proc. Acad. Nat. Sci. Phila., I, 1841, pp. 97-98.
Mus lecontii Aud. & Bach., Journ. Acad. Nat. Sci. Phila., VIII, 1842, p. 307 (Georgia,
taken by Major John Le Conte).
Mus humilis Aud. & Bach., Quad. N. Am., IT, 1851, pp. 103-106, Plate LXV.
Reithrodon lecontet Le Conte, Proc. Acad. Nat. Sci. Phila., VI, 1853, p. 413.
Reithrodon humilis Baird, Mamm. N. Am., 1857, p. 448.
Ochetodon humilis Coues, Proc. Acad. Nat. Sci. Phila., 1874, p. 185 [part].
Reithrodontomys humilis Rhoads, Proc. Acad. Nat. Sci. Phila., 1894, p. 161.
Reithrodontomys lecontii Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 116.
Reithrodontomys humilis dickinsont Rhoads, Amer. Nat., X XIX, 1895, p. 590 (Willow
Oak, Florida).
Reithrodontomys humilis Rhoads and Young, Brce! Acad. Nat. Sci. Phila., 1897, p. 309.
Reithrodontomys humulis Osgood, Proc. Biol. Soc. Wash., XX, 1907, pp. "49-50 (name
formally reinstated).

Type locality Charleston, 8S. C.

Disiribution.—Southeastern United States, east of the Alleghenies,
from southern Virginia to central Florida.

Characters—Size small; color dark brown; skull narrow, with
highly arched cranium and heavy rostrum.

Color.—General tone of upperparts about Prout’s brown, being a
mixture of blackish brown and pinkish cinnamon, usually darkest
along median line; underparts ashy, usually with a tinge of light
pinkish cinnamon; tail bicolor, fuscous or hair-brown above, grayish
white below; ears fuscous or fuscous-black; feet grayish white.
Immature specimens are more fuscous above with slight admixture
of brown.

Skull—Braincase narrow, highly arched; rostrum short and broad;
nasals broad, ending nearly on a line with end of premaxille; zygo-
mata parallel or slightly contracted anteriorly; palatal foramina
broadest in the middle, ending posteriorly about on a line with plane
of first molars; bulle ene small and elongated.

Measurements—Average of 5 adults from Georgia and South Caro-
lina: Total length, 120 (114-124); tail vertebra, 57 (53-60); hind
foot, 16 (15-17); ear, 9.5 (9-10). Average of 8 adults from Dismal
Swamp, Va.: 122; 56; 16; (ear) 9.6. Skull: (See table, p. 81).

Remarks.—This little species—the first member of the genus to be
described—has a rather extensive range in the South Atlantic States,
from southern Virginia to Florida. In Georgia, Alabama, and Florida

1 Small tubercles irregularly present in some species, particularly R. rufescens and R. dorsalis.

90 NORTH AMERICAN FAUNA. ~ ENo. 36.

the typical race shades insensibly into the subspecies merriami of the
Mississippi Valley. Specimens from North Carolina and Virginia, as
might be expected, exhibit the specific characters in their most pro-
nounced form, the color of the upperparts being uniformly brownish.
Specimens from Florida are intermediate but nearer on the whole to
humulis. In a series of three from Kissimmee one is clearly humulis,
while the others might without impropriety be referred to merriami.
The skulls, however, are nearer to humulis. The type of “‘dickonsoni,”’
from Willow Oak, is a very dark specimen, but is in the uniform sooty
pelage indicative of immaturity. Itis closely similar to numerous in-
dividuals of merriami examined from Alabama and Louisiana. A
single specimen from Tarpon Springs, however, can be exactly matched
by specimens of humulis from the coast of Georgia. In the absence

g
7 ot
ef 2S . Sie v
x oar 3S
WS e Notas ¥
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f \\ SON ex
7 -_--—<-
~—- 7
-_—. ww 44a S
s £ Faort™ c =

,
-

R.A. HUMULIS
R.H. PMPIGER
SSS a b.merniam

M1 -. a. ALBESCEYS
PA k.aariseus
Te

Fig. 1.—Distribution of Reithrodontomys humulis, R. albescens, and subspecies.

of suitable material the west Florida form is provisionally referred to
humulis, but a larger series may show it to be nearer merriamt.
Specimens ecamined.—Total number, 96, from the following locali-
ties:
Virginia: Dismal Swamp, 17.
North Carolina: Chapanoke, 1; Currituck, 4; Moran, 1; Raleigh, 53.
South Carolina: Georgetown, 1.
Georgia: Riceboro (Le Conte Plantation), 8.
Florida: Enterprise, 1;1 Gainesville, 4;! Kissimmee, 3; Sawgrass Island, Polk
County, 1; Tarpon Springs, 1;2 Willow Oak, Pasco County, 1.?
REITHRODONTOMYS HUMULIS IMPIGER Bangs.
SMALL-EARED Harvest Mouse.
Reithrodontomys lecontit impiger Bangs, Proc. Biol. Soc. Wash., XII, 1898, p. 167.
Type locality White Sulphur Springs, W. Va.
Distribution —Northern Virginia and mountains of West Virginia.

1 Collection Field Mus. Nat. Hist. 2 Collection Acad. Nat. Sci. Phila.

1914.) REITHRODONTOMYS HUMULIS GROUP. 2 |

Characters—Closely similar in size and color to humulis; ears
decidedly smaller.

Color—Not appreciably different from that of humulis.

Skull—Closely similar to that of humulis.

Measuremenis.—Average of 9 adults from type locality: Total
length, 116 (114-122); tail vertebra, 53 (50-57); hind foot, 15.7
(15-16.5); ear from notch, 8.7 (8-9). Skull: (See table, p. 81).

Remarks.—This subspecies is a rather poorly marked race of humulis
occupying the northern end of the range of the species. In the series
from the type locality—White Sulphur Springs, W. Va.—I have been
unable to see any color differences to distinguish it from humulis from
the coast of the Carolinas. Three specimens from the vicinity of
Washington, D. C., however, are considerably paler than the series of
topotypes, with which they agree in having very small ears. These
may represent an undescribed race, but it seems best not to name it
until more material can be secured.

Specumens examined.—Total number, 12, from the following locali-
ties:

West Virginia: White Sulphur Springs, 9.
Virginia: Alexandria, 1; Falls Church, 1; Fort Myer, 1.

REITHRODONTOMYS HUMULIS MERRIAMI Allen.
Merriam Harvest Mouse.
CELE he 2) Pls IV, fig. 25 Pl. Viils fic. 8)
Reithrodontomys merriami Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 119.

Type locality — Austin Bayou, near Alvin, Tex.

Disiribution.—Coast region of east Texas and southern Louisiana
north to northeastern Kentucky and West Virginia; east to Ala-
bama; limits of range imperfectly known.

Characters —Similar to humulis, but upperparts blacker and grayer
(less brownish); ears smaller and blacker.

Color—Adulis: Upperparts mixed blackish and light pinkish cin-
namon, the black prevailing and forming usually a well-defined stripe
in middle of back. Specimens in full, unworn pelage show an indis-
tinct lateral stripe of light pinkish cinnamon. Ears blackish brown;
tail fuscous above, grayish white below; underparts grayish, some-
times with a distinct wash of light pinkish cinnamon. Some speci-
mens in worn pelage have a broad median dorsal band of black and
searcely any cinnamon on the sides. Young: Upperparts fuscous-
black, practically without cinnamon hairs.

Skull——Practically identical in size and proportions with that of
humulis, but rostrum slightly heavier.

Measurements.—Average of 7 adults from type locality: Total
length, 113 (107-128); tail vertebre, 54 (51-60); hind foot, 16
(15.5-17). Skull (see table, p. 81).

99, NORTH AMERICAN FAUNA. [No. 36.

Remarks.—This subspecies is a slightly differentiated race of
humulis, with which it is connected by a complete series of inter-
grades. Indeed, occasional specimens of humulis from localities
within the range of the typical race are difficult to distinguish from
merrramt. The differences are perhaps most evident on comparing
individuals of the two forms in worn pelage, merriami being in that
condition decidedly sooty, while humulis is a warm brown. Three
specimens from Lexington, Ky., seem referable to this subspecies,
but one from Ceredo, in the extreme western corner of West Virginia,
is intermediate in color between merriami and humulis.

Specimens examined.—Total number, 37, from the following locali-
ties:

Texas: Austin Bayou near Alvin, 8; Richmond, 2.
Louisiana: Hackley, 1; Lafayette, 3; Mermenton, 3.7
Alabama: Barachias, 5; Carlton, i; Dean, 1; Jackson, 2; York, 7.

Kentucky: Lexington, 3.
West Virginia: Ceredo, 1.

REITHRODONTOMYS ALBESCENS ALBESCENS Cary.
Patup Harvest Mouse.
(Pl. I, fig. 4; Pl. IV, fig. 4.)

Reithrodontomys albescens Cary, Proc. Biol. Soc. Wash., XVI, 1903, p. 53.
Reithrodontomys montanus albescens Cary, N. Am. Fauna No. 33, 1911, p. 110.

Type locality Eighteen miles northwest of Kennedy, Nebr.

Distribution.—Sand-hill region of Nebraska and western South
Dakota; west to Loveland, Colo. |

Characters.—Similar to R. montanus, but paler, with a more dis-
tinct dorsal stripe; earssmaller; skull shorter and broader.

Color— Unworn winter pelage (November): Upperparts mixed
blackish and light buff, darkest on the median line and shading to
pure buff on the sides; ears buffy, usually with two rather large brown-
ish or blackish patches; tail sharply bicolor, dark hair-brown above,
white below; feet soiled whitish; underparts pure white. Worn
summer pelage (October): Upperparts mixed ochraceous-buff and
blackish, the buff prevailing, shading to light ochraceous-buff on sides.
A gray phase occurs in which the upperparts are mixed blackish and
pale mouse-gray, with very slight admixture of buff.

Skull—Shorter and relatively broader than that of montanus;
rostrum short and heavy; zygomata heavy and widely expanded.

Measurements—Average of 6 adults from Nebraska: Total length,
125 (121-129); tail vertebrae, 53.4 (50-56); hind foot, 16.7 (16.5-17).
Skull (see table, p. 81).

Remarks.—This handsome mouse is the palest member of the genus.
According to Cary it occurs only in sand-hills or on sandy land. At

1 Collection Field Mus. Nat. Hist.

1914.] REITHRODONTOMYS HUMULIS GROUP. 23

Kennedy and Neligh, Nebr., it has been taken in the same weed
patches with R. megalotis dychei. It may be distinguished from the
latter by its smaller size, paler and less ochraceous coloration, and
small ears with distinct black spots.

Although closely resembling montanus in color, it differs consider-
ably from it in cranial characters, and no evidence of intergradation
between the two species has been obtained. In southern Nebraska
it grades into the subspecies griseus.

Specimens examined.—Total number, 25, from the following locali-
ties:

Nebraska: Kennedy, 1; 18 miles northwest of Kennedy, 4; Neligh, 8; Niobrara,
1; Niobrara River, 10 miles south of Cody, 2; Verdigris, 4.

South Dakota: Belle Fourche River (15 miles from mouth), 1.
Colorado: Loveland, 4.1

REITHRODONTOMYS ALBESCENS GRISEUS Bailey.
Lirtte Gray Harvest Mouvuse.
(Rie fisra "Er TV, fig:’3:)
Reithrodontomys dychei Allen, Bull. Am. Mus. Nat. Hist., VIII, 1896, p. 67 (not of
VII, 1895, p. 120).
Reithrodontomys griseus Bailey, N. Am. Fauna No. 25, 1905, p. 106.

Type locality—San Antonio, Tex.

Disiribution.—Southern Nebraska, Kansas, Oklahoma, central and
western Texas, and eastern New Mexico. Limits of range imper-
fectly known.

Characters —Similar to albescens but colors darker and skull
narrower; much paler and grayer than merriami.

Color.—Upper parts mixed black and light ochraceous-buff, the
black predominating and usually forming an indistinct stripe along
the median line; lateral line of buff very faintly indicated; buff on
sides of head darker than in montanus; ears same color as back,
usually with a large blackish patch on exterior surface; tail sharply
bicolor, hair-brown above, grayish white below.

Skull—Similar to that of albescens, but zygomata weaker and less
widely expanded and rostrum narrower. Compared with merriami:
Braincase less elongated; rostrum slenderer; zygomata more widely
expanded posteriorly; interpterygoid fossa averaging narrower;
bullae less elongated. Compared with montanus: Braincase shorter
and relatively broader, with shorter rostrum; zygomata more
contracted anteriorly.

Measurements —Average of 9 specimens (mostly immature) from
type locality: Total length, 115 (107-120); tail vertebrae, 54 (48-61);
hind foot, 14.6 (14-16).2. Average of 2 adults from Clyde and San

1 Collection of G. S. Miller, jr.

2 The total length is probably less than in adult specimens and the foot measurements are apparently
smaller in some cases than they should be.

QA NORTH AMERICAN FAUNA, [No. 36.

Angelo, Tex.: 142; 60; 15. Average of 8 (subadult) from Alva, Okla.:
117; 51.5; 16. Skull: (See table, p. 81).
| Remarks.—Although clearly belonging in the same group with
fh. humulis merriami, this form appears not to intergrade with it. In
both skin and skull characters it agrees closely with albescens and
differs widely from merriami, its nearest relative geographically.
Specimens from Kansas and Oklahoma and one from Santa Rosa,
N. Mex., can not be distinguished from typical examples.
Specimens examvned.—Total number, 82, from the following locali-

ties:

Nebraska: London, 2

Kansas: Onaga, 18; Pendennis, 5; Trego County, 10.

Oklahoma: Alva, 12.!

Texas: Clyde, 1; Gainesville, 1; Mason, 9; San Angelo, 1; San Antonio, 22.

New Mexico: Santa Rosa, 1.

REITHRODONTOMYS MEGALOTIS GROUP.

REITHRODONTOMYS MONTANUS (Baird).

San Luis VALLEY Harvest Mouse.
(Pl.d, fie. 5; Pl. LY sfiewb))

Reithrodon montanus Baird, Proc. Acad. Nat. Sci. Phila., VII, 1855, p. 335; Mamm.
N. Am., 1857, p. 449.

Ochetodon montanus Coues, Proc. Acad. Nat. Sci. Phila., 1874, p. 186.

Reithrodontomys montanus Allen, Bull. Am. Mus. Nat. Hist., V, 1893, p. 80; VII,
1895, p..123; Cary, N. Am. ane No. 33; 1911) joemitise

Type locality —‘‘Rocky Mountains, latitude 38°” [upper end San -
Luis Valley, Colo., near San Luis Lakeet

Dist ition san Luis Valley, Colo.

Characters.—Size medium (smaller than R. m. megalotis, larger
than Rf. a. griseus); ears small; color slightly paler and less ochra-
ceous than in megalotis; skull similar to that of megalotis but smaller
and narrower. Compared with R. a. albescens and R. a. griseus:
No distinct blackish dorsal stripe; skull relatively longer and nar-
rower; tail slightly longer.

Color—Fresh winter pelage (October and November): Light buff,
clearest on sides and face, much mixed with blackish on dorsal sur-
face; black hairs most pronounced on hinder back; no distinct
median line of black; ears dark hair-brown, often clothed on inner
surface with ochraceous-buff hairs (a little darker than body hairs),
rarely with a distinct blackish area on lower outer margin; tail dis-
tinctly bicolor, dark hair-brown above, white beneath; feet and
underparts white. Compared with megalotis, the general tone is paler
and less intensely ochraceous; the sides light buff instead of ochra-
ceous-buff.

1 Collection Field Mus. Nat. Hist.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. 25

Skull.—Similar in shape to that of BR. m. megalotis, but smaller and
brainease relatively narrower; zygomata (in adults) with sides par-
allel. Compared with R. a. griseus: Longer and relatively narrower,
with longer rostrum.

Measurements—Average of 10 specimens (including adults and
subadults) from Medano Ranch, near Mosca, Colo.: Total length, 126
(118-139) ; tail vertebrae, 58 (51-64); hind foot, 17 (16-17); ear from
notch, 11.2 (11-12). Skull: (See table, p. 81).

Remarks.—This species, the second member of the genus to be
recognized, was described by Baird in 1855 irom a single specimen,
and until within the last few years has remained practically un-
known. Efforts were made by the Biological Survey field party
in 1904 to secure specimens from the San Luis Valley and a single
immature individual was caught at Del Norte by Vernon Bailey.
In the autumn of 1907, Merritt Cary trapped for about 10 days at
Medano Ranch, 15 miles northeast of Mosca, and succeeded in se-
curing a series of 20 specimens, including several adults, which for
the first time made possible accurate comparison and characteriza-
tion of the species. Most of the specimens taken by Cary were caught
in a grassy weed-patch on a broad sand-ridge extending through the
meadows and perhaps 6 feet above their level. None was taken
in wet situations.

Dr. J. A. Allen has determined the approximate type locality of
this species from a careful study of the records of Capt. Beckwith’s
expedition, on which the type was secured.1 The type specimen
(No. 13 of the expedition) was collected in August, 1853, in the
upper end of the San Luis Valley, at some point between Fort Massa-
chusetts and Sahwatch Creek. The route of the expedition was
northward along the east side of the valley to a point a little north
of 38° latitude, and thence westward across the valley not far
from the present town of Saguache.? The second camp of the party,
on the night of August 24, was on a small stream “‘nearly opposite
to Roubideau’s or Mosca Pass,’”’ about 21 miles from Fort Massa-
chusetts. The specimens collected by Cary in 1907 were taken
along the very creek on which the Beckwith party camped, now called
Medano Creek. The type may have been taken on this creek or at
one of the other camping places a few miles farther north.

Even with a series of 20 topotypes, the relationships of this species
are not entirely clear. This uncertainty is due in part to the fact
that the topotype series includes only two adults, one of which has
the skull broken, and in part to the failure to find in the series a
single skull that agrees with that of the type. The latter, an adult
with moderately worn teeth, is decidedly smaller and has a much

1See Bull. Am. Mus. Nat. Hist., VII, 1895, pp. 124-125.
2 See Expl. & Surv. Pac. R. R., Il, pp. 41-45, and map accompanying Vol. XI.

26 NORTH AMERICAN FAUNA. [No. 36.

shorter rostrum than the two adults taken by Cary at Medano Ranch,
thus closely agreeing with specimens of Ff. a. griseus from Texas.
The rostrum of this specimen is deflected markedly to the right,
indicating an injury during life, which may account for its small
size and peculiar characters. The original description of the color
accords perfectly with the series now in hand, so that there seems
no alternative but to consider the type skull aberrant, and to con-
tinue to use the name for the form represented by the modern series.
_ The species, although combining in a remarkable degree the char-
acters of the megalotis and albescens groups, seems not to be directly
connected with either of them. It is perhaps best placed in the
megalotis group, but seems not to intergrade with any member of it.!

Though but little smaller than &. m. megalotis, it is markedly
smaller than aztecus—the form ranging through northern New Mexico,
nearest to the home of montanus. It may be distinguished from both
megalotis and aztecus by smaller size, particularly of the ears and tail,
and by paler and less ochraceous coloration. Externally it much
resembles R. a. albescens but is somewhat darker and has larger ears
and feet. The nearest approach geographically of &. albescens griseus
to the range of montanus is at Santa Rosa, N. Mex., and the single
specimen taken there shows no departure from typical griseus.

Specimens examivned.—Total number, 21, from the following locali-
ties:

Colorado: Del Norte, 1; Medano Ranch (15 miles northeast of Mosca), 20.

REITHRODONTOMYS MEGALOTIS MEGALOTIS (Baird).
Desert Harvest Movse.
(Pi.1, fie.6; Bl. DV, fics 6: Pl. VIL tes. tea

Reithrodon megalotis Baird, Mamm. N. Am., 1857, p. 451.

Reithrodontomys megalotis Allen, Bull. Am. Mus, Nat. Hist., V, 1893, p. 79; VII,
1895, p. 125.

Reithrodontomys megalotis desertt Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 127
(Oasis Valley, Nev.).

Reithrodontomys megalotis sestinensis Allen, Bull. Am. Mus. Nat. Hist., XIX, 1903,
p. 602 (Rio Sestin, northwestern Durango).

Type locality —Between Janos, Chihuahua, and San Luis Springs,
N. Mex.

Distribution—From northern Nevada and southern Idaho south
to Zacatecas, Mexico; occupying the greater part of Nevada, Arizona,
and Utah (except eastern part) ; southern New Mexico; western Texas
(west of Pecos River); desert regions of southern and northeastern
California, northeastern Lower California, and northern Sonora;
and northern portion of Mexican table-land.

1Cary’s assignment of this species to the albescens group (N. Am. Fauna No. 33,1911, p. 108) was on the
authority of the present writer, who, after more detailed study, has reached the conclusions set forthabove.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. 27

Characters.—Size medium (larger than R. montanus and smaller
than R. f. fulvescens); general tone brownish buff above and white
below; ears usually without dark markings. |

20 15° (oy
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2| Z pe. éc MEGALOTIS
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ROO PR. 66 PENINSULAE
SATURATUS

ZACATECAE

10° 105°
Fic. 2.—Distribution of Reithrodontomys montanus, R. megalotis, and subspecies.

Color.— Unworn winter pelage: Upperparts mixed blackish brown
and light ochraceous-buff, darkest in middle of back, shading to
nearly pure buff on sides; feet and underparts white; tail hair-brown
above, whitish below; ears drab, usually with a tuft of ochraceous-
buff hairs at base. Unworn summer pelage: Similar to the fresh

28 NORTH AMERICAN FAUNA. [No. 36. —

winter pelage, but colors less strongly contrasted and black variations
of upperparts much reduced. Worn spring and summer pelage: Gen-
eral tone browner and colors less contrasted than in the winter
pelage, the buff on sides less pronounced and often lacking.

Skull.—Larger than that of montanus, with broader, flatter brain-
case. Compared with fulvescens and tenuis: Slightly smaller, with

longer palatal foramina and narrower interpterygoid fossa.
Measurements —Average of 6 adults from type region (Casas
Grandes, Chihuahua; Mexican boundary, 50 to 100 miles west of
HK] Paso; and Organ Mountains, N. Mex.): Total length, 140 (128-
145); tail vertebre, 71 (65-77); hind foot, 17.6 (17-18.5); ear from
notch, 12.5 (12-13.2). Average of 12 adults from Lone Pine, Cal.:
139; 70; 17.5; of 10 adults from Oasis Valley, Nev. (measured to
end of tail hairs): 1389; 72; 18.4. Skull: (See table, p. 81).

Remarks.—This subspecies has the widest range of any of the forms
of megalotis and is subject to considerable individual variation, both
in color and in size of skull. It intergrades with aztecus in central
New Mexico, with longicaudus in California, with nigrescens in south-
ern Idaho, and with cinereus in central Mexico. The present writer
is in accord with Dr. J. A. Allen, who revised the group in 1895, in
his failure to find any constant color differences between the series
from the deserts of Nevada and California and that from the type
region (southern New Mexico and northern Chihuahua), individuals
in comparable pelage from the two regions being practically indis-
tinguishable. Specimens from the Death Valley region of southern
California and Nevada have slightly larger skulls than those of the
typical form from Chihuahua and southern New Mexico but smaller
than those of aztecus. ‘These have been separated as a subspecies,
“desertt”’ by Dr. Allen on the basis of an alleged difference in the rel-
ative length of body and tail. A careful comparison of measurements
of adult specimens from the two regions, however, fails to show any
appreciable differences in proportion. The apparent differences
shown in Dr. Allen’s published measurements probably are due in
part to differences in methods of measurement and in part to the inclu-
sion of a greater number of immature examples in the series from
New Mexico and Utah. The cranial differences are considered too
slight and inconstant to warrant recognition of the form by name.

A large series from the Colorado River, Sonora, at Monument No.
204, Mexican boundary, are practically typical megalotis. Two in-
dividuals from Volcano Lake, head of Hardy River, Lower California,
are very small, but no smaller than an individual in the series from
Casas Grandes, Chihuahua, quite close to the type locality.

A large series from various places in northwestern Nevada and a
small series from Salt Lake Valley, Utah, are fairly typical megalotis,
having slightly smaller skulls than the Death Valley series and thus
showing no approach to aztecus.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. 29

Intergradation with longicaudus is perfectly shown by a large
series from the Mojave Desert in southern California. In color these
specimens are intermediate between the two races, but rather nearer
to megalotis, while in skull characters they most resemble longicaudus.
Some individuals in the series much resemble mgrescens, but are
generally more suffused with buff. A series from the south fork of
Kern River, Cal., are nearly typical megalotis but slightly browner.

In New Mexico the typical form occurs only in the extreme south-
ern part, near the Mexican boundary, a gradual increase in size of
skull being observed as we proceed northward toward the range of
aztecus. Southward over the Mexican table-land there is no percep-
tible change in color as far as Zacatecas. Specimens from north-
western Durango, described by Allen under the name “‘sestinensis,”
do not differ appreciably from the typical form.

Specumens examined.—Total number, 415, from the following
localities:

Idaho: American Falls, 2; Swan Lake, 1.

California: Amedee, 2; Argus Mountains, 1; Barstow, 12; Bishop Creek, 1;
Cartago, Owens Lake, 10; Colorado River (opposite Parker, Ariz.), 4; Emi-
grant Spring, 1; Fort Yuma, 1; Furnace Creek, Death Valley, 5; Grapevine
Spring, 4; Keeler, 4; Lone Pine, 30; Long Valley (Lassen County), 4; Mojave,
3;1 Olanche, Owens Lake, 22; Owens Valley, 8; Onyx, south fork of Kern
River, 19;1 Panamint Mountains, 7; Panamint Valley, 2; Pilot Knob, 8;
Resting Spring, 3; Saratoga Spring, 3; Shepherd Canon, Argus Mountains,
1; Tehachapi, 20;1 Twelve Mile Spring, Inyo County, 1; Victorville, 1.1) 2

Nevada: Ash Meadows, 30; Carson Sink, 1; Fallon, 5; Gardnerville, 1; Grape-
vine Mountains, 2; Oasis Valley, 16; Pahranagat Valley, 4; Pahrump Valley,

_ 11; Panaca, 1; Pine Forest Mountains, 7; Quinn River Crossing, 20; St.
Thomas, 1; Smoky Creek, 1; Vegas Valley, 5; Verdi, 3; Virgin Valley, 1.3

Arizona: Colorado River at Monument No. 204, Mexican boundary, 27; Fair-
bank, 2; Grand Canyon (Indian Gardens), 2; Lee’s ferry, 1; Parker, 5; St.
Johns, 2; Winslow, 17; Zuni River, 1.

New Mexico: Animas Valley,1; Deming,3; Dry Creek, Socorro County, 3; Fair-
view, 1; Gallo Canyon, 35 miles southeast of Corona, 1; Gila, Grant County, 1;
Glenwood, 1; Jicarilla Mountains, 1; Kingston, 2; Las Cruces, 3; Las Palo-
mas, 5; Mesa Jumanes, 2; Monument No. 15, Mexican boundary, 1; Monu-
ment No. 40, Mexican boundary, 3; Organ Mountains, 3; Pleasanton, 1;
Redrock, Grant County, 1; Roswell, 1; San Andreas Mountains, 2; San
Mateo Mountains, Socorro County, 2; Silver City, 3; Tularosa, 5.

Texas: Alpine, 1; 25 miles west of Fort Stockton, 1; Franklin Mountains, 1;
Guadalupe Mountains, 1; Pecos City, 1.

Lower California: Gardners Lagoon, 1; Seven Wells, 2.

Sonora: Cienega Well, 30 miles south of Monument No. 204, Mexican boundary, 1.

Chihuahua: Casas Grandes, 4; Chihuahua, 2.

Coahuila: Saltillo, 1; Sierra Guadalupe, 1.

Durango: Rio Sestin, 4.1

Zacatecas: Zacatecas, 2.

1 Approaching longicaudus. 8 Collection Mus. Vert. Zool., Univ. of Cal.
2 Received from Jos. Grinnell. 4Collection Amer. Mus. Nat. Hist.

30 NORTH AMERICAN FAUNA. | [No. 36.

REITHRODONTOMYS MEGALOTIS AZTECUS Allen.
Aztec Harvest Mouvss.
Reithrodontomys aztecus Allen, Bull. Am. Mus. Nat. Hist., V, 1893, p. 79.

Type locality —La Plata, N. Mex.

Distribution.—Northern New Mexico, northeastern Arizona, south-
eastern Utah, and western Colorado, north to Grand Junction and
Rifle.

Characters.—Similar to megalotis but with larger ears and skull.

Color.—Not appreciably different from that of megalotis; ears some-
times with irregular dusky blotches.

Skull_—Decidedly larger than that of megalotis.

Measurements.—Average of 7 adults from La Plata and Aztec,
N. Mex., and Noland Ranch, Utah: Total length, 144 (133-155); tail
vertebree, 68 (62-73); hind foot, 18 (17-19); ear from notch, 13.8
(12-15.5). Skull (ee table, p. 81).

Remarks.—This subspecies does not differ from megalotis in color,
but its skull is so much larger that it seems desirable to recognize the
race. The ears average larger, also, in the series from the type region.
Dr. Allen provided the form with a provisional name many years ago,
but later placed it in synonymy.

This race is connected with megalotis by a complete series of inter-
grades from New Mexico and Arizona. The specimens from Datil
and Manzano Mountains are too young to be satisfactorily identified,
but are provisionally referred to aztecus.

Specimens examined.—Total number, 143, from the following locali-
ties:

New Mexico: Aztec, 39; Espanola, 8; Farmington, 4; Fruitland, 6; Gallup, 1;
Guadalupita, 1; La Plata, 32; Las Vegas, 2; Manzano Mountains, 1; Rinco-
nada, 1; Rio Puerco, 4; Wingate, 2.

Arizona: Canyon de Chelly, 1.

Utah: Bluff City, San Juan River, 4; Noland Ranch, San Juan River, 5.

Colorado: Arboles, 1; Ashbaugh’s Ranch, Montezuma County, 2; Cortez, 1;
Grand Junction, 26; Rifle, 2.

REITHRODONTOMYS MEGALOTIS DYCHETI Allen.

PRAIRIE Harvest Mouse.
CPV) fe isk AV, toi)

Reithrodontomys dychet Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 120.

Reithrodontomys dychei nebrascensis Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 122
(Kennedy, Nebr.).

Reithrodontomys griseus Ruthven & Wood, Proc. Iowa Acad. Sci., XIX, 1912,
p. 204 (not of Bailey).!

Type locality Lawrence, Kans.

1 Misidentification of specimen by the present writer.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. | 31

Distribution.— Greater part of Kansas, Nebraska, Iowa, Missouri,
and South Dakota; southern North Dakota; southeastern Montana;
eastern Colorado and eastern Wyoming.

Characters.—Similar to megalotis, but black of upperparts more
extensive and ochraceous shades more intense; ears slightly smaller;
tail shorter.

Color.— Full winter pelage (topotypes): Upperparts mixed black
and light ochraceous-buff; in some specimens the median dorsal area
is noticeably darker, in oehers the color is nearly uniform over the
whole back; sides clear buff, the lateral line sometimes, though not
always, well-marked; ears hair-brown externally, thinly clothed with
buffy hairs on inner surface, and with a tuft of ochraceous-buff hairs
at anterior base; feet and underparts white; tail sharply bicolor,
dark hair-brown above, white beneath.

Skull—About same size as that of megalotis (smaller than that of
aztecus); rostrum shorter and broader.

Measurements.'—Type: Total length, 133; tail vertebra, 52; hind
foot, 15.5. Average of 6 nearly adult specimens from eastern
Nebraska (Neligh and Verdigris): Total length, 185 (130-142); tail
vertebre, 61 (57-65); hind foot, 17.5 (16.5-18); ear from notch, 10. 5
(10.3-11.1). Average of 8 adults from Kennedy, Nebr.: 133; 63;
17.5; 10.8. Skull: (See table, p. 81).

Remarks.—This form clearly belongs in the megalotis group and
apparently intergrades with aztecus, as indicated by specimens from
central Colorado (Loveland, Greeley, and Canon City). These aver-
age slightly paler and grayer than the typical form and their skulls are
intermediate in size, with somewhat longer rostri than in typical
dyche.

The series from Kennedy, Nebr., on which Dr. Allen based the sub-
species ‘‘nebrascensis”’ seem to be indistinguishable from specimens
of typical dychet in comparable pelage. At many localities in
Nebraska and Kansas ff. albescens griseus occurs with the present
species, and although occasional specimens are hard to distinguish by
color alone, yet in size and cranial characters the species are distinct.
Dr. Allen confused the two in his original description of dychet, the
series listed from Onaga, Kans., and London, Nebr., being referable
to griseus. The latter race may be distinguished from dychet by its
smaller size, shorter tail, and smaller skull with short rostrum and
short palatal foramina. In coloration dychei is more intensely ochra-
ceous than griseus and often not conspicuously darkened on the dorsal
area; griseus is distinctly grayer and the ochraceous-buff of the sides
is paler and less extensive.

1 The great variation in the measurements of this species given by Dr. Allen in the original description
indicates that probably many of the specimens were immature and the average therefore too small.

ao NORTH AMERICAN FAUNA. [No. 36,

Specumens examined.—Total number, 153, from the following locali-
ties:

North Dakota: Ellendale, 2; Fort Clark, 4; Hankinson, 4; Ludden, 2; Oakes, 1.

South Dakota: Clay County, 2.

Montana: Billings, 5; Fort Custer, 4.

Wyoming: Arvada, 4; Casper, 1; Meadow, 1; Pole Creek, Laramie County, 1;
Splitrock, 3; Sun, 3.

Colorado: Boulder, 1; Canon City, 5; Denver, 2; Golden, 2; Greeley, 1; Love-
land, 14; Valmont, 1.

Kansas: Cloud County, 2;! Lawrence, 9;? Neosho Falls, 1; Onaga, 18; Pen-
dennis, 5.

Nebraska: Alliance, 2; Beemer, 1; Callaway, 3; Cherry County, 1; 10 miles
south of Cody, 1; Columbus, 3; Ewing, 1; Glen, Sioux County, 1; Haigler,
2;3 Kearney, 2; Kennedy, 12; 18 miles northwest of Kennedy, 5; Neligh, 9;
Norfolk, 1; Two Mile Lake, Cherry County, 1; Valentine, 2; Verdigris, 2.

Iowa: Atlantic, 2; Hillsboro, 2; Palo Alto County, 1.+

Missouri: St. Louis, 1; Thayer, 2.

REITHRODONTOMYS MEGALOTIS NIGRESCENS subsp. nov.
Dusky Harvest Mouse.

Type from Payette, Idaho. No. 201616, U.S. Nat. Mus., Biologi-
cal Survey Collection, ¢ adult, June 9,1913; L.E. Wyman. Original
No. 98.

Distribution.—Eastern Oregon and western Idaho; north to Pres-
cott, Wash., south to Bieber, Cal.

Characters.—Similar to megalotis, but upperparts more blackish and
less buffy; grayer and less ochraceous than longicaudus.

Color.— Winter pelage: Upperparts mixed blackish and pale ochra-
ceous-buff, the black predominating on dorsal area; lateral line of
buff only faintly indicated; ears hair-brown, clothed with ochraceous
hairs; tail blackish brown above, white below; feet and underparts
white. Summer pelage (July): General tone of upperparts more
brownish and less blackish than in winter pelage.

Skull.—Closely similar to that of megalotis, but with slightly longer
nasals.

Measuremenis.—Average of 7 adults from Idaho: Total length, 144
(140-153); tail vertebrae, 68 (63-75); hind foot, 17 (146-18). Skull:
(See table, p. 81).

Remarks.—This race occupies the extreme northwestern part of the
range of the species, chiefly in a region of old lava beds where the soil
is richer and vegetation more abundant than in the more arid deserts
to thesouthward. Itis most nearly related to megalotis, but probably
intergrades also with longicaudus.

1 Collection Kansas Univ. Mus. fy 3 Collection Univ. of Nebraska.
2 Collection Am. Mus. Nat. Hist. 4 Collection Univ. of Michigan.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. 33

Specimens examined.—Total number, 36, from the following locali-
ties :
Washington: Prescott, 7.
Oregon: Narrows, Malheur County, 11; Vale, 1.
California: Bieber, 1; Brownell, 4.
Idaho: Nampa, 5; Payette, 5; Weiser, 2.

REITHRODONTOMYS MEGALOTIS LONGICAUDUS (Baird).
CALIFORNIA Harvest Mouse.

Reithrodon longicauda Baird, Mamm. N. Am., 1857, p. 451.

Ochetodon longicauda Coues, Proc. Acad. Nat. Sci. Phila., 1874, p. 186; Mon. N. Am.
Rodentia, 1877, p. 126.

Reithrodontomys pallidus Rhoads, Am. Nat., XX VII, 1893, p. 835 (Santa Ysabel,
Cal.).

ee ini longicauda Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 129.

Reithrodontomys klamathensis Merriam, N. Am. Fauna No. 16, 1899, p. 93 (Shasta
Valley, Cal.).

Reithrodontomys megalotis longicauda Grinnell, Proc. Cal. Acad. Sci., Ser. 4, III,
1913, p. 303.

Type locality Petaluma, Cal.

Distribution.—Greater part of western California, east to the foot-
hills of the Sierra Nevada, San Bernardino, and San Jacinto Ranges;
north to Grants Pass, Oreg., and south into northwestern Lower
California to about latitude 32°.

Characters —Slightly smaller than megalotis; colors darker and more
intense.

Color.— Fresh winter pelage: Upperparts mixed blackish and ochra-
ceous-buff, the black predominating on dorsal area, shading to nearly
pure ochraceous-buff on sides; lateral line of buff often well defined;
ears same color as back, clothed with scattering ochraceous hairs;
feet grayish white; tail sharply bicolor, hair-brown or fuscous above,
grayish white below; underparts grayish white, usually with a tinge
and often with a strong wash of ochraceous-buff. Fresh summer
pelage: Similar to the winter pelage, but decidedly paler and lacking
most of the black on the dorsal area.

Skull.—Similar to that of megalotis, but smaller.

Measurements.—Adult topotype: Total length, 152; tail vertebre,
75; hind foot, 17.5. Average of 13 specimens (mostly young adults)
from vicinity of San Francisco Bay: 139 (130-146); 73 (68-79);
17 (16-18). Skull: (See table, p. 81).

Remarks.—This form has a wide range on the Pacific coast and inter-
grades with megalotis at various points along the edge of the deserts.
Intermediate specimens have been examined from Barstow, Mojave,
Tehachapi, Santa Paula, Kern River, Bieber, etc. The form from
Shasta Valley described under the name “klamathensis”’ is likewise
intermediate between longicaudus and megalotis, specimens from there
agreeing exactly in color with summer specimens of longicaudus from

28657°—14—_3

34 NORTH AMERICAN FAUNA. [No. 36.

Petaluma, while their skulls can be matched very closely in the series
of megalotis. The type of “klamathensis”’ is a very old individual
and has a skull as large as that of &.m.aztecus. Skulls of topotypes,
however, do not differ appreciably from skulls of megalotis from the
Death Valley region. eithrodontomys “ pallidus’”’ of Rhoads is a pure
synonym of longicaudus, neither the type nor topotypes being distin-
guishable from typical specimens of this race. Color variation in
this subspecies is considerable and the winter and summer pelages
are strikingly different.

Specimens examined.—Total number, 555, from the following locali-
ties:

Oregon: Grants Pass, 1.1

California: Adobe Station, 1;? Alton Junction, Humboldt County, 3; Aptos, 15;
Armona, 1; Arroyo Seco, 10 miles south of Paraiso Springs, 4; Arroyo Seco
Canyon, near Pasadena, 1;3 Ballena, San Diego County, 3; Banta, 8; Bear
Valley, head of Carmel River, 2; Bear Valley, San Benito County, 1; Berg-
man, Riverside County, 1; Berkeley, 31; Boulder Creek, 2; Briceland, 2;
Burbank, 4; Calabasas, 1; Cameron’s Ranch, San Diego County, 1; Campo,
San Diego County, 1; Carlsbad, San Diego County, 3; Carpenteria, 2; Chico, 6;
Chinese Camp, 1; Church Ranch, 5 milesnorth of Tassajara Springs, 2; Dulzura,
1; El Nido, Jamul Creek, 9; Elsinore, 2; Eshom Valley, Tulare County, 1;
Fairfield, 3; Fort Bragg, 1; (old) Fort Tejon, 1; Freestone, 2; Fremont Peak,
Gabilan Range, 3; Fresno, 3; Fresno Flat, 4; Gaviota Pass, 5; Gilroy, 2; Glen
Ellen, 10; Hueneme, 3; Humboldt Bay, 3; Jackson, 1; Jacumba, 8; James-
burg, 5; Jolon, 4; King City, 1; Lagunitas, 1; Las Virgines Creek, 2; Layton-
ville, 1; Lebec, 1; Leesville, 6; Lemoore, 2; Los Banos, 1; Los Olivos, 2; Lower
Lake, 3; Lytle Creek, Los Angeles County, 2; Martinez, 1; Marysville, 6; Marys-
ville Buttes, 12; Mendota, 2; Milpitas Ranch, south base Santa Lucia Peak, 3;
Milquatay Valley, 1; Modesto, 1; Mono Flats, Santa Ynez River, 2; Montalvo,
3; Monterey, 16; Morro, 2; Mountain Spring, San Diego County, 3; Mount
George, 2; Mount St. Helena, 2; Nelson, Butte County, 2; Nicasio, 5; Novato,
3; Oakland, 1; Oceanside, 1; Orosi, 2; Pacheco Pass, Santa Clara County, 1;
Pacific Grove, 3; Pacific Ocean, near Mexican boundary, 6; Palo Alto, 1;
Paraiso Springs, 8;? Paso Robles, 5; Petaluma, 6;! Petrolia, 9; Pine Valley,
Monterey County, 1; Point Reyes, 19; Porterville, 1; Posts, 5; Pozo, 3;?
Radec, Riverside County, 2; Riverside, 5; Rockport, 5; Salinas, 4; San
Bernardino, 13;? San Bernardino Mountains (altitude 5,200 feet), 6; San
Diego, 6; San Emigdio Canyon, 7; San Fernando, 7; San Jacinto, 1; San Luis
Obispo, 5;? San Marcos, 1; San Mateo, 15; San Pasqual Valley, 1; San Pedro,
1; San Simeon, 5; Santa Barbara, 2; Santa Maria, 3; Santa, Monica, 4; Santa
Paula, 17;? Santa Ynez Mission, 4; Santa Ysabel, 5; Santiago Springs, San
Luis Obispo County, 1;? San Ygnacio Valley, San Diego County, 1; Shasta
Valley, 7;? Soledad, 2;? Stanford University, 22; Strawberry Valley, San
Jacinto Mountains, 3; Sur, 6; Sur River, 6; Tecate Valley, 3; Tecelote Canyon,
1; Tehama, 4; Tejon Canyon, 3; Temescal, 2; Three Rivers, 1; mouth Tia
Juana River, 1; Tracy 10; Twin Oaks, 1; Ventura River, 2; Walnut Creek,
20; Warren’s Ranch, Riverside County, 1; Westport, 2.

Lower California: El Rayo, Hanson Laguna Mountains,1; La Huerta, Hanson
Laguna Mountains, 1; Nachoguero Valley, 2; Rancho Viejo, 15 miles east of
Alamos, 1; San Isidro Ranch, near monument No. 250, Mexican boundary, 5.

1 Collection Field Mus. Nat. Hist. 2A pproaching megalotis. 3 Collection of Joseph Grinnell.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. 30

REITHRODONTOMYS MEGALOTIS PENINSUL& Elliot.
PENINSULA Harvest Mouse.

Rhithrodontomys peninsule Elliot, Pub. Field Columb. Mus., Zool. Ser., IIi, 1903,
p. 164.

Type locality San Quintin, Lower California.

Distribution.—West coast of Lower California, between latitude 30°
and 31°; southern limit of range not definitely known.

Characters —Similar to longicaudus, but with larger ears and
longer tail.

Oolor—Fresh pelage (October to January): Similar to corre-
sponding pelage of longicaudus, but averaging richer and redder, and
less brownish on the back; underparts white with a tinge of ochraceous-
buff. Worn summer pelage: Specimens not appreciably different from
longicaudus in same pelage.

Skull.Similar to those of megalotis and longicaudus.

Measurements.—Average of 11 specimens (adult and subadult)
from type locality: Total length, 154 (140-170); tail vertebre, 84
(78-91); hind foot, 17.4 (16-18). Skull: (See table, p. 81).

Remarks.—This form has a rather restricted range on the Pacific
coast of Lower California. It is closely related to longicaudus, with
which it intergrades in the vicinity of San Telmo.

Specimens exramined.—Total number, 32, from the following locali-
ties:

Lower California: Pozo Luciano (northwest slope San Pedro Martir Moun-
tains), 1; Rosario, 20;! San Quintin, 8; San Telmo, 1; Socorro, 2.

REITHRODONTOMYS MEGALOTIS CINEREUS Merriam.
AsHuy Harvest Mouse.

Reithrodontomys saturatus cinereus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 556.

Type locality.—Chalchicomula, Puebla, Mexico.

Distribuiion.—Southern portion of Mexican table-land in the States
of Hidalgo, Puebla, and Tlaxcala.

Characters.—Slightly larger and darker than megalotis with much
darker ears and tail; paler than saturatus.

Color.—Upperparts mixed black and light ochraceous-buff; darkest
in the median line where the black sometimes appears as a broad
band; sides light ochraceous-buff; underparts grayish white with a
tinge of ochraceous-buff and scmetimes with one or more small
blotches of the same color, chiefly in the pectoral region; ears fuscous,
with a large blackish patch on the inferior inner margin and another
on the superior outer margin; tail, fuscous, darker than in megalotis.

Skull.—Practically the same as that of saturatus, possibly a little
shorter.

1 Including 9 from Collection Mus. Comp. Zool.

36 NORTH AMERICAN FAUNA. [No. 36.

Measuremenis.—Average of 5 adults from type locality: Total
length, 148 (142-151); tail vertebre, 74 (71-78); hind foot, 19
(18.5-19.5). Skull: (See table, p. 81).

Remarks.—This form is intermediate between megalotis and satu-
ratus, being slightly larger externally than megalotis, with skull about
the size of that of saturatus. One of the topotype series matches
megalotis very closely in color, but the others are much darker; all
have darker ears, tail, and underparts.

The range of cinereus is apparently not extensive and is nearly
surrounded by that of saturatus, at least on the east, south, and west.

Specimens examined.—Total number, 11, from the following locali-
ties in Mexico:

Puebla: Chalchicomula, 5; Mount Orizaba, 1.

Tlaxcala: Apixaco, 2; Huamantla, 1.
Hidalgo: Real del Monte, 2.

REITHRODONTOMYS MEGALOTIS SATURATUS Allen & Chapman.
Dusky Harvest Mouse.
(Pld, fie-'8; Pl. WV, fie 58)

Reithrodontomys saturatus Allen and Chapman, Bull. Am. Mus. Nat. Hist., 1X, 1897,
p. 201.

Type localhty.—Las Vigas, Vera Cruz.

Distribution.—From Jalisco (Ocotlan), Hidalgo, and Vera Cruz
south to Oaxaca; altitudinal range approximately from 6,000 to
10,000 feet.

Characters.—Larger and darker (blacker and more intensely
ochraceous) than cinereus.

Color.—Upperparts mixed black and ochraceous-buff, the black
predominating; sides with less black but without a distinet ochra-
ceous line; underparts grayish white, often strongly suffused with
ochraceous-buff; ears blackish, or fuscous with a large blackish patch
on the inner posterior margin; tail sharply bicolor, fuscous-black
above, whitish beneath; feet grayish white.

Skull.—Similar in proportions to that of megalotis but decidedly
larger; zygomata parallel or slightly contracted anteriorly; closely
similar to those of tenwis and difficilis, but averaging larger, with
larger bulle and longer palatal foramina.

Measurements.—Average of 7 adults from type locality: Total
length, 162 (158-169); tail vertebre, 87 (80-95); hind foot, 19
(18-19.5). Skull: Gee table, p. 81). .

Remarks.—This is the largest and darkest form in the megalotis
group. It occupies the humid mountain slopes of southern Mexico,
intergrading with cinereus along the edge of the table-land and with
zacatece in Michoacan. Its range meets that of R. fulvescens

1914,] REITHRODONTOMYS MEGALOTIS GROUP. 37

dificilis in Vera Cruz, and the two may occur together in some
localities. They are closely similar in general appearance, but
saturatus may be distinguished by its blacker ears and back, whiter
underparts, and larger skull. Specimens from Oaxaca City are redder
than typical specimens, with less black on the dorsal area, but two
from the mountains west of Oaxaca are typical.
Specimens ecamined.—Total number, 68, from the following locali-
ties in Mexico:
Vera Cruz: Huauchinango, 1; Las Vigas, 11; Motint Orizaba, 2;! Perote, 6;
Xuchil, 7.1
Hidalgo: Tulancingo, 1.
Mexico: Salazar, 6; Toluca Valley, 2; Volcan Toluca, 1.
Morelos: Huiitzilac, 3.
Michoacan: Nahuaitzin, 9.
Jalisco: Ocotlan, 3.
Oaxaca: Mount Zempoaltepec, 4; mountains 15 miles west of Oaxaca, 2; moun-
tains near Ozolotepec, 1; Oaxaca City, 6; Tamazulapam, 1; Tlapancingo, 2.

REITHRODONTOMYS MEGALOTIS ALTICOLUS Merriam.
Cerro San Feuip—E Harvest Mouse.
Celene 2 aE EV, fies 2.)
Reithrodontomys megalotis alticolus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 556.

Type locahty.—Cerro San Felipe, Oaxaca, Mexico (altitude 10,000
feet).

Distribution.—Known only from vicinity of type locality.

Characters.—Similar to saturatus; tail shorter; skull with larger
bullz and more inflated braincase.

Color.—Upperparts mixed black and ochraceous-buff, darkest
on the dorsal area; ears fuscous, with a blackish patch on lower
inner margin; feet buffy white; ankles with a dusky streak; tail hair-
brown above, grayish white beneath.

Skull.Similar in size and proportions to that of saturatus; brain-
case more inflated (subglobular); bulle larger and more inflated;
nasals narrowed to a point posteriorly.

Measurements.—Type (3 ad.): Total length, 153; tail vertebre,
75; hind foot, 19. Average of two adults from La Parada, Oaxaca:
152; 78.5; 18.5. Skuil: (See table, p. 81).

Remarks.—This subspecies is a slightly differentiated form, very
closely related to saturatus, from which it differs chiefly in shorter
tail and in slight cranial characters. It seems to be confined to the
Cerro San Felipe and its environs. Specimens from La Parada, at
the base of the mountain, are not quite typical.

Specumens examined.—Total number, 3, from the following localities
in Mexico:

Oaxaca: Cerro San Felipe, 1; La Parada, 2.

i Collection Field Mus. Nat. Hist.

38 NORTH AMERICAN FAUNA. [No. 36.

REITHRODONTOMYS MEGALOTIS ARIZONENSIS Allen.
CuiIrRIcAHUA Harvest Mouss.

Reithrodontomys arizonensis Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 134.

Type locality —Rock Creek, Chiricahua Mountains, Ariz. (altitude
about 8,000 feet).

Distribution Known. only from the type locality.

Characters —About the size of megalotis; tail averaging slightly
longer; colors much darker and more ochraceous. Very similar to
longicaudus, but a little redder on the head, ears blacker, and tail
paler (hoary gray instead of brown).

Color—Upperparts ochraceous-buff, heavily mixed with black;
front and sides of face nearly pure buff; ears dark hair-brown, with
darker patches on both inner and outer margins; feet white; ankles
dusky; underparts white, with an ochraceous patch on the breast
between the fore legs; tail mouse-gray above, grayish white below,
clothed all around with scattered whitish hairs.

Skull_—Closely similar to that of megalotis.

Measurements —Average of 4 adults from type locality: Total
length, 149 (145-152); tail vertebrae, 78 (74-80); hind foot, 17
(16-18); ear, 13 (12.5-14). Skull: (See table, p. 81).

Remarks.—This subspecies, as noted by the original describer,
bears a surprising resemblance to Rk. megalotis longicaudus of Cali-
fornia, but the ranges of the two forms are separated by an extensive
area occupied by R. m. megalotis, and the present form is probably
more Closely related to R. m. zacatecee of the mountains of western —
Mexico, the range of which is known to extend north at least to
southern Chihuahua. The latter differs from arizonensis chiefly
in its more ochraceous underparts.

No intergrades between arizonensis and megalotis are known,
but quite probably such will later be found. Likewise the present
form is so close to zacatece that the two will probably be found
to intergrade. Mr. W. W. Price, who collected the type series,
thus describes its habitat:

Five specimens of this species were trapped on Rock Creek, in the Chiricahua
Mountains, July 7-8, at an elevation of about 8,000 feet. Two were in rocks and

dry soil away from the bed of the creek, and the others were caught under logs and
brush near the water.!

Specimens examined.—Your, from type locality.

1 Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 235.
2Two in Am. Mus. Nat. Hist., 2 in Field Mus. Nat. Hist.

1914.) REITHRODONTOMYS MEGALOTIS GROUP. 39

REITHRODONTOMYS MEGALOTIS ZACATECA Merriam.

MountTAIN Harvest Mouse.
(Ply Ts fig. 11 Pb gh, fie. 1.)

Reithrodontomys megalotis zacatece Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 557.

Reithrodontomys megalotis obscurus Merriam, Proc. Wash. Acad. Sci., III, 1901,
p. 558 (Sierra Madre, near Guadalupe y Calvo, Chihuahua).

Reithrodontomys colimz Allen, Bull. Am. Mus. Nat. Hist., XXII, 1906, p. 249 (not
of Merriam; specimens from Volcan de Fuego, Jalisco).

Type locality Valparaiso Mountains, Zacatecas, Mexico.

Distribution—Mountains of Western Mexico, from southern
Chihuahua to Michoacan.

-Characters—Very similar to arizonensis, but with darker under-
parts; smaller and paler than saturatus, with less black on upperparts.

Oolor.—Adults: Upperparts ochraceous-buff, heavily mixed with
black, the ochraceous color most pronounced on head and sides;
the black predominating on dorsal area but not forming a distinct
band; underparts washed with ochraceous-buff, this color usually
most intense between the fore legs; ears fuscous or dark hair-brown,
usually with a blackish patch on lower inner margin; feet whitish
or buffy white, ankles dusky; tail sharply bicolor, dark hair-brown
above, grayish white beneath. Yowng: Paler and grayer on upper-
parts; sides with a well-marked lateral line of light ochraceous-buff;
underparts whiter.

Skull—Similar to that of megalotis but smaller; rostrum slenderer,
narrowed at the tip; zygomata slightly narrower anteriorly; nasals
narrowed to a point posteriorly, ending on a line with premaxille.

Measurements —Average of 3 adults from type locality: Total
length, 154; tail vertebre, 84; hind foot, 18.5. Average of 3 adults
from Guadalupe y Calvo, Chihuahua: 160; 85; 18. Skull: (See
table, p. 81).

Remarks.—This subspecies is a small, dark-colored race of megalotis
inhabiting the mountains of western Mexico. No specimens have
been seen indicating intergradation between megalotis and zacatece,
but very likely such material may in the future be secured. Inter-
eradation with saturatus is indicated by specimens from the State of
Mexico (Salazar, Toluca Valley, etc.) which resemble saturatus in color
but have smaller ears and skulls intermediate in size between the two
forms. From Nahuatzin, Michoacan, specimens nearly typical of both
forms are athand. Some are larger with larger, darker ears and large
skulls (as in saturatus), but at least 2 adult individuals, by reason
of small size, small skull, small ears, and ochraceous underparts,
must be referred to zacatecze. Specimens from Patamban, Michoacan,
average more intensely ochraceous, especially on the underparts,
and have more black on the ears, but are otherwise typical. The

40 NORTH AMERICAN FAUNA. [No. 36. °

series from Guadalupe y Calvo, Chihuahua, forming the basis of
‘fobscurus’”’, is too slightly different to warrant recognition. The
coloration of the two series is practically the same, and the skulls of
‘fobscurus’”’ average only slightly larger.

This subspecies bears a striking resemblance to longicaudus from
California, but differs in having ochraceous underparts, slightly
longer tail, and a little more ochraceous color on the head.

Specimens examined.—Total number, 27, from the following locali-
ties in Mexico:

Chihuahua: Sierra Madre, near Guadalupe y Calvo, 3.
Durango: El Salto, 2.
Zacatecas: Valparaiso Mountains, 14.

Michoacan: Nahuatzin, 2; Patamban, 4.
Jalisco: Volcan de Fuego, 2.!

REITHRODONTOMYS AMOLES sp. nov.

QUERETARO Harvest Mouse.

Type from Pinal de Amoles, Queretaro, Mexico. No. 81234, U.S.
Nat. Mus., Biological Survey Collection, @ adult, September 20, 1896;
E. W. Nelson and EK. A. Goldman. Original No. 10169.

Characters.—Similar in color to &. m. saturatus, but very much
smaller.

Color.—Worn pelage: Upperparts mixed blackish and ochraceous-
buff; underparts whitish, strongly tinged on pectoral region with
ochraceous-buff; tail fuscous above, grayish white below; hind feet
whitish; fore feet white with a pronounced dusky stripe on upper
surface; ears dark brown (much mutilated, and size not known).

Skull—Similar to that of R. m. megalotis but much smaller;
zygomata parallel; bulle smaller and rather flat.

Measurements.—Type: Total length, 154; tail vertebrae, 83; hind
foot 16.5. Skull: (See table, p. 81.)

Remarks.—On geographical grounds this form ought to be close to
R. m. saturatus, but it differs markedly in size from that race. It is
even smaller than R. m. zacatece of the Sierra Madre, which it some-
what resembles in color.

Specumen examuned.—One, the type.

REITHRODONTOMYS CATALINA Elliot.
CATALINA Harvest Mouse.

Rhithrodontomys cataling Elliot, Pub. Field Columb. Mus., Zool. Ser. III, 1903, p. 246.
Reithrodontomys megalotis cataline Grinnell, Proc. Cal. Acad. Sci., Ser. 4, III, 1913,
p. 304.

Type locality.—Santa Catalina Island, Cal.
Distribution.—Santa Catalina Island, Cal.

1 Collection Am. Mus. Nat. Hist.

1914.] REITHRODONTOMYS MEGALOTIS GROUP. Aj

Characters—Sumilar to longicaudus but larger and slightly paler.

Color.— Worn spring pelage:' Upperparts mixed light ochraceous-
buff and blackish brown, darkest on back, but without a distinct
dorsal stripe; underparts white, tinged with buff in pectoral region;
tail dark hair-brown above, white beneath; ears hair-brown, clothed
with ochraceous hairs.

Skull_—tLarger than that of R. m. longicaudus, but not otherwise
different.

Measuremenis.—Average of 10 adults from type locality: Total
length, 169 (165-175); tail vertebre, 94 (90-99); hind foot, 18.6
(18-19.5). Skull: (See table, p. 81).

Remarks.—So far as known this is the only insular species in the
genus. Mr. C. P. Streator, who collected a series in 1892, found the
mice abundant on the island in brush and cactus.

Specimens examined.—T wenty-eight, from type locality.

REITHRODONTOMYS RAVIVENTRIS RAVIVENTRIS Dixon.
RED-BELLIED Harvest Mouse.
(PINT fie 9° PY: TV, fie. 9.)
Reithrodontomys raviventris Dixon, Proc. Biol. Soc. Wash., X XI, 1908, p. 197.

Type locality Redwood City, Cal.

Distribution.—Salt marshes of San Francisco Bay, Cal.

Characters —Similar to BR. megalotis longicaudus but upperparts
darker and underparts reddish; skull larger; tail slightly shorter.

Color.—Upperparts mixed black and pinkish cinnamon, the black
predominating on dorsal area; sides pale tawny in some individuals;
underparts pinkish cinnamon (rarely with a small white spot on chin) ;
ears black or fuscous on both surfaces, with a tuft of ochraceous hairs
at anterior base; hind feet and tail usually very dark, varying from
fuscous to clove-brown or sepia, the tail slightly paler beneath, but
feet usually darker beneath; toes whitish; front feet sepia, often
tinged with buffy white.

Skull—Decidedly larger than that of R. m. longicaudus; with rela-
tively shorter rostrum; nasals and palatal foramina shorter; zygo-
mata more widely expanded anteriorly.

Measurements.—Average of 21 from type locality: Total length,
130.7 (120-142); tail vertebre, 64.8 (56-74); hind foot, 16.6 (15-18).
Average of 8 from Melrose Marsh, Alameda County: 137; 66; 17.7.
Skull: (See table, p. 81).

Remarks.—This species is remarkable on account of its peculiar
characters and limited distribution. It is the darkest form found
in the United States and the only one having reddish underparts.
In color it most nearly resembles R. australis of Costa Rica, but the
underparts are even darker than in that species.

1 Fresh pelage not seen.

42, NORTH AMERICAN FAUNA. [No. 36.

It is apparently confined to the salt marshes in the southern part of
San Francisco Bay and although in many places its range abuts on
that of longicaudus, no evidence of intergradation between them has
been discovered. Two specimens—one from Palo Alto and one from
Berkeley—have whitish underparts but in other respects are typi-
cal ravwentris. Intergradation with halicwtes seems probable. The
pelage of both of these marsh forms is longer and seemingly thicker
than that of longicaudus.

Specumens ecamined.—Total number, 44, from the following locali-
ties :

California: Berkeley, 2; Elmhurst, 1;1 Melrose Marsh, Alameda County, 13;}
Palo Alto, 4;2 Redwood City, 24.}

REITHRODONTOMYS RAVIVENTRIS HALICGTES Dixon.
PretatumMA Marsa Harvest Mousse.
(Pl. I, fig. 10; Pl. IV, fig. 10.)
Reithrodontomys halicetes Dixon, Univ. Cal. Pub. in Zool., V, 1909, p. 271.

Type locality.—Salt marsh 3 miles south of Petaluma, Cal.

Distribution.—Salt marshes of San Pablo Bay, Suisun Bay, and the
lower San Joaquin and Sacramento Rivers.

Characters.—Similar in color to raviventris but larger; underparts
white; ears and feet paler; tail more distinctly bicolor. Compared
with R. m. longicaudus: Decidedly larger and darker, with a large
white patch on the throat; skull larger, with more widely spreading
zygomata.

Color.—Fresh pelage: Upper parts ochraceous-buff, heavily mixed —
on the back with black; sides ochraceous-buff without a well-defined
lateral line; underparts white (the bases of hairs plumbeous), some-
times irregularly blotched with ochraceous-buff; throat and sides of
mouth pure white to base of hairs; sides of nose and eye ring blackish;
ears and upper surface of tail fuscous or fuscous-black; underside of
tail dull grayish white with a buffy tinge; feet white or buffy white.
Worn pelage: Decidedly more ochraceous, the black tips of the hairs
seemingly worn off.

Skull—sSimilar to that of raviventris; larger than that of A. m.
longicaudus; zygomata usually (at least in adults, widely expanded
anteriorly.

Measurements.—Average of 13 from type locality: Total length, 156
(149-64); tail vertebree, 82 (75-85); hind foot 17.7 (17-19). Skull:
(See table, p. 81).

Remarks.—This form, although living only a short distance from
raviventris and under seemingly identical conditions, is readily sep-
arable from it. Nor is it in any sense a connecting form between

1 Collection Mus. Vert. Zool., Univ. of Cal. 2 Three from Collection Mus. Vert. Zool.

1914.] REITHRODONTOMYS FULVESCENS GROUP. 43

raviventris and longicaudus, being larger than either with upperparts
fully as dark as raviventris. The underparts are whiter than in
longicaudus, the latter usually being more or less tinged with pale
ochraceous-buff. The close resemblance between halicetes and raw-
ventris in skull characters and the fact that nearly half of the speci-
mens of halicetes examined from Petaluma are more or less suffused
beneath with ochraceous (though never so completely as in raviven-
tris) leads to the belief that the two are subspecifically related.
Specimens of halicetes in worn pelage are much redder (less blackish)
than those in fresh pelage, thus rather closely resembling in color
certain specimens of longicaudus.

Mr. Joseph Dixon, who collected the type series of this mouse,
states:

This mouse seems to be restricted to the salt marsh, its range being coextensive with
that of the ‘‘pickle grass” (Salicornia). Diligent search and trapping failed to reveal
its presence outside the Salicornia and no specimens of R. longicauda could be caught
in the Salicornia. The harvest mice use the runways of Microtus extensively.'

Specimens examined.—Total number, 56, from the following locali-
ties:
California: Brentwood, 1; Cordelia, Solano County, 6;? Grand Island (2 miles
north of Knights Landing, Yolo County), 1;? Grizzly Island, Solano County,
27;? Petaluma, 21.”

REITHRODONTOMYS FULVESCENS GROUP.
REITHRODONTOMYS FULVESCENS FULVESCENS Allen.

Sonoran Harvest Mouse.

Reithrodontomys mexicanus fulvescens Allen, Bull. Am. Mus. Nat. Hist., VI, 1894,
p) o9.
Reithrodontomys fulvescens Allen, Ibid., VII, 1895, p. 188.

Type locality —Oposura, Sonora, Mexico.

Mstribution.—Mountainous parts of southern Sonora, western Chi-
huahua, and northern Durango.

Characters. —Size medium—a little larger than R. m. megalotis, with
decidedly longer tail; prevailing color of upperparts ochraceous-buff
with a pronounced lateral line.

Color.—Upperparts light ochraceous-buff, sparingly mixed with
blackish brown; buff color most strongly marked on sides where it
frequently forms a pronounced lateral line, unmixed with brown; front
and sides of face tinged with grayish; underparts white, sometimes
faintly tinged with pale buff; feet buffy white; ears hair-brown
externally, clothed on inner surface with ochraceous-tawny hairs;
tail hair-brown above, grayish white beneath.

Skull—Similar to that of R. m. megalotis but slightly larger; inter-
pterygoid fossa broader.

1 Univ. of Cal. Pub. in Zool., V, 1909, p. 271. 2 Collection Mus. Vert. Zool., Univ. of Cal.

44 NORTH AMERICAN FAUNA. [No. 36. .

Measurements.—Type: Total length, 183; tail vertebra, 102; hind
foot, 19. One specimen (subadult) from Providencia Mines, Sonora:
158.5; 84; 20.5. Average of 4 (adult and subadult) from Casas
Grandes, Chihuahua: 165 (157-174); 93 (84-97); 20. Skull: (“ee
table, p. 81).

Remarks.—The mice of this group may be distinguished from R. m.
megalotis by their greater size, longer tails, and more intensely ochra-
ceous coloration. The present form resembles megalotis more closely
than do any of the other subspecies and certain specimens of the two
forms are colored almost alike on the back. The sides of fulvescens,
however, are more extensively buffy and less mixed with brown than
those of megalotis. The skulls of fulvescens may usually be distin-
guished from those of any member of the megalotis group by the
ereater breadth of the interpterygoid fossa.1 The ranges of the two
species meet along the eastern border of the Sierra Madre in Chi-
huahua, but there is no indication of intergradation between them.

This subspecies was the first member of the group to be described
and the third distinctively Mexican species to receive recognition.
Although described in 1894, it is still imperfectly known, only a small
number of specimens having been collected in the type region. Atthe
time of naming this form Dr. Allen considered it a subspecies of mext-
canus? and noted its close relationship to the Rio Grandeform. Later
he accorded it specific rank. The abundant material now available
from Mexico shows clearly that fulvescens, tenuis, and wntermedius are
closely related subspecies, connected through central Mexico by a per-
fect series of intergrades. The present form apparently intergrades,
also, with toltecus, as indicated by a specimen from Inde, Durango,
which agrees with fulvescens in color, but has a long tail like toltecus,
and skull intermediate in size between the two forms. The series
from Casas Grandes, Chihuahua, agrees with the type series in ex-
ternal characters, except for a more pronounced grayish wash on
the head and shoulders, but the single adult in the series has a some-
what larger skull than any in the Oposura series. In this character
it is matched by a specimen from Parral, Chihuahua.

Specuomens examined.—Total number, 12, from the following locali-
ties in Mexico:

Sonora: Oposura, 3;? Providencia Mines, 2.4

Chihuahua: Casas Grandes, 5; Parral, 1.
Durango: Inde, 1.°

1 Skulls of R£. m. saturatus are sometimes difficult to distinguish by this character from those of R&. f. tenuis.
2The name mczicanus at that time was applied to all the mice of this group from Texas and Mexico.

3 Collection Am. Mus. Nat. Hist.

4 Collection Field Mus. Nat. Hist.

56 Approaching toltecus.

1914.] REITHRODONTOMYS FULVESCENS GROUP. 45

REITHRODONTOMYS FULVESCENS TENUIS Allen.
Mexican Harvest Mouse.
(CPE Tne SURI NG fe 72)
Reithrodoniomys tenuis Allen, Bull. Am. Mus. Nat. Hist., XII, 1899, p. 15. of

Reithrodontomys griseoflasus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 558
(Ameca, Jalisco).

Type locality —Rosario, Sinaloa, Mexico.

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Fig. 3.—Distribution of Reithrodontomys fulvescens and subspecies.

Mstribution.—Greater part of Mexico, except extreme northern and
southern portions; from southern Sonora, Durango, Zacatecas, San
Luis Potosi, Nuevo Leon, and central Tamaulipas, south to southern
Jalisco, Michoacan, and the coast region of Vera Cruz.

Characters—Similar to fulvescens, but colors darker and more
intensely ochraceous.

46 NORTH AMERICAN FAUNA. [No. 36.

Color.—Upperparts and sides rich ochraceous-buff; head and
shoulders strongly ochraceous, without trace of grayish; back more
heavily mixed with black than in fulvescens; ears slightly darker;
underparts usually grayish white, sometimes faintly tinged with buff;
tail hair-brown above, grayish white below.

Skull_—Closely similar to that of fulvescens.

Measurements.—Average of 3 adults from Mazatlan, Sinaloa: Total
length, 169 (163-177); tail vertebre, 98 (96-102); hind foot, 20.8
(20.5-21). Average of 5 adults from Chacala, Durango: 177; 105;
21.3. Skull: Gee table, p. 81).

Remarks.—This is the most widely distributed member of the
fulvescens group and over its extensive range shows remarkably little
variation. Specimens from opposite sides of the continent can not
be distinguished by characters either of skin or skull. There is con-'
siderable individual variation in both color and cranial characters, but
none which is correlated with distribution.

The subspecies intergrades with fulvescens in southern Sonora, with
intermedwus in Nuevo Leon and Tamaulipas, with dificilis in Vera
Cruz and with ¢oltecus in Michoacan. Direct comparison with the
type has not been possible, but specimens in the Biological Survey
Collection from Mazatlan, forwarded to Mr. G. S. Miller, jr., while he
was in London, were compared by him with the type in the British
Museum and found to agree very closely with it. Mr. Miller states:
“‘T should say there is no question of the identity of the Mazatlan
specimens with tenuis.” In making comparisons these specimens
have been chiefly used.

Specimens from Ameca, Jalisco, forming the basis of ‘‘griseoflavus,”’
show in comparison with tenuis a very slight cranial difference, con-
sisting of a more inflated braincase. In this character they resemble °
helvolus, although on geographical grounds they should be approach-
ing toltecus. In color and size they agree closely with tenuis. The
type is decidedly more grayish on the back and less intensely buffy
on the sides than is usual in this subspecies, but two topotypes agree
perfectly with the Mazatlan specimens of tenuis. A series from Cha-
cala, Durango, is a little darker and more richly colored, as well as
slightly larger than the typical form. Specimens from the arid coast
region of Vera Cruz (Carrizal, Santa Maria, and Catemaco) are clearly
intermediate between tenuis and dificilis, the ears and tails being
noticeably darker than in tenwis but the underparts and backs paler
than in dtfficilis.

Specimens from as far north as Cerro de la Silla and Santa Catarina,
Nuevo Leon, are referable to tenuis, though those from Santa Cata-
rina might almost as well be considered intermedius. Specimens
from Acambaro, Michoacan, are intermediate in size between tenuis
and toltecus.

1914.] REITHRODONTOMYS FULVESCENS GROUP. 47

Specimens examined.—Total number, 127, from the following local-
ities in Mexico:

Sinaloa: Sinaloa, 2; Altata, 1; Culiacan, 1; Mazatlan, 4; Rio Mazatlan, 1.

Sonora: Alamos, 6.!

Jalisco: Ameca, 3; Atemajac, 2; Estancia, 1;? Etzatlan, 6;? Lagos, 1; Las
Canoas, 2;? Los Masos, 1;? Mascota, 1; Ocotlan, 2; Plantinar, 2; San Sebas-
tian, 8;3 Talpa, 1; Zapotlan, 7.

Tepic: Tepic, 2.

Durango: Chacala, 8; Durango, 3.

Zacatecas: Berriozabal, 1; Valparaiso, 5.

San Luis Potosi: Hacienda La Parada, 3.

Queretaro: Jalpan, 2; Tequisquiapam, 2.

Michoacan: Acambaro, 2.

Morelos: Cuernavaca, 1.*

Guanajuato: Santa Rosa, 1; Silao, 2.

Vera Cruz: Carrizal, 4; Catemaco, 1; San Carlos, 2;> Santa Maria, 4.

Tamaulipas: Alta Mira, 11; Hidalgo, 9; Jaumave, 1; Victoria, 5.

Nuevo Leon: Cerro de la Silla, 3; Santa Catarina, 3.

REITHRODONTOMYS FULVESCENS INTERMEDIUS Allen.
Rio GRANDE Harvest MOUSE.

Ochetodon mexicanus Allen, Bull. Am. Mus. Nat. Hist. III, 1891, p. 223 (not Reith-
rodon mexicanus Sauss.)

Reithrodoniomys mexicanus intermedius Allen, Bull. Am. Mus. Nat. Hist., VII, 1895,
p. 186.

Reithrodontomys laceyi Allen, Bull. Am. Mus. Nat. Hist., VIII, 1896, p. 235 (Watson’s
Ranch, 15 miles south of San Antonio, Tex.).

Reithrodontomys intermedius Bailey, N. Am. Fauna No. 25, 1905, p. 104.

Type locality —Brownsville, Tex.

Distribution.—Southern Texas and adjacent parts of Mexico from
Del Rio to Brownsville; east to Bexar and Bee Counties; north to
Wichita Mountains, Okla.

Characters —Very similar to tenuis but averaging duller and less
intensely ochraceous above, and sides paler. Compared with fulves-
cens: Upper parts, particularly head and shoulders, deeper ochraceous.

Color— Winter pelage (February): Ground color of upperparts
light ochraceous-buff, brightest on sides, strongly mixed with blackish
brown on the back; ears hair-brown, usually tinged with ochraceous
on inner surface; tail hair-brown above, grayish white beneath; feet
white; underparts white, sometimes faintly tinged with buff. Sum-
mer (worn) pelage: Decidedly redder on back and sides, some speci-

mens approaching dull orange-cinnamon. Young: Colors grayer and
less ochraceous.

1 Approaching fulvescens.

2 Collection Am. Mus. Nat. Hist.

3 Approaching nelsoni.

* This specimen seems best referable to this subspecies, although on geographical grounds it should
beeither foltecus or helvolus.

5 Collection Field Mus. Nat. Hist.

48 NORTH AMERICAN FAUNA. [No. 36. _

Skull—Very similar to that of tenuis, but braincase averaging
slightly larger. :

Measurements.—Average of 8 adults from Rio Grande Valley
(Matamoras and Camargo, Tamaulipas): Total length, 169 (160-181);
tail vertebrae, 98 (88-103); hind foot, 20.8 (20-21.5). Skull: (See
table, p. 81).

Remarks.—This subspecies is most nearly related to tenuis, than
which it has a much less extensive range. The differences between
untermedius and tenuis are really very slight, consisting in a more
intense suffusion of ochraceous-buff in éenuis, especially noticeable on
the sides. Specimens from Bexar and Kerr Counties—the type region
of “laceyt’’—average slightly grayer and less ochraceous than speci-
mens from the mouth of the Rio Grande, but individuals in each
series may be matched by those from the other and the differences
seem too slight to warrant recognition of the form. If both inter-
medius and ‘‘laceyi’’ were to be recognized, the former would be
nothing more than a series of intermediates between fenwis on the
one side and ‘‘aceyz”’ on the other.

Specomens examined.—Total number, 65, from the following locali-
ties:

Oklahoma: Mount Scott, 9.

Texas: Brownsville, 26; Corpus Christi, 2; Del Rio, 2; Lacey’s Ranch, near
Kerrville, 4; Padre Island, 1; Rio Grande City, 1; San Antonio, 3; San
Diego, 5; Santo Tomas, 2.

Tamaulipas: Camargo, 4; Matamoros, 6.

REITHRODONTOMYS FULVESCENS AURANTIUS Allen.

GoLDEN Harvest Mouse.

Reithrodontomysmexicanus aurantius Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 137.

Reithrodontomys chrysotis Elliot, Field Columb. Mus., Zool. Ser., I, 1899, p. 281
(Dougherty, Okla.).

Reithrodontomys aurantius Bailey, N. Am. Fauna No. 25, 1905, p. 105.

Type locality.—Lafayette, La.

Disiribution.—Louisiana (west of the Mississippi River), southern
and east-central Arkansas, eastern Texas, and eastern Oklahoma;
north to southwestern Missouri (Carthage). Confined to Lower
Austral Zone.

Characters.—Colors decidedly richer and darker than in interme-
dws; skull slightly larger.

Color.—Adults in unworn pelage: Upperparts varying from pinkish
cinnamon to ochraceous-tawny, usually heavily mixed with blackish
brown; sides of head and body usually rich ochraceous or tawny but
without a pronounced lateral line where the color of the sides meets
that of the underparts; dark markings of back frequently forming a

1914.] REITHRODONTOMYS FULVESCENS GROUP. 49

distinct median band; ears dull sepia, clothed on inner surface with
tawny hairs; underparts grayish white, often with a distinct tinge of
pale buff. Immature pelage: Decidedly more grayish, often lacking
entirely the bright tawny shades. Variation: There is considerable
variation in color in this subspecies, even among adult specimens.
This consists both in the intensity of the ochraceous shades and in the
amount of blackish suffusion on the back and sides. Some individ-
uals have the whole back heavily sprinkled with blackish hairs,
while others show only a rather narrow band down the median line.
A specimen from Matagorda Island (March 31) is quite exceptional
in being intensely tawny over the entire upperparts with only faint
indications of black hairs on the back, thus closely resembling speci-
mens of Peromyscus nuttalli. Other individuals from the same island
are normal in color.

Skull_—_Very similar to that of intermedius, but braincase averag-
ing a little broader.

Measurements.—Adult from Houma, La.: Total length, 176; tail
vertebre, 100; hind foot, 20. Average of 5 adults from Velasco,
Tex.: 170; 97; 21. Average of 9 young adults from Sour Lake, Tex.:
162 (154-170); 89 (83-94); 20 (19-21). Skull: (See table, p. 81).

Remarks.—This subspecies is a well-marked race, occupying the
humid or Austroriparian division of the Lower Austral Zone, west of the
Mississippi River. Although it shows considerable individual varia-
tion and occasional specimens—chiefly immature ones—are hardly
distinguishable from specimens of intermedius, the general intensity
of coloration shown by any series of specimens makes identification
possible at a glance. The type and topotypes of ‘‘chrysotis’”’ have’
been examined and found to agree perfectly with specimens of auran-
tius from eastern Texas.

Specomens eramined.—Total number, 88, from the following local-
ities:

Louisiana: Avery, 5; Belcher, 1; Foster (5 miles east of Shreveport) 1; Houma,

1; Iowa Station, 1; Lafayette, 2; Lecompte, 1 (skull); Mer Rouge, 4; Natchi-
toches, 1.

Arkansas: Beebe, 2; Delight, 4.

Missouri: Carthage, 2.

Oklahoma: Dougherty, 3; Stilwell, 3.

Texas: Barnard Creek, west of Columbia, 6; East Caranchua Creek, Matagorda
County, 2; Elliott, Matagorda County, 1; Hempstead, 8; Joaquin, 7; Mata-
gorda, 5; Matagorda Island, 5; Nacogdoches, 2; Selkirk Island, Matagorda
County, 1; Sour Lake, 10; Texarkana, 1; Velasco, 9.

1 Collection Field Mus. Nat. Hist,
28657 °—]4-4

50 NORTH AMERICAN FAUNA. [No. 36. -

REITHRODONTOMYS FULVESCENS DIFFICILIS Merriam.
ORIZABA Harvest Mousse.
GPL fer 8s rio las)

? ? Reithrodon sumichrasii De Saussure, Rev. Mag. Zool., 2d Ser., XIII, 1861, p. 3.1
Ochetodon mexicanus Coues, Proc. Acad. Nat. Sci. Phila., 1874, p. 186; Mon. N. Am.
Rodentia, 1877, pp. 128-130. (Not Reithrodon mexicanus Sauss.)
Reithrodontomys mexicanus Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 135; IX,
1897, p. 199 (not Retthrodon mexicanus Sauss).
Reithrodontomys difficilis Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 556.

Type locality.—Orizaba, Vera Cruz.

Distribution.—Interior mountain slopes along the southern end of
the Mexican table-land in the States of Vera Cruz and Puebla.

Characters.—Similar to tenuis, but colors decidedly darker; ears and
tail blacker, and underparts more ochraceous.

Color.—Upperparts pinkish cinnamon mixed with blackish brown,
in some specimens the brown prevailing, in others the cinnamon;
sides of face and body usually clear pinkish cmnamon, though in
certain worn or immature specimens this color is nearly absent;
ears fuscous or sepia, usually with more or less tawny hairs on inner
surface; tail fuscous above, grayish white below; underparts grayish,
with a strong wash of light pinkish cinnamon, the latter color pre-
vailing in the majority of the individuals; feet grayish white; ankles
dusky.

Skull—Closely similar to that of tenwis; braincase averaging a
trifle broader. Compared with R. megalotis saturatus: Skull shorter
and relatively broader, with shorter nasals and broader interptery-
goid fossa.

Measurements.—Average of 5 (subadult) from type locality:
Total length, 170 (162-177); tail vertebre, 96.6 (88-101); hind foot,
19.5 (19-20). Adult from Maltrata, Vera Cruz: 172; 97; 21. Skull:
(See table, p. 81).

Remarks.—This is the darkest of the races of fulvescens. Although
rather restricted in range, it is a well-marked form, easily distin-
guished from tenuis by its darker colors, but intergradation is clearly
shown by specimens from the coast region of Vera Cruz (Carrizal
and Santa Maria). From R. megalotis saturatus, the range of which
is. adjacent to that of dificilis in Vera’Cruz, it differs in more tawny
coloration, ochraceous instead of grayish underparts, and smaller
and. paler ears.

1 When the type of R. ‘‘sumichrasti” can be compared with modern material it may be possible to
identify the species and use the name. I have seen photographs of the type skull, which clearly show it to
belong in the typical subgenus, and the original description agrees best with the present form; but as no

definite type locality is assigned and the description is inadequate for subspecific determination, it seems
best to let the name remain in synonymy.

1914.] REITHRODONTOMYS FULVESCENS GROUP. 51

Specimens examined.—Total number, 36, from the following local-
ities in Mexico:
Vera Cruz: Malirata, 1; Orizaba, 18; Mirador, 2; Jalapa, 14.1
Puebla: Tehuacan, 1.

REITHRODONTOMYS FULVESCENS TOLTECUS Merriam.
TottEc Harvest Movusse.
CPE he sO; Biey, fig. 10.)

Reithrodontomys levipes toltecus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 555.
Rhithrodontomys inexspectatus Elliot, Field Columb. Mus., Zool. Ser., III, 1903, p.
145 (Patzcuaro, Michoacan).

Type locality—Tlalpam, Federal District, Mexico.

Distribution.—Table-land region of southern Mexico, from southern
San Luis Potosi to Michoacan and the Valley of Mexico.

Characters.—Similar to tenuis, but decidedly larger and slightly
darker.

Color—Upperparts mixed black and rich ochraceous-buff, with a
cinnamon tinge, the black usually showing a tendency to form a median
band on the back from nose to tail; sides pure ochraceous-buff or
sparingly mixed with black; underparts with a slight tinge of pale
buff; tail fuscous or hair-brown above, grayish white below.

Skull—Similar in shape to that of tenuis but decidedly larger;
braincase somewhat more inflated, and evenly rounded; rostrum
long and relatively slender; zygomata nearly parallel to axis of skull.

Measuremenis.—Average of 2 adults from type locality: Total
length, 193 (189-196); tail vertebrz, 106 (104-108); hind foot, 21.5
(21-22). One adult from Patzcuaro, Michoacan: 180; 105; 21.
Average of 2 adults from Rio Verde, San Luis Potosi: 192; 110;
21.8. Skull: (See table, p. 81).

Remarks.—This is a well-marked form of the fulvescens group,
occupying the higher parts of the Mexican table-land. Itis poorly
represented in the material at hand, and its characters and exact
distribution are not well known. Skulls from the same or near-by
localities show an unusual amount of variation in size.

Intergradation apparently takes place between toltecus and tenuis
and between foltecus and helvolus, but the material at hand is
too scanty to show this clearly. Specimens from Rio Verde, San
Luis Potosi, and from Zamora and Los Reyes, Michoacan, are con-
sidered intermediates between this form and tenuis, and specimens
from Huajuapam, Oaxaca, and Chilpancingo, Guerrero, interme-
diates between it and helvolus. The series from Los Reyes is very
puzzling. In color the specimens are all exactly alike, being a little
darker and redder than either tenuis or typical foltecus. One speci-

1 Collection Am. Mus. Nat. Hist,

52 NORTH AMERICAN FAUNA. [No. 36.

men, an adult male, equals toléecus in size and agrees perfectly with it
in skull characters. Several other skulls in the series, however,
equally old, are very much smaller—as small, indeed, as typical
tenuis, while between the two extremes is a nearly perfect series of
intergrades.

Two specimens from Patzcuaro, Michoacan, including the type of
“inexspectatus,’’ do not differ appreciably from ¢oltecus. The topo-
type has the underparts strongly suffused with ochraceous and has
no white on the tail. This species differs widely in skull characters
from levipes, with which it was originally associated as a subspecies.

Specumens exramined.—Total number, 30, from the following locali-
ties in Mexico:

Mexico: Tlalpam, 3.

Hidalgo: Marques, 1 (skull); Zimapan, 1.

San Luis Potosi: Rio Verde, 6.

Michoacan: Los Reyes, 13; Patzcuaro, 2;1 Zamora, 4.

REITHRODONTOMYS FULVESCENS HELVOLUS Merriam.
Oaxaca Harvest Mouss.
CELe tig 9s" Pile Vi ceo)
Reithrodontomys griseoflavus helvolus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 554.

Type locality Oaxaca City, Oaxaca, Mexico.

Distribution.—Interior plateau of Oaxaca, Guerrero, and Puebla.

Characters—About the size of toltecus but differing in paler colors,
larger ears, and smaller skull; very similar to tenuis, but color of back
and sides more pinkish (less ochraceous) and belly whiter.

Color.—Upperparts light ochraceous-salmon, sparingly lined with
black but without a distinct median band; underparts white, often
with a slight yellowish tinge, but never (Gn type series) with any
suffusion of ochraceous. Ears dark hair-brown, clothed on inner sur-
face with ochraceous hairs; tail fuscous or hair-brown, soiled whitish
below.

Skull—Smaller than that of toltecus and slightly larger than that
of tenuis; braincase rounded and moderately inflated, narrowed pos-
teriorly; rostrum rather short; interpterygoid fossa actually and
relatively broader than in either foltecus or tenuis.

Measurements.—Average of 11 adults from type locality: Total
length, 189 (181-200); tail vertebrae, 110 (104-116); hind foot, 20.5
(20-21); ear, 13.2 (12.5-13.7). Skull: (See table, p. 81).

Remarks.—This subspecies agrees in size with foltecus, but most
nearly approaches ¢enuwis in color and cranial characters. No inter-
mediate specimens between helvolus and tenuis have been examined,
however, and our present knowledge of the distribution indicates a

1 Including type of ‘‘inerspectatus”’ from Collection Field Mus, Nat. Hist,

1914.] REITHRODONTOMYS FULVESCENS GROUP. 53

gap between the ranges of these two forms, but further collecting in
Michoacan and Guerrero may show that they intergrade. From
toltecus this form differs, as already pointed out, in having a decidedly
smaller skull, larger ears, and whiter belly. A specimen from Hua-
juapam, Oaxaca, is intermediate in characters between the two forms.
Specimens examined.—Total number, 22, from the following locali-

ties in Mexico:

Oaxaca: Huajuapam, 1; Oaxaca, 18; Yalalag, 1.

Guerrero: Tlalixiaquilla, 1; Tlapa, 1.

REITHRODONTOMYS FULVESCENS CHIAPENSIS subsp. nov.

CHIAPAS HARVEST MOUSE.

Type from Canjob, Chiapas, Mexico. No. 132865, U.S. Nat. Mus.,
Biological Survey Collection, ¢ adult, May 2, 1904; Nelson and
Goldman. Original No. 16741.

Distribution.—Highlands of Chiapas.

Characters.—Sunilar to helvolus but darker and smaller, with smaller
ears.

Color.—Upperparts ochraceous-salmon heavily lined on back with
black hairs; lateral line of salmon moderately well defined; ears fus-
cous; underparts grayish white; tail fuscous above, soiled whitish
below; feet grayish; ankles fuscous.

Skull.—Not appreciably different from that of helvolus.

Measurements —Average of 8 adults from type locality: Total
length, 169 (163-183); tail vertebre, 96 (92-102); hind foot, 20.1
(20-20.5); ear, 14. Skull: (See table, p. 81).

Remarks.—This is a small dark form apparently most nearly related
to helvolus. Insize and general color it resembles tenuis rather closely,
butis darker, especially ontheheadand ears. Itisalmostasdarkabove
as difficilis, but has whiter underparts. It closely resembles aurantius
also, but the underparts are more nearly pure white (never tinged with
buff), and the general tone of the upperparts is slightly paler. The
skull of chiapensis averages a little broader with flatter braincase and
shorter nasals than that of aurantius.

Specumens examined.—Total number, 17, from the following locali-
ties in Mexico:

Chiapas: Canjob, 9; Comitan, 4; San Bartolome, 3; San Vicente, 1.
REITHRODONTOMYS FULVESCENS NELSONT subsp. nov.

NELSoN Harvest Mouse.

Type from Colima, Colima, Mexico. No. 33493, U.S. Nat. Mus.,
Biological Survey Collection, ? adult, Mar. 9, 1892; E. W. Nelson.
Original No. 2050.

Distribution.—Coast region of Colima (and Jalisco 2).

54 NORTH AMERICAN FAUNA. [No. 36.

Characters.—Smilar to tenuis, but smaller; colors brighter and more
intensely ochraceous.

Color.—Adults: Upperparts varying from deep ochraceous-buff to
pinkish cinnamon, mixed on top of head and back with black; under-
parts suffused (sometimes heavily) with pinkish buff; ears dusky
hair-brown; tail dark hair-brown or fuscous above, grayish white
below; fore feet pale buff; hind feet soiled whitish. Young: Colors
less intensely ochraceous; underparts whiter.

Skull——Similar to that of tenuis, but smaller; zygomata more
contracted anteriorly.

Measurements —Type: Total length, 173; tail vertebrae, 95; hind
foot, 19. Average of 5 from type locality: 166; 92; 19.2. Skull:
(See table, p. 81).

Remarks.—This subspecies is a small, bright-colored form occupying
the coast plain of Colima and perhaps of adjacent States. Specimens
from San Sebastian and Etzatlan, Jalisco, are intermediate between
this form and tenuis, but seem to be nearer to the latter. A single
specimen from Acaponeta, Tepic, is very small, and clearly referable
to nelsoni, although on geographical grounds it might be expected to
be tenuis.

Specimens examined.—Total number, 8, from the following localities
in Mexico:

Colima: Colima, 7.
Tepic: Acaponeta, 1.

REITHRODONTOMYS FULVESCENS MUSTELINUS subsp. nov.

BUFF-BELLIED Harvest Mouse.

Type from Llano Grande, Oaxaca, Mexico. No. 71549, U.S. Nat.
Mus., Biological Survey Collection, @ adult, February 18, 1895;
K. W. Nelson and E. A. Goldman. Original No. 7483.

Distribution.—Coast region of Oaxaca and Guerrero.

Characters.—Similar to helvolus but colors darker and richer; un-
derparts buffy.

Color.—Upperparts varying from ochraceous-salmon to rich pinkish
cinnamon, heavily mixed on head and back with black; underparts
strongly suffused with pinkish buff; ears hair-brown; tail fuscous
above, whitish beneath; fore feet buffy white; hind feet grayish
white.

Skull.—Similar in size and proportions to that of helvolus; nasals
longer, narrowed to a point posteriorly; palatal foramina short, not
reaching plane of molars; bulle small.

Measurements.—Type: Total length, 184; tail vertebrz, 105; hind
foot, 20. Adult from Chilpancingo, Guerrero: 198; 110; 20. Skull:
(See table, p. 81).

1914.] REITHRODONTOMYS FULVESCENS GROUP. 55

Remarks.—This subspecies is most nearly related to helvolus, dif-
fering from it in much deeper coloration. It resemples A. levipes
rather closely in external appearance, but differs widely from itin
skull characters. A specimen from Chilpancingo, Guerrero, has a
longer skull than the type, somewhat suggesting the skull of toltecus.

Specimens examined.—Total number, 4, from the following locali-
ties in Mexico:

Oaxaca: Llano Grande, 2.
Guerrero: Acapulco, 1; Chilpancingo, 1.

REITHRODONTOMYS AMENUS Elliot.
TEHUANTEPEC Harvest Movse.
(BEAT fie iP lVetie 19
Rhithrodontomys amenus Elliot, Proc. Biol. Soc. Wash., XVIII, 1905, p. 234.

Type locality.— Reforma, Oaxaca, Mexico.

Distribution.—Known only from the type locality.

Characters.—Size very small; tail moderate; similar in color to
R. f. helvolus but upperparts deeper ochraceous.

Color.—Upperparts bright ochraceous-buff, becoming tawny-ochra-
ceous on middle of the back, where indistinctly lined with black;
underparts and feet white; tail hair-brown above, paler below.

Skull.—Small, with short heavy rostrum; braincase moderately flat
and narrowed posteriorly; zygomata slightly contracted anteriorly;
bullz small and flat; interpterygoid fossa very broad; palatal foramina
long, reaching behind plane of molars.

Measurements.—Type: Total length, 141; tail vertebre, 81; hind
foot, 18.5. Skull: (See table, p. 81).

Remarks.—This species, known from only a single specimen, seems
not to be closely related to any of the recognized forms. It most
nearly resembles #. f. helvolus in color, but differs widely from it in
size and cranial characters, being decidedly smaller than any other
species in southern Mexico. It inhabits the low coast plain bor-
dering the Gulf of Tehuantepec in southern Oaxaca.

Specumen examined.—One, the type.

REITHRODONTOMYS OTUS Merriam.
BiG-EARED Harvest MOUSE.
(Pla fics 2) PLAVa fie. 2 3)
Reithrodontomys levipes otus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 555.

Type localitySierra Nevada de Colima, Jalisco, Mexico (altitude
6,500 feet).

Distribution.— Known only from the type locality.

Characters.—Similar to R. f. toltecus, but tail longer and ears larger;
colors more ochraceous and less blackish.

56 NORTH AMERICAN FAUNA. [No. 36.

Color.—Upperparts pinkish cinnamon with an ochraceous tinge,
mixed with brownish black; underparts heavily washed with light
pinkish cinnamon; sides without a distinct lateral line of ochraceous;
ears between fuscous and hair-brown with scattering ochraceous hairs
on inner surface; front feet dull buffy white; hind feet grayish, with
a tinge of buff; ankles dark hair-brown; tail hair-brown above,
soiled whitish below.

Skull.—Similar in size and shape to that of #. f. toltecus but brain-
case flatter and zygomata narrowed anteriorly; nasals and pre-
maxille as in toltecus; audital bulle also similar (arger than in R.
levipes); foramen magnum very large; molars with simple enamel
pattern, as in the fulvescens group.

Measurements.—Type: Total length, 202; tail vertebre, 120; hind
foot, 22. Skull: (ee table, p. 81).

Remarks.—This seems to be a well-marked species, but as only a
single specimen is known its relationships are not clear. In skull
characters it resembles R. f. toltecus rather closely and differs from
R. levipes in having a longer, narrower skull, with larger bulle and
no subsidiary enamel loops on the upper molars, being therefore in
the typical subgenus. The ground color of the upperparts is similar
to that of toltecus, but is a little duller and the blackish median
band is lacking; the underparts are decidedly more ochraceous.
There is no evidence to show that it intergrades with foltecus, so for
the present it seems best to consider it a distinct species. The
simple ename! pattern of the molars shows it to be not closely related
to levipes, which is in the subgenus Aporodon.

Specimen examined.—One, the type.

REITHRODONTOMYS RUFESCENS GROUP.
REITHRODONTOMYS RUFESCENS RUFESCENS Allen & Chapman.
RUFESCENT Harvest MovwsE.

Reithrodontomys rufescens Allen & Chapman, Bull. Am. Mus. Nat. Hist., IX, 1897,
p. 199.

Type locality —Jalapa, Vera Cruz, Mexico.

Distribution.—Mountain slopes of eastern Mexico in the States of
Queretaro, Hidalgo, Puebla, Vera Cruz, and Oaxaca.

Characters.—Larger than Rf. fulvescens difficilis and much darker;
sides lacking the bright ochraceous line characteristic of the fulves-
cens group; tail blackish, unicolor.

Color.—Fresh pelage (April specimens, Jalapa, Vera Cruz): Upper-
parts tawny, strongly mixed with black over the entire dorsal area from
nose to tail; sides nearly pure tawny; ears varying from fuscous to
fuscous-black; feet buffy white washed with fuscous; tail fuscous,
nearly unicolor, clothed with scattered grayish hairs; underparts

—— ee

1914.] REITHRODONTOMYS RUFESCENS GROUP. 57

heavily washed with pinkish cmnamon. Worn pelage July specimens,
Jico, Vera Cruz): Underparts somewhat paler; ears and tail blacker.

Skull—tLarger than that of &. fulvescens difficilis, with longer and
relatively slenderer rostrum; braincase usually subglobular (occasion-
ally moderately flattened) ; meds long and ceed posteriorly to a
point, ending about on a line with premaxille; zygomata moderately
contracted anteriorly; audital bulle relatively small; palatal foramina
long, reaching to or beyond plane of first molars. Harnel pattern of
upper molars in some specimens simple, in others showing incomplete
subsidiary loops and small accessory tubercles.

Measurements.—Average of 10 adults from Jico and Jalapa, Vera
Cruz: Total length, 176 (168-182); tail vertebree, 98 (93-104); hind
foot, 20 (18-21). Skull: (ee table, p. 81).

Remarks.—This subspecies is one of the darkest forms in the genus.
It has a rather limited range in the humid mountainous parts of
eastern Mexico, intergrading with luteolus in Oaxaca and Guerrero.
Its range overlaps slightly that of R. fulvescens difficilis (both species
occurring at Jalapa, Vera Cruz), but it does not intergrade with any
member of the fulvescens group. It is readily distinguished from
them by its dark tawny coloration and unicolor tail.

In dental characters this species is quite inconstant, as pointed out
above; it is considered an aberrant member of the subgenus
Reithrodoniomys, bridging the gap between it and Aporodon in much
the same way that levipes does.

Specimens examined.—Total number, 15, from the following locali-
ties in Mexico:

Vera Cruz: Jalapa, 10;! Jico, 8.
Puebla: Huauchinango, 3.

Queretaro: Pinal de Amoles, 1.
Oaxaca: Reyes, 1

RETTHRODONTOMYS RUFESCENS LUTEOLUS subsp. nov.
YELLOW Harvest Mowusz.
CEE ie OP iV fie. 6.)

Type from Juquila, Oaxaca, Mexico (altitude 5,000 feet). No.
71558, U.S. Nat. Mus., Biological Survey Collection, ? adult, Feb. 28,
1895; EK. W. Nelson and E. A. Goldman. Original No. 7579.

Distribution.—Mountains of Oaxaca and Guerrero.

Characters.—Similar to rufescens but colors brighter and less
blackish; ears larger and tail longer; skull with flattened braincase;
molars without accessory cusps.

Color.—Upperparts and sides rich ochraceous-buff, varying to
pinkish cinnamon, more or less darkened on the back with blackish

1 Collection Am. Mus. Nat. Hist.

58 NORTH AMERICAN FAUNA. [No. 36.

brown; underparts light pinkish cmmamon; tail usually bicolor,
fuscous above, soiled whitish below; fore and hind feet whitish,
tinged with buff; ankles dusky; ears fuscous on both surfaces.

Skull—Similar to that of rufescens but averaging broader; brain-
case very flat; nasals decidedly narrow posteriorly, ending on a line
with premaxille; audital bulle rather small (as in rufescens).

Measurements.—Average of 8 adults from type locality: Total
length, 181 (169-199); tail vertebre, 103 (92-112); hind foot, 20.4
(20-21). Skull: (See table, p. 81).

Remarks.—This subspecies is a well-marked race of rufescens,
occupying the mountains of the west coast of southern Mexico. It
closely resembles #. colume nerterus in color, but differs from it

cute}

RC
NK a e sess
iS a

Spee MN

RAS
Boe a

4 a
Seater) fat

Sy AEVTHRODONTOMVS PR. RUFESCENS cose: AGUA
WK R. LUTEOLUS

DORSALIS

A. AUSTRALIS

A. MODESTUS

COLIMAE

ALLEM/

Fic. 4. Distribution of Reithrodontomys rufescens, R. dorsalis, F. australis, R. colime, #. dileni,
and subspecies.

in skull characters. Additional material, however, may show that
nerterus intergrades with the present form.

Specimens from Omilteme, Guerrero, are slightly darker than the
type series and one of them has a unicolor tail, as in rufescens. These
specimens, as also one from the mountains west of Oaxaca City,
are considered intermediate between rufescens and luteolus. Speci-
mens from the type locality of this form show no approach to the
subgenus Aporodon, but in the series from Omilteme is one specimen
having well-developed subsidiary tubercles, as is frequently seen in
the subspecies rufescens.

Specimens examined.—Total number, 15, from the following locali-
ties in Mexico:

Oaxaca: Juquila, 11; mountains 15 miles west of Oaxaca, 1.
Guerrero: Omilteme, 3.

1914.] REITHRODONTOMYS RUFESCENS GROUP. 59

REITHRODONTOMYS ALLENI sp. nov.

ALLEN Harvest Mouse.

Type from mountains near Ozolotepec, Oaxaca, Mexico (altitude
10,000 feet). No. 71563,U.8S. Nat. Mus., Biological Survey Collection,
¢ adult, Mar. 27, 1895; E. W. Nelson and E.A.Goldman. Original
number 7749.

Distribution —Known only from the type locality.

Characters.—Externally similar to R. c. colime but back darker,
underparts whiter, and tail longer; skull apparently nearest to that
of R. rufescens uteoras.

Color —Upperparts mixed black and light pinkish cinnamon; darkest
along the median line; sides without a lateral line, less richly colored
than in colime and much less so than in luteolus; underparts white;
ears pale fuscous; tail fuscous above, slightly paler beneath; front
feet buffy white; hind feet grayish white, washed with hair-brown;
ankles dusky.

Skulli—Apparently similar to that of luteolus, but smaller; bulle
similar; rostrum short and broad; nasals longer than premaxill and
narrowed to a point at posterior end; zygomata standing out rather
squarely; molars without accessory enamel loops.

Measurements.—Type: Total length, 182; tail vertebre, 100; hind
foot, 19. Skull: (ee table, p. 81).

Remarks.—This species, although living at a high altitude and
somewhat resembling in color one of the members of the chrysopsis
group, nevertheless seems to be most nearly related to R. rufescens
luteolus, which occupies the lower foothill country below the range of
allen. The skull of allent resembles that of luteolus in having small
bullz and long nasals narrowed posteriorly. Externally allen is
paler than luteolus and has white underparts. Additional material
may show it to be a subspecies of the latter. The species is named
for Dr. J. A. Allen, in recognition of his extensive work on the genus.

Specumen examined.—One, the type.

REITHRODONTOMYS COLIMA COLIMA: Merriam.

Comma Votcano Harvest Mouse.
(Plat fies 5: Pl. ‘Vi, fig. 5)
Reithrodontomys colimx Merriam, Proc. Wash. Acad. Sci., IIT, 1901, p. 551.

Type locality Sierra Nevada de Colima, Jalisco, Mexico (altitude,
12,000 feet).

Disiribution.—Known only from the type locality.

Characters.—Externally similar to R.¢. chrysopsis, with long, woolly
pelage, but smaller and paler; skull similar to that of BR. c. tolucz, but
smaller and with simple enamel pattern.

1Type and only specimen badly broken; shape of braincase not known.

60 NORTH AMERICAN FAUNA. [No. 36.

Color.—Upperparts dull ochraceous-buff (slightly paler than in
chrysopsis), sparingly mixed with black on the back, the median
dorsal band only faintly indicated; sides paler but without lateral
line of buff; ears fuscous; underparts pinkish buff; fore feet buffy
white, washed with brownish; hind feet hair-brown, becoming white
toward the toes; tail pale fuscous above, grayish white beneath.

Skull——Similar in shape to that of 2. c. toluce, but slightly smaller,
with less inflated braincase; rostrum a little shorter and broader;
zygomata standing out a little more squarely anteriorly; nasals
long, ending on plane of premaxille; interpterygoid fossa rather
wide (as in chrysopsis); bulle intermediate in size between those of
chrysopsis and toluce; palatal foramina short and widely open;
enamel pattern of upper molars simple, with only a trace of a sub-
sidiary enamel loop.

Measurements.—Average of 2 specimens from type locality: Total
leneth, 165.5; tail vertebrae, 90; hind foot, 20. Skull: (See table,
DySl):

Remarks.—This species resembles externally the members of the
chrysopsis group, but the absence of sudsidiary loops on the upper
molars makes it necessary to place it in the rufescens group in the
typical subgenus. It is apparently a connectant species between
the two subgenera.

Specumens eramined.—Two, from type locality.

REITHRODONTOMYS COLIMA NERTERUS Merriam.
Corma Harvest Mouse.

Reithrodontomys colimx nerterus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 551.

Type locality.—Sierra Nevada de Colima, Jalisco, Mexico (altitude,
6,500 feet).

Disiribution.—Known only from the type locality.

Characters.—Similar to colimx, but ochraceous color more intense,
pelage shorter, and tail longer; skull slightly smaller.

Color.—Upperparts deep ochraceous-buff, nearly pure on sides,
mixed with black on median dorsal area; underparts heavily washed
with light ochraceous-buff; ears fuscous, shading to fuscous-black;
tail indistinctly bicolor, pale fuscous above, dusky drab below; feet
buffy white, shaded with dusky; ankles dark hair-brown.

Skull.—Slightly smaller than that of colume but of essentially similar
shape; nasals considerably shorter; interpterygoid fossa relatively
wide; enamel pattern of upper molars simple. Compared with
luteolus: Braincase narrower and higher; rostrum slenderer; nasals
broader at posterior end; bulle smaller but more inflated (rounder) ;
palatal foramina shorter. Compared with R. r. rufescens: Braincase
higher; rostrum and nasals shorter; bulle more inflated.

1914.] REITHRODONTOMYS RUFESCENS GROUP. 61

Measuremenis.—Type (@ ad.): Total length, 190; tail vertebrae,
110; hind foot, 20; ear from notch, 14.5; topotype (¢ ad.): 182;
99; 21. Skull: (See table, p. 81). |

Remarks.—This form closely resembles &. rufescens luteolus, both
in color and in the character of the pelage, which is shorter and less
wooly than in R. colume columx, which lives at higher altitudes.
Compared with luteolus the present form is a slightly paler shade of
ochraceous-buff. Its skull characters indicate close relationship to
colime and necessitate its separation from luteolus, but additional
material may serve to connect the two groups, making luteolus a sub-
species of colume.

Specimens examined.—Two, from type locality.

REITHRODONTOMYS DORSALIS Merriam.
BLACK-BACKED Harvest Movsz.
CREOLE figs 4D 5 figs 4)
Reithrodontomys dorsalis Merriam, Proc. Wash. Acad. Sci, III, 1901, p. 557.

Type locality.—Calel, Guatemala.

Disiribution.—Highlands of Chiapas and Guatemala.

Characters—Smaller than &. rufescens luteolus, with blacker back
and paler sides; skull with higher braincase.

Color.—Fresh pelage (January): Upperparts ochraceous-buff,
mixed with black, with a well-defined median band or stripe of black;
sides varying from ochraceous-buff to pinkish cinnamon, sometimes
with a rather pronounced lateral line next to the belly; underparts
washed with light ochraceous-buff; ears fuscous-black, darker than
in luteolus; tail bicolor, fuscous above, grayish white beneath; hind
feet grayish white; front feet buffy with a dusky stripe reaching half-
way to the toes. Worn pelage: General tone redder (pinkish cinna-
mon to pale tawny); black dorsal area much less clearly defined.

Skull.—Resembling that of R. r. rufescens in general shape, but
averaging smaller, with larger bulle. Compared with R. r. luteolus:
Smaller; braincase decidedly more inflated; interpterygoid fossa
much narrower. Compared with saturatus: Larger, with heavier
rostrum and more widely expanded zygomata. ‘The upper molars
in a majority of the specimens examined have a simple enamel pat-
tern, but some specimens show an incomplete subsidiary loop and
well-developed accessory tubercles on the outer border, thus com-
pletely bridging the gap between the two subgenera.

Measurements.—Average of 10 specimens (adult and subadult) from
type locality: Total length,167.5 (160-175); tail vertebrz, 90 (82-96) ;
hind foot, 19.3 (19-20). Skull: (ee table, p. 81).

Remarks.—This species, although clearly belonging to the rufescens
group, seems not to intergrade with luteolus, its nearest neighbor on

62 NORTH AMERICAN FAUNA. [No. 36.

the north. The low country on the Isthmus of Tehuantepec appar-
ently is an effective barrier separating the ranges of these two moun-
tain-loving species. |
The species exhibits considerable individual and seasonal variation
incolor. Specimens in worn pelage very much resemble similar speci-
mens of rufescens, but the ears and tail are paler.
Specomens examuned.—Total number, 154, from the following
localities:
Guatemala: Calel, 23; Hacienda Chancol, 29; Jacaltenango, 1; Todos Santos,
10; Volcan Santa Maria, 4; Zunil, 3.
Chiapas: Canjob, 2; Comitan, 27; Pinabete, 8; San Cristobal, 5; Tenejapa, 3;
Teopisca, 6; Tumbala, 33.

REITHRODONTOMYS AUSTRALIS AUSTRALIS Allen.
Trazu Harvest Mouse.
GEE TW es 3. Pl Vi aoa)

Reithrodontomys australis Allen, Bull. Am. Mus. Nat. Hist., VII, 1895, p. 328.
Reithrodontomys australis vulcanius Bangs, Bull. Mus. Comp. Zool., XX XIX, 1902,
p. 38 (Volcan de Chiriqui, Panama).
Type locality.—Volcan de Irazu, Costa Rica.
Mstribution.—Mountains of Costa Rica and western Panama.
Characters.—Similar to &. dorsalis, but slightly smaller, with
smaller ears; tail less sharply bicolor; ochraceous colors slightly
deeper and black dorsal band less clearly defined.
Color.—Upperparts ochraceous-buff, varying to pale orange-
cinnamon, rather heavily mixed on back with black; median dorsal
band usually very indistinct; underparts washed with light pinkish
cinnamon; ears pale fuscous to fuscous-black; tail fuscous above,
grayish ate beneath; feet buffy white, shaded with hair-brown;
wrists and ankles hei heoae.
Shghtly smaller than that of dorsalis, with flatter brain-
case; bulle rather flat, averaging smaller than in dorsalis; ascend-
ing arms of premaxille projecting a little beyond end of nasals;
upper molars simple, without accessory tubercles.
Measuremenis.—Type: Total length, 169 (160-179); tail verte-
bre, 84.5 (82-92); hind foot, 18.5 (18-19). Skull: (See table, p. 81).
Remarks—This species belongs in the rufescens group, being
apparently the most southerly ranging member of the group and of
the subgenus. It closely resembles dorsalis both in color and cranial
characters, and additional material from Central America may make
it necessary to unite the two as a single species.
The type of “‘vulcanius” from Panama has been examined and
found to agree closely with topotypes of australis in the Biological
Survey Collection.

1914.] SUBGENUS APORODON. 63

Specimens examined.—Total number, 18, from the following locali-
ties:
Costa Rica: Volcan de Irazu, 17.
_ Panama: Volcan de-Chiriqui, 1.}

REITHRODONTOMYS AUSTRALIS MODESTUS Thomas.
Nicaragua Harvest Mouss.
Reithrodontomys modestus, Thomas, Ann. Mag. Nat. Hist., Ser. 7, XX, 1907, p. 163.

Type locality —Jinotega, Nicaragua.

Distribution.—Known only from vicinity of type locality.

Characters.—Closely similar to australis, but underparts whiter and
tail darker.

Color.—Upperparts mixed black and dull ochraceous-buff, with
a fairly well-defined median band of blackish; underparts grayish
white, rarely with a faint tinge of light ochraceous-buff and a pectoral
spot of the latter color; tail distinctly bicolor, except at the tip,
fuscous above, soiled whitish below; ears fuscous; hind feet grayish
white; ankles dusky; front feet buffy, with a dusky stripe.

Skull.Not seen. Apparently slightly smaller than that of aus-
tralis, with smaller audital bulle.

Measurements.—Type: “‘Head and body, 59; tail, 70; hind foot
(eu), 16.5; ear, 12.5.’ Skull: (Gee table, p. 81).

Remarks.—This form appears to be very closely related to aus-
tralis, but the material at hand is too scanty to show clearly its
characters. The color description above is from specimens taken by
Mr. William B. Richardson at San Rafael del Norte and kindly
loaned by Dr. J. A. Allen of the American Museum of Natural His-
tory. These differ from Thomas’s description of the type in having
ochraceous instead of ‘‘drabby”’ sides and pectoral spot, and tail
distinctly whiter beneath. The Richardson specimens are without
complete skulls, so a study of the cranial characters has not been
possible. Mr. W. H. Osgood, who compared the type specimen with
material from the Biological Survey Collection, notes that its skull
is slightly smaller than that of australis, the premaxille shorter, and
the audital bullz somewhat smaller.

Specumens examined:

Nicaragua: San Rafael del Norte, 8.?
Subgenus APORODON nobis.

Type.—Reithrodontumys tenuirostris Merriam.

Subgeneric characters —Upper molars with subsidiary enamel loops
in the outer primary reéntrant angles, these loops in most species
reaching the outer border of the tooth and appearing, when viewed

1 Collection Mus. Comp. Zool. 2 Collection Am. Mus. Nat. Hist.

64 NORTH AMERICAN FAUNA, [No. 36.

in profile, as prominent accessory tubercles; in other species (chrys-
opsis group) the enamel loops sometimes do not reach the outer
border of the tooth and the accessory tubercles are often absent or
much reduced.

Remarks.—The subgenus Aporodon, while nee shane set off from
Reithrodontomys by any constant external characters, differs so
widely in the molar pattern as described above that its segregation
seems desirable.

It includes several well-marked groups, some of which possess
striking characters, both cranial and external, while other species
resemble the typical subgenus in all but the tooth characters.

The tenuirostris-microdon group (including also creper) and the
mexicanus group (including milleri, séderstrémi, and gracilis) show the
greatest amount of differentiation from typical Reithrodontomys.
These agree in having the outer wall of the anteorbital foramen rela-
tively narrow (usually much narrower than width of interpterygoid
fossa), broad interpterygoid fossa, short palatal foramina, unicolor
tail, and dense pelage (rather woolly in most species) of a uniform
tawny or ochraceous color, without pronounced darker grizzling. In
all species in these groups the subsidiary enamel loops of the upper
molars are well-developed and in unworn specimens appear as promi-
nent tubercles.

The chrysopsis group approaches the typical subgenus in having
the outer wall of anteorbital foramen broader, interpterygoid fossa
narrower and palatal foramina longer. The subsidiary enamel loops
of the upper molars are always well developed, but are usually (except
in perotensis) not continuous to the outer edge of the tooth and the
accessory tubercles are absent or much reduced. ‘The pelage is long
and full, and somewhat’more silky than in tenwirostris; the upper-
parts are more or less varied with black, and the tail is bicolor.

R. levipes and R. hirsutus seem to be aberrant members of the
subgenus, agreeing in the character of the pelage with the members
of the typical subgenus, but having the subsidiary enamel loops of
the upper molars well developed.

REITHRODONTOMYS LEVIPES GROUP.
REITHRODONTOMYS LEVIPES Merriam.
SAN SEBASTIAN Harvest Mouse.
(PI. TIT Mig) 2: eRe tie! 2: PI Vil fesesmes)
Reithrodontomys levipes Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 554.

Type locality.—San Sebastian, Jalisco, Mexico.

Distribution Known only from the type locality.

Characters —About the size of R. fulvescens toltecus; color more
decidedly tawny than any members of the fulvescens group; skull
relatively short and broad.

1914.] REITHRODONTOMYS LEVIPES GROUP. 65

Color.—Upperparts ochraceous-salmon, sparingly mixed on back
with black; general tone of sides between ochraceous-salmon and
ochraceous-orange; underparts strongly suffused with light pinkish
cinnamon; fore and hind feet grayish white, sometimes tinged with
the color of the sides; ears pale fuscous, with ochraceous hairs on
inner surface; tail pale fuscous above, grayish white below. Com-
pared with toltecus, the ochraceous color is more intense, there is
much less darkening on the middle of the back, and the underparts
are more intensely buffy.

Skull.—About the size of that of &. f. toltecus but shorter and
broader; braincase rather flat; zygomata narrowed anteriorly; nasals
short; ascending arms of premaxille extending about 1 mm. beyond
end of nasals; audital bulle very small; interpterygoid fossa broad;
palatal foramina short and widely open. First and second upper
molars with accessory enamel loops in primary reéntrant angles.

Measurements.—Average of 3 adults from type locality: Total
length, 190 (188-192); tail vertebre, 110.5 (110-111); hind foot,
20.8 (20.5-21). Skull: (See table, p. 81).

Remarks.—This species is remarkable not only for its peculiar
characters but because of its (seemingly) restricted range. It resem-
bles somewhat in color and character of pelage certain members of
the fulvescens group, but its skull characters place it in the subgenus
Aporodon. It seems to be most nearly related to the much larger
hirsutus, known only from Ameca, Jalisco.

At the type locality of levipes occurs another much smaller species—
R. fulvescens tenuis—a member of the typical subgenus. In the series
referred to this species are three specimens which combine in a
remarkable manner the characters of the two species. Externally
they differ very little from lewipes except in being somewhat less
intensely ochraceous, both above and below, and in having rather
shorter tails (103 and104mm.). Theirskulls, however, are decidedly
narrower than those of levipes and about intermediate in size between
skulls of the latter and of tenuis. The upper molars have small acces-
sory tubercles present in the principal angles, but the enamel pattern
is practically the same as in tenuwas. Another much smaller (adult)
skull in the series shows a strong tendency to develop the accessory
enamel loop characteristic of the subgenus Aporodon. This skull
(No. 88056, U. S. Nat. Mus.) is smaller than those of typical tenuis.
Anomalous as this situation may be, there seems to be no other expla-
nation than that these specimens are hybrids between levipes and
tenuis.

Specumens examined.—Three, from type locality.

28657°—14—_

66 NORTH AMERICAN FAUNA. [No. 36.

REITHRODONTOMYS HIRSUTUS Merriam.
Giant Harvest Movs.
(Plg i figs di yRiWV Gate, go)
Reithrodontomys hirsutus Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 553.

Type locality—Ameca, Jalisco, Mexico.

Distribution.—Known only from the type locality.

Characters.—Size very large; colors much as in KR. f. tenuis, paler
than in £. levipes.

Color—Upperparts ochraceous-buff, brightest on sides, sparingly
mixed on the back with blackish brown; underparts grayish white,
usually with a distinct tinge of light buff; fore feet whitish, washed
with light buff; hind feet grayish white, tinged with dusky; ankles
fuscous; tail pale fuscous above, grayish white below; ears brownish
drab.

Skull.—Large and robust with a well-defined supra-orbital bead
which extends back to the parietals; braincase rather flat and some-
what narrowed posteriorly; zygomata contracted anteriorly; rostrum
and nasals short; ascending arms of premaxille extending back of
end of nasals; audital bulle very small; palatal foramina short and
wide; upper molars with subsidiary enamel loops.

Measuremenis.—Average of 5 adults from type locality: Total
length, 211 (203-233); tail vertebree, 127 (122-143); hind foot, 21.8
(21-22). Skull: (See table, p. 81).

Remarks.—This species is one of the largest in the genus, being
exceeded in size of skull only by R. tenuirostris, a widely different |
species. So far as known, it has no near relative except dewpes, and
from that it differs both in size and color. It agrees with it, how-
ever, in every important cranial character—heavy rostrum, short
and wide palatal foramina, broad interpterygoid fossa, and small
bulle. |

In color, hirsutus bears a remarkable resemblance to &. f. tenuas,
which inhabits the same region, but is hardly more than half the size
of the present species.

Specimens examined.—Six, from type locality.

REITHRODONTOMYS CHRYSOPSIS GROUP.
REITHRODONTOMYS CHRYSOPSIS CHRYSOPSIS Merriam.
Voutcano Harvest Movwse.

(Pl. Ill, fig 3; BIS Vi fe: 35 PE Vil eso

Reithrodontomys chrysopsis Merriam, Proc. Biol. Soc. Wash., XIII, 1900, p. 152.
Type locality —Mount Popocatepetl, Mexico (altitude 11,500 feet).

1914.] REITHRODONTOMYS CHRYSOPSIS GROUP. 67

Distribution.—High mountains around the valley of Mexico; Mount
Patamban and Mount Tancitaro in Michoacan. Altitudinal range
from 9,000 feet to 13,500 feet (timber line).

Characters —Size large (almost equaling &. hirsutus); tail long;
pelage very long, soft, and silky; ears black or blackish; tail bicolor;
skull with rounded and much inflated braincase; bulle large.

Color.—Upperparts a rich shade of ochraceous, between ochraceous-
buff and orange-buff, rather heavily mixed on back with black, the
latter color usually forming an indistinct median band; general tone
of sides near pinkish cinnamon; ears fuscous-black; tail sharply
bicolor, fuscous above, grayish white beneath; feet grayish white;
ankles fuscous; underparts strongly suffused with light pinkish cin-
namon.

Skull.—Of large size (considerably exceeding that of R. r. rufescens);
braincase subglobular, usually much inflated, sometimes moderately

R‘C. CHRYSOPSIS
f “@ TOLUCAE
“<C. ORIZABAE
“« PEROTENSIS
“© LEVIPES

“ HIRSUTUS
Fic. 5.—Distribution of Reithrodontomys chrysopsis, R. perotensis, R.levipes, R. hirsutus, and subspecies.
flattened; zygomata slender, strongly contracted anteriorly; outer
wall of anteorbital foramen broad; rostrum slender, narrowing
gradually to the tip; audital bulls very large and moderately inflated;
interpterygoid fossa relatively narrow (compared with R. levipes);
palatal foramina long (as in rufescens); upper molars with sub-
sidiary enamel loops in primary angles, but these usually not reaching
outer border of tooth; accessory tubercles absent or much reduced.

Measurements.—Type: Total length, 194; tail vertebre, 108; hind
foot, 21. Average of 5 specimens from Ajusco, Salazar, and Huit-
vilac: Total length, 185 (177-188); tail vertebre, 106 (98-111);
hind foot, 20.8 (20-21). Skull: (See table, p. 81).

Remarks.—This handsome and striking species is an inhabitant of
the upper slopes of the mountains about the valley of Mexico and on
more or less isolated mountains in western Michoacan. It is the
first-described and best-known member of a subalpine group of
species found on most of the higher mountains of southern Mexico

68 NORTH AMERICAN FAUNA. [No. 36. -

and is apparently the only form having an extensive range. Speci-
mens from Mount Tancitaro and Patamban show no characters to
distinguish them from the typical form. From R. f. toltecus, which
occupies the valley of Mexico, chrysopsis differs in larger size, richer
colors, longer pelage, and blacker ears, as well as in cranial char-
acters.

On Mount Popocatepetl, Nelson and Goldman found this species
occurring sparingly in grassy places in the open forest from the base
of the mountain up to 12,000 feet. On Mount Patamban it was most
numerous at about 11,000 feet in the upper part of the Canadian
Zone, a few ranging to the extreme summit above 12,000 feet. In the
upper part of its range it was found among grass and brush.

Specumens ecamined.—Total number, 25, from the following locali-
ties in Mexico:

Mexico: Ajusco (11,000 feet), 3; Mount Iztaccihuatl (13,500 feet), 1; Mount
Popocatepetl (11,500 feet), 3; Salazar (9,000 feet), 1.

Morelos: Huitzilac (10,000 feet), 1.
Michoacan: Mount Tancitaro (10,000-12,000 feet), 7; Patamban (11,000 feet), 9.

REITHRODONTOMYS CHRYSOPSIS TOLUCH Merriam.
ToLucaA Harvest Mousse.
Reithrodontomys chrysopsis toluce Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 549.

Type locality.—North slope of Volean Toluca, Mexico (altitude,
11,500 feet).

Distribution.—Known only from the type locality.

Characters.—Similar to chrysopsis, but slightly smaller and much |
less intensely ochraceous; skull slenderer.

Color—Worn pelage: Upperparts mixed black and ochraceous-
tawny with a well-defined black median band, as in chrysopsis, but
lacking almost entirely the bright golden color shown by the latter;
ears fuscous; front feet buffy white, with a band of hair-brown
reaching half way to the toes; hind feet soiled whitish, tinged with
dusky; ankles fuscous; ring around eye blackish; underparts grayish
with scarcely a trace of buff; tail hair-brown above, grayish white
below.

Skull.—Slightly smaller than that of chrysopsis; braincase nar-
rower; rostrum slenderer, much narrowed at the tip; bulle much
smaller; subsidiary enamel loops of upper molars low and slightly
developed; accessory tubercles absent.

Measurements.—Type: Total length, 180; tail vertebrze, 98; hind
foot, 21. Skull: (See table, p. 81).

Remarks.—This form, known from only a single specimen, has
rather pronounced characters. On account of the worn condition
of the type specimen, the colors can not be satisfactorily described.
Specimens in fresh pelage will doubtless prove less different from

1914.] REITHRODONTOMYS CHRYSOPSIS GROUP. 69

chrysopsis than is this worn individual, but the underparts apparently
are distinctly whiter.
Specumen examvned.—One, the type.

REITHRODONTOMYS CHRYSOPSIS ORIZABZ Merriam.
Mount OrizABA Harvest Mouse.
Reithrodontomys orizabe Merriam, Proc. Wash. Acad. Sci., ITI, 1901, p. 550.

Type locality—Mount Orizaba, Puebla, Mexico (altitude, 9,500
feet).

Distribution.—Known only from the type locality.

Characters._-Externally the same as chrysopsis; skull narrower and
bull smaller.

Color.—As in chrysopsis.

Skull.i—Apparently smaller than that of chrysopsis, but length of
skull and shape of zygomata not known; braincase narrower, audital
bullz decidly smaller; molar series and nasals shorter; subsidiary
enamel loops of m? well developed, of m+ present but shorter. Com-
pared with toluce: Braincase and bulle similar; rostrum broader.

Measurements.—Type: Total length, 182; tail vertebrae, 105; hind
foot, 20. Skull: (See table, p. 81).

Remarks.—This form so closely resembles chrysopsis in color that
it seems best to consider it a subspecies of the latter; in cranial
characters it more nearly resembles toluce. Additional material is
necessary to determine its true relationship.

Specumen examined.—One, the type.

REITHRODONTOMYS PEROTENSIS Merriam.
PreroteE Harvest Mouse.
Reithrodontomys perotensis Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 550.

Type locality.—Cofre de Perote, Vera Cruz, Mexico (altitude, 9,500
feet).

Distribution.—Known only from the type locality.

Characters.—Externally similar to chrysopsis, but color of upper-
parts less intensely ochraceous; skull smaller with much smaller
bullee.

Color.—Upperparts cinnamon with strong tinge of ochraceous,
extensively mixed with black and with a distinct median band of
the latter color; ears fuscous, somewhat paler than in chrysopsis;
underparts strongly washed with light pinkish cinnamon; tail pale
fuscous above, slightly whitened beneath.

Skull.—Smaller than that of R. c. chrysopsis, with flatter braincase
and smaller bullae; zygomata squarely spreading anteriorly, nearly

1 The type and only known specimen is badly broken.

70 NORTH AMERICAN FAUNA. [No. 36. .

parallel to axis of skull; subsidiary enamel loops in upper molars well
developed, reaching panes border of tooth.

Measuremenis.—Type: Total length, 176; tail vertebre, 119; hind
foot, 19. Skull: (ee table, p. 81).

Remanka Dine species Aftes widely in cranial characters from the
other members of the chrysopsis group. Indeed, its skull somewhat
resembles that of ff. megalotis saturatus (which occupies the low
country around the base of the Cofre de Perote), especially in the
widely spreading zygomata and in the shape and size of the audital
bulle. It differs from saturatus, however, in having a much broader
and flatter braincase, longer a eign dees rostrum, buffy instead of
white underparts, and in dental characters.

Specumen examuned.—One, the type.

REITHRODONTOMYS MEXICANUS GROUP.
REITHRODONTOMYS MEXICANUS MEXICANUS (De Saussure).
SaussuRE Harvest Movusz.

(PIOTIT, figs 45° Pl. V1, fig. 4; Pl Vil, ies. 45 9ssO ae)
Reithrodon mexicanus De Saussure, Rev. Mag. Zool., 2d ser., XII, 1860, p. 109.
Ochetodon mexicanus Coues, Proc. Acad. Nat. Sci. Phila., 1874, p. 186.
Reithrodontomys costaricensis jalape Merriam, Proc. Wash. Acad. Sci., III, 1901,
p. 552 (Jalapa, Vera Cruz).
Reithrodontomys cherriet jalapx Miller, Bull. 79, U.S. Nat. Mus., 1912, p. 180.

Type locality Mountains of Vera Cruz, Mexico.
Mstribution.—From Jalapa, Vera Cruz, south to Chiapas and
Guatemala. |

Characters.—Size rather large (about like #. f. toltecus); tail long,
concolor; color uniform tawny above, white below; hind feet, ears,
and tail Fon skull short, broad, and flat.

Color. —Upperparts varying from tawny to pinkish cinnamon, spar-
ingly darkened on the back with blackish hairs; general tone of back
varying from cinnamon-brown to Prout’s brown; an indistinct black-
ish ring around eye; tail nearly unicolor, fuscous or clove-brown to
fuscous-black, in some specimens slightly paler beneath; upper sur-
faces of both fore and hind feet dark hair-brown; toes usually buffy
or grayish white; ears fuscous or fuscous-black; is of nose, upper
lips, and litderparts white.

Skull.—Short and relatively broad, with very short, broad rostrum
and short nasals; braincase squarish, rather flat, depressed poste-
riorly; zygomata slender, contracted anteriorly; nasals ending behind
rather squarely, on a line with ends of premaxille or slightly anterior
to them; outer wall of anteorbital foramen narrow; interpterygoid
fossa broad; palatal foramina short, ending at or slightly in front of
plane of first molars; audital bulle small.

1914.] REITHRODONTOMYS MEXICANUS GROUP. val

Measurements.—Adult male (Jalapa, Vera Cruz): Total length, 197;
tail vertebre, 119; hind foot, 21. Average of 2 adults from Toton-
tepec, Oaxaca: 193; 117; 21. Average of 3 adults from Jacalte-
nango, Guatemala, and Tumbala, Chiapas: 194; 117; 20.2. Skull:
(See table, p. 81).

Remarks.—This species, although described over 50 years ago, is
still imperfectly known, and the name mezicanus has been misapplied
by all authors since De Saussure. Three distinct species of Reithro-
dontomys occur in the mountains of Vera Cruz and a fourth (R.
megalotis saturatus) not far away on the table-land. These (using the
modern names) are Rf. rufescens rufescens, R. fulvescens difficilis, and
R. “cherrva jalapex.’”*

The original description? is so complete and agrees so perfectly with
the last of these that there appears to be not the slightest doubt that
the species now known as “jalapx”’ should be referred to mexicanus.
The combination of tawny upperparts, white belly, dark feet, and
unicolor tail is possessed by no other species in this region.

Through the kindness of M. Maurice Bedot, director of the Geneva
Museum, who has furnished the Biological Survey with photographs
and measurements of the type skull, I am able to present additional
evidence corroborative of the above decision. The photographs
- clearly show the relatively narrow outer wall of the anteorbital fora-
men and the broad interpterygoid fossa, characteristic of the sub-
genus Aporodon, and the measurements agree closely with those of
specimens of “‘jalapx’’ from Jalapa, Vera Cruz.

The type of &. mexicanus, with the skull inside the skin, was bor-
rowed in 1890 from the Geneva Museum and examined in Washington
by Drs. Allen, Merriam, and True. Comparison was made with a
specimen ® (No. 7007a, U.S. Nat. Mus.) from Tehuacan, Puebla, at that
time practically the only available specimen from southern Mexico.
Dr. Allen in referring this specimen to mexicanus stated that it agreed
with the type.* Dr. Merriam’s notes, however, taken at the same

time, indicate some important differences. These are as follows:
No. £19. This specimen is the type of De Saussure’s description. Measure-
ments.—Hind foot, 18.5. Ear from crown, 10; from anterior base, 13. The upper-

1 All of these have been taken at Jalapa, Vera Cruz.

2 Following is a translation of the pertinent portions:

“‘The size of this animal is nearly exacily that of the European field-mouse (Mus silvaticus), though its
forms are somewhat more thick set. * * * Tail very long, its length exceeding that of the body and
head. The color of the pelage is tawny brown, on the sides becoming absolutely tawny, or even orange-
tawny. The tawny color becomes fainter toward the line where it comes in contact with the white of the
belly. The lips, lower part of cheeks, chin, throat, and entire underparts are almost pure white, here and
there slightly washed with a tawny tint, especially on the breast and throat. * * * The hairs are slate
gray, the tips only passing into russet, or white. The ears are brown * * *. The fore feet are white,
except above, as far as the root of the digits, where they aregray. The tail is blackish, scaly, unicolor, and
covered with rather dark gray hairs; it is especially hairy toward the tip; at its base the hairs are scanty
and very short; but they become longer toward the tip.”

8 Referred by the writer to R&. fulvescens difficilis.

4 Bull. Am. Mus. Nat. Hist., VII, 1895, p. 136.

a2 NORTH AMERICAN FAUNA. [No. 36. .

parts are so thoroughly suffused with fulvous that it can almost be called a
‘‘red mouse.’’ The upper surface of the hind foot is dark brown to base of
toes. The toes are whitish. The skull has never been removed from the skin
There is a skin of this species or a closely related subspecies in the United States
National Museum (No. 7007a) collected at Tehuacan, Puebla, Mexico, by Sumi-
chrast. Upperparts rusty fulvous, but not quite so deep or bright as in the mounted
specimen [the type]. The only departure of importance from the type is the color
of the upper surface of the hind foot, which is soiled whitish instead of dark brown,
and the color of the underside of the tail, which is whitish instead of being concolor
with the upper surface.

These differences—whiter feet and bicolor tail—are sufficient to
show that the Tehuacan specimen can not be referred to mexicanus.
No mention is made, either by Allen or Merriam, of the color of the
underparts, but taking into consideration the soiled and faded condi-
tion of the Tehuacan specimen (and probably, also, of the type) this
omission is not strange.

On the evidence of this determination the name mexicanus was used
by Allen for the dark form of the fulvescens group later described
from Orizaba by Merriam as R. diffictlis, but, as shown above, it prop-
erly applies to the species now under consideration—a member of the
subgenus Aporodon.

The species exhibits considerable individual variationin color. One
specimen from Totontepec, Oaxaca, is fairly typical, while another
from the same place is considerably darker, with darker ears and tail.
A specimen from Jacaltenango, Guatemala, is considerably paler
than Vera Cruz specimens, but agrees with them in other characters.

Intergradation with goldmani on the north and with cherrw in Central
America seems fairly certain to be established.

Specimens examined.—Total number, 10, from the following locali-
ties in Mexico and Guatemala:

Vera Cruz: Jalapa, 2.1

Oaxaca: Totontepec, 2.

Chiapas: Comitan, 1; Tenejapa, 1; Tumbala, 3.
Guatemala: Jacaltenango, 1.

REITHRODONTOMYS MEXICANUS GOLDMANI Merriam. -
GoLtpmMan Harvest Mouse.
(PE Tete GPloV 1s fig.163)
Reithrodontomys goldmani Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 552.

Type locality —Metlaltoyuca, Puebla, Mexico (altitude 800 feet).
Distribution.—Known only from northern Puebla.
Characters.—Similar to mexicanus, but smaller and paler.
Color.—Upperparts pinkish cinnamon, faintly darkened on top of
head and back with brownish hairs; general tone of back about
snuff-brown; sides of nose, upper lips, and underparts white, the

1 One in Collection Am. Mus. Nat. Hist.

1914.] REITHRODONTOMYS MEXICANUS GROUP. 73

latter with a faint tinge of pale buff; ears dark hair-brown; tail
fuscous, nearly unicolor or slightly paler beneath; fore and hind feet
grayish white, tinged with dusky; ankles hair-brown.

Skull—tLonger and relatively narrower than that of mexicanus;
braincase flattened, and narrowed posteriorly; rostrum and nasals
longer, the latter narrowed to a point posteriorly; audital bull
slightly larger; palatal foramina wide.

Measurements.—Type: Total length, 190; tail vertebre, 109; hind
foot, 21.5. Skull: (See table, p. 81).

Remarks.—This subspecies is the most northerly ranging member
of the group. It occupies the low, arid, coast region of northern
Puebla and probably adjacent States. It is distinctly paler than
mexicanus and the skull of the type shows pronounced characters,

NY

nee

4%,

oI

Livia
2,

M. MEXICANUS
M. GOLDMANI
M. CHERRI/
GRACILIS

FIG. 6.—Distribution of Reithrodontomys mexicanus, R. gracilis, and subspecies.

some of which, however, may not prove to be constant. Two speci-
mens from Huauchinango, Puebla (altitude 5,000 feet) are inter-
mediate between goldmani and mezicanus, the skulls being nearer to
the latter form, the skins nearer goldmani.
Specimens examined.—Total number, 3, from the following local-
ities in Mexico:
Puebla: Huauchinango, 2; Metlaltoyuca, 1.

REITHRODONTOMYS MEXICANUS CHERRII (Allen).
CENTRAL AMERICAN Harvest Mouse.
GPIe TIT fie’ 5; PIS Viv fies)
Hesperomys ( Vesperimus) cherrii Allen, Bull. Am. Mus. Nat. Hist., III, 1891, p. 211.
Sitomys cherriet Allen, Ibid., V, 1893, p. 238.

Reithrodontomys costaricensis Allen, Ibid., VII, 1895, p. 139 (La Carpintera, Costa Rica).
Reithrodontomys cherriei Osgood, Proc. Biol. Soc. Wash., XX, 1907, p. 50 (type fixed).

14. WORTH AMERICAN FAUNA. [No. 36.

Type locality.—San. Jose, Costa Rica.

Distribution.—Costa Rica and Chiriqui, Panama.

Characters.—Similar to mexicanus, but colors brighter (more tawny,
less blackish); ears paler; skull larger.

Color.—Upperparts varying from bright tawny to hazel, more or
less mixed with black on dorsal area; ears dusky hair-brown, scantily
haired; underparts white (rarely washed with yellowish buff); tail
fuscous, unicolor, clothed with short, bristly hairs; fore and hind feet
pale sepia, broadly edged with whitish; toes buffy white.

Skull—Larger than that of mexicanus, with longer rostrum; brain-
case more inflated.

Measuremenits.—Average of 7 adults from Costa Rica (San Pedro
and La Carpintera): Total length, 191 (182-198); tail vertebra, 111
(103-123); hind foot, 20 (19.5-20.5). Average of 13 adults from
Boquete, Chiriqui: Total length, 207 (192-227); tail vertebre, 126
(115-140); hind foot, 20 (19-22). Skull: (See table, p. 81).

Remarks.—Through an unfortunate mixing of skulls this species
was originally described as a Hesperomys |[=Peromyscus] and a few
years later, before the mistake was discovered, redescribed as
Reithrodontomys ‘ costaricensis,’’ by which name it has usually been
known. Osgood has shown the pertinence of the original description
to the present species and has selected a type specimen from the
original series.+

The species is one of the larger members of the genus, though not so
large as R.t. tenwirostris and R. hirsutus. Intergradation with mezi-
canus seems probable, though not perfectly shown by the material
at hand. A specimen from Jacaltenango, Guatemala, is somewhat
intermediate in characters, but apparently nearest to meaxicanus.
Specimens from Boquete, Chiriqui, have somewhat longer tails than
the series from Costa Rica. A single specimen from Nicaragua is
much less tawny than the type series, somewhat resembling the Gua-
temala specimen.

Specimens examined.—Total number, 21, from the following locali-
ties:

Costa Rica: La Carpintera, 3;? San Jose, 2;2 San Pedro, 3.?
Panama: Boquete, Chiriqui, 13.3

REITHRODONTOMYS MILLERI Allen.
CoLoMBIAN Harvest Mouse.
Reithrodontomys millert Allen, Bull. Am. Mus. Nat. Hist., XX XI, 1912, p. 77.

Type locality —Munchique, Cauca, Colombia (altitude, 8,325 feet).
Distribution.—Known only from Colombia (altitude 6,000 to 10,300
feet).

1 Proc. Biol. Soc. Wash. XX, 1907, p. 50. 8 Eleven in Collection Mus. Comp. Zool.
2 Collection Am. Mus. Nat. Hist.

1914.] REITHRODONTOMYS MEXICANUS GROUP. 15

Characters.—Paler and less tawny than R. m. cherrii; tail shorter.

Color.—Upperparts varying from light ochraceous-salmon, exten-
sively mixed with brownish to nearly pure tawny, with slight admix-
ture of brown; lateral line only faintly indicated; underparts white,
sometimes with a faint tinge of light buff; ears and tail dark hair-
brown, the latter almost unicolor, but slightly paler beneath; feet
hair-brown, sometimes edged with grayish white.

Skull—vVery similar to that of R. m. cherrii, with short, heavy
rostrum, broad frontals, and short palata! foramina.

Measurements.—Ten adults: Total length, 181 (169-190); tail
vertebrae, 107 (98-116); hind foot, 19 (18-20). Skull: (ee table,
p- 81).

Remarks.—This species is rather closely related to R. m. cherrn,
from which it differs in browner, less tawny coloration and shorter
tail. No intermediate specimens are known, but further collecting in
Panama may result in securing such. 3

Specimens examined.—Total number, 7, from the following locali-
ties :!

Colombia: Cocal, 1; El Roble (7,000 feet), 5; La Guneta, 1; Munchique, 5; San
Augustin, Huila (10,300 feet), 1.

REITHRODONTOMYS SODERSTROMI Thomas.
Ecuapor Harvest Movuss.

Reithrodontomys séderstr6mi Thomas, Ann. Mag. Nat. Hist., ser. 7, I, 1898, p. 451.

Type locality. Quito, Ecuador.

Distribution.—Known only from the type locality and from Valle
de las Papas, southern Colombia.

Characters — Apparently in the cherrit group and closely related
to miller; colors less tawny above and more buffy or fawn-colored
below; hind foot, ear, and skull larger.

Color.—* Dull grayish fawn, not nearly so rufous as in costaricensis
[cherrit]. Brighter lateral line little developed. Under surface not
sharply defined, its color much less bright than in costaricensis, the
tips of the hairs more or less buffy or fawn-colored. Hars thinly
haired, brown, little darker than the general color. Hands and feet
white, without darker markings on the metapodials. Tail pale brown
above and below, the tip white.’’? A series of 4 specimens from Valle
de las Papas, central Andes, Huila, Colombia, may be described as
follows: Upperparts mixed blackish brown and ochraceous-salmon;
ears clove-brown; tail dark mummy-brown; hind feet soiled whitish,
with a broad band of mummy-brown reaching nearly to the toes;

1 All in Collection Am. Mus. Nat. Hist.
2 Thomas, loc. cit.

® Compared by Dr. J. A. Alien and Mr. Oldfield Thomas with the type and 12 topotypes in the British
Museum and said to agree perfectly with them.

76 NORTH AMERICAN FAUNA. [No. 36.

front feet similar, but with less brown; underparts white, strongly
tinged with pale ochraceous-salmon. With regard to the color of
the hind feet in the typical series, Mr. Thomas writes me as follows:

Of 20 specimens, 12 have large metapodial patches [of brown], 7 have smaller ones
down to quite minute ones (often not the same on both sides) and the type has prac-
tically none at all—though even there a few hairs are dark on each foot, making a
minute spot.

Skull.—Similar to that of miller, but larger. ‘‘Low, with a flat
superior profile. Palatal foramina short, barely reaching backward
to the level of thefront of m!.”’! More recently Mr. Thomas has com-
pared the topotype series with specimens of R. m. cherrw and states :?
“‘T can find no skull difference of any importance or constancy.”

Measurements.—Type (from dry skin): ‘‘ Head and body, 72; tail
(tip doubtfully perfect), 83; hind foot (wet) without claws, 19; ear
(wet), 14. Skull: Back of parietal to nasal tip, 21.1; greatest breadth,
11.5; nasals 8.9 x 2.6; interorbital breadth, 3.7; palate from hense-
lion, 9.2; diastema, 5.9; palatal foramina, 4.1 x 1.6; upper molar
series, 3.8.’% Skull of fully adult topotype (measured by Mr.
Thomas): Greatest length 24.2; breadth of braincase, 11.5; length
of nasals, 9.1; width of outer wall of anteorbital foramen, 1.9.

Remarks.—This species, so far as known, is the most southerly
ranging member of the genus. I have not been able to examine the
type series of 20 specimens in the British Museum, but I have seen a
small series from southern Colombia, kindly loaned by Dr. Allen,
and which both he and Mr. Thomas consider typical of this species.
From an examination of this series it is clear that sdderstrémi is a
member of the mexicanus group and rather closely related to miller,
but whether it is connected with the latter by intermediate forms
can not at present be determined.

Its habits are described by Mr. L. Séderstrém, the collector of the
type series, as follows: “‘Feeds on flowers and seeds in the gardens.
Comes out from among the climbing plants every evening at about
py mae?

Specumens examined.—Four, from Valle de las Papas, Huila, Co-
lombia (central Andes, altitude 10,000 feet).*

REITHRODONTOMYS GRACILIS Allen & Chapman.
Yucatan Harvest Movuse.
CPM tie: ofits Vil fig g7.3)

Reithrodontomys mezxicanus gracilis Allen and Chapman, Bull. Am. Mus. Nat. Hist.,
IX, 1897, p. 9.

Type locality.—Chichen Itza, Yucatan, Mexico.
Distribution.— Yucatan and Campeche.

1 Original description by Thomas. 3 Thomas, loc. cit.
2 In epistle, Oct. 21, 1913. 4 Collection Am. Mus. Nat. Hist.

1914.] REITHRODONTOMYS MEXICANUS GROUP. Tl

Characters.—Size small; similar in color to R. m. goldmam, but
brighter; skull much smaller with narrow braincase.

Oolor.—Upperparts pinkish cinnamon, sparingly lined on dorsal
surface with blackish brown; color brightest on sides next to the
belly, sometimes forming an indistinct lateral line (as in the fulvescens
group); ears dark hair-brown; tail fuscous, slightly paler beneath;
fore feet buffy white; hind feet grayish white; ankles fuscous;
underparts white, sometimes with a slight yellowish cast; less white
on sides of nose than in mezicanus and goldmani.

Skull.—Decidedly smaller than that of either mexicanus or gold-
mani; similar in size and general proportions to that of difficilis, but
differing in the characters of the subgenus; rostrum short and broad;
braincase narrow and moderately flat; nasals short; zygomata nearly
parallel to axis of skull (much less contracted anteriorly than in
mexicanus); palatal foramina very short; bulle rather small.

The outer wall of the anteorbital foramen, although much nar-
rower than in skulls of R. fulvescens difficilis, is relatively broader
than in the other members of the subgenus and may slightly exceed
the width of the interpterygoid fossa. ‘The subsidiary enamel loops
are present on the upper molars, but the accessory tubercles are
rather low.

Measuremenits.—Type (immature): Total length, 165; tail verte-
bre, 98; hind foot, 16. Average of 2 topotypes (immature): 169;
100; 18. A somewhat older (subadult) specimen from Yohaltun,
Campeche: 191; 113; 20. Skull: (See table, p. 81).

Remarks.—This is the smallest member of the mezxicanus group.
Externally it bears a striking resemblance to R. fulvescens tenuis,
differing chiefly in darker and more nearly unicolor tail and more
uniform ochraceous color (less grizzled with black) on the upper-
parts. In skull characters the two differ widely, gracilis clearly be-
longing in the subgenus Aporodon.

From R. m. mexicanus, its nearest relative, it differs in much
smaller size, paler coloration, and narrower skull. Judging from the
very limited material at hand, it seems to be specifically distinct
from the other members of the group. One specimen from Yohaltun,
Campeche, differs from the type series in being somewhat darker,
with blacker, more nearly unicolor tail.

Specimens examined.—Total number, 9, from the following locali-
ties in Mexico:

Yucatan: Chichen Itza, 6;! Progreso, 1.
Campeche: Apazote, 1; Yohaltun, 1.

1 Four in Collection Am. Mus. Nat. Hist.

73 NORTH AMERICAN FAUNA. [No. 36.

REITHBODONTOMYS TENUIROSTRIS GROUP.
REITHRODONTOMYS TENUIROSTRIS TENUIROSTRIS Merriam.
Wooty Harvest Mouse.

Reithrodontomys tenuirostris Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 547.

Type locality—Todos Santos, Guatemala (altitude 10,000 feet).

Distribution —Known only from the type locality.

Characters Size very large (equaling Fk. hirsutus, except in
length of tail); pelage long, soft, and wooly; color rich tawny; tail
unicolor; skull with much swollen braincase and narrow rostrum.

Color —Adults: Upperparts deep tawny, darkest on the dorsal
area where the color becomes hazel; an indistinct blackish ring around
eye; underparts light pinkish cinnamon; ears and tailfuscous; ankles
and hind feet clove-brown; tips of toes whitish; upper surface of fore
feet hair-brown; toes buffy. Young: Upperparts Prout’s brown, with
a tinge of tawny; underparts paler than in the adult; toes buffy white.

Skull——Braincase broad, much inflated, depressed posteriorly;
anterior portion of frontals abruptly depressed, forming a shallow
sulcus at posterior end of nasals; rostrum long and narrow; nasals
narrowed to a point posteriorly, ending on a line with premaxille;
zygomata slender, slightly contracted anteriorly; palatal foramina
relatively short, not reaching plane of first molars; interpterygoid
fossa broad (about as wide as outer wall of anteorbital foramen) ;
bulle small and rather flat.

Measurements —Type (¢ ad.): Total length, 210; tail vertebre,
124; hind foot, 23. Skull: (See table, p. 81).

Remarks.—This is one of the largest and most striking members
of the genus. Resembling R. m. mexicanus in the color of the upper-
parts, it is readily distinguished byits cinnamon belly. Its skull is
so peculiar that it is placed in a different group from the latter.

Specimens examined.—Two, from type locality.

REITHRODONTOMYS TENUIROSTRIS AUREUS Merriam.
CaLEL Harvest Mouse.
(PI, 10s fig, 83 PliVi fie. 8: Ply VIL fea)
Reithrodontomys tenuirostris aureus Merriam, Proc. Wash. Acad. Sei., III, 1901, p. 548.

Type locality —Calel, Guatemala (altitude 10,200 feet).

Distribution—Known only from the type locality.

Characters —Slightly smaller and paler than tenwirostris.

Oolor—Adulis: Upperparts ochraceous-tawny moderately dark-
ened on dorsal area with black; underparts light ochraceous-bufi
(a little yellower than in tenuirostris); ears fuscous-black; tail fus-
cous, slightly paler beneath; fore feet dark hair-brown; hind feet
clove-brown, both edged with white; toes whitish. Young: Upper-

1914.] | REITHRODONTOMYS TENUIROSTRIS GROUP. 79

parts Prout’s brown, faintly tinged with ochraceous; underparts
white.

Skull—Slightly smaller than that of tenutrostris, with flatter brain-
case and more inflated bull; palatal foramina longer and slightly
broader, extending behind plane of first molars.

Measuremenis.—Adult female: Total length, 196; tail vertebree, 112
hind foot, 22.5. Skull: (See table, p. 81).

does zenuirosiris and only a short distance away. With the very

limited amount of material at hand, it is impossible to say whether

the characters distinguishing the two forms will prove to be constant.
Specimens examined.—Two, from type locality.

REITHRODONTOMYS CREPER Bangs.
Cuirieur Harvest Mouse.
Reithrodontomys creper Bangs, Bull. Mus. Comp. Zool., XX XIX, 1902, p. 39.

Type locality.— Volcan de Chiriqui, Panama (altitude, 11,000 feet).

Distribution.—Known only from the type locality.

Characters.—Size large; colors very dark (similar to R. a. australis
above, but darker beneath); skull similar to that of R. t. tenuirostris,
but Persie

Color..—Type: Upperparts mummy-brown, varying on sides to
Mars brown; sides of face blackish; underparts russet; ears dark
sepia; tail dusky—almost clove-brown—all around, except terminal
fourth which is whitish; feet whitish, palest on toes.

Skull.—Similar in shape to that of R. ¢. tenwirostris, but smaller;
braincase narrowed posteriorly; palatal foramina expanded in the
middle, narrowing anteriorly; zygomata much contracted anteriorly.
Compared with #. ¢. aureus: Skull slightly shorter and much nar-
rower; bullz smaller.

Measurements.—Type (? ad.): Total length, 215; tail vertebre,
130; hind foot, 23. Skull: Gee table, p. 81).

Remarks! ies very distinct species is known only from a single
specimen, “caught on the cold barren summit of the Volcan de
Chiriqui’.? It seems to be nearest related to A. tenutrostris, from
Guatemala.

Specumen examined.—One, the type.®

1Comparisons with Ridgway’s *‘ Nomenclature of Colors’’ (1886).
2 Bangs, loc. cit.
2 Collection Mus. Comp. Zool.

80 NORTH AMERICAN FAUNA. [No. 36.

REITHRODONTOMYS MICRODON MICRODON Merriam.

SMALL-TOOTHED Harvest Mouse.
(Pl. III, fig. 9; Pl. VI, fig. 9.)

Reithrodontomys microdon Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 548.
Type locality—Todos Santos, Guatemala.
Distribution.—Highlands of Guatemala; limits unknown.
Characters.—Closely similar in color to R. t. tenuirostris, but very

much smaller.

Color.—Adults: Upperparts tawny, brighter on the sides next to
the belly, darkened on back and head with black, where the tone
becomes hazel; blackish ring around eye; underparts light pinkish
cinnamon; ears and tail fuscous, the latter slightly paler beneath;
hind feet dark hair-brown edged with whitish; toes white; fore feet
buffy white with a patch of dusky. Subadults: Upperparts browner
(about Prout’s brown) and less mixed with tawny; underparts white.

Skull_—Similar in general shape to that of R. t. tenuirostris, but
very much smaller; braincase moderately inflated, narrowed and
depressed posteriorly; zygomata slender, decidedly contracted ante-
riorly; rostrum narrow; palatal foramina ending on plane of front
molars; interpterygoid fossa very broad; bulle rather large and
inflated; outer wall of anteorbital foramen very narrow.

Measuremenis.—Type (¢ ad.): Total length, 185; tail vertebre,
113; hind foot, 21. Topotype (¢ subadult): 180; 112; 21. Skull:
(See table, p. 81).

Remarks.—This little species is in every way a miniature of the large
yenutrostris, which lives in the same region.

Specimens examined.—Total number, 3, from the following locali-
ties:

Guatemala: Todos Santos, 2; Volcan Santa Maria 1.

REITHRODONTOMYS MICRODON ALBILABRIS Merriam.
WHITE-LIPPED Harvest Mouse.

Reithrodontomys microdon albilabris Merriam, Proc. Wash. Acad. Sci., III, 1901, p. 549.

Type locality—Cerro San Felipe, Oaxaca, Mexico (altitude, 10,000
feet).

Distribution.—Known only from the type locality.

Characters.—Similar to microdon, but paler; underparts white;
closely similar in color to R. mexicanus goldmani.

Color.—Upperparts pinkish cinnamon, darkened on dorsal area with
black, where the tone becomes snuff-brown; blackish ring around eye;
ears, tail, and hind feet fuscous, the last edged with whitish; toes
whitish; fore feet buffy, with a dusky patch; underparts white.

Skull_—Very similar to that of microdon; braincase slightly broader
and more inflated; interpterygoid fossa narrower; bulle slightly
smaller; nasals ending on a line with premaxille; zygomata decidedly

1914.]

CRANIAL MEASUREMENTS.

31

contracted anteriorly; palatal foramina short, not reaching plane of

molars.

Measuremenis.—Type (¢ ad.): Total length, 187; tail vertebre,
117; hind foot, 20. Skull: (See table below).
Remarks. ap his form is closely related to microdon, differing chiefly

in paler coloration.

Specimen examined.—One, the type.

Average cranial measurements of Reithrodontomys.

Species.

humulis humulis....------
humulisimpiger....-------
humulis merriami....-----
albescens albescens ..-..-----
. albescens griseus ..-.-------
SILGIEEILUIS al ses yee <2
. megalotis megalotis...------
. megalotis aztecus...-.------
. megalotis dychei....-------
. megalotis nigrescens...-----
. megalotis longicaudus .-.---
. megalotis peninsulz....-.-
. megalotis Cinereus...-.-----
. megalotis saturatus......-.-
. megalotis aliicolus.....-----
. megalotis arizonensis......-
. megalotis zacatece....-.-.-

bay Pd Bo Ey Ed Ed EB by by By yb Bs DB

R. fulvescens toltccus.........-
R. fulvescens helvolus.....--.-
R. fulvescens chiapensis.......
R. fulvescens nelsoni.........-
R. fulvescens mustelinus.....-
DIOS at eens esos cna

TUSESCENS TUSESCENS..-.-----
rufescens luteolus.......---
ele eee ale te

pry RRA

RR. chrysopsis chrysopsis....-.-
R. chrysopsis tolucz........--
F. chrysopsis orizabz.....--.-
Pe ENOLETESES orae So ik Ss
FR. mericanus mezicanus ..-.-.
R. merxicanus goldmani.....-.
R. mexicanus cherrii.......-.-

R. tenuirostris tenwuirostris.....
R. tenuirostris aureus..-...-.-
ik CIEE Re oe eee
R. microdon microdon...-...-

R. microdon albilabris.........

Number averaged.

Localities.

Greatest length.

Breadth of brain-
case.

Length of nasals,

of anteorbital for-

Width of outer wall
amen,

i | | | |

North Carolina; South Carolina; Georgia. -.| 19.4

West Virginia: White Sulphur Springs ORS 19.4
Texas: Austin Bayou; Louisiana: Lafayette.) 19

Nebraska; South Dakar... Sue melas 19.5
Texas: San Antonio, San Angelo.......---- 19.3
Colorado: San Luis Valley TO AEL Te ae hn Relate IX 20

New Mexico (southern); Chihuahua. ....... 20. 7
New Mexico: La Plata, Aztec, Guadalupita.| 22.1
Kansas: Lawrence, Onaga...........-.----- | 20.6
Idaho: Payette, Weiser, Nampa.........-.- 20.8

California: Vic cinity of San Francisco Bay--| 20.2

Colimas; Colima fe pee oan ae Su eee

OaxacacsRielormiane eee so ie ee mune ae
Jalisco: Sierra Nevada de Colima..........-
Vera Cruz: Jalapa, Jico.........------------
Oaxacas Jug UUAs se hse eee a eee

Lower California: San Quintin, Rosario. .-.--- 20.7
PucplasChalehicomulassc: jess asec eee 21.6
Mera Cruz das ivicasan tes fbi cose ne ye a 22.5
Oaxaca: Cerro San’ Felipe: : 2.0622. 22 22. 4
Arizona: Chiricahua Mountains...........- 20. 4
Zacatecas: Valparaiso Mountains..........-. 20.5
Queretaro: Pinal de Amoles.............-..- 19.8
California: Catalina Island...............-.-- 21.9
California: San Francisco Bay.........-.-.- 21.2
California: Petaluma (marshes)............- 20. 8
Sonora: Oposura, Providencia Mines. -.-..-- 21.6
Sinaloa: Mazatlan, Sinaloa, Escuinapa..... 21.9
Texas: Brownsville; TPamawlipases Messe 21.9
Texas and Louisiana........--.---.----.--- 22.2
Mera CruzsOrizabas Jalapa- 28.25.5258: 21.8
Mexico: Tlalpam, TD) Dea ee SRR 23.4
OamacatO axa cael weakest vee ae tes Sees 22. 4
Chig nas: C any OD esse peer whens es ec 22.4

2

3

5

8

6

9

bw mwre by bo
She sco sauoes eo

Oaxaca: Mountains near Ozolotepec.....-.-|....--|.-.-.-.

J Bisco: Sierra Nevada de Colima..........- aS A
Ginter: 0 aU aT NER 0a TNE Ee Ne 23
Costa RicasVolean raz Ws 252 se ae 2250
INicaraguaAsdinotesales ass oes ee as 21
JaliscouSaniSebastzanes Mee oe ee eee PPE
AISCO ANN ECA eRe ie Sate a eae Ne 24,9
Mexico (State) and Morelos.....---.--.----- 24.3
Mexico: Violean@olucavsss = 222s 55g ee 24
iPueplias Mount: Onzabpar soe one ee a eee (eae
Vera Cruz: Cofre de Perote......---.---.--- Fa
Vera Cruz; Oaxaca; Guatemala......-.--..- 23.1
Puebla: Metlalioyuca.........-.--.------:- 24
Costa Rica: La Carpintera, San Pedro....-- 2351
Colombia: Munchique..-....-.-...--------- 22.

2,
BE CURd ORO Wit OMe ncn gas acer eee eee 24.
Yucatan: Chichen Itza; Campeche: Yohal- | 21.
5
5)
o:

tun.
Guatemala: Todos Santos. ....-...-5-....-- 2b:
Guatemala Calel. jyevse sel! wey Sha ee 25:
Panama: Volcan de Chiriqui..............- 2
vata: Todos Santos, Volcan Santa | 22.3
aria
Oaxaca: Cerro Sam Melipesasa. 25 -s5-eene- 22.3

28657°—14——_6

a

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EE 2 2 8 2 ee 2 5 ae 6S SS 52
D NT OVO BD CO HAD 09 C1 CO CO CH RCI DD COOH OO HW ODD OD

ee

Kr OTH CO CLV OVW NO CIM COM DI OVOUST OR A OO CHO CODD WO PP

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eee eo ee cre ee OO ea an Tioe nee foe) a aS) SS ETSI OD) GOO OST 20) 00/0900 SSI5S OO) S31 0051! 99:00) 00 20/5100 51 OO ST SST) S22

CO CNOCOM | Ort ROT BOO OCOD OR IOC

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MOWATODOWNNO OFM NMR ON FIT

MN NNN ER ENNENNNNEN EVN NEN NN NNNEREENNNENENNNNENNEE
Th OOOCOHI NOH NEE ONO HHO

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reat 15

12.

PLATE If.
[Twice natural size.]

Reithrodontomys humulis humulis. Raleigh, N. C. (No. 189304, U. 8. Nat.
Mus., Merriam Coll.)

. Reithrodontomys humulis merriami. Austin Bayou, Tex. (No. 44849, U.S.

Nat. Mus., Biological Survey Coll.)

. Retthrodontomys albescens griseus. Type, San Antonio, Tex. (No. 87852,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys albescens albescens. Type, Kennedy, Nebr. (No. 116358,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys montanus. Medano Ranch, Costilla County, Colo. (No.

150714, U. S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys megalotis megalotis. Dry Creek, Socorro County, N. Mex.

(No. 158319, U. S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys megalotis dychet. Onaga, Kans. (No. 147132, U. S. Nat.

Mus., Biological Survey Coll.)

. Reithrodontomys megalotis saturatus. Las Vigas, Vera Cruz. (No. 54434,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys raviventris raviventris. Berkeley, Cal. (No. 93208, U. S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys raviventris halicetes. Type, Petaluma, Cal. (No. 7146;

Mus. Vert. Zool., Univ. California. )

. Reithrodontomys megalotis zacatece. Valparaiso Mountains, Zacatecas. (No.

91912, U. S. Nat. Mus., Biological Survey Coll.)
Reithrodontomys megalotis alticolus. Type, Cerro San Felipe, Oaxaca. (No.
68392, U. S. Nat. Mus., Biological Survey Coll.)

82

North American Fauna No. 36, U.S. Dept. Agr. Biological Survey.

5 18s
Den
Gs >
v

_R

.h. humulis.

. h. merriami.

. a. griseus.

. a. albescens,

SKULLS OF REITHRODONTOMYS.

. R. montanus.
6. R. m. megalotis.
7. R.m. dychei.
8. R.m. saturatus,

RR
5 dae
R
R

PLATE lI.

. Ir. raviventris.
. vr. haliecetes
.m. zacatece.
. m. alticolus.,

Fig.

(7)

Or

10.

PLATE Te

[Twice natural size.]

. Reithrodontomys amenus. Type, Reforma, Oaxaca. (No. 14064, Field Mus.

Nat. Hist.)

. Reithrodontomys otus. Type, Sierra Nevada de Colima, Jalisco. No. 45705,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys australis australis. Volcan Irazu, Costa Rica. (No. 116614,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys dorsalis. Type, Calel, Guatemala. (No. 77009, U. S. Nat.

Mus., Biological Survey Coll.)

. Reithrodoniomys colime colime. Type, Sierra Nevada de Colima, Jalisco.

(No. 45706, U. S. Nat. Mus., Biological Survey Coll.)

. Reithrodoniomys rufescens luteolus. Type, Juquila, Oaxaca. (No. 71558,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens tenuis. Mazatlan, Sinaloa. (No. 96713, U. S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens dificilis. Orizaba, Vera Cruz. (No. 58268, U. S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens helvolus. Type, Oaxaca, Oaxaca. (No. 68387,

U.S. Nat. Mus., Biological Survey Coll.)
Reithrodontomys fulvescens toltecus. Type, Tlalpam, D. F., Mexico. (No.
50746, U. S. Nat. Mus., Biological Survey Coll.) i

84

SKULLS OF REITHRODONTOMYS.

. i. difficilis.
. f. helvolus.
. ft. toltecus.

. R. ec. ecolime.
R. r. luteolus.
. f. tenuis.

. amocenus.
. otus. 5B.
. a. australis. 5 LR
. dorsalis.

we ee Saran | ~ ¥ * hs

PLATE III.

[Twice natural size.]}

. 1. Reithrodontomys hirsutus. Type, Ameca, Jalisco. (No. 82200, U. 8S. Nat.

Mus., Biological Survey Coll.)
2. Reithrodontomys levipes. Type, San Sebastian, Jalisco. (No. 88057, U. S.
Nat. Mus., Biological Survey Coll.)
3. Reithrodontomys chrysopsis chrysopsis. Type, Mount Popocatepetl, Mexico.
(No. 52031, U. S. Nat. Mus., Biological Survey Coll.)
4. Reithrodontomys mexicanus mexicanus. Jalapa, Vera Cruz. (No. 108538,
U. 8. Nat. Mus., Biological Survey Coll.)
5. Reithrodontomys mexicanus cherrii. La Carpintera, Costa Rica. (No. 7898,
Am. Mus. Nat. Hist.)
6. Reithrodontomys mexicanus goldmani. Type, Metlaltoyuca, Puebla. (No.
93096, U. 8. Nat. Mus., Biological Survey Coll.)
7) Reithrodontomys gracilis. Chichen Itza, Yucatan. (No. 108143, U.S. Nat.
Mus., Biological Survey Coll.) |
8. Reithrodontomys tenuirostris aureus. Type, Calel, Guatemala. (No. 76939,
U.S. Nat. Mus., Biological Survey Coll.)
9. Reithrodontomys microdon microdon. Type, Todos Santos, Guatemala. (No.
76923, U. 8. Nat. Mus., Biological Survey Coll.)

86

North American Fauna No, 36, U.S. Dept. Agr. Biological Survey.

SKULLS OF REITHRODONTOMYS,

1. R. hirsutus. 4. R. m. mexicanus, 7. R. gracilis.
2. R. levipe 5. R. m. cherrii. 8. R. t. aureus.
3. R. c. chry 31 6. R. m. goldmani. 9. R. m. microdon.

ns:

wey iaiaesba rete ; :
AV. (ty He aty i j ars aren: :

Tat

be: bre 4 e
> ; 7
S a 4
« AY A

Fig. 1.

PLATE IY.

[Twice natural size.]

Reithrodontomys humulis humulis. Raleigh, N. C. (No. 189304, U. S. Nat.
Mus., Merriam Coll.)

. Reithrodontomys humulis merriami. Austin Bayou, Tex. (No. 44849, U.S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys albescens griseus. Type, San Antonio, Tex. (No. 87852,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys albescens albescens. Type, Kennedy, Nebr. (No. 116358,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys montanus. Medano Ranch, Costilla County, Colo. (No.

150714, U. 8. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys megalotis megalotis. Dry Creek, Socorro County, N. Mex.

(No. 158319, U. 8S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys megalotis dychei. Onaga, Kans. (No. 147132, U. S. Nat.

Mus., Biological Survey Coll.)

. Retthrodontomys megalotis saturatus. Las Vigas, Vera Cruz. (No. 54434,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys raviventris raviventris. Berkeley, Cal. (No. 93208, U. S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys raviventris halicetes. Type, Petaluma, Cal. (No. 7146,

Mus. Vert. Zool., Univ. California.)

. Reithrodontomys megalotis zacatece. Valparaiso Mountains, Zacatecas. (No.

91912, U. S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys megalotis alticolus. Type, Cerro San Felipe, Oaxaca. (No.

68392, U. S. Nat. Mus., Biological Survey Coll.)

88

Biological Survey. PLATE IV.

North American Fauna No. 36, U. S. Dept. Agr.

OF REITHRODONTOMYS.
2. ©. raviventris.
. Tr. hbaliecetes.
v. M. Zacatece.
4. m. alticolus.

.h. humulis. ». R. montanus.

. h. merriami. 3. R. m. megalotis.
2. a. griseus. . R. m. dychei.

. a. albescens. . R. m. saturatus,

fre
patsy ay
>

.

Fig. 1.

10.

PLATE V.
[Twice natural size.]

Reithrodontomys amenus. Type, Reforma, Oaxaca. (No. 14064, Field Mus.
Nat. Hist.)

. Reithrodontomys otus. Type, Sierra Nevada de Colima, Jalisco. (No. 45705,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys australis. Volcan Irazu, Costa Rica. (No. 116614, U. S.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys dorsalis. Type, Calel, Guatemala. (No. 77009, U. S. Nat.

Mus., Biological Survey Coll.)

; appears colimx colimx. Type, Sierra Nevada de Colima, Jalisco.

(No. 45706, U. 8. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys rufescens Theos Type, Juquila, Oaxaca. (No. 71558,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens tenuis. Mazatlan, Sinaloa. (No. 96713, U.S. Nat.

Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens difficilis. Orizaba, Vera Cruz. (No. 58268, U. 8.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys fulvescens helvolus. Type, Oaxaca, Oaxaca. (No. 68387,

U.S. Nat. Mus., Biological Survey Coll.)
Reithrodontomys fulvescens toltecus. Type, Tlalpam, D. F., Mexico. (No.
50746, U. S. Nat. Mus., Biological Survey Coll.)

90

North American Fauna No. 36, U.S. Dept. Agr. Biological Survey. PLATE V

SKULLS OF REITHRODONTOMYS.

. amcenus, 5. R. ec. colime. . f. difficilis.
. otus. 6. R.r. lutolus. . f. helvolus.
. australis. 7. R. f. tenuis. . f. toltecus.

. dorsalis.

Fie. 1.

is PLATE VI.
[Twice natural size.]

Reithrodoniomys hirsutus. Type, Ameca, Jalisco. (No. 82200, U. S. Nat.
Mus., Biological Survey Coll.)

. Reithrodontomys levipes. Type, San Sebastian, Jalisco. (No. 88057, U. 8.

Nat. Mus., Biological Survey Coll.)

. Reithrodontomys chrysopsis chrysopsis. Type, Mount Popocatepetl, Mexico.

(No. 52031, U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys mexicanus mexicanus. Jalapa, Vera Cruz. (No. 108538,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys mexicanus cherrvi. La Carpintera, Costa Rica. (No. 7898,

Am. Mus. Nat. Hist.)

. Reithrodontomys mexicanus goldmani. Type, Metlaltoyuca, Puebla. (No.

93096, U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys gracilis. Chichen Itza, Yucatan. (No. 108143, U.S. Nat.

Mus., Biological Survey Coll.)

. Reithrodontomys tenwirosiris aureus. Type, Calel, Guatemala. (No. 76939,

U.S. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys microdon microdon. Type, Todos Santos, Guatemala. (No.

76923, U.S. Nat. Mus., Biological Survey Coll.)
92

North American Fauna No. 36, U. S. Dept. Agr.

Biological Survey, PLATE VI.

SKULLS OF REITHRODONTOMYS.
1. R. hirsutus. 4. R. m. mexicanus. 7. R. gracilis.
2. R. levipes. 5. R. m. cherrii. 8. R. t. aureus.
3. R. ec. chrysopsis. 6. R. m. goldmani. 9, R. m. microdon.

Fies. 1, 7.

3,5.

4,9, 10, il.

PLATE VII.
[Teeth about 10 times natural size; skulls twice natural size.]
Reithrodoniomys (Reithrodontomys) megalotis megalotis.
Fig. 1. Side view of upper molars. (No. 167353, U.S. Nat. Mus.,
Biological Survey Coll.)

Fig. 7. Worn crowns of upper molars. (No. 58090, U. S. Nat. Mus.,
Biological Survey Coll.)

. Reithrodontomys (Aporodon) tenuirostris aureus. Side view of upper

molars. (No. 77008, U.S. Nat. Mus., Biological Survey Coll.)
Reithrodontomys (Aporodon) levipes.
Fig. 3. Worn crowns of lower molars. (No. 88057, U. 8. Nat. Mus.,
Biological Survey Coll.)
Fig. 5. Worn crowns of upper molars. (No. 88057, U. 8. Nat. Mus.,
Biological Survey Coll.)
Reithrodontomys (Aporodon) mexicanus mexicanus.
Fig. 4. Worn crowns of upper molars. (No. 68684, U. S. Nat. Mus., —
Biological Survey Coll.)
Figs. 9,10, 11. Skull of type. (No. 253, Geneva Museum.)

. Reithrodontomys (Aporodon) chrysopsis chrysopsis. Worn crowns of upper

molars. (No. 125883, U. 8. Nat. Mus., Biological Survey Coll.)

. Reithrodontomys (Reithrodontomys) humulis merriami. Worn crowns cf

upper molars. (No. 178261, U. S. Nat. Mus., Biological Survey Coll.)

North American Fauna No. 36, U. S. Dept. Agr. Biological Survey.

PLATE VII.

MOLAR TEETH AND SKULLS OF REITHRODONTOMYS.

1. R. m. megalotis. 4, R.m. mexicanus. 7. R.m. megalotis.
2. R. t. aureus. 5. R. levipes. 8. R. h. merriami.
3. R. levipes. 6. R. ¢c. chrysopsis. 9, 10. 11. R. m. mexicanus.

INDEX.

[New names in bold=face type; synonyms in ifalics; pages containing the principal reference to a subject
in bold-faee figures.]

A. E.

Acknowledgments, 9-10. Economic status, habits and, 10.
albescens, Reithrodontomys albescens, 10, 22=23s

Reithrodontomys montanus, 22. FP.
albilabris, Reithrodontomys microdon, 80-81. Food, 11.
alleni, Reithrodontomys, 59. fulvescens, Reithrodontomys fulvescens, 43-44.
alticolus, Reithrodontomys megalotis, 13, 37. Reithrodontomys mezicanus, 43.
amoenus, Reithrodontomys, 55.

Fhithrodontomys, 55. G.
amoles, Reithrodontomys, 40. Generic characters, 13.
Aporodon, subgenus, 10, 13, 58, 63-64, 71, 72, 77. | Genus Reithrodontomys, 13.

subgenus, species, list of, 15. goldmani, Reithrodontomys mexicanus, 11, 72-73:

type localities, list of, 15. gracilis, Reithrodontomys, 64, 75-77.

arizonensis, Reithrodontomys megalotis, 38. Reithrodontomys mexicanus, 76.
Arvicola, 7. griseoflavus, Reithrodoniomys, 45, 46.
aurantius, Reithrodontomys fulvescens, 12, 48-49. | griseus, Reiihrodontomys, 30.

Reithrodontomys mexicanus, 48. Reithrodontomys albescens, 23-24.
aureus, Reithrodontomys tenuirostris, 78-79. Groups, genus Reithrodontomys, 14-16.

australis, Reithrodontomys australis, 62-63.

aztecus, Reithrodontomys megalotis, 8, 13, 30. me

Habits and economic status, 10.

~: haliccetes, Reithrodontomys raviventris, 42-43.
carolinensis, Reithrodon, 8. helvolus, Reithrodontomys fulvescens, 52-53.
cataline, Reithrodontomys, 40-41. Reithrodontomys griseofiavus, 52.
Reithrodontomys megalotis, 40. Hesperomys, 8,74.
Ehithrodontomys, 40. [Peromyscus], 74.
Characters, generic, 13. ( Vesperimus) cherrti, 73, 74.
cherriei, Reithrodontomys, 73. hirsutus, Reithrodontomys, 64, 66.
Sitomys, 73. History and material, 7.
cherrii, Hesperomys ( Vesperimus), 73, 74. humilis, Mus, 7, 19.
Reithrodontomys mexicanus, 73-74. Ochetodon, 8, 13, 19.
chiapensis, Reithrodontomys fulvescens, 53. Reithrodon, 8, 19.
chrysopsis, Reithrodontomys chrysopsis, 64, 66- Reithrodontomys, 19.
68. humulis, Mus, 7, 19.
chrysotus, Reithrodontomys, 48, 49. Reithrodontomys humulis, 13, 19-20.
cinereus, Reithrodontomys megalotis, 35-36. I
Reithrodontomys saturatus, 35. ;
colimz, Reithrodontomys, 39. impiger, Reithrodontomys humulis, 20-21.
Reithrodontomys colime, 59-60. Reithrodontomys lecontii, 20.
costaricensis, Reithrodontomys, 73, 74. inerspectatus, Rhithrodontomys, 51, 52.
Cranial measurements, table of, 81. intermedius, Reithrodontomys fulvescens, 11, 47-
creper, Reithrodontomys, 79. 48.
D Reithrodontomys mezicanus, 47.
deserti, Reithrodontomys, 9. ts
Reithrodontomys megalotis, 26, 28. jalapx, Reithrodontomys cherriei, 70.
dickinsoni, Reithrodontomys humilis, 19, 20. Reithrodontomys cherrii, 71.
difficilis, Reithrodontomys fulvescens, 50-51, 71. Reithrodontomys costaricensis, 70, 71.
Distribution, 10. K
dorsalis, Reithrodontomys, 61-62. ‘
dychei, Reithrodontomys, 23. Key to species and subspecies, 16.
Reithrodontomys megalotis, 11, 12, 30-32. klamathensis, Reithrodontomys, 33, 34.

95

96 NORTH AMERICAN FAUNA. [No. 36

L. Reithrodon—Continued.
lecontei, 19.
longicauda, 8, 33.
megalotis, 8, 13, 26.
meticanus, 8, 47, 50, 70.
montanus, 8, 24.
sumichrasti, 8, 50.
Reithrodontomys, genus, 8, 10, 13.
genus, groups, 14-16.
chrysopsis group, 15, 64, 66-70.
fulvescens group, 15, 43-56.
humulis group, 14, 19-24.
levipes group, 15, 64-66.
megalotis group, 14, 24-43.

laceyi, Reithrodontomys, 47, 48.
lecontei, Mus, 8.

Reithrodon, 19.
lecontii, Mus, 7, 13, 19.

Feithrodontomys, 19.
levipes, Reithrodontomys, 64-65.
longicauda, Ochetodon, 8, 13, 33.

Reithrodon, 8, 33.

Reithrodontomys, 33.

megalotis, 33.

longicaudus, Reithrodontomys megalotis, 33-34.
luteolus, Reithrodontomys rufescens, 57-58.

M. mexicanus group, 15, 64, 70-77.
; rufescens group, 15, 56-63.
Measurements, table of cranial, 81. tenuirostris group, 16, 64, 78-81.
megalotis, Reithrodon, 8, 13, 26. genus, key to species, 16.
Reithrodontomys megalotis, 8, 12, 13, 26-29. species, list of, 14.
merriami, Reithrodontomys humulis, 12, 21-22. type localities, list of, 14.
mexicanus, Ochetodon, 8, 13, 47, 50, 70. subgenus, 10, 19, 64.
Reithrodon, 8, 47, 50, 70. species, list of, 14.
Reithrodontomys, 50. type localities, list of, 14.
mexicanus, 9, 64, 70-72. Reithrodontomys albescens albescens, 10, 22-23.
microdon, Reithrodontomys microdon, 64, 80. albescens griseus, 23-24.
milleri, Reithrodontomys, 64, 74-75. alleni, 59.
modestus, Reithrodontomys australis, 9, 63. amcenus, 55.
montanus, Ochetodon, 8, 13, 24. amoles, 40.
Reithrodon, 8, 24. australis australis, 62-63.
Reithrodontomys, 8, 10, 13, 24-26. modestus, 9, 63.
Mus, 7, 8. vulcanius, 62.
humilis, 7, 19. cataline, 40-41.
humulis, 7, 19. cherriei, 73.
lecontei, 8. jalape, 70.
lecontii, 7, 13, 19. cherrii jalape, 71.
mustelinus, Reithrodontomys fulvescens, 54-55. chrysopsis chrysopsis, 64, 66-68.
N. orizabe, 69.
tolucee, 68-69.
nebrascensis, Reithrodontomys, 9. chrysotis, 48, 49.
Reithrodontomys dychei, 30, 31. colime, 39.
nelsoni, Reithrodontomys fulvescens, 53-54. colime, 59-60.
nerterus, Reithrodontomys colime, 60-61. nerterus, 60-61.
nigrescens, Reithrodontomys megalotis, 32-33. costaricensis, 73, 74.
O. jalapx, 70, 71.
creper, 79-
obscurus, Reithrodontomys megalotis, 39, 40. dorsalis, 61-62.
Ochetodon, 8, 13. deserti, 9.
humilis, 8,13, 19. dychei, 23.
longicauda, 8, 13, 33. nebrascensis, 30, 31.
merxicanus, 8, 13, 47, 50, 70. fulvescens aurantius, 12, 48-49.
montanus, 8, 13, 24. chiapensis, 53.
sumichrasti, 8, 13. difficilis, 50-51, 71.
orizabee, Reithrodontomys chrysopsis, 69. fulvescens, 43-44.
otus, Reithrodontomys, 55-56. helvolus, 52-53.
Reithrodontomys levipes, 55. intermedius, 11, 47-48.
Pp. mustelinus, 54-55.
nelsoni, 53-54.
pallidus, Reithrodontomys, 9, 33, 34. tenuis, 9, 45-47.
Pelages, 12. toltecus, 51-52.
peninsule, Reithrodontomys megalotis, 35. gracilis, 64, 76-77.
Fhithrodontomys, 35. griseoflavus, 45, 46.
perotensis, Reithrodontomys, 64, 69-70. helvolus, 52.
R. griseus, 30.
hirsutus, 64, 66.
raviventris, Reithrodontomys raviventris, 10, 41- humilis, 19.
AZ. dickinsoni, 19, 20.
keithrodon, 8, 13. humulis humulis, 13, 19-20.
carolinensis, 8, impiger, 20-21.

humilis, 8 , 19. merriami, 12, 21-22.

1914.]

Reithrodontomys—Continued.

klamathensis, 33, 34.
laceyi, 47, 48.
lecontii, 19.
impiger, 20.
levipes, 64-65.
otus, 55.
toltecus, 51.
longicauda, 33.
megalotis alticolus, 13, 37.
arizonensis, 38.
aztecus, 8, 13, 3G.
caialing, 40.
cinereus, 35-36.
deserti, 26, 28.
dychei, 11, 12, 20-32.
longicauda, 33.
longicaudus, 33-24.
megalotis, 8, 12, 138, 26-2.
nigrescens, 32-33.
obscurus, 32, 40.
peninsule, 25.
saturatus, 36-37, 71.
sestinensis, 26, 29.
zacatece, 39-40.
mezicanus, 50.
auranius, 48.
cherrii, 73-74.
Sulvescens, 43.
goldmani, 11, 72-72.
gracilis, 76.
intermedius, 47.
mexicanus, 9, 64, 70-72.
microdon albilabris, 80-81.
microdon, 64, 8O.
mnilleri, 64, 74-75.
montanus, 8, 10, 13, 24-26.
albescens, 22.
nebrascensis, 9.
otus, 55-56.
pallidus, $, 33, 34.
perotensis, 64, GS-7.
raviventris haliccetes, 42-43.
Taviveniris, 10, 41-42.

28687°—14——-7

INDEX. 97

Reithrodontomys—Continued.
rufescens luteclus, 57-58.
rufescens, 56-57, 71.
saturatus cinereus, 35.
soderstromi, 9, 10, 64, 75-76.
sumichrasti, 9.
tenuirostris aureus, 78-79.
tenuirostris, 13, 63, 64, 78-
Ehithrodontomys, 13 (footnote).
amenus, 55.
cataline, 40.
inexspectatus, 51, 52.
peninsula, 35.
rufescens, Reithrodontomys rufescens, 66-57, 71.

Ss.

saturatus, Reithrodontomys megalotis, 36-37, 71.
sestinensis, Reithrodontomys megalotis, 26, 29.
Sitemys cherriei, 73.
sdderstr6mi, Reithrodontomys, 9, 10, 64, 75-76.
Species and subspecies, key to, 16.

list of, 14.
Subgenus 4poredon, 63.

RKeithrodontomys, 19.
sumichrasti, Ochetodon, 8, 13.

Reithrodon, 8, 50.

RKeithrodontomys, 9.

Ae

tenuirostris, Reithredontomys tenuirostris, 13, 63,
4, 78.
tenuis, Reithrodontomys fulvescens, 9, 45-47.

| toltecus, Reithrodontomys fulvescens, 61-62,

Reithrodontomys levipes, 51.
toluce, Reithrodontomys chrysopsis, 68-69.
Type localities, list of, 14.

Vi

( Vesperimus) Eesperomys cherrii, 73, 74.
vulcanius, Reithrodontomys australis, 62.

Le
zacatece, Reithrodontomys megalotis, 39-40.

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- U.S. DEPARTMENT OF AGRICULTURE

BUREAU OF BIOLOGICAL SURVEY
HENRY W. HENSHAW, Chief

NORTH AMERICAN FAUNA |

No: 37

[Actual date of publication, April 7, 1915]

REVISION OF THE AMERICAN MARMOTS

BY

ARTHUR H. HOWELL

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY

; WASHINGTON
GOVERNMENT PRINTING OFFIOE

1915

| Survey

Ica

Biolog

North American Fauna No. 37, U. S. Dept. Agr.

[Omer ‘Wady ‘quoyy ‘oouor0,y ‘ayry wo]
(VHYOHdOSON SINLNAAIAV14 VLOWHVI]) LOWYVI] GSTLNVI-N3q07105)

peo DEPARTMENT OF AGRICULTURE

BUREAU OF BIOLOGICAL SURVEY
HENRY W. HENSHAW, Chief

NORTH AMERICAN FAUNA

INTO. & «

[Actual date of publication, April 7, 1915]

REVISION OF THE AMERICAN MARMOTS

BY

ARTHUR H. HOWELL

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY

WASHINGTON
GOVERNMENT PRINTING OFFIOE

1915

LETTER OF TRANSMITTAL.

ae ee

Unirep States DEPARTMENT OF AGRICULTURE,
Bureau or BIOLOGICAL SURVEY,

Washington, D. C., October 21, 1914.
Sir: | have the honor to transmit herewith for publication as
North American Fauna No. 37 a revision of the American marmots,

by Arthur H. Howell, assistant biologist of the Biological Survey.
Although marmots are found over most of the United States and
Canada, the relationships and ranges of the several species until now
have been very imperfectly known. The present report furnishes
descriptions of 26 forms and contains a series of maps showing their
geographic distribution. In many localities marmots are a decided
pest to agriculture, especially in the East, both because they are
destructive to crops and because their burrows seriously interfere

with farming operations. Moreover, marmots are known to carry |

the germs of Rocky Mountain spotted fever and other diseases. The
animals are hence of considerable economic importance.
Respecttully,
Henry W. HEnsHaw,
Chief, Biological Survey.
Hon. Davin F. Houston,

Secretary of Agriculture.
2

CONTENTS.

Page
Memeo IO WPI oe oi wo ss Sh oe dn See oe ee eee be sae ees oe 5
Pure te anoeMOUleNClALUTC - .. = 60s ed io ee ote be nck ei dee eens 5
Mena reMatpE aise ce eee ee et ek kk ee eB. di
ele? ee See a Sa AS torte wie S's Ha ee Hae se ie as ch was 8
ESTEISCRIEL 2 6 Soc 6 A Ce RE Se et es rem inane ee ean ieee en 9
SUS CICTGR TICS 505.5 Ao = ee ee er a Para oy ee en ap ey er 10
TSG Gs 5 Js Soe Ode a See eRe ee peor ee ee eh i are 12
POO Ge occ serge ea SSC ee a at Haan a i pe eMC ret 12
EXGPEGTTPNYS (SNS eae Ra ee et Pe nm eee rea 13
ME eelOrssiearelers Of Ciscase... 2 0i.--220- 2220. 9. oot oe 14
SSE PSL UE CLIC PS ifs i ao, aaa ere ye ree re nee i
EE 2S GING! TCL th 3 Gee CO ene er pe ate elie pen ae nate 16
Vie nist ate oe es Oe ae es a eo ae ee iu
Material examined and acknowledgments.......-...........2..2..220--. iy
Explanaior ot cranial measurements .....--..-------.....2--2ceceeeese 18
“PETG: LUGE TACIT: “oily: Sk Ae See pe one re Ae eey re eer iL
SDE: oo oacd eco SSG BEGG Oe a eo See OO Ce Rete aeenn ersten es eas 18
List of American species and subspecies, with type localities.............. 19
Kexgia american species and subspecies.....-....------2------- 202-2222 20
Descriptions of American species and subspecies.............-...---..--. 21
ER GE GL ID ONEROUS oo alee Selene) eine seek Ac So ate nen lc aoe ae 21
(Ezbleiol cranial measurements-.. 2.02. yess 52 ee ee 35
EVOHEATOUMILOUIVERTTIS STOWP 2% ono cpa ce arse oes ea ieee a 36
(Pable.on cranial measurements.s2 <2. 2... 62 yee ne = 225i o- 55
HEC OVC LG OU STOMP pee Ss). Se. MS eo Gertie See ce ee 56
abiciou cramio’ measurements: 02.) ee ee 72
iPeaplamaulomots platesian 25.2 Soe. So Sees Se See 74
HSEEe Neen se eh Pee SES ee Ae cing ieid cee 2 SS Rie sisjm ee wisiclaye ones ss 79

PLATE I,
UE

LLEUSTRATIONS:

PLATES.
[Plate I, frontispiece; Plates II-XV, following page 77.]

Golden-mantled marmot ( Marmota flaviventris nosophora).
Skins of Marmota olympus and M. fiaviventris obscura, showing molt.

III. Hind feet of American marmots, showing sole pads.

IV. Crania and mandiblesof Marmota flaviventris dacota, M.monaxmonaz, M. cali-
gata oxytona, M. flaviventris sierrae, M. monax rufescens, and M. caligata
caligata.

V. Skulls (upper views) of Marmota monax monax, M. monax rufescens, M.
monax preblorum, and M. monax ignava.

VI. Skulls (upper views) of Marmota monazx canadensis, M. monax petrensis, M.
monaz ochracea, and M. flaviventris warreni.

VII. Skulls (upper views) of Marmota flaviventris flaviventris, M. flaviventris
parvula, M. flaviventris avara, and M. flaviventris engelhardti.

VIII. Skulls (upper views) of Marmota flaviventris nosophora, M. flaviventris
luteola, M. flaviventris dacota, and M. flaviventris obscura.

IX. Skulls (upper views) of Marmota olympus, M. caligata oxytona, M. van-

couverensis, and M. caligata caligata.

X. Skulls (upper views) of Marmota caligaia cascadensis, M. caligata nivaria, M.
caligata okanagana, and M. caligaia oxytona.

XI. Skulls (lower views) of Marmota monax monaz, M. monaz rufescens, M.
monaxz preblorum, and M. monax canadensis.

XII. Skulls (lower views) of Marmota monax ignava, M. flaviveniris warreni, M.
monax ochracea, and IM. caligata nivaria.
XIII. Skulls (lower views) of Marmota flaviventris flaviventris, M. flaviveniris par-
vula, M. flaviventris avara, and M. flaviventris engethardit.
XIV. Skulls (lower views) of Marmota jflaviventris nosophora, M. flaviveniris
dacota, M. flaviventris luteola, and M. flaviventris obscura.
XV. Skulls (lower views) of Marmota olympus, M. vancouverensis, M. caligata
caligata, and M. caligata ccscadensis. —
TEXT FIGURES.
Page.
Hig. 1. Distribution of the Marmota monax group--......-2- 4252552 eee eee 28
2. Distribution of the Marmota flaviwventris group -- =. 22232222 eee 38
58

3. Distribution of the Marmota caligata sroup...-.---+~-22-2-==-- eee

4

No. 87, NORTH AMERICAN FAUNA, April, 1915,

REVISION OF THE AMERICAN MARMOTS.

——

By ArtHur H. Howe...

INTRODUCTION.

The American marmots, more often called woodchucks or ground
hogs, are among the best known of our native wild mammals. They
naturally divide into three distinct groups: (1) The woodchucks
(Marmota monax group) of eastern United States and Canada; (2) the
yellow-footed marmots (MV. flaviwentris group) of western United States
and southern British Columbia; and (3) the hoary marmots (W. caligata
group), chiefly restricted to the higher mountains of western North
America. The present paper is a revision of the American species
only, as it was not possible at this time to include a discussion of the
Kurasian forms of the genus.

HISTORY AND NOMENCLATURE.

Widely distributed, in many places abundant, diurnal in habit, and
frequently destructive to crops, the eastern species (monaz) quickly
attracted the attention of the early settlers, and through the medium
of skins or captive animals sent to Europe soon became known to
naturalists.

The first reference in literature to the American marmots dates
from 1703, when Baron La Hontan published a very brief account of
the woodchuck of eastern Canada, based on his observations in the
region about Lake Champlain.1 The name “siffleur’’ which he ap-
plied to the animal was carried by the French voyageurs throughout
the northwestern fur countries and still is current among the French
Canadians of the Hastern Provinces.

Catesby, in 1743, gave a brief description of the eastern woodchuck
under the name of “The Monax,’” and a few years later, in 1747,
Edwards published a more extended account with a very inaccurate
figure of the animal under the title of “The Monax or Marmotte of
America.’? Edwards’s description and figure were copied by many

e 5 ies 7
subsequent authors and furnished also the basis of the first technical
SEI se ee Ee

1La Hontan, Baron de. Voyages dans l’Amerique, 1703, p. 81. os
2Catesby, Mark. Nat. Hist. of Carolina, etc., 1, 1743, App., D. XXvill.
8 Edwards, George. Nat. Hist. Uncommon Birds, II, 1747, p. 104, pl. civ.

6 NORTH AMERICAN FAUNA. [No. 37.

name applied to the species— Mus monaz Linnaeus. While the figure
is wholly unrecognizable, the description is sufficiently clear to war-
rant the use of the name bestowed by Linnaeus.

Erxleben, in 1777, named the Canadian woodchuck (now recog-
nized as a subspecies of monaz) “‘Gls’’ canadensis,’ and Pallas in the
following year renamed it “‘ Mus” empetra,? both descriptions being
based on the ‘Quebec marmot” of Pennant.’ Blumenbach, in 1779,
named the genus Marmota,® and Schreber the following year intro-
duced the name Arctomys monaz on a plate evidently copied from
Edwards’s figure of ‘The Monax.’’* Schreber’s generic name, though
of later date than Blumenbach’s, received general acceptance and
continued in common use for the marmots until the early years of the
present century, when the name Marmota was restored as the proper
appellation of the genus.” In 1788, Gmelin proposed the name
Arctomys pruinosa,® based on the hoary marmot of Pennant, and
this name was used for that animal until 1888, when Tyrrell showed
that it had been incorrectly applied and proposed to use in its place
Arctomys caligata Exschscholtz, first described in 1829.°

Sabine, in 1822, published an account of the American marmots,
recognizing three species: monaz, empetra (=canadensis), and prui-
nosa (=caligata).° His descriptions of monex and pruinosa were
copied from previous authors, but that of empetra was drawn from
a specimen in the British Museum, and furnished apparently the
first correct description of any American marmot. The first accurate
drawing of an American species is that of the Canadian woodchuck
published by Richardson in the Fauna Boreali-Americana (1829).

In 1836 King, recognizing clearly that pruinosa of Gmelin was
not applicable to the hoary marmot, but overlooking Eschscholtz’s
name caligata, proposed the name Arctomys okanaganus for a marmot
of this group obtained in southern British Columbia, and gave a very
full and accurate description and a good figure of the animal, drawn
from a living specimen which he sent to the Zoological Gardens in
London." His name, however, was not accepted by zoologists, and
pruinosa continued in use for many years. The specimen taken by
King was seen by Audubon in London, and furnished the basis of his

1The use of the name ‘‘monax” by both Catesby and Edwards independently (Edwards states that he
had never seen an account of the animal) indicates that it came from the vernacular—a theory which is
strengthened by the fact that the animal is still called ‘‘moonack” in parts of southern Virginia.

2Erxleben, J.C. P. Syst. Anim., Mamm., 1777, p. 363.

3Pallas, P.S. Nov. Spec. Glir., 1778, p. 74.

4 Pennant, Thomas. Syn. Quad., 1771, p. 270, Plate 24, fig. 2.

5 Blumenbach, J. F. Handb. der Naturgesch., I, 1779, p. 79.

6Schreber, J.C. D. von. Siugthiere, pl. eeviii, 1780; text, IV, 1782, p. 737.

7Trouessart, E. L. Cat. Mamm., Suppl., 1904, p. 343.

- 8Gmelin, J. F. Syst. Nat., 1, 1788, p. 144.
9Tyrrell, J. B. Proc. Can. Inst.; 3d Ser., VI, 1888, p. 88.

10Sabine, Joseph. Trans. Linn. Soc. London, XIII, 1822, pp. 579-591.
uKing, R. Narr. Journ. to Shores of Arctic Ocean, II, 1836, pp. 232-248.

1915.1 INTRODUCTION. 7

figure of the hoary marmot in the ‘‘Quadrupeds of North America,”
but the illustration is colored much too brown to represent the animal
correctly. In the work referred to’! the authors gave also an ex-
tended account and a good drawing of the eastern woodchuck, and
a shorter account, with a figure, of the yellow-bellied marmot of
western North America, first described by them in 1841 under the
name Arciomys flaviventer. This is the first appearance in literature
of the widely distributed group of yellow-footed marmots, and the
work of Audubon and Bachman was the first treatise in which all
three groups of American species were recognized.

Little advance in knowledge of the genus was made for nearly half
a century, and the three groups (monax, flaviveniris, and caligata) con-
tinued to be known by only a single species in each. The Canadian
woodchuck, so clearly described by Sabine and Richardson, was con-
sidered by both Baird and Allen to be identical with monaz (of which
it is, indeed, a subspecies) and the names based on it (canadensis
Erxleben, empetra Pallas, melanopus Kuhl) were placed by them in
synonymy, but were later revived by several authors.

In 1889 Merriam described dacota, a member of the flaviveniris
group, from the Black Hills, S. Dak.; and in 1898, olympus, a member
of the hoary marmot group, from the Olympic Mountains, Wash. In
1899 Bangs named ignava from Labrador and avara from southern
British Columbia; in 1905 J. A. Allen described engelhardti from
Utah; in 1909 Heller proposed vigilis for a hoary marmot from
Alaska; in 1911 Swarth named ochracea from Alaska and vancowver-
ensis from British Columbia; and in 1912 Hollister proposed the name
sibila for a hoary marmot from the northern Rocky Mountains, but
as this name was found to be preoccupied the species was renamed
by him oxytona in 1914. The present writer, after making a prelimi-
nary study of the group, added, in 1914, 10 new forms to the 13 cur-
rently recognized.? Two additional new races are here described and
one old name (okanagana) is revived, making a total of 26 forms rec-
ognized in this revision.

VERNACULAR NAMES.

The marmots of the monaz group are known in the Northern States
as woodchucks, and in the Southern States as ground hogs, the EKuro-
pean name marmot being practically unknown in eastern North
America. In eastern Canada, among the French Canadians, the
name ‘‘siffieur”’ is current, and in central Canada the Cree Indian

1Audubon & Bachman. Quad. N. Am., I, 1841, pp. 16-24, pl. ii; III, 1854, pp. 17-20, pl. ciii; pp. 160-162,
pl. exxxiv.
2Rhoads, 8. N., Proc. Acad. Nat. Sci. Phila., 1897, p. 30; Allen, J. A., Bul. Amer. Mus. Nat. Hist., X,

1898, p. 456; Preble, E. A., N. Am. Fauna No. 22, 1902, p. 47; Ibid., No. 27, 1908, p. 159.
3Howell, A. H. Proc. Biol. Soc. Washington, XX VII, 1914, pp. 13-18.

8 NORTH AMERICAN FAUNA. ENON 37.

name ‘‘wenusk”’ is generally used. In southern Virginia, as I am
informed by Edward A. Preble, the woodchuck is locally known as
“moonack,” which is probably a corruption of the original name
‘“monax’’ used by both Catesby and Edwards.

The yellow-footed marmots (flaviventris group) are commonly
called woodchucks or rockchucks, more rarely ground hogs or marmots.

The hoary marmots (caligata group) are most often known as
ground hogs, whistlers or “‘siffleurs,” sometimes as whistling “pigs,”
whistling marmots, or ‘‘ badgers.”’

HABITS.

The eastern woodchucks live for the most part in pairs or family
groups, the yellow-footed marmots in more or less scattered colonies,
while the hoary marmots are more strongly gregarious. All the
species live in burrows which they dig for themselves. In regions
where rock piles, rock ledges, or stone walls occur the burrows are
usually excavated underneath or among rocks, but natural openings
in cliffs are often utilized for dens.

Eastern woodchucks (monaz group), while preferring rocky blufis
or stone walls for a habitation, often live in meadows devoid of rocks
and where the burrows are surrounded by an abundant growth of
grass or clover.

Yellow-footed marmots (flaviventris group) usually live either on
rocky hillsides, in the crevices of cliffs, or beneath rock piles in
meadows. They frequently make their burrows beneath unoccupied
buildings, but are never found far from hills, and are often abundant
in the higher parts of mountains.

Hoary marmots (caligata group), when living at timber line in the
mountains, as is their invariable habit in the southern part of their
range, are always found in or about rock slides, but in Alaska and
northern British Columbia, where they frequently descend to low
altitudes, they often make their burrows in grassy flats or on open
hillsides.

All the species are mainly terrestrial, but the eastern woodchucks
occasionally climb into trees and bushes. They are not at home,
however, in such situations, and as a rule may easily be dislodged.
The tree-climbing habit appears to be more strongly marked in the
woodchucks of the Mississippi Valley than in those inhabiting the
Atlantic States. Charles Aldrich, of Webster City, Iowa, has re-
corded an instance of a woodchuck ascending an oak tree to a height
of 40 feet,1 and Dr. F. W. Langdon states that in Ohio he has seen
one descend the perpendicular trunk of a large sugar maple, head
jirst2 In Minnesota and Wisconsin, as I am informed by Vernon

1 Aldrich, Charles. Am. Naturalist, XV, 1881, p. 737.
2 Langdon, F. W. Journ. CincinnatiSoc. Nat. Hist., III, 1880, p. 305.

1915.] INTRODUCTION. 9

Bailey and H. H. T. Jackson, it is a common occurrence for wood-
chucks to take to trees when pursued by dogs.

The eastern woodchuck is mainly diurnal, but occasionally is
found abroad at night also, especially by moonlight. Merriam says
of it:

In summer, throughout the farming districts, they commonly leave their burrows
early in the morning, late in the afternoon, and during moonlight nights, but may
sometimes be found abroad at all hours. As autumn approaches, and they become

more and more fat and sleepy, they usually appear only in fine weather, and then
but for a few hours in the hottest part of the afternoon.

The yellow-footed and hoary marmots are fond of sunning them-
selves on projecting points of rock where they are safe from attack
and may overlook a wide stretch of country. During cloudy and
stormy weather they are less active and spend a larger part of the
time in their burrows.

All the American marmots when alarmed utter a loud, shrill
whistle, a habit apparently most pronounced in the hoary marmot,
whose notes are much stronger than those of the smaller species
and capable of being heard at a distance of more than a mile. This
habit has given the name ‘whistler’? to the hoary marmot and
‘‘siffeur’’ to the woodchuck of eastern Canada.

BURROWS.

Although originally living in the woods, the eastern woodchuck
prefers clearings for its abode, and, as a result of an abundance of
easily obtained food, is now probably much more numerous than in
primitive times. Its burrows are commonly excavated in the face
of a bluff, in a grassy meadow, or underneath a stone wall, a stump,
or the roots of a tree.

Merriam states that the burrows are of two principal types—

* * * the first slopes at a moderate angle from the surface and has a mound of
dirt near its entrance; the other is more or less vertical for several feet (often a metre
or more) immediately below the surface, and no loose earth can be found in its neigh-
borhood. * * * Asa rule they [the galleries| slant abruptly downward from the
entrance to a depth of from three to four feet * * * , whence inclining slightly
upward and usually curving to one side, they extend horizontally for a varying dis-
tance (commonly from 10 to 25 feet) * * *. Two or more short lateral branches
are generally given off from the main gallery, and lead, sloping upward and then
downward, to the more or less circular chambers that contain the animals’ nests. It
has been my invariable experience to find these chambers above the level of the
bottom of the entrance incline, and I have seen one that was within a foot and a hali
* * * of the surface. The nest itself is usually composed of dry grasses and
leaves and rarely exceeds a foot in diameter. * * * ‘The main gallery or one of
its branches commonly terminates in a slight excavation, which is found to contain
the animal’s excrement.”

1 Merriam, C. H. Mamm. of the Adirondacks, Trans. Linn. Soc. N. Y., I, 1884, p. 146.
2Tbid., pp. 148-149.

10 NORTH AMERICAN FAUNA. [No. 37.

William Hubbell Fisher excavated and measured nine burrows of
the woodchuck in Lewis County, N. Y., and has published a detailed
description of them, with diagrams. The longest burrow which he
examined measured (including side branches) 44 feet 94 inches; the
shortest, 6 feet 84 inches; the deepest burrow was 49 inches below ~
the surface; the shallowest, 23 inches.*

No description of the burrows of the yellow-footed marmot has
come to the writer’s notice, and the only known account of those of
the hoary marmots is that given by William H. Wright in his work
on the grizzly bear. Describing the manner in which a grizzly had
opened up a den of these marmots, he says:

The den ran in under several layers of loose flat rocks, some of which were two or
three feet long by half as many wide, and several inches thick. These he had
ripped out easily and thrown down hill, and the dirt and small bowlders had been
hurled out and now covered the snow ail about for a space of ten or twelve feet.

On the rocks and snow were large spots and blotches of blood, telling of the feast
that had rewarded his labors, and that there had been more than one marmot was
shown by the numerous tracks, These animals had burrowed down some six or seven
feet into the side of the mountain, and under a large flat stone they had scooped.
out a little cave, some three feet in diameter, where they had a soft bed of grasses
that they had carried in. When the grizzly broke his way into their home there
had been a great rush for freedom.

The marks in the snow indicated that all the marmots had been
devoured by the bear.’

HIBERNATION.

All the species hibernate for periods varying from 4 to 6 months.
Merriam states that in New York, along the western border of the
Adirondacks, the woodchuck usually goes into winter quarters
between the 18th and 25th of September and reappears the middle
or latter part of March; in early springs following mild winters, he
adds, ‘““woodchucks occasionaily appear in February, but reenter
their burrows and again become dormant if the temperature falls.”’ 3
Bachman states that he once observed a woodchuck in New York
State on October 23 sunning himself at the mouth of his burrow, and
also in the same State saw one killed by a dog on March 1.4 Extreme
dates of occurrence for this region are: February 22, Adirondack
Mountains, N. Y., specimen in the Merriam collection; and Novem-
ber 20, Fort Miller, N. Y., one seen by Dr. E. A. Mearns.> In the
more southern States, hibernation covers a shorter period, as indicated
by the occurrance of the animals at the base of Roan Mountain,
N.C., as early as February 7 and as late as October 23. Other dates

1 Fisher, W. H. Jour. Cincinnati Soc. Nat. Hist., XVI, 1893, pp. 105-123.
2 Wright, W. H. The Grizzly Bear, London, 1909, p. 82.
3 Merriam, C. H. Mamm. of the Adirondacks, Trans. Linn. Soc. N. Y., II, 1884, pp. 143-144.

4 Audubon & Bachman. Quad. N. Am. I, 1849, p. 20.
5 Mearns, E. A. Bul. Amer. Mus. Nat. Hist., X, 1898, p. 337.

1915.] - INTRODUCTION. iar

of late occurrence are as follows: October 6, Teslin Lake, Yukon;
October 17, Dowagiac, Mich.; October 25, Johnson County, Iowa.
Hahn states that in southern Indiana woodchucks usually retire about
the middle of October and begin to clean out and enlarge their
burrows during the last days of February.*

The yellow-footed marmots go into hibernation between the
middle of August and the first of October, the date varying with the
altitude and local conditions. Individuals living in the valleys
retire earlier than those living higher up in the mountains. Warren
states that m Gunnison County, Colo., this species dens up about
the first of October, but individuals are sometimes seen much later.?
Allan Brooks states (in epistle) that at Okanogan Landing, British
Columbia, practically all these marmots disappear before the middle
of August, but he has occasionally seen their tracks as late as early
October. In the mountains of Montana and Wyoming this species
usually enters hibernation during the last of August or the first of
September; Biological Survey field parties have never found the
animals later than the first week in September. In the Bitterroot
Valley, Mont., the first one seen in the spring in 1910 was on March
24, and by April 1 they were numerous; in 1911 one was seen there
by Bernard Bailey, on March 13. In western Oregon extreme dates
of occurrence are: February 4 (Klamath Lake), and September 25
(Mount Hood).

The hibernating period of the hoary marmot begins the last of
September or first of October. Near Tatletuey Lake, British Colum-
bia, Edward A. Preble found the species still active a few hundred feet
above timber line on September 23 and 25, but none was seen after
the latter date. Like the yellow-footed species, the hoary marmots
retire earlier in the valleys than in the mountains. Heller states
that at Valdez Narrows, Alaska, the species went into hibernation
about the middle of September.®

Bachman thus described his observations of a pair of hibernating
marmots:

In the summer of 1814, in Rensselaer County, in the State of New York, we marked
a burrow which was the resort of a pair of marmots. In the beginning of November
the ground was slightly covered with snow, and the frost had penetrated to the
depth of about an inch. We now had excavations made in a line along the burrow
or gallery of the marmois, and at about twenty-five feet from the mouth of the hole;
both of them were found lying close to each other in a nest of dried grass, which did not
appear to have been any of it eaten or bitten by them. They were each rolled up,
and looked somewhat like two misshapen balis of hair, and were perfectly dormant.

We removed them to a haystack, in which we made an excavation to save them from
the cold. One of them did not survive the first severe weather of the winter, having,

1 Hahn, W.L. Mamm. of Indiana, 1909, pp. 481-482.
2 Warren, E.R. Mamm. of Colorado, 1910, p. 148.
8 Heller, Edmund. Univ. of California Pub. Zool., V, 1910, p. 339.

119 NORTH AMERICAN FAUNA. [No. 37.

as we thought on examining them, been frozen to death. The other, the male, was
now removed to a cellar, where he remained in a perfectly dormant state until the
latter part of February, when he escaped before we were aware of his reanimation.
We had handled him only two days previously, and could perceive no symptoms of
returning vivacity.'

The following account of a yellow-footed marmot (Marmota flavi-
ventris subsp.) found in midwinter in the Silver Mountain tunnel at
Ophir, Colo., probably indicates a common method of hibernation
in that species:

* * %* [Helhad packed in grass fora nest, and taken up his winter quarters. He was
rolled up like a ball, with his forepaws over his eyes; we pulled his paws away, and
his eyes were closed; all our efforts to awake him were futile; he would yawn like a

boy that had been disturbed when sleeping soundly, return his paws to his eyes, and
curl himself up in his original position.?

BREEDING.

The eastern woodchuck usually produces from 4 to 6 young at a
birth. Bachman states, however, that on two occasions he counted
7 and on another 8 young in a litter, and H. H. T. Jackson informs
me that he once saw a litter of 9. In New York State, according to
Merriam, this species brings forth its young the last of April or first
of May. In the Southern States they are born somewhat earlier.

The yellow-footed marmots breed at about the same season as
their eastern relatives and produce from 38 to 8 young ata birth. In
the Bitterroot Valley, Montana, 5 females collected between April 8
and April 16 were pregnant, the number of embryos being in most
cases 5 or 6 (in one case 3). Young marmots were out in numbers
in that region on May 30. Warren states that an individual of this
species collected at Sulphur Springs, Colo., on May 4, contained 8
embryos.”

The hoary marmots probably breed somewhat later than their
smaller relatives, but little information on this point is available. <A
female specimen of Marmota caligata mvaria, taken May 27, 1895, near
St. Marys Lake, Mont., contained 5 embryos. Swarth states that
in southern Alaska young individuals of MM. c. caligata were seen
running about in the middle of June, but on Vancouver Island, dur-
ing the first three weeks of July, no young ones [of J. vancouverensis]
had yet emerged from the burrows.*

FOOD.

The principal food of the eastern woodchuck is clover, alfalfa, and
grass, and-the animals do considerable damage to these crops both

1 Audubon & Bachman. Quad. N. Am., I, 1849, p. 22.
2Osborn, S. E. The Observer, III, 1892, p. 32.

3 Warren, E.R. Mamm. of Colorado, 1910, p. 148.

4Swarth, H.S. Univ. of California Pub. Zool., X, 1912, p. 90.

1915.] INTRODUCTION. 13

by consuming the forage and by trampling down much that they
do not eat. Cultivated crops and orchard trees are occasionally
injured by them, but the damage usually is not serious. HEyvermann
and Clark state that in Indiana woodchucks—

* * * sometimes damage young corn plants and occasionally feed on the leaves

of pumpkin, squash, and bean vines. They sometimes visit the kitchen garden
and do more or less damage to the cabbage heads and celery. !

Brooks, writing of the woodchuck in West Virginia, says:

* * * Feeds on corn in the roasting-ear, which it procures by breaking down the
stalks; is also fond of pumpkins, young beans, grass and other cultivated crops. Fre-
quently gnaws and scratches the bark of young fruit trees.”

Hahn mentions capturing a woodchuck in a sassafras tree and
finding its stomach gorged with sassafras leaves.* Dr. Witmer Stone
states that the woodchuck sometimes eats cantaloupes.

The food of yellow-footed marmots is similar to that of eastern
woodchucks, but probably includes a larger proportion of wild plants
and less grass and clover. Vernon Bailey found in the stomachs of
the Black Hills marmot flowers, leaves, and green seeds of various
plants, including Astragalus bisulcatus and Sedum douglasu. In dry
excrement of marmots at Spokane Bridge, Wash., he found the seeds
of Amelanchier alnifoha and Rubus nutkanus. Birdseye states that
the marmots in the Bitterroot Valley, Montana, feed on timothy,
clover, alfalfa, dandelions, and other native plants. THe says:

In hayfields they consume a very considerable amount of feed; and beans, carrots,

potato vines, cabbage, and other garden truck are almost sure to suffer whenever
woodchucks have access to them.+

Little is known definitely concerning the food habits of hoary
marmots, but they doubtless feed, like the other species, on grass
and the tender leaves and stems of native plants.

ECONOMIC STATUS.

As already shown, the food habits of the American marmots
make the animals a decided pest wherever they occur about culti-
vated lands. In this respect the woodchucks of the Eastern
States are the chief offenders, and the farmers of that region wage
constant warfare on the animals, with indifferent success. Where
the woodchucks occupy mowing lands they not only consume con-
siderable grass and tread down much more which can not be cut by
a mowing machine, but their burrows and mounds make it difficult
and dangerous to operate a mower. Horses sometimes are injured
by stepping into the holes made by woodchucks, and the knives of

1Evermann & Clark. Proc. Washington Acad. Sci., XVII, 1911, p. 13.
2 Brooks, F. E. Report W. Va. Board Agr. for 1910 (1911), p. 15.

3 Hahn, W. L. Mamm. of Indiana, 1909, p. 482.
4 Birdseye, Clarence. Farmers’ Bul. 484, U. 8S. Dept. Agr., 1912, p. 28.

14 NORTH AMERICAN FAUNA. [No. 87. _

the machines are liable to be dulled or broken by running into piles
of earth or rocks. Vernon Bailey states that he has seen an acre of
oats on a hillside in New York State almost ruined by a family of
woodchucks, their trails having broken down most of the grain which
they had not cut to eat.

The fur of the American marmots is not at present used com-
mercially, but the hides of hoary marmots are employed to some
extent by the Indians of western Canada for making robes. The fur
of this animal is fully as good as that of the European and of some of
the Asiatic species, all of which figure extensively in the fur trade.
Furthermore, the American animal is larger than most of the Eura-
sian species, and the color pattern of its skin is such as to make a
very handsome natural fur. Ii a sufficient number of these pelts
could be secured in late autumn when they are in prime condition
there would seem to be every reason for utilizing them in the fur
trade. In parts of Alaska and northern British Columbia the
animals are very abundant, and Indians of that region annually cap-
ture large numbers both for fur and flesh. The yellow-footed
marmots are considerably smaller than the hoary marmots, but their
pelage is long, full, and silky, and could readily be made into a very
attractive fur. The pelage of the eastern woodchuck is much coarser
and thinner and has never been considered of much value for fur;
the hides, however, are tough and durable and might be utilized for
some kinds of leather. Farmers’ boys often tan the skins of the
eastern woodchuck and make them into shoe strings, whip lashes, ball
covers, or mit facings.

The flesh of marmots is said to be palatable and in certain regions
of the Northwest furnishes an important food supply for the native
Indians.

MARMOTS AS CARRIERS OF DISEASE.

For many years the opinion has generally prevailed that the mar-
mots of central Asia are in some way concerned in the spread of the
plague, and this theory was accepted even as late as 1911 by the
members of the International Plague Conference held at Mukden in
April of that year. Accounts have frequently been published in
medical literature of the appearance of the disease among ‘‘tar-
bagan’”’?? hunters under circumstances pointing strongly to the
possibility of infection from handling the animals or consuming their
flesh, but unfortunately none of these accounts shows bacteriological
evidence of the existence of the plague among marmots, nor do they
demonstrate conclusively that the disease is contracted in any way
from the animals.”

1 This is the name applied in medical literature to the marmots of Asia. It is said to be of Mongolian

origin. The Russian name of the animal is ‘‘churok.”’
2 See especially an article by Dr. Frank G. Clemow, Jour. Trop. Medicine, February, 1900, pp.169-174.

e

1915.1 INTRODUCTION. 15

The last great outbreak of pneumonic plague in Manchuria, during
the winter of 1910-11, was supposed to have started among tar-
bagan hunters in the town of Manchouli, a station on the Trans-
Siberian Railway, whence it spread rapidly southward along the rail-
way to Harbin and Mukden. Efforts were made during the progress
of this epidemic to locate the disease among the native marmots,!
and a number of individuals captured near Mukden were inoculated
experimentally by Prof. Zabolotny and found susceptible to plague
infection. One individual suffering from the disease was brought to
him, this being the only known instance of the natural appearance of
plague among marmots.? More recently a systematic effort has been
made by the Chinese Government to discover the disease in nature,
but the evidence obtained is wholly of a negative character.’

The experimental evidence just mentioned and the close relation-
ship of marmots to ground squirrels, which are known to carry plague
on the Pacific coast of North America, warrant looking upon these
animals with suspicion whenever they occur in a plague-infested region.

One American species—the golden-mantled marmot (Marmoia flavi-
veniris nosophora)—is known to assist in spreading the deadly spotted
fever by serving as host for the fever tick (Dermacentor venustus), both
inadult and nymphal stages. Over 200 ticks have been taken from a
single wild marmot, and when in captivity the species has been shown
to be susceptible to spotted fever.t Since this marmot is not utilized
for fur there is relatively little danger of the spread of the disease to
other regions through the medium of the animals, but every effort
should be made to exterminate them where they occur about culti-
vated lands or in the vicinity of dwellings. Suggestions for destroy-
ing marmots are given in Farmers’ Bulletin 484, United States De-
partment of Agriculture.

EXTERNAL CHARACTERS.

x

In the American marmots the body is thickset and clumsy; the
head short and broad; the legs short and stout; and the tail rather
short (about one-fifth to one-third of the total length), densely haired,
and slightly flattened. The nose is broad and blunt, covered with
hair to the edge of the nostrils, and the ears are short, broad, rounded,
and well-haired. ‘Tufts of long black bristles grow from the side of
the head, one directly behind the nose, another underneath and
between the eye and ear, and a third shorter tuft over the eye; small
scattered bristles also are found under the chin. The eyes are rather

1 The marmot of Manchuria, although generally referred to in plague literature as Arciomys bobac, is in

all probability Marmota sibirica, specimens of which, collected at Urga, Mongolia, are in the U.S, Na-
tional Museum.

2Strong, R. P. Rept. Intern. Plague Conference held at Mukden, April, 1911. Manila, 1912,

3 Dr. Wu Lien Teh (G. L. Tuck). The Lancet, London, Aug. 23, 1913, pp. 529-535.

4 Birdseye, Clarence. Farmers’ Bul. 484, U.S. Dept. Agr., 1912, p. 28.

16 NORTH AMERICAN FAUNA. [No. 37.

small and nearly or quite circular. The feet are robust, with stout,
slightly curved fossorial claws, those on the fore feet somewhat
heavier; the thumb of the front foot (in the American species) is
rudimentary and often very small, but bears a broad, flat nail; the
third digit is the longest, the second and fourth subequal, the fifth
decidedly shorter; the palm is naked, bearing 3 pads at the bases
of the digits and 2 larger posterior ones; on the hind foot the third
digit is slightly longer than the subequal second and fourth, the fifth
and first successively much shorter; the sole is naked except at the
heel and bears 6 pads, 4 at the bases of the digits and 2 posterior to
them. The mammae number 5 pairs, except in the monaz group, in
which there are usually but 4 pairs. In M. monaz, according to
Baird, ‘‘there is a short, shallow cavity between the muscles of the
jaw and the cheek, attaining a depth perhaps of half an inch or less
and occupying the place of the internal pouch of Tamas and Sper-
mophilus [Citellus].’’ 3
PELAGE AND MOLT.

The pelage of the American marmots consists of hairs of two kinds—
a dense, soft, and somewhat woolly underfur, confined chiefly to the
back and sides; and longer, somewhat coarser hairs covering the
whole body, intimately mixed on the back with the underfur and
projecting beyond the tips of the latter. The underfur is of two
colors, usually some shade of gray or dark brown tipped with a
lighter color, generally a shade of gray, buff, cinnamon, or reddish
brown. The long hairs are also of two colors, usually a shade of
brown, hazel, or black, tipped with a lighter shade—buff or white.
The tips of the underfur usually show through the long hairs, and
form an important element in the general color tone of the pelage.
The hairs on the head and feet are shorter than on the rest of the body,
and are unicolor to the roots; those on the tail are long and coarse,
their bases of a darker shade than the tips. The hair on the under-
parts is shorter and sparser than on the back and is without underfur.

The pelage is renewed annually in summer, usually in August,
sometimes in July or even June. Individuals taken in early spring,
soon after emerging from hibernation, are usually in full, long pelage,
but by midsummer the pelage often becomes very much worn and
faded. There seems to be no uniform method of molting, the new
pelage sometimes appearing in patches on various parts of the body.
In a specimen of Marmota olympus, taken August 28, the new pelage
was coming in in large patches, the molt being farthest advanced on
the middle of the back, with streaks of new hair along the sides of the
neck and body. (See Pl. II.) In another individual of the same
species, taken August 18, new hair was coming in on the rump and

1 Baird, S. F. Mamm. N. Am., Rept. Expl. and Surv. R. R. Pacif., VIII, 1857, p. 340. The author has

had no opportunity to examine specimens of any of the groups in the flesh, so does not know whether this
character is possessed by all.

1915.] INTRODUCTION. 17

hinder back, the old intimately mixed with the new. In animmature
individual of M. caligata cascadensis, taken August 5, new pelage
covered all of the body except the rump and tail. A somewhat
similar condition is shown by an adult individual of M. monaz preb-
lorum, taken July 11, the new pelage coming in over the entire
upperparts, farther advanced on the fore back and shoulders. In
aspecimen of I. flaviventris nosophora, taken June 7, the molt was
just beginning in two patches on the hinder back. In a specimen of
M. f. obscura, taken July 27, new pelage appears in the form of a
band on the middle of the back reaching from the top of the head
totherump. (See Pl. IT.)

MELANISM.

Melanism is most strongly developed in the subspecies Marmota
caligata vigilis, occupying the region around Glacier Bay, Alaska. In
this race some individuals are entirely black except for a few grayish
hairs on the sides and neck, and small whitish patches on the under-
parts and nose. No purely black specimens of Jf. monaz have been
seen, but a melanistic phase is rather common in New York and
New England. The darkest specimen seen (from Lake George, N. Y.)
is dark blackish brown ali over, except the head and face, which are
mixed mummy brown and benzo brown. Other specimens from
New York State are dark chestnut-brown. Vernon Bailey states that
he has seen a very few black individuals in Minnesota.

A dark phase occurs also in WU. flaviveniris luteola, but this can
hardly be said to be melanistic (see description under that species).
A curious specimen of WU. f. avara from Pullman, Wash., has the tips
of the hairs of the head, fore back, and most of the underparts dark
blackish brown, the hirider back being pale grayish mixed with dark

_ brown.

MATERIAL EXAMINED AND ACKNOWLEDGMENTS.

The present revision is based on a study of 1,051 specimens, 722
of which are contained in the U. S. National Museum, including
the Biological Survey and the Merriam collections; the remainder
(329 specimens) have been borrowed from other museums and from
private collections. For the loan of this material I desire to extend
my thanks to the following: Dr. J. A. Allen, of the American Museum
of Natural History; Messrs. Samuel Henshaw and Outram Bangs, of
the Museum of Comparative Zoology; Dr. Witmer Stone, of the
Academy of Natural Sciences, Philadelphia; Dr. W. J. Holland and
Mr. W. E. Clyde Todd, of the Carnegie Museum; Dr. A. G. Ruthven,
of the University of Michigan; Messrs. Charles B. Cory and W. H.
Osgood, of the Field Museum of Natural History; Mr. C. D. Bunker,
of the Kansas University Museum; Prof. C. C. Nutting, of the Uni-
versity of Iowa; Mr. J. D. Figgins, of the Colorado Museum of Natural

69168°—15—2

18° NORTH AMERICAN FAUNA. (No. 37.

History; Dr. Joseph Grinnell, of the Museum of Vertebrate Zoology,
University of California; Mr. P. A. Taverner, of the Victoria Memorial
Museum; Mr. Arthur H. Helme, of Miller Place, N. Y.; Mr. H. H. T.
Jackson, of the Biological Survey; and Mr. Edward R. Warren, of
Colorado Springs, Colo.

EXPLANATION OF CRANIAL MEASUREMENTS.

Measurements of skulls of marmots, in millimeters, have been taken
as follows:

Condylo-basal length—From posterior border of condyle to most
anterior point of premaxillae.

Palatal length—From posterior border of palate (disregarding
median process) to most anterior point of premaxillae.

Postpalatal lengih—From posterior border of palate to inferior lip
of foramen magnum. .

Length of nasals—From most anterior point to most posterior
point.

Zygomatic breadth.—Greatest breadth across zygomata. |

Breadth across mastoids.—Greatest breadth across mastoid processes.

Least interorbital breadth—Shortest distance across frontals in
front of postorbital processes.

Breadth of rostrum.—Greatest breadth of rostrum at most anterior
points of maxillae.

Mazillary tooth row.—Alveolar length of maxillary molar-premolar
tooth row.

Genus MARMOTA Blumenbach.

Glis Erxleben, Syst. Regni Anim., I, 1777, p. 358 (part). (Not Glis Brisson, 1762.)

Marmota Blumenbach, Handb. der Naturgesch., I, 1779, p. 79. Type, Marmota
alpina [= Mus marmota Linnaeus].

Arctomys Schreber, Saéugthiere, 1780, Plate CCVIII. Type, Mus monaz Linnaeus.

GROUPS.

The American marmots embrace three very distinct groups as
follows: (1) The monax group—all the eastern woodchucks, the
Canada woodchuck, the British Columbia woodchuck and the ochra-
ceous woodchuck of Alaska and northern British Columbia; (2) the
flaviventris group—all the yellow-footed marmots; and (3) the cali-
gata group—the hoary marmots, including the species caligata,
olympus, and vancouverensis. ‘The characters of the various groups
are given in connection with the technical descriptions of the species.

A detailed study of Old World forms was not possible in the present
revision, but, so far as known, none of the American species has any
very near relative in Eurasia.1 Several groups are represented there,
some of the species resembling the caligata group in certain skull
characters, others apparently being more nearly related to the monaz

1 The species living in eastern Siberia has not been seen.

1915.] LIST OF SPECIES AND SUBSPECIES. 19

group. Marmcta marmota, the type of the genus, though widely
differing from the American species in the greater depth and convex-
ity of the cranium, resembles monazx in the characters of the basi-
occipital, the shape of the postorbital processes, the relatively wide

interorbital region, and the nearly parallel maxillary tooth rows.

Most of the Asiatic species, however, apparently are more closely
related to MM. caligata than to either of the other American groups.
Theskulls of ali American species, contrasted with those of the Eura-
sian species, show a marked flattening of the cranium, the dorsal
outline of the skull being decidedly more convex in practically all
Old World forms than in those of America. All the Eurasian species
differ also in coloration from the American forms. All the American
species possess a small rudimentary thumb bearing a flat nail—a
character present in most of the Asiatic species, but absent in J.
marmota of Europe.

List of American Species and Subspecies, with Type Localities.

Marmota monax group:

Marmota monax monaz (Linnaeus)..-... Maryland.
monax rufescens Howell.......--..- Elk River, Minn.
monax preblorum Howell. ......... Wilmington, Mass.
monas wgnava (Bangs). .---+....-.--- Black Bay, Labrador.
monax canadensis (Erxleben).....- ‘Canada et ad fretum Hudsonis”—
fixed at Quebec, Quebec.
mowae petrensis Nobis... -.-..--.-- Revelstoke, British Columbia.
monaz ochracea Swarth.....-......- Head of Fortymile Creek, Alaska.

Marmota flaviventris group:
Marmota flaviventris flaviventris (Audu-

emer hachmManm) 2.002.220. els ‘‘Mountains between Texas and Cali-

fornia ”’—fixed on Mount Hood, Oreg.

flaviventris avara (Bangs) ....-....- Okanogan, British Columbia.

flavwentris sierrae nobis ....-..-..-- Mount Whitney, Cal.

flaviventris parvula Howell ........ Jefferson, Toquima Range, Nev.

flaviventris engelhardtit Allen ....-.-.- Beaver Mountains, Utah.

flaviventris nosophora Howell. ...-..- Willow Creek, 7 miles east of Corvallis,
Mont.

flaviventris dacota (Merriam) ....... Custer, S. Dak.

flaviventris luteola Howell ........-. Voods P. O., Medicine Bow Mountains,
Wyo.

flaviventris warrent Howell ......-- Crested Butte, Colo.

flaviwentris obscura Howell .......-. Wheeler Peak, N. Mex.

Marmota caligata group: ;
Marmota caligata caligata (Eschscholtz) . Bristol Bay, Alaska.

caligataargilis Heller... .2.--.--- Glacier Bay, Alaska.
caligata sheldoni Howell.......-.--.- Montague Island, Alaska.
caligata oxytona Hollister..........- Head of Smoky River, Alberta.
caligata okanagana (King).......-- Gold Range, British Columbia.
caligata nivaria Howell............- Mountains near St. Marys'Lake, Mont.
caligata cascadensis Howell..-...-.--- Mount Rainier, Wash.
Marmota olympus (Merriam).....-.----- Olympic Mountains, Wash.
Marmota vancouverensis Swarth......-- Mount Douglas, Vancouver Island, Brit-

ish Columbia,

20 NORTH AMERICAN FAUNA. . [No. 37.

Key to American Species and Subspecies.
[Based on typical adults.]

a. Upperparts mainly black and white (shaded with cinnamon-bufi on rump).
b. Underparts mainly white.
c. Size small; condylo-basal length of 2 skull less than 90 mm....sheldoni (p. 62).
c’. Size large; condylo-basal length of 9 skull more than 90 mm.
d. Skull shorter; condylo-basal length in 9 less than 98 mm...caligata (p. 59).
d’, Skuil longer; condylo-basal length in 9 more than 98 mm. -nwaria (p. 66).
b’. Underparts d was (blackish brown, soiled whitish, or clay color).
c. Skull coe, short and broad; condyle barca length in ¢ less than 101
mm.; in 9 less than 98 mm.
d. Nasals shorter, rarely extending back of posterior ends of premaxillae.
okanagana (p. 64).
d’. Nasals longer, always extending back of posterior ends of premaxillae.
: vigilis (p. 61).
ce’. Skull relatively long and narrow; condylo-basal length in § more than 101
mm.; in 2 more than 95 mm.
d. Upperparts whiter; skull relatively broader (ratio of zygomatic breadth to

cond ylo-basal lensth 64-67)+.- 52:27 a2 cascadensis (p. 67).
d’. Upperparts blacker; skull relatively narrower (ratio of zygomatic breadth
to cond ylo-basal length: 62-64). - 2: -= =. S-2.2- = ee eee oxytona (p. 63).

a’. Upperparts mainly brownish, yellowish, drab, or buffy.
6b. Upperparits of solid colors (not grizzled).
c. Upperparts brownish drab, buffy, or russet......-.-222-.-2-2 olympus (p. 69).
c’. Upperparts dark vandyke brown, blackish brown, or black.
a Posterior border of nasals deeply emarginate; color dark vandyke brown.
vancouverensis (p. 70).
d’. Posterior border of nasals not deeply emarginate; color blackish brown or

black <2 220s. cc. ss-ns Moa yes ee. i vigils (p. 61).
6’. Upperparts of mixed colors (grizzled).

c. Sides of neck with conspicuous buffy patches.
d. Underparts distinctly reddish.
es. Orowir chestnuts.) aes eee eee Ay ee warrent (p. 52).
e’. Crown blackish brown.
f. Long hairs on fore back with a subterminal band of black or brown.

nosophora (p. 46).
J’. Long hairs on fore back without a subterminal band of black or brown.

dacota (p. 49).
d’. Underparts not distinctly reddish.

e. Size smaller; condylo-basal length of ? skull 71-80 mm.; of ¢ skull

85-87 .1
jf. Fore back with a distinct buffy mantle.
g. Underparts darker; size smaller. —- - 3222223325 parvula (p. 44).
g’. Underparts paler; size larger......-. 5. <2 22e53=eeeee avara (p. 41).
J’. Fore back without a distinct buffy mantle........-- engelhardtt (p. 45).
e’. Size larger; condylo-basal length of 9 skull 77-90 mm.; of ¢ skull 84-97
mm.
f. Face markings and neck patches reduced ........-..---- obscura (p. 53).
jf’. Face markings and neck patches prominent.
g. Hind feet darker (hazel or chestnut-brown)......----- luteola (p. 50).

g’. Hind feet lighter (tawny to light buff).
h. Larger; condylo-basal length of ¢ skull more than 90 mm.; of 2

more thani$4 mm?) 1952. 5 sense tee eee eee flavientris (p. 39).
h’. Smaller; condylo-basal length of ¢ skull less than 90 mm.; of 2
less than 84 mim: 24. S24 520 2a. eee sterrae (p. 43).

1 Adult male of parvula and engelhardti unknown.

1915.] MARMOTA MONAX GROUP. D1

ce’. Sides of neck without conspicuous buffy patches.
7S 0, COINPSE VER ST Va ges 2 ee ee ee ene ee ochracea (p. 34).
d’, Tail dark brown or black.
e. Colors pale (underfur on back light pinkish cinnamon to light gray at
tips; belly not deep red).
jf. Skull larger; condylo-basal length more than 90 mm...-.- monax (p. 22).
f’. Skull smaller; condylo-basal length less than 90 mm.preblorum (p. 27).
e’. Colors dark (underfur on back pinkish cinnamon to orange-cinnamon at
tips; belly deep red).
f. Size smaller; condylo-basal length of ¢ skull less than 82 mm. TOELe
less than 80 mm......... .----canadensis oe 30).
f’. Size larger; soncsiba lengthy of . dail more than 82 mm.; of 9
more than 80 mm.
g. Nasals relatively short and wide; width at posterior end in males
maGSEC Mpls Tee Pin eS Se te ignava (p. 29).
g. Nasals relatively long and narrow; width at posterior end in males
less than 11 mm.
h. Size larger; nasals longer, extending considerably back of posterior

enevonpremiaxilaye: 200523. o.5.2 5252 22st St rufescens (p. 25).
h’. Size smaller; nasals shorter, extending little, ii any, back of pos-
feniomend ot premaxillae «5... 2.6. ose 5 petrensis (p. 33).

DESCRIPTIONS OF AMERICAN SPECIES AND SUBSPECiES.

Marmota monax Group.

[Characters under species.]
MARMOTA MONAX (LINNAEUS).

[Synonymy under subspecies.]

External characters.—Size medium; tail relatively short (about 20
to 25 per cent of total length); ears large; posterior pad on sole of
hind foot oval in shape and situated near middle of sole (see Pl.
Ii, fig. 3); mammae—P. 2, A. 3, I. +=8; head without white
markings (except around nose); sides of neck same color as upper-
parts; feet black or dark brown; fore legs overlaid with deep reddish-
colored hairs (hazel to burnt sienna); tail black, dark brown,or (in
ochracea) pinkish cinnamon.

Cranial characters.—Skull with superior outline nearly straight, the
occipital region slightly depressed and rostrum considerably pee not
abruptly depressed from about posterior border of premaxillae; brain-
case broad and noticeably flattened; interorbital region relatively
broad; postorbital processes heavy, projecting nearly at right angles
to axis of skull or slightly forward; width of nasals at posterior end
usually decidedly greater than width of nasal branches of premaxillae;
temporal ridges often not meeting in old age, or sometimes meeting
but not united (rarely fused into a sagittal crest) ; floor of basi-occipital
nearly flat, bordered on each side by low processes which converge
posteriorly and sometimes meet near inferior lip of foramen magnum;

1Audubon and Bachman give the weight of one as 9 pounds 11 ounces; a writer in Forest and Stream,
LV, p. 424, gives the average weight of 158 as 10 pounds; heaviest, 133 pounds.

99 NORTH AMERICAN FAUNA. | [No. 87.

palate abruptly truncated at posterior border; interpterygoid fossa
relatively wide; palatal foramina contracted anteriorly; molar teeth
heavy; maxillary tooth rows approximately parallel; anterior face of
incisors yellowish white or ivory yellow to pale orange-yellow.
Color.—General tone of upperparts grayish- or reddish-brown, or
Gn ochracea) vinaceous-cinnamon, grizzled with white, light buff, or
cinnamon-buff; underfur of upperparts at base dark mouse gray to
blackish brown, succeeded by pallid neutral gray, light ochraceous-
buff, pinkish-, vinaceous-, or orange-cinnamon; long hairs dark
chestnut-brown, blackish brown, or (in ochracea) orange-cinnamon,
tipped with light buff, pale ochraceous-buff, or white; top of head and
face varying from dark hair-brown or benzo brown to vandyke- or
clove-brown; sides of face ight buff, ochraceous-buff, or white, mixed
with brown; borders of nose, lips, and chin, white or buffy white;
underparts buffy white, light ochraceous-buff, pinkish cinnamon,
tawny, hazel, or burnt sienna, more or less varied with brown, the
bases of hairs often blackish brown or black; fore legs and feet black,
blackish brown, dark chestnut-brown, fuscous, or hazel Gin ochracea),
the legs and thighs (except in ochracea) overlaid with burnt sienna,
Sanford’s brown, or tawny; hind legs and feet similar, but thighs
often paler, sometimes overlaid with pinkish cinnamon; tail black or
blackish brown, vandyke- or clove-brown, or (in ochracea) pinkish
cinnamon, more or less grizzled with buffy white or cinnamon-buff;
ears drab-gray on both surfaces, often clothed with fuscous hairs.
Geographic distribution.—From eastern Alaska (Fortymile Creek),
Great Slave Lake, York Factory (Hudson Bay), East Main and
Hamilton Rivers, Quebec, south to northern Idaho in the Rocky
Mountains, and east of the Great Plains to northern Arkansas, north-
ern Alabama, and southern Virginia; west in the United States to
eastern parts of North Dakota, Kansas, and Oklahoma. (See fig. 1.)

MARMOTA MONAX MONAX (LinnagEwvs).
SOUTHERN WOODCHUCKE.
(Pl. IV, fig..2; Pl. V, fig. 1: Plo xX fesse)

[ Mus] monaz Linnaeus, Syst. Nat., ed. 10, I, 1758, p. 60.

[Glis] monax Erxleben, Syst. Regn. Anim., 1777, p. 361.

Arctomys monaxz Schreber, Siugthiere, IV, 737, 1782; Plate CCVIITI, 1780.
[ Marmota] monax Trouessart, Cat. Mamm., Suppl., 1904, p. 344.

Type locality — Maryland.

Distribution—Middle eastern United States from Pennsylvania,
New Jersey(?), Ohio, Indiana, Illinois, and Iowa, south to the north-
ern parts of South Carolina, Georgia, Alabama, and Arkansas; west
to eastern Kansas.

Characters.—Size large; colors pale, the underfur grayish white;
underparts little, if any, darker than upperparts; skull massive.

1915.) MARMOTA MONAX GROUP. 23

Color.—General tone grayish brown above, with a faint buffy tinge;
underiur blackish brown at base succeeded by pallid neutral gray,
faintly (rarely strongly) tinged with light ochraceous-buff, strongest
on hinder back; long hairs blackish brown, broadly tipped with white

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p22? petrensis |
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| 4. ignava

Po”? tutescens
16.» preblorum |

Fic. 1.—Distribution of the Marmota monaz group. Unshaded areas within the range of the group
indicate lack of definite knowledge as to the subspecies occurring there.

or buffy white; top of head and face varying from benzo brown to
clove brown; sides of nose and borders of lips and chin buffy white;
sides of face mixed brown and buffy whité; feet and legs black or
blackish brown (rarely fuscous or dark chestnut-brown), the hairs on
legs tipped with tawny or hazel; tail blackish brown, sparingly

24 NORTH AMERICAN FAUNA. | [No. 37.

grizzled with buffy white; underparts buffy white or light ochraceous-
buff, the bases of hairs blackish brown.

Skull.—Largest of the group; relatively long and narrow; sagittal
crest well developed (in adults) ; interorbital region broad; nasals long
and broad, usually squarely truncated posteriorly; premaxillae nar-
row; palate long, extending from 2 to 4 mm. behind plane of molars;
palatal foramina rather narrow; interpterygoid fossa broad; audital
bullae moderately inflated.

Measurements.—Adult male from Gunston, Va: Total length, 665;
tail vertebrae, 153; hind foot, 88. Adult male from Sandy Spring,
Md.: 573; 145; 82. Average of 4 adult females from District of
Columbia and Virginia: 557; 139; 83. Skull: Adult male:! Condylo-
basal length, 97.3-102.5 (average 97.8); palatal length, 55-59.5
(57.4); postpalatal length, 34.5-37.7 (86.2); length of nasals, 39.7—41.8
(40.8); zygomatic breadth, 63-69.2 (65.9); breadth across mastoids,
44.5-50.2 (46.9); least interorbital breadth, 25-29.2 (27.1); breadth
of rostrum, 20.6-23.7 (21.7); maxillary tooth row, 21-21.9 (21.3).
Adult female:? Condylo-basal length, 90.4-91.8 (91.2); palatal length
50.2-56 (53.1); postpalatal length, 33.7-36.3 (34.9); length of nasals,
36.8-40.6 (38.7); zygomatic breadth, 59.5-62 (61.2); breadth across
mastoids, 44-45.8 (44.7); least interorbital breadth, 23.5-26.5 (24.9);
breadth of rostrum, 19.7-21.3 (20.5); maxillary tooth row, 20.5-22.4
(2112).

Remarks.—The southern woodchuck is a rather large animal of
massive skull and pale color. With a wide range in the Middle
States, it grades into a smaller and darker form (rufescens) in the
southern parts of New York, Michigan, and Wisconsin. Specimens
from northern Illinois (Lake Forest and Willow Springs) and northern
Indiana (Rose Lawn and Lake Maxinkuckee) are typical of monaz, but
those from Delavan, Wis., and southwestern Michigan are inter-
mediate between monax and rufescens. Specimens from Marble Cave,
Mo., and from Johnson County, Iowa, are typical, but a small series
from the vicinity of Lawrence, Kans., seems to indicate the presence
there of a large form, skulls of females, especially, being much larger
than those of females from Maryland and Virginia and equaling in
size skulls of males from that region. The hind feet also average
about 16 per cent longer in the Kansas animal (3 specimens). Addi-
tional material from that region may require naming the form.

Specumens examined.—Total number, 75, as follows:

Alabama: Ardell, 3.

District of Columbia: Washington, 6.
Georgia: Young Harris, 1.

1 Six specimens from Maryland, Virginia, West Virginia, and Pennsylvania.
2 Five specimens from Virginia, District of Columbia, and Pennsylvania.

1915.] MARMOTA MONAX GROUP. 95

Hlinois: Cook County, 1; Lake Forest, 1;! Ozark, 2;! West Northfield, 1; Willow
Springs, 1.1

Indiana: Lake Maxinkuckee, 1; Marion County, 1;2 Rose Lawn, 1.

Iowa: Iowa City, 4;3 Johnson County, 4;3 Wall Lake, 1; no specific locality, 3.8

Kansas: Douglas County (near Lawrence), 6;* Lawrence, 3.4

Kentucky: Mammoth Cave, I.

Maryland: Plummer Island, 2; Sandy Spring, 1; Simpsonville, 1; near Wash-
imeton, D. C., 1.

Michican: Cass County, 1;° Dowagiac, 1.°

Missouri: Marble Cave, 2.

Worth Carolina: Magnetic City (base of Roan Mountain), 7; Roan Mountain
(altitude 3,000-4,000 feet), 5.

Ohio: Hicksville, 1.1

Pennsylvania: Beaver, 2;° Chester County, 3; Fair Oaks, 1;° Kennett Square, 1;7
Landenberg, 1;’7 Marpie, 1;7 Meadville, 1; Round Island, Clinton County, 2.’

Tennessee: Dover, Stewart County, 1; Duck River, 6 miles southwest of
Waverly, 2; Highcliff, 1.

Virginia: Bluemont, 1; Clarke County, 1; Doswell, 1; Fairfax County (Potomac
River), 3; Fincastle, 1; Fredericksburg, 2; Gunston, 1; Peaks of Otter, 1;
Washington, 2.

West Vireinia: Franklin, 3; Jobs Knob, 1; North Mountain, Hardy County, 1;
Rowleysburg, 1.

MARMOTA MONAX RUFESCENS Howe tt.
RUFESCENT WOODCHUCK.
CE oLVestiowo: bl Vite. 2sPlox fe?)
Marmoita monaz rufescens Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 13.

Type locality. — Elk River, Minn.

Nsirvbution.—EKastern North Dakota, central and southern Minne-
sota, Wisconsin, and Michigan, southern Ontario, greater part of New
York (including Long Island), and higher parts of western Massa-
chusetts.

Characters.—Similar to monaz but colors much redder, both above
and below, the underfur on back pinkish cimnamon instead of light
buff; similar in color to zgnava, but averaging paler; skull decidedly
smaller than that of monaz, but much larger than that of canadensis.

Color.—Underfur of upperparts varying from fuscous-black to dark
mouse gray at base, succeeded by a broad area of light pinkish cinna-
mon shading to pinkish cinnamon on hinder back and to orange-
cinnamon on hind legs; long hairs dark chestnut-brown subterminally,
tipped with hght buff; top of head and face benzo brown to clove
brown; sides of face light buff, more or less mixed with brown; fore
legs black, overlaid with burnt sienna; hind legs similar but usually
less intensely red; feet black or blackish brown; tail vandyke brown

1Collectiov Field Mus. Nat. Hist. 8 Collection Univ. of Michigan.
2 Coliection Amer. Mus. Nat. Hisé- 5 Collection Carnegie Museum.
3 Collection Univ. of Iowa. 7 Collection Acad. Nat. Sci. Philadelphia.

4Collection Kansas Univ. Mus.

26 NORTH AMERICAN FAUNA, [No. 37.

to clove brown or black; underparts varying from burnt sienna to
Sanford’s brown, more or less mixed with tawny, ochraceous-buff,
and black (general tone of under parts usually reddish but sometimes
mixed brown and buff with little red); tail vandyke brown to clove
brown or black. Variation: Specimen from Essex County, N. Y.: Very
dark chestnut-brown above (the underfur pinkish cinnamon), moder-
ately grizzled on fore back with buffy white; hinder back solid brown.

Skull.—Similar to that of monaz, but decidedly smaller and
relatively broader across zygomata; much larger than that of cana-
densis; slightly larger than that of wgnava, but narrower across
orbits and with narrower nasals.

Measurements.—Adult male: Total length, 520-582 (average,
548); tail vertebrae, 135-155 (143); hind foot, 81-85 (83). Adult
female:? Total length, 545-608 (571); tail vertebrae, 145-170 (156);

hind foot, 838-89 (85). Skull: Adult male:? Condylo-basal length,
' 81.3-94.4 (88.6); palatal length, 50.7-54.5 (51.8); postpalatal
length, 32-36.4 (33.5); length of nasals, 33.9-39.8 (86.8); zygo-
matic breadth, 53.4-64.7 (60.2); breadth across mastoids, 40.3-46
(43.2); least interorbital breadth, 21.7-25.4 (23.8); breadth
of rostrum, 18-21.2 (19.4); maxillary tooth row, 18.7—21.8 (20.1).
Adult female:* Condylo-basal length, 84-88.9 (85.8); palatal
length, 49.1-52.4 (50.6); postpalatal length, 31.5-31.8 (31.6); length of
nasals, 33.4—-39.1 (35.6); zygomatic breadth, 57.5-62 (59.9); breadth
across mastoids, 40.6—48.8 (42.4); least interorbital breadth, 22.2-26.3
(23.6); breadth of rostrum, 18.2-20.4 (19); maxillary tooth row,
Po 937):

Remarks.—The woodchucks of the Northern States, from Minne- —
sota to New York, differ from typical monax of the Middle States
in having smaller skulls and darker colors, the underfur especially
being more strongly reddish. The race shows a strong tendency
toward melanism, particularly in New York and southern Ontario.
The New York series differs from the Minnesota series in somewhat
longer and relatively narrower skulls, being, therefore, more like
monaz, but the differences between the two extremes are too slight
and inconstant to warrant recognition of another form. Specimens
from the Catskill Mountains are fairly typical of rufescens, but those
from the Hudson Valley and Lake George are clearly intermediate,
the skulls being almost typical of monaz and the skins. of rufescens.
These show no approach to the New England form (preblorum).
Two skulls (without skins) from Easthampton, Mass., are fairly
typical of rufescens, indicating that this race probably occupies the
higher parts of western Massachusetts. A specimen from Lake of
Bays, Ontario (east side of Georgian Bay), considered intermediate
between rufescens and canadensis, is very deep red below—fully

1 Five specimens from Elk River and Fort Snelling, Minn. 3 Eleven specimens from Minnesota.
2 Five specimens from Fort Snelling, Minn. 4 Five specimens from Minnesota.

1915.] MARMOTA MONAX GROUP. 7.

as dark as ignava. The northern limit of the range of rufescens is
assumed to be about the latitude of Ottawa, but no material is
avallable from that part of Ontario, excepting the specimen men-
tioned above.

Specimens examined.—Total number, 231, as follows:

Massachusetts: Easthampton, 2.

Michigan: Ann Arbor, 3;1 Au Sable River, Oscoda County, 1;1 Genesee, 1;
Rush Lake, Huron County, 1; Sand Point, Huron County, 1.1

Minnesota: Elk River, 10; Fort Snelling, 17;? Princeten, 1.

New York: Adirondack Mountains, Essex County, 76 (skulls only); Ardsley, 1;
Amber, 2; Brantingham, 1; Croton Lake, 1; Dutchess County, 1;? Elizabeth-
town, 2; Essex County, 8; Halcotisville, 10;* Hastings, 1;° Highland Falls,
1; Lake George, 11; Leyden, 2; Locust Grove, 15; Lyons Falls, 5; Miller
Place, 4;+ Orange County, 1; Owego, 1; Oyster Bay, 2; Peterboro, 4; Piseco,
1; Schroon,1; Schroon Lake, 4; Sing Sing, 4; Suffolk County, 2;%° Troy, 3;
Tupper Lake, 1;° Wells, 3; no specific locality, 7.

North Dakota: Fargo, 3;7 Grafton, 1; Leonard, 1. §

Ontario: Lake of Bays, 1; Lorne Park, 7.

Wisconsin: Bridgeport, 1; Delavan, 2; Milton, 2;° Racine, 1.

MARMOTA MONAX PREBLORUM Howe tt.
New ENGitaAND WooDcHUCKE.
(Lelie OLD bss Stee 39 bo stools tcciers Se ol el es @] Geo sega)
Marmota monax preblorum Howell, Proc. Biol. Soc. Washington, XX VIT, 1914, p. 14.

Type locality.— Wilmington, Mass.
Distribution.—Southern New England, from Connecticut to central
Vermont and New Hampshire and southern Maine.
Characters.—Size medium (smaller than rufescens, larger than
canadensis); colors pale (redder than monax, but red not so dark
as in canadensis or rufescens); skull smaller and relatively narrower
than that of rufescens. |
Color.—Adult: Underfur on upperparts pinkish cimnamon to light
pinkish cinnamon, the bases of hairs fuscous-black; long hairs
blackish brown, extensively tipped with white or light buff; top of
head and face dark hair-brown to clove brown; sides of face light
buff; fore legs burnt sienna or Sanford’s brown, the bases of hairs
often black; hind legs somewhat paler, shading to pinkish cinna-
mon; feet black or blackish brown; tail clove brown to black,
much mixed with cinnamon; underparts pinkish cinnamon or San-
ford’s brown, varied with light buff. Young (specimen from Saun-
1 Collection Univ. of Michigan.
2 Three in collection Field Mus. Nat. Hist.; five in Amer. Mus. Nat. Hist.
3 Collection Amer. Mus. Nat. Hist.
4 Collection A. H. Helme, Miller Place, N. Y.
5 Collection Field Mus. Nat. Hist.
6 Collection Mus. Comp. Zool.
7 Two in coliection Fargo College; one in N. Dak. Agr. College.

8 Collection N. Dak. Agr. College.
9 Collection H. H. T. Jackson, Washington, D. C.

a

98 NORTH AMERICAN FAUNA. [No. 87.

derstown, R. I.): General tone, both above and below, pinkish buff
(the hairs extensively tipped with that color and the bases of same
shade), becoming pinkish cinnamon on hinder back; subterminal
band of each ‘hair blackish brown; fore legs Sanford’s brown; hind
legs tawny; top of head hair-brown.

Skull.—Similar to that of rufescens, but smaller and relatively
narrower, especially the rostrum and interorbital region; bullae
smaller; much larger than that of canadensis, with longer, slenderer
rostrum and longer nasals.

Measurements.—Adult male: Total length, 418-608 (average 515);
tail vertebrae, 105-149 (120); hind foot, 75-80 (77.7). Adult female:?
465-600 (547); 100-157 (141); 69-88 (77). Skull: Adult male:*
Condylo-basal length, 84-89 (86.2); palatal length, 49.9-51.6 (50.7);
postpalatal length, 31.2-33.4 (32); length of nasals, 34.3-37.7 (35.9);
zygomatic breadth, 57.3-60.4 (58.7); breadth across mastoids,
41.38-44.3 (42.8); least interorbital breadth, 20.7-22.5 (22.1);
breadth of rostrum, 17.2-18.6 (18.2); maxillary tooth row, 18.2-
20.2 (19.1). Adult female:* Condylo-basal length, 79.1-86.6 (83);
palatal length, 47.7-51.6 (49.5); postpalatal length, 28-31.8 (30.1);
length of nasals, 33.7-36.8 (85.6); zygomatic breadth 54-58 (56.6) ;
breadth across mastoids, 37.7-42 (40.3); least interorbital breadth,
19.8-22.6 (21.6); breadth of rostrum, 16-19.5 (18); maxillary tooth
row, 18-20.4 (19.3).

Remarks—The New England woodchuck is noticeably smaller
and paler than rufescens, and larger and paler beneath than canaden-
sis. In skull characters it most resembles rufescens and probably
intergrades with it where their ranges meet, but no intermediate
specimens have been examined. A specimen from Liberty Hill,
Conn., has somewhat redder underparts than the Massachusetts
series, its skull being fairly typical of preblorum. Specimens of
rufescens from the Hudson Valley, however, approach monaz rather
than preblorwm in skull characters. Intergradation with canadensis
also undoubtedly occurs, but material from northern New England
is needed to show where the two forms come together. Specimens
from Rutland, Vt., agree in skull characters with preblorum, but one
of the two skins examined is somewhat redder below than in Massa-
chusetts examples.

Specimens examined.—Total number, 38, as follows:

Connecticut: East Wallingford, 1;° Liberty Hill, 1.°
Maine: Eliot, 1; Norway, 1.6

1 Seven specimens from eastern Massachusetts.

2 Nine specimens from eastern Massachusetts and southern New Hampshire.
8 Five specimens from Wilmington and Wareham, Mass.

4 Four specimens from Wilmington and Lunenburg, Mass.

5 Collection Am. Mus. Nat. Hist.

6 Collection Mus. Comp. Zool.

1915.] MARMOTA MONAX GROUP. 29

Massachusetts: Essex County, 1; Haverhill, 1;1 Lunenburg, 4; Newtonville,
2;1 Sherborn, 1; Springfield, 2;! Wareham, 2;1 Wayland, 1;1 Wilmington, 8;
mebimn, iG

New Hampshire: Charlestown, 2; Gauipee. 2; Webster, 3.7

‘Rhode Island: Saunderstown, 1.

Vermont: Ruiland, 3.”

MARMOTA MONAX IGNAVA (Bangs).
LABRADOR WOODCHUCK.
@iye ae 4 Pl Mt fe 1)

Arctomys ignavus Bangs, Proc. New England Zool. Chib, 76 1899; p-3:
[Arctomys monaz] ignavus Elliot, Synop. Mamm. N. Am., Field Columb. Mus., Zool.

pee, IT, 1901, p.105.

[ Marmota monaz] ignavus Trouessart, Cat. Mamm., Suppl., 1904, p. 344.
Marmota ignava Miller, Bul. 79, U.S. Nat. Mus., 1912, p. 292.

Type locality.— Black Bay, Straits of Belle Isle, Labrador.

Nstribution.—Known only from vicinity of type locality; probably
north to Hamilton Inlet.

Characters.—Size much larger than canadensis, nearly equaling
rufescens; similar in color to rufescens (much darker than canadensis) ;
skull short and broad with very broad nasals.

Coior—Underfur of upperparts blackish brown at base (a shade
darker than in canadensis), succeeded by a broad area of orange-
cinnamon; long hairs blackish brown subterminally, tipped with
pale ochraceous-buff or buffy white; top of head and face vandyke
brown or clove brown; sides of nose and borders of lips and chin
buffy white; sides of face ight ochraceous-buff more or less mixed
with brown; feet and legs black, or very dark brown, the legs and
thighs overlaid with burnt sienna; tail blackish brown, usually with
little or no white grizzling; underparts burnt sienna mixed with
_black, varying to tawny and in some individuals more or less mixed
with pinkish cinnamon or pale buff; in others, mixed blackish brown
and buff below without any red.

Skull—similar to that of rufescens, but shorter and. relatively
broader; nasals shorter and relatively ante posteriorly; premaxil-
lae averaging narrower; bullae smaller and less inflated; sagittal
crest more prominent; incisors with numerous shallow longitudinal
grooves on outer face. Compared with preblorum: Skull about same
length, but relatively broader; rostrum broader; nasals shorter and
broader posteriorly; molars heavier. Compared with canadensis:
Skull much larger, with ee heavier sagittal crest and less
inflated bullae.

Measurements —Adult male:* Total length, 480-562 (average,
536); tail vertebrae, 111-155 (137); hind foot, 74-86 (80.8). Adult

1 Collection Mus. Comp. Zool.

2 Collection Amer. Mus. Nat. Hist.
3 Seven specimens from vicinity of type locality.

30 NORTH AMERICAN FAUNA. [No. 37.

female: 496-556 (528); 102-147 (126); 75-80 (78.6). Skull:
Adult male: 1 Condylo-basal length, 82.5-87.4 (85.1); palatal length,
48.5-51.2 (50); postpalatal length, 30.6-33 (31.7); length of nasals,
31.5-35.6 (84.1); zygomatic breadth, 57.5-63.4 (60.5); breadth
across mastoids, 40.6—-44 (41.7); least interorbital breadth, 24-26.8
(25); breadth of rostrum, 18—20.3 (19.2); maxillary tooth row, 19.9-
20.8 (20.4). Adult female: ! Condylo-basal length, 79.7-84.4 (82.2);
palatal length, 47-49.4 (48.2); postpalatal length, 29.2-31.7 (80.6);
length of nasals, 31.38-34.2 (82.6); zygomatic breadth, 55.4-58.7
(57.1); breadth across mastoids, 39-41.5 (40.6); least interorbital
breadth, 22.5-25 (23.8); breadth of rostrum, 17.6-19.7 (18.7); max-
illary tooth row, 19-20.4 (19.9).

Remarks.—The Labrador woodchuck is a strongly marked form of
the monaz group, much larger and darker than canadensis, its nearest
neighbor. Indeed, it might be considered a distinct species were it
not practically certain that the ranges of zgnava and canadensis are
contiguous and that more material from the region between Murray
Bay and the Straits of Belle Isle would show intergradation. In
color and size the present form more closely resembles rufescens than
canadensis and the skull is more like that of preblorum than that of
any other form, but zgnava is much darker than preblorum. The
subspecies is reported to be common on the coast in the vicinity of
Black Bay, and is said to be found about the head of Hamilton
Inlet,? but the limits of its range are not known. Stearns reports
woodchucks ‘‘common at Mingan, growing scarce toward Bonne
Esperance.” ®

Specumens ecamined.—Total number, 15, as follows:

Labrador:* Ailik, Peter’s Cove, 1; Black Bay, 4; L’ Anse au Loup, 10.

MARMOTA MONAX CANADENSIS (ERxLeBeEy).
CANADA WoopcHUCK.
(PL Vi, fies 1; PL er fie 74)

[Glis] canadensis Erxleben, Syst. Regn. Anim., 1777, p. 363.

Mus empetra Pallas, Nov. Spec. Quad., Gli. Ord., 1778, p. 75.

Arctomys sibila Wolf, Linne’s Natursyst., II, 1808, p. 481. (Name proposed to include
Arctomys empetra Pallas and Arctemys pruinosa Gmelin, supposed to be the same).

Arctomys melanopus Kuhl, Beitrage, 1820, p. 64.

Arctomys marmota canadensis Kuhl, Beitrage, 1820, p. 64.

Arctomys empetra Sabine, Trans. Linn. Soc. London, XIII, 1822, p. 584; Richardson,
Fauna Boreali-Americana, I, 1829, p. 147.

Arctomys monax melanopus hanes Brae! Acad. Nat. Sci. Philadelphia, 1S97, pa 30:

Arctomys monax canadensis Allen, Bul. Amer. Mus. Nat. Hist., X, 1898, p. 456.

| Marmota monazx] canadensis isamiacns Cat. Mamm., Suppl., 1904, p. 344.

1 Six specimens from vicinity of type locality.

2Low,A.P. Ann. Rept. Geol. Surv. Canada, VIII, 1895 (1897), p. 320L.
3 Stearns, W.A. Proc. U.S. Nat. Mus., VI, 1883, p. 115.

4 All incollection Mus. Comp. Zool.

1915.] se MARMOTA MONAX GROUP. 311

Type locality —‘‘Canada et ad fretum Hudsonis”—here fixed at
Quebec, Quebec.

Disiribution.—Greater part of interior of Canada, from Great Slave
Lake and York Factory south to southern Alberta (Red Deer), cen-
tral Saskatchewan (Cumberland House), northern Minnesota, north-
ern Wisconsin, northern Michigan, central Ontario, southern Quebec,
New Brunswick, and Nova Scotia; northern and eastern limits of
range in Quebec unknown.

Characters.—Size small; sexes about same size; colors strongly ae
dish, above and below; Jol small without pronounced sagittal crest.

Soe —Underfur on upperparts blackish brown at base, succeeded
by pinkish cmnamon or light pinkish cinnamon; long hairs blackish
brown subterminally, tipped with white or pinkish buff; top of head
and face hair-brown, sometimes shading to clove brown; sides of
face light buff; feet and legs black, blackish brown, or fuscous, the
legs and thighs overlaid with burnt sienna; tail blackish brown, con-
siderably grizzled with cinnamon-buff or light buff; underparts deep
tawny or burnt sienna sometimes varied with buff and moderately
mixed with black. Melanistic specimens are rarely found, but one
from Aitkin, Minn., is glossy blackish brown all over.

Skull.—Smallest of any member of the group; shorter and rela-
tively broader than that of preblorum, with short, broad rostrum;
much smaller than that of ignava, with sagittal crest only slightly
developed; nasals narrowed posteriorly; bullee relatively large,
smoothly rounded, and considerably inflated.

Measurements.—Adult male: Total length, 510-515 (average 513);
tail vertebrae, 108-109 (108.5); hind foot, 74-78 (76);1 average of
three adult males from Mackenzie and Riperta: 500-124; 75s - Adult,
female:? 508-560 (536); 131-140 (186); 69-76 (73). Skull: Adult
male:* Condylo-basal length, 75—-80.8 (78.1); palatal length, 43.7—48.2
(45.7) ; postpalatal length, 27.5-29.6 (28.7); length of nasals, 29.9-32
(31.3); zygomatic breadth, 53-56.6 (54.1); breadth across mastoids,
36.4-39.4 (38); least interorbital breadth, 18.4-22.7 (20.7); breadth
of rostrum, 15.1-18 (16.6); maxillary tooth row, 18.2-19.3 (18.8).
Adult female: + Condylo-basal length, 77.4-80 (78.9); palatal length
45.6-47.2 (46.2); postpalatal length, 28.2-30.5 (29.3); length of
nasals, 30.5-34.5 (32.6) ; zygomatic breadth, 52.8—58.2 (54.8) ; breadth
across mastoids, 36.5-39.8 (37.8); least interorbital breadth, 19.8-22
(20.9); breadth of rostrum, 14.5-17.8 (16.3); maxillary tooth row,
18.1-19.2 (18.6).

Remarks.—The Canada woodchuck has the most extensive dis-
tribution of any of the American forms and over the greater part of
_ 1 Two specimens from Murray Bay, Quebec.

2 Three specimens from Quebec and Ontario.

3 Eight specimens from Mackenzie, Alberta, Manitoba, Quebec, Nova Scotia, and northern Wisconsin.
4 Seven specimens from Mackenzie, Manitoba, Ontario, Nova Scotia, and Quebec.

o2 NORTH AMERICAN FAUNA. [No. 87.

its range shows comparatively little variation. Although markedly
smaller than rufescens it intergrades with that form wherever their
ranges meet, intermediate examples having been examined from
Tower and Two Harbors, Minn., and Lake of Bays, Ontario. Inter-
gradation with ignava seems highly probable, though not shown by
the materialin hand. Murray Bay specimens have very much smaller
skulls than typical zgnava from the Labrador coast, and no specimens
have been examined from the intervening region. Two specimens
from Nova Scotia show slight approach to ignava, the skins being
the same color except that the bases of the hairs on the back are
browner. The skull of one of these specimens is typical of canadensis,
the other is a little larger, with somewhat broader nasals and a well-
marked sagittal crest.

A specimen from Porcupine Mountains, Mich., resembles cana-
densis in general coloration, but is extensively mixed with black
both above and below. Its skull is slightly longer than skulls of
typical specimens from Quebec. A specimen from Mount Mansfield,
Vt., and one from Columbia Falls, Me., (both without skulls), are pro-
visionally referred to canadensis, the former agreeing in color with
the typical form, the latter with the dark Nova Scotia form. Speci-
mens from southern Mackenzie and northern Alberta are practically
identical in coloration with the Quebec series, but their skulls average
longer and narrower, thus showing approach to ochracea. A single
young specimen in very worn pelage from near the head of Finlay
River, British Columbia, seems referable to canadensis, but more ma-
terial. from that region may necessitate its reference to ochracea.
Wocdchucks of this group are reported by Edward A. Preble as
occurring at Fort Grahame, on Finlay River, and at Hudson’s Hope.

Specumens examined.—Total number, 45, as follows:

Alberta: Athabaska River (near Fort McMurray), 1; McLeod River, 1; Peace
River Landing, 2;1 Red Deer, 1; South Edmonton, 1.

British Columbia: Finlay River (near head), 1.

Mackenzie: Fort Liard, 1; Fort Simpson, 5; Little Buffalo River, 1.

Maine: Columbia Falls, 1.

Manitoba: Oxford House, 1; Trout Lake, 4; York Factory, 1.

Michigan: Porcupine Mountains, 1;? Whitefish Point, Chippewa County, 1.?

Minnesota: Aitkin, 1;? Tower, 1; Two Harbors, 1.

New Brunswick: Arthurette, 2;* Scotch Lake, 1.

Worth Dakota: Pembina, 1.

Nova Scotia: Newport, 2.4

Ontario: Devils Portage, Mattagami River, 1;> James Bay, 1; Moose River
(near Hudson Bay) 2.4

Quebec: Murray Bay, 7.?

Vermont: Mount Mansfield, 1.

Wisconsin: Conover, 1.?

1 Collection Victoria Mem. Mus. 4 Collection Amer. Mus. Nat. Hist.
2 Collection Field Mus. Nat. Hist. 5 Collection Carnegie Mus.
3 Collection Univ. of Michigan.

1915.] MARMOTA MONAX GROUP. 30

MARMOTA MONAX PETRENSIS susspr. Nov.
BrRitTIsH COLUMBIA WOODCHUCK.
CBI Vas fie, 2.)

Type from Revelstoke, British Columbia. Adult 3, No. 203532,
U. S. Nat. Mus., Biological Survey collection. Collected May 12,
1890, by W. Spreadborough; original number, 170.

- Distribution.—Interior ranges of southern British Columbia and
adjacent parts of United States, from Barkerville, British Columbia,
south to Thompson Pass, Idaho.

Characters.—Similar to canadensis, but skull larger and relatively
longer.

Color.—Practically the same as canadensis.

Skull.—Similar to that of canadensis, but larger and relatively
longer; nasals rather short, projecting but little back of posterior
ends of premaxillae; bullae rather small. Compared with rufescens:
Smaller with shorter nasals and smaller, rounder bullae.

Measurements.—Adult male (type): Total length, 540; tail verte-
brae, 127; hind foot, 76; specimen from Barkerville, British Columbia:
460; 106; 72. Adult fernale (specimen from Barkerville, British
Columbia): 505; 125; 68. Skull: Adult male (type): Condylo-basal
length, 86; palatal length, 50.3; postpalatal length, 31.8; length of
nasals, 32.5; zygomatic breadth, 57.4; breadth across mastoids, 42;
least interorbital breadth, 23; breadth of rostrum, 18.6; maxillary
tooth row, 20. Adult female (specimen from Barkerville, British
Columbia): Condylo-basal length, 79; palatal length, 46.5; post-
palatal length, 29.5; length of nasals, 32.4; zygomatic breadth, 53.2;
breadth across mastoids, 36; least interorbital breadth, 19.4; breadth
of rostrum, 17.6; maxillary tooth row, 18.8.

Remarks.—This is a slightly differentiated form of monaz, occu-
pying an area somewhat segregated from the rest of the species.
It is known at present from only a few specimens and the limits of
its range can not definitely be determined. It probably occurs
sparingly throughout the heavily timbered mountain sides of the
northern Rocky Mountains in Montana, Idaho, and British Co-
lumbia, but whether confined entirely to the west slope or not (as
seems probable) has not yet been ascertained. This subspecies has
no direct connection with monaz or rufescens, its range being separated
from theirs by an extensive area of plains, but it undoubtedly inter-
grades with ochracea on the north and possibly with canadensis
through some of the passes of the mountains in southern British
Columbia.

Specumens examined.—Total number, 6, as follows:

British Columbia: Barkerville, 2; Glacier, 2; Revelstoke, 1.
Idaho: Thompson Pass, 1.

1 From dry skin.
69168°—15——3

34 NORTH AMERICAN FAUNA. [No. 37. .

MARMOTA MONAX OCHRACEA Swarts.
OcHRACEOUS WoODCHUCKE.
(PIV; fig. 3: Pll xt fievo.)
Marmota ochracea Swarth, Univ. of California Pub. Zool., VII, 1911, p. 208.

Type locality.—Head of Fortymile Creek, Alaska.

Distribution.—Interior mountain ranges of Yukon and northern
British Columbia, from Fortymile Creek south to the Babine Moun-
tains (and Stuart Lake ?).

Characters.—Similar to canadensis, but paler below and hairs on
back extensively tipped with ochraceous, the underfur more vina-
ceous; tail pinkish cinnamon all around; skull longer and relatively
narrower.

Color.—Underfur of upperparts dark mouse gray at base, suc-
ceeded by a broad area of vinaceous-cinnamon, the latter shading
toward the tips into orange-cinnamon, most intense on hinder back;
long hairs orange-cinnamon, then blackish brown, broadly tipped
on fore back with ochraceous-buff and on hinder back with light
ochraceous-buff; top of head dark hair-brown; sides of nose and face
hight ochraceous-buff; underparts tawny shading to hazel; legs hazel;
feet fuscous or fuscous-black with scattering hazel hairs; tail pinkish
cinnamon all around, the tip clove brown. Variation: Young speci-
men from type locality: Darker below than adult, and feet more
varied with hazel. Immature specimen from Pike River, British
Columbia: Upperparts cinnamon (without plumbeous bases to the
hairs), tipped with pinkish buff; underparts cinnamon-rufous.

Skull.'—Longer and relatively narrower than that of canadensis,
with narrower rostrum and interorbital region; bullae broader.

Measurements. ?—Skull: Subadult (male?) from Babine Mountains,
British Columbia: Condylo-basal length, 81.2; palatal length, 48;
postpalatal length, 30; length of nasals, 33.4; zygomatic breadth,
52.7; breadth across mastoids, 39.8; least interorbital breadth, 19.8;
breadth of rostrum, 17.2; maxillary tooth row, 20.

Remarks.—This subspecies is the most northerly ranging member
of the group. It occurs west of the continental divide in Yukon
and northern British Columbia, but the limits of its range are not
known. Only six skins have been seen. A specimen from near
Teslin Lake, southern Yukon, is slightly darker below than the type
(being uniform kaiser brown); the fore back is darker ochraceous,
and the head blackish brown. Another specimen from Pike River,
British Columbia, is dark blackish brown all over. A series of skulls
without skins from Stuart Lake, British Columbia, is provisionally
referred to this race.

1 No skulls from the type region available; description based on subadult specimen from Babine

Mountains, British Columbia.
2 No external measurements available.

1915.] MARMOTA MONAX GROUP.

30

Specimens examined.—Total number, 15, as follows:

Alaska: Fortymile Creek (at head), 2.1

British Columbia: Babine Mountains, 1 (skull); Pike River, Atlin, 2;2 Stuart
Lake, 8 (skulls); Tacla Lake, 1.

Yukon: Thirty Mile Mountains, near Teslin Lake, 1.8

Cranial Measurements of the Marmota monax Group.

| Pt oa hk) la :
S| lela 8 ed Bale
i 5 | |x my | &
= | a S18 lel SS
| 3 |€/2/2/5 jegiss| 2/8
No. Species and locality. 2)6¥lala|s i|8ei8s| 2/2] Remarks.
% |2 Sas = oF a eB
| SB IS(218/8is le lela
Lule isleifieg lle
= 3 Ba Ss
& SJAla AINE BR 2S |
ST rr a (ees ees a ee Ge Se ee eee
Marmota monar monaz. | | | | |
| get
114009 | Montgomery Co.,Md........--. G | 102. 5 59. 5/37. 7/41. 5)69. 2/50. 2)27.3/23.721.9) Old.
175010 | Plummers Island, Md. ---.-.-.-.-- of 97.556.7/37 |41.8/66 |46. 5/25.9/21.2/21.2) Aduit.
143962 | Peaks of Otter, Va.....--------- $ | 98.1/58.1/36 |40. 5/65. 7/47. 2/27. 2/20. 7,21. 2 Do.
58695 | Roan Mountain, N.C......-.--. S | 94 |57.3/32. 6/38. 7]... ./42.9|24. 4/19. 521.6 Do.
4 447 | Douglas County, Kans......--.- S | 98.7.58.8/36.5/43 |... ./45.7|27.2/19. 7/20. 6| Subadult
125331 | Bluemont, Va........-.- eee 2 | 93.756 35.3140. 6/61. 9145. 8/26. 5/19.9'21.2} Adult.
gee) Washington, D.C_...-...------- Q | 90.458. 4/33. 7/38 [59.5 44. 3/23. 5[21. 3/20. 5 DOs aes
bak epee County, Kans..-.-..--. é pepe ger ee Hag a eee oes ge Be.
eR eee ae se eae? 5. 8/56. : .3/48.5 : P Oo.
616029 | Lake Forest, Ill..........------- Q | 92 |54 |34.2138. 662.746.4126 |22. 5121.6 Do.
Marmota monaz rufescens. | | | |
186521 | Elk River, Minn.........------- | g | 87.2\50.9/32 ee eee 18.7/20.4| Adult; type.
| 2 ee EEE |e ese oe
6 1780 G0--eeeceeceeeee| & | 94434 si3e:4lae lei.zi44’ 25. alai.i19 | Do.
191244 | Locust Grove, N. Y...-.--------| 3 90 |53. 2/32. 2/36. 5|64.5144 |25.5|21 |20. 6 Do.
7) | Adirondack Mountains, N.Y..... g¢ | 91.654. 2/33.7/38. 4/61. 4 44. 1/24. 4/19.8120.8) = Do.
191339 | Elk River, Minn......- seeeeez--) Q | 84.2/49.1)31. 5/34. 5]57. 5/40. 6 22. 5/18. 3 20 | Do.
ee ee Moan =. ----- eae oe a0" | Ae
237) 7 ee Oo ee OSS oe Ae eee ae 2 50. 3/31. 5/33. 4/59. 2 : 6 O.
191250 | Locust Grove, N. Y......-.----- 2 | 86.3/51 32-337 3/60 42 Bs 17.5|20.2 Do.
(8) | Adirondack Mountains, N.Y... Q | 86.851. 4/31.2/37. 1/59.8)/41.9)23. 5/19..2/20. 3 Do.
111091 | Peterboro, N. Y.......--------- Q | 85 HUI ES OBL 22. 8/19. 5/20. 3 Do.
Marmota monaz preblorum. | | | | | |
aes enuetany Mies a g 85.3)50. 5)32 [34. 8157. 4143. 7122. 8118. of18.2} Adult; type.
78354 |..... Cas. ee S$ | 86.1)51. 5/31. 2/36. 3/58. 2/42. 7/22. 5/17. 2/18.3) Adult.
a ee es Ses 3 | 84 |49.9131. 2/36. 2/57.3/41.3/20. 7/18. 2 19.5) Do.
gett Ossipce N. H........----------- S | 85.4)49. 7/31. 6/36 60. 4/43. 6|23 |19.5/18.9| Do.
69235 | Rutland, Vt_........----------- g | 85 (50.5/31 87.958 |42.2)23.5|18.5|20.5) Do.
Ss eee GANG ee ge
78359 |... - Gere ee 2 22S =~-- = -- . 8/49. 9]31. 836. 8157. 8 6/18,
96083 | Lunenburg, Mass.....-...-.---- | 2 | 86.6)51.6)30. 7/36. 5/56. 7/42 |21.4/19.5/20.4) Do.
96142 }_.... LT SS ae ee eee | Q | 81.6/48.7/30 (35.3/58 |41.522.6/18.3)19.6) Do.
j | : |
Marmota monat ignava. | | | |
shee L’Anse au Loup, Labrador ...-- | 3 eral ape Ober 63. 4/41. 7/26. 8 18. $100 5 paul.
[al sae TS aie Aline a a ae 3 2.7/48.7/31 |33.2/59 41. 2/24. 3]18. 5/2 0.
98873 |..... io a | g | 86.5/51 32 |35.4/60. 5142) [24 20.3/20.7, Do.
98876 |_...- “Ce apa at ie ic aap | Q | 84.4)49. 1/31. 7:33. 6/58. 7/41.3/25 |19. 7/19 Do.
9 8875 |..--- (C3 a ee | Q | 84 |49.4:30. 6.34. 2/58. 6)41. 5)24. 3/18. 8)19. 6 Do.
27968 | Black Bay, Labrador..........-) Q | 80.6/47.7.29.731.5 55.840 [23 |17.6)19.5) Do.
| H
| Marmota monar canadensis. | | | | |
$ 7607 | Murray Bay, Guepec vest: ee 77.3)44 29.6 30.5 56. 639. 4|22. 7/17. 4|19.1 Adult.
Be ee ee ee) 25 307/28: 832 (55.26.21 Ate 18,7 0.
6 16209 | Newport, Nova Scotia........-. aac 76. 4/44. 5/28. 7 32 53 '38. 2120. 5|15. 8118.6 Subadult
191344 | Carberry, Manitoba............. | g | 79.2/46.3|27.5:31. 7)... -/36.5|20.5]15.1]19 | Adult.
177740 | Smith Portage, Mackenzie....... | go | 78.3 45.8|28.631 |53.3138 |20.5|17.2|18.3 Do.
57602 | Murray Bay, Quebec............ | Q | 77.6)/45.6/28.8 33.5\56.638 |22 |17.8|19 Do.
5 7608 |..-.. da eee pee yaa. ass | Q | 78.746 |29.632 |53.239 |20.4/17. 118.4) Subadult
110192 | Oxford House, Manitoba........, Q | 79.7/47 |28.433.3'54.737.2)21. 1/15. 9|18.7| Adult.
177378 | Little Buffalo River, Mackenzie.| Q | 80 pee eee 16.6|18.1| Do.
6 16208 | Newport, Nova Scotia .......... Q | 79.6/46 |30.3.33.3'59.2/39.8)22 |17.2|19.2) Do.

1 Collection Mus. Vert. Zool., Univ. of California.
2Collection Provincial Mus., Victoria, B. C.

3 Collection Victoria Mem. Mus.

¢ Collection Kansas Univ. Mus.

® Collection Field Mus. Nat. Hist.

6 Collection Amer. Mus. Nat. Hist.

7 Average of 10 adults, Merriam collection.
& Average of 7 adults, Merriam collection.
$ Collection Mus. Comp. Zool.

36 NORTH AMERICAN FAUNA. [No. 37. .

Cranial measurements of the Marmota monax group—Continued.

| 4 Baie id Sap |i lees
| »| |¢|/8/8 fe | 8/2
| S| || S| ele deena
| a |4/8/2/2ls.ig8/3/8
3 c ~ = tay | {t= nD Es} = ro)
No. | Species and locality. BS) 2) | ae) eats |S See ol eedRemarks,
3/2/38) 2/8 lem ekla | 2
> IS/8/Sidis le |els
5 ue) Ts & mi ols 3 3 |
oO (oe) [sj (=) oO a = fas}
oD) Oo SPA PRIN IA IA ele
a
Marmota monax petrensis. | | |
203532 | Revelstoke, British Columbia....; ¢ 86 |50. 3/31. 8/32. sl 442 (23 |18.6/20 | Adult; type.
101295 | Barkerville, British Columbia--.| @ 79 146. 5/29. 5/32. sig 2)'36 |19. 4|17. 6/18. 8) Adult.
Marmota monaz ochracea. | |
202785 | Babine Mountains, British Co- | |
lumbia -..... = ale 2 eieleteie late ath .---| [2] | 81.2/48 (30 (33. 4/52. 7:39. 8/19. 8)17.2)/20 | Subadult
77143 | Stuart Lake, British Columbia..| ¢ 80. 6/47. 6/29. 5 30. 6/55. 6 38. 2/20. 6)18.8/19 | Adult.
Uae ees COE, ero Seema i tea 80. 2/47. 3/30. 5 32. 4 ----/38 119. 8)17. 2|17.6 Do.
CB aacoe GOs so 82555 Se ee ee a Q?]| 76.5/45 |28.932.4/52 |36.7/22 |16.8/18.6 Do.
COW eseee MO nec eels ce sirelre eee eee | [Q2?] | 77.1/45. 4/29 Hee 4/51.3 36. 2/19. 6/17. 4)... Do.

Marmota flaviventris Group.
[Characters under species. ]}
MARMOTA FLAVIVENTRIS (Aupuson & BacuMan).
[Synonymy under subspecies. ]

External characters.—Size variable (small to medium);! tail rela-
tively long (about 25 to 30 per cent of total length); ears small; sole
pads as in the monaz group (see PL. III, fig. 2); mammae: P. 2; A. 2;
T.4=10; head usually with white markings between eyes (absent or
reduced in obscura); sides of neck with conspicuous buffy patches;
feet varying from light buff to hazel or dark brown (never black);
tail mixed hazel and dark brown (fading to clay color).

Cramal characters—Skull similar in general outline to that of
monax; interorbital region relatively narrower; postorbital processes
longer and slenderer, projecting slightly back of a lme drawn across
their bases at right angles to axis of skull; nasals decidedly narrowed
posteriorly, where their width is equal to or less than that of nasal
branches of premaxillae; temporal ridges usually united in old age to
form a well-defined but rather low sagittal crest; floor of basi-occipital
with a median subcircular depression, bounded laterally by two low
processes which converge and unite into a ridge near the inferior lip of
the foramen magnum; posterior border of palate beveled at an ob-
tuse angle; interpterygoid fossa relatively narrow; palatal foramina
usually contracted posteriorly or of equal width throughout; molar
teeth light; maxillary tooth rows slightly divergent anteriorly; ante-
rior face of incisors yellowish white to zinc orange.

1 Two adult male specimens of M. flaviventris nosophora weighed, respectively, 10 and 12 pounds; two
adult males of M. flaviventris dacota, 8 and 17 pounds, the latter being very fat.

1915.} MARMOTA FLAVIVENTRIS GROUP. 37

Color.—General tone of upperparts vinaceous- or orange-cinnamon,
hazel, chestnut-brown, or vandyke brown; underfur of upperparts
at base mouse gray, fuscous, bister, clove brown, or blackish brown,
succeeded by pinkish-, ochraceous-, or cinnamon-buff, pinkish-, vina-
ceous-, or orange-cinnamon, cartridge- or tilleul-buff, buffy white,
hazel, or pale russet; long hairs chestnut-brown, olive-brown, hazel,
or black, tipped with light buff, ochraceous- or cinnamon-buff, or
white; top of head and face-cinnamon-drab, chestnut, bay, chestnut-,
vandyke-, or clove-brown, or black; sides of face chestnut-brown or
blackish brown, more or less mixed with white or-buff; face usually
more or less extensively marked with a band or patch of white or
buff between the eyes (nearly obsolete in obscura); borders of nose,
lips, and chin white or ochraceous-buff; sides of neck usually with
conspicuous patches of ochraceous-buff or cinnamon-buff (nearly
obsolete in obscura); underparts ochraceous-buff, ochraceous-tawny,
hazel, chestnut-, kaiser-, or blackish-brown, often varied with
russet, Sanford’s brown, tawny, or pale buff; fore legs ochraceous-
buff, tawny, hazel, russet, or kaiser brown, the feet chestnut-brown,
russet, auburn, pinkish cinnamon, or cinnamon-buff; hind legs
ochraceous-buff, tawny, or hazel (rarely brownish), the feet vary-
ing from light buff or pinkish cinnamon to russet, hazel, chestnut-
brown, or blackish brown; tail above, hazel, tawny, chestnut-brown,
blackish brown, or (in faded pelage) clay color; beneath, blackish
brown or chestnut-brown; ears usually some shade of buff, bordered
with dark brown.

Geographic distribution—From the interior valleys of southern
British Columbia south in the Great Basin to the Toyabe Mountains,
Nev., and Parawan Mountains, Utah; in the Cascade-Sierra system,
from Mount Hood, Oreg., to vicinity of Owens Lake, Cal.; and in
the Rocky Mountain system, from Flathead Lake, Mont., to the
Pecos River Mountains, N. Mex.;' east to the Black Hills, S. Dak.;
confined to mountains, foothills, and rocky canyons, not occurring
on the plains proper. (See fig. 2.)

Remarks — Marmota flavwentris usually may be readily distin-
guished from J. monax by the more ochraceous coloration, by the
buffy, hazel, or tawny legs and feet (the latter usually black or black-
ish brown in monazx), and by the presence of white or buffy face
markings; occasional specimens of MM. flaviventris obscura and WM.
flaviventris luteola rather closely resemble certain specimens of the
monaz group, but always lack the deep reddish hairs on the fore legs.

1 Formerly to the Manzano and Datil Mountains (vicinity of Old Fort Tularosa).

Ye,
oe
; S
‘D, =| - D f
ee

2\
oF

7
a a
Li
tA
:
Ye}
t a
“a
=e
ES
2k
A teael

i

au

44
(=

| 2. M. flaviventris flaviventris
+9 Bierrae
avara

Fig. 2.—Distribution of the Marmota flaviventris group. Unshaded areas within the range of the
group indicate a lack of definite knowledge as to the subspecies occurring there.

1915.] MARMOTA FLAVIVENTRIS GROUP. 89
MARMOTA FLAVIVENTRIS FLAVIVENTRIS (Aupunon & BacuMay).
YELLOW-BELLIED MARMOT.

(Pl. III, fig. 2; Pl. VII, fig. 1; Pl. XIII, fig. 1.)

Arctomys flaviventer Audubon & Bachman, Proc. Acad. Nat. Sci. Philadelphia, 1841,
p. 99; Quad. N. Am., III, 1853, p. 160, Pl. CKXXIV.
[ Marmota] flaviventer Trouessart, Cat. Mamm. Suppl., 1904, p. 344.

Type locality—‘‘ Mountains between Texas and California’’—
here fixed on Mount Hood, Oreg. (See remarks following, p. 40.)

Distribution —The Cascade Range in Oregon and the northern
Sierra in California, south to Lake Tahoe.

Characters.—Size large; general tone of upperparts russet, grizzled
with white, the fore back sometimes overlaid with a rather indistinct
buffy mantle; underparts ochraceous-buff to hazel; feet ochraceous
or hazel; skull massive.

Color—Underfur of upperparts deep mouse gray at base, suc-
ceeded by a broad area of buffy white, the latter shading on hinder
back to pinkish cinnamon (sometimes to hazel); long hairs chestnut-
brown or clove brown subterminally, tipped on fore back with light
buff, on hinder back with white; top of head and face chestnut-
brown to blackish brown, sometimes varied with cinnamon-drab, with
an indistinct band of buffy white or ochraceous-buff across face in
front of eyes; sides of face mixed chestnut-brown and cinnamon-buff;
sides of neck with large patches of light ochraceous-buff; fore legs
and feet tawny to hazel; hind feet light ochraceous-buff more or less
varied with light hazel; tail hazel (fading to clay color), the bases of
the hairs and under surface natal brown, varying to chestnut-brown
or blackish brown; underparts ochraceous-buff, varying to hazel,
becoming light ochraceous-buff around base of tail, the bases of some
of the hairs indistinctly chestnut-brown; sides of nose, lips, and chin
white, often varied with ochraceous-tawny around nose and edged
with hazel or kaiser brown on throat.

Skull—Largest of any member of the group; relatively long and
narrow; nasals long, and narrow posteriorly; nasal branches of pre-
maxillae broad; postorbital constriction broad; rostrum long and nar-
row; palatal foramina narrow.

Measurements.—Adult male from Donner, Cal.: Total length, 700;
tail vertebrae, 180; hind foot, 90. Adult female from Crater Lake,
Oreg.: 640; 170; 80. Adult female from Donner, Cal.: 600; 170; 85.
Skull: Adult male:! Condylo-basal length, 92.5-97.1 (average 94.5);
palatal length, 52.3-55 (53.8); postpalatal length, 37.6-38.2 (37.9);
length of nasals, 42.5-42.6; zygomatic breadth, 62.3; breadth across
mastoids, 41.1-46.2 (43.9); least interorbital breadth, 21.2-23 (22.1);
breadth of rostrum, 22.3-23 (22.8); maxillary tooth row, 19.5-

1 Three specimens from Mount Hood, Oreg., and Fort Crook and Donner, Cai.

AO NORTH AMERICAN FAUNA. [No 3%; |

20.9 (20.1). Adult female:' Condylo-basal length, 82-87 (84.2);
palatal length, 45.8-48.7 (47.6); postpalatal length, 30.6-34.5 (84.1);
length of nasals, 34.5-37.2 (85.9); zygomatic breadth, 54.4-58.4
(55.9); breadth across mastoids, 38.5-40.7 (39.8); least interorbital
breadth, 17.3-19.3 (18.5); breadth of rostrum, 17.8-21.4 (19.8);
maxillary tooth row, 19.5-22.1 (20.5).

Remarks.—This species was described in 1841 and the name has
been in use ever since in a broad sense for the marmots of the Sierra-
Cascade range and, by most authors, for the Rocky Mountain forms
as well. The original describers’ statement as to the source of the
type specimen—‘‘mountains between Texas. and California’’—is
not only too indefinite to be of use, but the evidence points to
its being incorrect. The type specimen, as stated by Audubon
and Bachman, was obtained by David Douglass and (the skin
only) is still in a fair state of preservation in the British Museum,
but unfortunately is unaccompanied by data to show where it was
collected. Douglass’s journal has been published,? and from it we
learn that he spent a considerable time at Fort Vancouver near the
mouth of the Columbia River, whence he made journeys up the Co-
lumbia- and Willamette Valleys and through eastern Washington
and southern British Columbia. He spent some time, also, on the
coast of California, below Monterey, but apparently never visited
either the Sierra or the Cascades proper. His journal does not men-
tion the capture of any marmots.? He might easily have secured the
type specimen on his journey through eastern Washington, in which
case it would be the form now known as avara, or he might have
obtained it from Indians or travelers who had visited Mount Hood—
a comparatively short distance from his headquarters—in which case
it would be referable to the form which now bears the name flaw-
veniris. In order to settle the question a specimen from Mount
Hood and one from Okanogan, British Columbia, were submitted to
Oldfield Thomas, who compared them with the type in the British
Museum. He states as follows:

The underside of forearms [in the type] are about as rufous and its rump as brown
and finely speckled as the Oregon specimen, but it has got distinctly the broadly buffy
mantle on the fore back so marked in the British Columbia specimen. The type is
a good adult skin which has borne remaking very well. The skull, unfortunately,
is not in existence. ;

The buffy mantle of which Mr. Thomas speaks is not a diagnostic
character, since it appears in both the British Columbia and Cascade
forms. ‘The rufous color of the fore legs, however, is diagnostic,

1 Seven specimens from Crater Lake, Oreg., and northern parts of Sierra Nevada, Cal.

2 Hooker, W. J. Companion to Botanical Mag., II, 1836, pp. 79-182.

3 The statement [p. 92] that a curious species of Arctomys was secured and other references to ‘‘Arctomys
brachyurus”’ [pp. 101-115] doubtless refer to species of either Citellus or A plodontia.

1915.} MARMOTA FLAVIVENTRIS GROUP. 4]

and, except for size and cranial characters, is the only sure distin-
guishing characteristic of the two forms. It seems clear, therefore,
that the name flaviventris must be applied to the darker of the two
forms found in Oregon and Washington, and I have therefore selected
Mount Hood, Oreg., as the type locality. The series available from
the Cascades is small and contains only one adult male skulli—an old
weathered specimen picked up on Mount Hood. Several adult fe-
males in the collection from Crater Lake and the Klamath Lake
region, however, agree essentially with specimens from Fort Crook
and the northern Sierra of California. South of Lake Tahoe flaw-
ventris grades imperceptibly into the subspecies sierrae, occupying
the southern end of the Sierra Nevada, in the Mount Whitney region.
Intergradation with avara occurs in the region east of Klamath Lake
and probably all along the east base of the Cascades.
Specumens examined.—Total number, 48, as follows:
California: Donner, 17; Emerald Bay, 1; Fort Crook, 3; Glen Alpine Springs,
El Dorado County, 2;! Hope Valley, Alpine County, 2; Lassen Creek, Warner
Mountains, 1; Mount Lassen, 2; Pine Creek, Lassen County, 1.
Nevada: Mount Siegel, Douglas County, 1; Winters Mine, Douglas County, 1.?
Oregon: Crater Lake, 2; Fort Klamath, 3; Klamath Lake, 6; Linkville, 2; Mount
Hood, 3; Summer Lake, 1.

MARMOTA FLAVIVENTRIS AVARA (BaAnGs).
PatitiIp YELLOW-BELLIED MARMOT.
(Ee, Viilbw hic. 3-7 Plis OTT fo.)

Arctomys flaviventer avarus Bangs, Proc. New England Zool. Club, I, 1899, p. 68.
[ Marmota flaviventer] avarus Trouessart, Cat. Mamm., Suppl., 1904, p. 344.

Type locality —Okanogan, British Columbia.

Mstribution.—Interior valleys and foothills of southern British Co-
lumbia and eastern Washington and Oregen.

Characters —Simnilar to flaviventris, but smaller and colors paler,
especially the underfur and fore legs; upperparts more extensively
overlaid with buff. |

Color.—General tone of upperparts chestnut-brown, heavily griz-
zled with buff; underfur at base deep mouse gray (varying to pale
fuscous) succeeded by buffy white, the latter shading to light pinkish
cinnamon on hinder back; long hairs blackish brown or clove brown
subterminally, extensively tipped on fore back with warm buff and
on hinder back with light buff; head and face blackish brown or dark
chestnut-brown, with an indistinct narrow band of white or buff
across face in front of eyes; sides of neck warm buff; fore legs and
feet ochraceous-buff; hind legs, hind feet, and buttocks light ochra-
ceous-buff, the feet varying to hazel; tail hazel, the hairs tipped

1 Collection Mus. Vert. Zool., Univ. of California. 2 Collection Field Mus. Nat. Hist.

49, NORTH AMERICAN FAUNA. [No. 37.

with warm buff, their bases chestnut-brown; underparts ochraceous-
buff or light ochraceous-buff, the bases of the hairs on belly chestnut-
brown; sides of nose, lips, and chin white, bordered on throat with
hazel.

Skull.—Similar to that of flaviventris, but decidedly smaller; nasals
shorter, ending but little back of posterior border of premaxillae.

Measuremenis.—Adult male: Total length, 495-660 (average
554); tail vertebrae, 149-178 (165); hind foot, 76-80 (78). Skull:
Adult male: ? Condylo-basal length, 85.7-86.5 (86.1); palatal length,
47.9-48.8 (48.3); postpalatal length, 34-34.8 (84.4); length of nasals,
36.6-37.6 (37.1); zygomatic breadth, 55.6-56.7 (56.1); breadth across
mastoids, 39.1-42.2 (40.6); least interorbital breadth, 17.8-19.5
(18.6); breadth of rostrum, 19.4-20 (19.7); maxillary tooth row,
19-19.3. Adult female:* Condylo-basal length, 76.4-79.4 (77.9);
palatal length, 42.6-44.6 (43.6); postpalatal length, 30.3-31 (80.7);
length of nasals, 31.5-33.5 (32.8); zygomatic breadth, 52.4-52.7
(52.5); breadth across mastoids, 37.6-38.6 (38); least interorbital
breadth, 17.1-18 (17.6); breadth of rostrum, 16.8-18.3 (17.3); max-
illary tooth row, 17.8-19.1 (18.5).

Remarks.—This is a small, pale race of flaviventris living in the low
foothills of eastern Oregon and Washington and southern British
Columbia. It intergrades with flaviveniris along the eastern base of
the Cascades, in Oregon; with nosophora in northwestern Montana;
and probably with parvula in northern Nevada. Specimens from
Spokane Bridge, Wash., are a little darker than the typical form,
and one from Elgin, Oreg., has a slightly larger skull. Specimens
from Rockland, Wash., are intermediate between avara and flavi-
ventris. Melanistic individuals are rare in this race; one from Pull-
man, Wash., has the fore back solid black, faintly grizzled with
white, and the underfur pale gray; the hinder back is gray, the hairs
being black subterminally and tipped with white; the belly and feet
are black varied with buff; and the tail is black varied with tawny and
white.

Specumens examrned.—Total number, 53, as follows:

British Columbia: Ashcroft, 8; Cascade, 6;* Midway, 9;*)5® Nicola Valley, 1;6
Okanogan, 9; Penticton, 1;4 Vernon, 1.
Oregon: Elgin, 2; Harney, 2; Guano Creek, Lake County, 1.

Washington: Cheney, 1; Diamond, 1; Douglas, 1; Pullman (14 miles south-
west), 1; Rockland, 4; Spokane Bridge, 5.

1 Four specimens from Okanogan, British Columbia; and Douglas, Wash.
2 Two specimens from Okanogan and Cascade, British Columbia.

3 Four specimens from southern British Columbia.

4 Collection Victoria Mem. Mus.

5 Collection Field Mus. Nat. Hist.

§ Collection Amer. Mus. Nat. Hist.

1915.] MARMOTA FLAVIVENTRIS GROUP. 43

MARMOTA FLAVIVENTRIS SIERRAE sussp. nov.
SouTHERN SreRRA MARMOT.
(Pl. IV, fig. 4.)

Type from head of Kern River, Mount Whitney, Cal. (altitude,
9,300 feet). Adult ?, No. 39984, U.S. Nat. Mus., Biological Survey
collection. Collected Sept. 3, 1891, by Vernon Bailey; original
number, 3242.

Distribution.—Higher parts of the southern Sierra Nevada om
upper Kern River north to vicinity of Mono Lake.

Characters —Similar to flavwentris, but colors redder and buffy
mantle reduced or obsolete; skull smaller.

Color.—General tone of upperparts vinaceous-cinnamon to mikado
brown, narrowly grizzled with white; underfur at base pale bister
succeeded by pale pinkish buff, shading at tips into vinaceous-
cinnamon and on hinder back to orange-cinnamon or mikado brown;
long hairs chestnut-brown or blackish brown subterminally, tipped
with white or buffy white; top of head and face dark cinnamon-
drab to clove brown, with a rather large patch of white in front of
eyes; sides of neck warm buff; fore legs and feet hazel varied with
light ochraceous-buff; hind feet similar, or sometimes light buff, varied
with brown; tail hazel, the bases of hairs light bister; underparts
light ochraceous-buff, much mixed on sides and belly and bordered
on throat with mikado brown.

Skull.—Similar to that of flaviventris, but decidedly smaller; very
similar to that of avara but averaging slightly larger, and nasals longer.

Measurements—Adult male topotype: Total length, 600; tail
vertebrae, 200; hind foot, 83. Adult male:' 600; 165; 80. Adult
female:? 555-645 (average 593); 167-186 (176); 75-80 (77). Skull:
Adult male:* Condylo-basal length, 79.1-85.5 (average 84.7);
palatal length, 44-50.5 (47.7); postpalatal length, 30.8-35 (83);
length of nasals, 32.6-39 (36.3); zygomatic breadth, 52-58.3 (55.8);
breadth across mastoids, 37.5-43.6 (40.8); least interorbital breadth,
17-20.3 (18.1); breadth of rostrum, 17-20 (18.6); maxillary tooth
row, 20.2-20.7 (20.4). Adult female: ? Condylo-basal length, 77.1-
83.2 (80.1); palatal length, 43.2—47.2 (45.5); postpalatal length, 28.7-
32.2 (30.5); length of nasals, 30.7-36.5 (34.2); zygomatic breadth,
51.4-55.4 (53.6); breadth across mastoids, 37-40 (38.4); least inter-
orbital breadth, 15-18.7 (17.2); breadth of rostrum, 17.4—20.2 (18.4);
maxillary tooth row, 19-20.2 (19.7).

Remarks.—By reason of the scarcity of specimens from the Cas-
cades (the type region of flaviventris) it is impossible satisfactorily to

1 One specimen from East Fork Kaweah River, Cal.

2 Five specimens from southern Sierra Nevada, Cal.
3 Six specimens from southern Sierra Nevada, Cal.

44 NORTH AMERICAN FAUNA. [No. 37.

characterize this southern race. The available specimens, however,
show a decided reduction in size of skull southward, the small form
apparently being confined to the high Sierra south of Mono Lake.
Intergradation with flavwentris occurs in the region between Mono
Lake and Lake Tahoe, and with parvula in the White Mountains,
on the boundary between California and Nevada.

Specumens examined.—Total number, 47, as follows:

California: Big Meadows, Tulare County, 3; Bishop Creek (altitude, 8,000
feet), 2; Cannell Meadows, Tulare County, 2;! Cottonwood Lakes, Inyo
County, 3;! Independence Lake, 1;! Kaweah River (East Fork), 3; Lake
Tenaya, 1; Menache Meadows (near Olancha Peak), 1; Mount Lyell, 1;
Mount Whitney, 9; Mulkey Meadows (15 miles south of Mount Whitney),
1; Owens River (near Mammoth Pass), 1; Round Valley (12 miles south
of Mount Whitney), 2; San Joaquin River (near head), 4; Siberian Outpost,
Tulare County, 1;! Tuolumne Meadows, 10; Whitney Creek, Tulare County,
1;1 Whitney Meadow, Tulare County, 1.!

MARMOTA FLAVIVENTRIS PARVULA Howe t.
NEVADA MARMOT. .
(PI Vil; figs2; PP Xie foe 2))
Marmota flaviventer parvula Howell, Proc. Biol. Soc. Washington, XXVII, 1914, p. 14.

Type locality.—Jefierson, Toquima Range, Nye County, Nev.
(about 10 miles north of Belmont).

Distribution.—Toyabe and Toquima Ranges, Nev.; and White
Mountains, Cal. (occurring from about 7,800 to 10,000 feet altitude);
probably occupies also other desert ranges in central Nevada.

Characters.—Similar to avara, but smaller and colors darker; in
color resembling sierrae, but upperparts less reddish and overlaid with a
buffy mantle; skull smaller than that of avara, with narrower rostrum.

Color.—Underfur of upperparts slaty fuscous at base, succeeded
by a broad area of cartridge buff on fore back and by light vinaceous-
cinnamon on hinder back; long hairs dark brown subterminally,
tipped on fore back with warm buff and on hinder back with white;
top of head and face dark vandyke brown; a band of buff or buffy
white across face in front of eyes; sides of nose and lips and a large
patch on chin white varied with buff; sides of neck with a conspicu-
ous patch of warm buff or cinnamon-buff; underparts ochraceous-
tawny, shading to russet on abdomen and throat; fore legs tawny or
russet, tipped with ochraceous-buff; fore and hind feet varying from
light pinkish cinnamon to russet; tail above, dark chestnut-brown
varied with tawny and grizzled with buff; beneath, blackish brown.

Skull.—Similar to that of avara, but decidediy smaller, with
rostrum narrowed anteriorly.

1 Collection Mus. Vert. Zool., Univ. of California.

1915.] MARMOTA FLAVIVENTRIS GROUP. 45

Measurements.—Adult female:1 Total length, 470-500 (average,
480); tail vertebrae, 130-150 (141); hind foot, 70. Skull: Adult
female:? Condylo-basal length, 71.3-73 (72.2); palatal length,
39.441 (40.2); postpalatal length, 27-28.6 (27.8); length of nasals,
29.4-30.5 (30); zygomatic breadth, 48.6—-49 (48.8); breadth across
mastoids, 34.2-34.3; least interorbital breadth, 15.5-15.6; breadth
of rostrum, 17.2-17.4 (17.3); maxillary tooth row, 17.7-18.6 (18.2).

Remarks.—This is the smallest of the races of flaviventris. It most
resembles typical flaviveniris in color, but is even smaller than avara.
Its small size was noted in the field by Vernon Bailey, who collected
the type series. The range of the subspecies is not definitely known
but probably extends at least to the Ruby Mountains where, on top
of one of the peaks, Mr. Bailey found evidences of the presence of
marmots.

A small series in badly worn pelage from White Mountains, Cal., is
provisionally referred to this race, the specimens being intermediate
in size between parvula and sierrae, and agreeing fairly well in color
with the former, except for a stronger suffusion of red. One adult
female skull agrees with skulls of parvula while a subadult male agrees
equally well with comparable specimens of sverrae.

Two very young specimens from Mountain City, northeastern
Nevada, are also provisionally referred to this race.

Specimens examined.—Total number, 16, as follows:

_ Nevada: Arc Dome, Toyabe Range, 6; Jefferson, Toquima Range, 1; Mountain

City, 2.
California: White Mountains (altitude 9,300-10,000 feet), 7.

MARMOTA FLAVIVENTRIS ENGELHARDTI Atizen.
ENGELHARDT Marmot.
(Pl Vil, fig A; Pl. XII, fig.;4.)
Marmota engelhardti Allen, Mus. Brooklyn Inst. Arts & Sci., Sci. Bul., I, 1905, p. 120.

Type locality—Briggs [=Britt’s] Meadow, Beaver Range, Utah
(altitude 10,000 feet).

Distribution.—Beaver and Parawan Mountains, southern Utah;
also Midvale, Idaho; exact limits of range unknown.

Characters.—Similar to flaviventris but smaller; underparts and
hind feet darker (redder); buffy patches on sides of neck less exten-
sive; larger than parvula, with darker feet and underfur; skull sim-
ilar to that of avara but bullae larger.

Color.—General tone of upperparts vandyke brown, grizzled with
buffy white; underfur fuscous at base succeeded by pinkish buff or
pinksh cinnamon, the latter shading on sides to Rood’s brown; long
hairs blackish brown, tipped with light buff or buffy white; top of

1 Three specimens from Toyabe and Toquima Ranges, Nev. 2 Two specimens from same localities.

46 NORTH AMERICAN FAUNA. [No. 37.

head and face blackish brown, with an irregular white patch in front
of eyes; sides of head mixed brown and buffy white; sides of neck
with a small area of ochraceous-buff; legs and feet hazel; tail above,
dark clove brown, tipped with hazel; beneath, blackish brown; under-
parts hazel or ochraceous-tawny, the bases of hairs blackish brown;
sides of nose, lips, and chin, white.

Skull—Similar to that of avara but audital bullae averaging
larger and more inflated.
- _Measurements.—Adult female topotype: Total length, 525; tail
vertebrae,110;? hind foot,75. Adult female from Parawan Mountains,
Utah: 554; 154; 68. Immature female topotypes:* 485-535 (average
507); 189-163 (151); 71-77 (73). Skull: Adult female:* Condylo-
basal length, 76.8-80 (78.4); palatal length, 43.5-46.1 (44.8); post-
palatal length, 29.5-30.7 (30.1); length of nasals, 31-34.6 (32.8);
zygomatic breadth, 52.3-54 (53); breadth across mastoids, 36.5-37.7
(37.2); least interorbital breadth, 16.3-18 (17.1); breadth of rostrum,
17.8-18 (17.9); maxillary tooth row, 18.8-19 (18.9).

Remarks.—This race is an intermediate form connecting flaviventris
and nosophora, darker beneath than the former but not so reddish as
the latter and lacking also its buffy mantle. It is about the size of
avara with a somewhat shorter tail, but much darker in color. A
specimen from Parawan Mountains, Utah, is paler beneath than the
type and topotypes, the underparts (except throat) and hind feet
being ochraceous-buff varied with brownish. An immature specimen
from Midvale, Idaho, provisionally referred to this form, agrees with
it in color, except that the underfur on the shoulders is paler (light
buff), and in skull characters so far as they can be determined.

Specimens ecamined.—Total number, 12, as follows:

Idaho: Midvale, 1.
Utah: Beaver Mountains, 6;° Parawan Mountains, 5.

MARMOTA FLAVIVENTRIS NOSOPHORA HoweEtt.
GOLDEN-MANTLED Marmot.
(Pl. I; Pl. VIII, fig. 1; Pl. XIV, fig. 1.)
Marmotaflaviventer nosophora Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 15.

Type locahty——Willow Creek, 7 miles east of Corvallis, Mont.
(altitude 4,000 feet).

DistributionRocky Mountain region of Montana, Idaho, and
Wyoming, from Flathead Lake, Mont., south to the Wasatch Moun-
tains, Utah, and east to the Bighorn Mountains, Wyo.; altitudinal
range from about 3,000 to 11,800 feet.

1 No adult males examined.

2 Apparently abnormally short.

8 Three specimens.

4 Three specimens from Beaver and Parawan Ranges, Utah.
5 Including type in collection Amer. Mus. Nat. Hist.

1915.] MARMOTA FLAVIVENTRIS GROUP. 47

Characters.—About the size of engelhardti, but with longer tail;
colors much more ochraceous above and redder below, the fore part
of back overlaid with a mantle of golden buff. Compared with
dacota: Slightly smaller, with upperparts less extensively reddish
and more mixed with black, and underfur decidedly paler; skull
averaging smaller, with relatively slenderer rostrum and much smaller
palatal foramina. |

Color—Underfur of upperparts at base blackish brown on fore
part of body, becoming fuscous on hinder parts, succeeded by a
broad area of whitish buff (tilleul buff of Ridgway) shading (on
hinder back) to pinkish cinnamon or pale russet; long hairs black
subterminally, broadly tipped on fore part of back with warm- or
ochraceous-buff and on hinder part with white or buffy white; top
and sides of head blackish brown, with a conspicuous band of white
or buffy white across face in front of eyes; sides of face mixed with
cinnamon or white; sides of nose, lips, and chin white or buffy white;
sides of neck with ochraceous-buff patches behind ears; fore legs
kaiser brown; hind legs and rump warm buff; hind feet hazel to
russet; tail chestnut-brown or blackish brown, varied with hazel or
cinnamon-buff (fading to dull cinnamon or clay color); underparts
hazel shaded with kaiser brown, becoming bright chestnut on throat
and sometimes on belly. Variation: A dark, brownish color-phase
occurring rarely, and seemingly most frequent at high altitudes, may
be described as follows: General tone of upperparts dark brown
grizzled with white; underfur at base mouse gray (shading on hinder
back to fuscous) succeeded by buffy white (shading on hinder back
to ight pinkish cinnamon); long hairs blackish brown, tipped with
white; top and sides of head blackish brown; underparts mixed
blackish brown and pinkish buff in about equal proportions; throat
shaded with russet; feet and tail blackish brown; legs brownish
mixed with ochraceous-tawny. (Specimen from Lake Fork, Wind
River Mountains, Wyo.; altitude 10,600 feet.)

Skull—FKemales averaging larger than those of engelhardti, with
broader rostrum and interorbital region and smaller bullae. Com-
pared with dacota: Smaller, with narrower rostrum and interorbital
region, and much smaller palatal foramina.

Measurements—Adult male: Total length, 590-600 (average
595); tail vertebrae 159-170 (165); hind foot, 78-79 (78.5). Old male
from Pryor Mountains, Mont.: 670; 165; 96. Adult female:? 534—
591 (565); 145-175 (165); 75-85 (78). Skull: Adult male:# Condylo-
basal length, 86.7-94.2 (90.6); palatal length, 47.7-53 (50.7); post-
palatal length, 34.2-37.2 (36.2); length of nasals, 35.1—-40.4 (87.8);
zygomatic breadth, 55.7-63.7 (59); breadth across mastoids, 41.3-45

1 Two specimens from Bitterroot Valley, Mont.

2 Seven specimens from Bitterroot Valley, Mont.
3 Six specimens from Montana, Idaho, and Wyoming.

48 NORTH AMERICAN FAUNA. [No. 37.

(43); least interorbital breadth, 17.8—22.2 (20.1); breadth of rostrum,
19-20.5 (19.8); maxillary tooth row, 19.3-20.8 (20). Adult female:!
Condylo-basal length, 78.2-83.8 (80.4); palatal length, 44.1-47-(45.1);
postpalatal length, 29.4~33.5 (381.2); length of nasals, 30.8-34.8 (82.4);
zygomatic breadth, 53.2-54.5 (53.9); breadth across mastoids, 37.4—
41.6 (89); least interorbital breadth, 18.5-19.7 (19.1); breadth of ros-
trum, 17-19.3 (18.3); maxillary tooth row, 19.7—20.5 (20.1).

Remarks.—This subspecies, one of the handsomest members of the
flavweniris group, is abundant and generally distributed in the
northern Rocky Mountain region of Montana, Idaho, and Wyoming.
Closely related to dacota of the Black Hills, it probably intergrades
with that race in central Wyoming. Specimens from the northern
limit of its range (Horse Plains and Weeksville, Mont.) are consider-
ably paler than the typical form, showing approach to avara. The
southern limits of the range of nosophora are not known, only a few
specimens having been seen from the Wasatch Mountains, and none
from the Uinta Mountains, Utah. Intergradation with engelhardti
probably occurs where their ranges meet.

A brown phase of this subspecies, having the tips of the hairs
white instead of cinnamon-buff and the underparts mixed brown and
buff instead of red, occurs in some localities with the normal phase.
Several immature specimens, varying somewhat in color, have been
examined from near timberline in the Wind River Mountains, Wyo.
An adult female in very worn pelage from timberline in the Bear-
tooth Mountains, Mont., is similar to those from the Wind River
Mountains, but the brown of the upperparts is paler, evidently faded,
and the underparts are mixed chestnut and black, shaded with >
tawny. Some specimens in this phase somewhat resemble ex-
ternally the members of the monaz group but the skulls are typical
of nosophora.

This marmot serves as a host for the Rocky Mountain fever tick
(Dermacentor venustus) and thus aids in the dissemination of the
deadly spotted fever, particularly along the western side of the
Bitterroot Valley, Mont., where the disease occurs in its most virulent
form.

Specimens examined.—Total number, 62, as follows:

Idaho: Bear Lake (east side), 1; Bridge, 1; Conant Creek (upper Snake River),
1; Grace, 2; Henry Fork of Snake River, 1; Irwin (20 miles northwest), 1;
Island Park, Snake River, 1; Moody Creek (upper Snake River), 1; Preuss
Mountains, 1; Sawtooth National Forest, 2; Teton Basin, 1.

Montana: Bass Creek (in mountains northwest of Stevensville), 3; Beartooth
Mountains, 3; Bozeman, 1; Como Lake, 1; Florence, 2; Horse Plains, 4;

Jardine, 1; Pryor Mountains, 2; Ross Fork (15 miles above Darby), 2; Weeks-
ville, 1; Willow Creek (in mountains east of Corvallis), 3.

1 Five specimens from Bitterroot Valley, Montana.

1915.] MARMOTA FLAVIVENTRIS GROUP. 49

Utah: Blacksmith’s Fork (near head), 1; Laketown, 1; Park City, Wasatch
Mountains, 3.

Wyoming: Bighorn Mountains (Trapper’s Creek), 6; Fremont Peak, 2; Jackson,
2; Kendall (12 miles north), 1; Lake Fork, Wind River Mountains, 3; Little
Sandy Creek, 3; Lost Cabin (15 miles northwest), 1; Pahaska, 1; Salt River
Mountains (10 miles southeast of Afton), 1; Sheridan, 1.

MARMOTA FLAVIVENTRIS DACOTA (MERRIAM).
Brack Hints Marmot.
(PI LV; fig: 1; Pl VIIE, fie. 3+ Pl. XIV; fie 2.)

Arctomys flaviventer Grinnell, Ludlow’s Black Hills of Dakota, 1875, p. 82. (Not
of Audubon & Bachman).

Arctomys dacota Merriam, N. Am. Fauna No. 2, 1889, p. 8.

[ Marmota] dacota Trouessart, Cat. Mamm., Suppl., 1904, p. 344.

Type locality Custer, S. Dak.

Distribution —Black Hills, S. Dak., and Bear Lodge Mountains,
Wyo., southwest to Bridger Pass, Wyo.

Characters.—Size large (equaling flaviventris); color similar to
that of nosophora but underfur redder and less mixed with black;
skull large, with broad rostrum and very large palatal foramina.

Color—General tone of upperparts orange-cinnamon overlaid
with warm buff; underfur at base fuscous or dark mouse gray,
succeeded by pinkish cinnamon or vinaceous-cinnamon on fore back
and by orange-cinnamon on hinder back, darkening to kaiser brown
on sides (sometimes vinaceous-cinnamon to roots on fore back);
long hairs on fore back bright cinnamon-buff at tips with an indis-
tinct subterminal band of hazel or chestnut-brown; on hinder back
chestnut-brown or blackish brown, tipped with white; buttocks
warm buff; top of head and nose blackish brown with a band of
yellowish white across face in front of eyes; sides of nose, lips, and
chin, white or yellowish white; fore legs kaiser brown, feet russet;
hind feet hazel; tail above, hazel mixed with chestnut-brown; beneath,
blackish brown; underparts kaiser brown shaded with ochraceous-
buff.

Skull_—Similar to that of nosophora but larger, with broad rostrum
and broad palatal foramina.

Measurements.—Adult male:' Total length, 610-680 (average
643); tail vertebrae, 178-200 (185); hind foot, 81-92 (85). Adult
female: ? 525-627 (602); 130-188 (168); 79-84 (81). Skull: Adult
male:? Condylo-basal length, 89.4-95.7 (92.5); palatal length,
50.8-52.2 (51.6); postpalatal length, 35-38.2 (36.6); length of nasals,
37.8-41.1 (39); zygomatic breadth, 59.5-61.4 (60.7); breadth across

1 Five specimens from Black Hills, S. Dak., and Bear Lodge Mountains, Wyo.

2 Five specimens from Black Hills, 8. Dak.
3 Three specimens from Black Hills, 8. Dak., and Bear Lodge Mountains, Wyo.

69168°—15——-4

50 NORTH AMERICAN FAUNA. [No. 37. -

mastoids, 42.3-44.7 (43.3); least interorbital breadth, 21.8-23.2
(22.6); breadth of rostrum, 22-23.6 (23); maxillary tooth row,
21-21.7 (21.4). Adult female:' Condylo-basal length, 81.4-84.5
(83.1); palatal length, 45.4-47.7 (46.8); postpalatal length, 31.6-33
(32.4); length of nasals, 31.5-35.5 (33.9); zygomatic breadth, 54.2-
56.7 (55.5); breadth across mastoids, 39.4-42.3 (40.5); least inter-
orbital breadth, 19.6—22.5 (20.6); breadth of rostrum, 20—-22.3 (20.9);
maxillary tooth row, 20.1-20.5 (20.4).

Remarks.—This form is the brightest of all the races of the species,
the red and yellow shades being most pronounced and the blacks
and browns reduced to a minimum. It is abundant in the Black
Hills and apparently ranges through eastern Wyoming to Bridger
Pass, intergrating with luteola in the Laramie Mountains.

Specomens examined.—Total number, 19, as follows:

South Dakota: ‘‘Black Hills,’ 1; Custer, 8; Savoy, 5; Tigerville (near Hill

City). 1:
Wyoming: Bear Lodge Mountains, 1; Bridger Pass, 3.

MARMOTA FLAVIVENTRIS LUTEOLA Howett.
Park MARMOT.
GPl VILL fig. 25 (Pl XV ties)

Arctomys flaviventer Allen, Bul. Essex Inst., VI, 1874, p. 57. (Not of Audubon &
Bachman.) :

Marmota flaviventer Warren, Colorado Coll. Pub., Sci. Ser., XI, No. 46, 1906, p. 243.
(Not of Audubon & Bachman.)

Marmota engelhardti Cary, N. Am. Fauna No. 33, 1911, p. 98. (Not of Allen.)

Marmota flaviventer luteola Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p.15.

Type locality— Woods P. O., in Medicine Bow Mountains, Wyo.
(altitude about 7,500 feet).

Distribution —Mountains of northern Colorado and southeastern
Wyoming, from Park County, Colo., (and probably Fremont County)
north to the Laramie Mountains, Wyo.

Characters—About the size of dacota and similar in color to it
and nosophora, but underparts yellowish instead of deep red, and
fore back overlaid with white instead of warm buff; skull similar
to that of nosophora, but averaging narrower.

Color—Normal phase: Underfur of upperparts dark mouse gray
at base (shading on hinder back to deep dusky drab), succeeded on
fore back by a broad area of warm buff and on hinder back by
ochraceous-buff; long hairs blackish brown or dark chestnut-brown,
tipped with light buff or buffy white; top of head and nose blackish
brown, with a rather large patch of white or ochraceous-buff between
eyes; sides of face mixed brown and buff; sides of nose, lips, and
chin, white, yellowish white, or ochraceous-buff; sides of neck warm

1 Five specimens from Black Hills, S. Dak.

1915.] MARMOTA FLAVIVENTRIS GROUP. 51

buff (the underfur ochraceous-buff) ; feet hazel or tawny; tail above,
mixed hazel and chestnut-brown; beneath, blackish brown; under-
parts ochraceous-buff (bases of hairs brownish) varied with tawny
along sides; rump and buttocks warm buff. Dark phase (specimen
from Boulder County, Colo., altitude, 8,000 feet): Underfur cmnamon-
buff becoming dark cinnamon on hinder back; entire body extensively
mixed with dark chestnut-brown hairs, sparingly tipped on back
with light buff; feet blackish brown shaded with tawny; light face-
markings reduced.

Skull.—Very similar to that of nosophora, but averaging relatively
narrower, especially rostrum and interorbital region; bullae smaller.

Measurements.—Adult male:+ Total length, 600-650 (average
623); tail vertebrae, 182-220 (200); hind foot, 86-90 (88). Adult
female:? 552-618 (579); 1387-192 (171); 78-85 (81). Skull: Adult
male:? Condylo-basal length, 86-92.5 (88.3); palatal length, 48.4—
51.1 (49.5); postpalatal length, 33.5-36.5 (35.2); length of nasals,
36-41 (38.4); zygomatic breadth, 55.7-60 (57.9); breadth across
mastoids, 39-43.4 (41.5); least interorbita] breadth, 17.3-21.8 (19.2);
breadth of rostrum, 18.9—20.8 (19.6); maxillary tooth row, 19.9-21.4
(20.5). Adult female:* Condylo-basal length, 78.3-85 (81.4); pal-
atal length, 43.7-47.6 (45.7); postpalatal length, 30-33.3 (81.5);
length of nasals, 32.5-35 (33.7); zygomatic breadth, 51.7-54.9 (53.3);
breadth across mastoids, 37.5-40.5 (38.6); least interorbital breadth,
16.4-17.8 (16.9); breadth of rostrum, 17.3-18.3 (17.8); maxillary
tooth row, 19.3-21:4 (20.2).

_Remarks.—In studying the marmots of this and related races in

Colorado several puzzling problems have been encountered, the ma-
' terial at present available being insufficient satisfactorily to work out
the characters and exact relationships of the forms. The specimens
exhibit considerable individual variation, and, in addition to the dark
phase already described, a light phase, eharacterized by less yellow-
ish underparts and white tips to the hairs above, occurs in the same
localities with the normal phase. Specimens in this light phase are
known from Meeker, North Park, and Boulder County (altitude
10,300 feet). They rather closely resemble engelhardit externally,
except that the underfur and feet are paler; the skulls, however, are
typical of luteola.

The series from Laramie Mountains, Wyo., is intermediate between
luteola and dacota, the pelage averaging redder and the skulls
relatively shorter and broader than in typical luteola. Most of
this series have large ochraceous-buff face markings. The series
from Sulphur Springs, Colo., shows intergradation with warreni, the

1 Three specimens from southern Wyoming and northern Colorado.
2 Six speeimens from northern Colorado.

3 Four specimens from Laramie and Medicine Bow Mountains, Wyo., and Mount Lincoln, Colo.
4 Five specimens from northern Colorado.

59 NORTH AMERICAN FAUNA. [No. 37..

specimens being considerably redder than typical Juteola; the skulls,
however, are typical.
Specimens examined.—Total number, 42, as follows:
Colorado: Boulder County (altitude 8,000-11,000 feet), 4;1 Coulter, 1; Elk-
head Mountains, 1; Estes Park, 1; Lake John, 1;? Longs Peak, 1; Meeker,
2; Middle Park, 1; Mount Lincoln, 6; North Park, 1;* Pikes Peak, 1;?
Sheephorn Pass, 4;7 Steamboat Springs (18 miles below), 2;? Sulphur
Springs, 5.°
Wyoming: Laramie Mountains, 7; Riverside, 2; Sherman, 1; Woods P. O., 1.

MARMOTA FLAVIVENTRIS WARRENI Howe tt.
WARREN’S Marmot.
(PI-Vi, fic. 4° Pl eh fier)
Marmota flaviventer warreni Howell, Proc. Biol. Soc. Washington, X XVII, 1914, p. 16.

Type locality. Crested Butte, Colo.

Distribution.—Western Colorado, from Garfield County south to
Saguache County; exact limits of range unknown.

Characters.—Size large (equaling dacota) ; colors deep red with little
buff; skull similar to that of obscura, larger than that of dacota or
luteola.

Color.—General tone of upperparts hazel, the underfur at base
between mouse gray and fuscous, succeeded by pinkish cinnamon
or cinnamon; long hairs hazel subterminally, tipped with a small
area of buffy white; top and sides of head dark chestnut or bay;
sides of neck cinnamon-buff; underparts hazel, varied with ochra-
ceous-tawny, becoming chestnut on lower abdomen and Sanford’s
brown on throat; lips soiled whitish, bordered with cream-buff or
pinkish cinnamon; fore legs hazel, shading to auburn on feet; hind
feet and toes hazel, varied with cinnamon; tail dark chestnut-brown
at base, shading to hazel or tawny; under surface blackish chestnut-
brown.. |

Skull—(Known only from females): Larger than that of dacota
with longer, slenderer rostrum and narrower palatal foramina;
very similar to that of obscura, but averaging longer and relatively
narrower.

Measurements.—Adult female (type): Total length, 565; tail verte-
brae, 131; hind foot, 82. Skull:® Adult female: Condylo-basal
length, 83.3-89.3 (average 85.9); palatal length, 47—49.3 (47.9); post-
palatal length, 33.6-36.9 (35.5) length of nasals, 36.8-39.2 (38.1);
zygomatic breadth, 57-59 (57.7); breadth across mastoids, 42.9-

1 Two in collection Field Mus. Nat. Hist.

2 Collection E. R. Warren, Colorado Springs, Colo.
3 Collection Mus. Comp. Zool.

4 Collection Colo. Mus. Nat. Hist.

5 Three in collection E. R. Warren; one in collection Amer. Mus. Nat. Hist.
$ Three specimens from Crested Butte and Mud Springs, Colo.

“4915.1 MARMOTA FLAVIVENTRIS GROUP. 58

44.7 (43.7); least interorbital breadth, 19-21.2 (20.2); breadth of
rostrum, 18.7-20.3 (19.6); maxillary tooth row, 14.9-20.5 (17.4).

Remarks.—This form most nearly resembles lJuteola in color, but is
much redder (less yellowish). Its skull also is much larger, agree-
ing rather with that of obscura. It is known from only a few speci-
mens and its range has not been definitely determined. It probably
will be found throughout west-central Colorado and adjacent parts
of Utah. In both size and color it differs markedly from engelhardti,
but additional material may show that the two forms intergrade.
Intergradation with luteola is indicated by a series of specimens from
Sulphur Springs (referred to luteola), and with obscura by a specimen
from Florida, Colo. (referred to obscura).

Specimens examined.—Total number, 11, as follows:

Colorado: Crested Butte, 4;! Cochetopa Pass (9 miles south), 1; Mud Springs,
‘Garfield County, 5;? Sapinero, 1.

MARMOTA FLAVIVENTRIS OBSCURA Howe tt.
Dusky Marmot.
(PIO 1h, fig? 2-Pl Vill, fic. 4; Pl. XIV? fig. 4.)
Marmota flaviventer obscura Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 16.

Type locality Wheeler Peak, 5 miles south of Twining, N. Mex.
(altitude, 11,300 feet).

Distribution.—Upper slopes of high peaks in northern New Mexico
and southern Colorado, from Pecos Baldy, N. Mex., north to Sierra
Blanea, vicinity of Fort Garland, and to San Juan Range near Osier,
Colo. (formerly in the Manzano and Datil Mountains, N. Mex.); occurs
in Hudsonian and upper Canadian Zones from about 9,600 feet alti-
tude to the summits of the peaks (13,300-13,700 feet).

Characters.—Size large (exceeding dacota and equaling flaviventris) ;
sexes about same size; taillong; colors dark brown mixed with white,
with relatively little of the buff or tawny shades of other races; face
usually without white markings; skull similar to those of warreni and
dacota.

Color.—Adults: General tone of upperparts dark brown, grizzled
with white, becoming cinnamon on hinder back in some individuals;
underfur clove brown succeeded by pinkish buff, shading in some
specimens to pinkish cinnamon on hinder back and rump; long hairs
dark chestnut-brown, finely grizzled with white; head and face dark
chestnut-brown or black, grizzled with white on sides of face, rarely
with a whitish band across nose; sides of nose, lips, and chin, white
or buffy white; feet dark chestnut-brown often extensively mixed
with white or buffy hairs, or sometimes cinnamon-buff shaded with
dark brown; tail chestnut-brown, shading to blackish brown beneath;

1 Three in collection E. R. Warren.
2 Four in collection E. R. Warren; one in Amer. Mus. Nat. Hist.

54 NORTH AMERICAN FAUNA. [No. 37. -

underparts mixed dark chestnut-brown or blackish brown and pale
buff (tilleul buff of Ridgway) in varying proportions, buff usually
most pronounced in the median line; chin, and sometimes throat,
shaded with tawny or bay. Young (Osier, Colo.): General tone
above, clove brown (becoming slightly more tawny on hinder back)
sparingly grizzled with white; underparts mixed blackish brown and
light buff; tail snuff brown above, pale clove brown below; hind feet
same color as under side of tail, overlaid with light buff.

Skull.—Similar to that of dacota, males about the same size or
slightly smaller, females larger; nasals (in females) averaging longer;
palatal foramina narrower; interpterygoid fossa broader; postorbital
constriction narrow; similar to that of warren. but averaging shorter
and relatively broader with slightly broader premaxillae.

Measurements.—Adult male:! Total length,.645-664 (average 655) ;
tail vertebrae, 180-220 (204); hind foot, 90-92 (90.7). Adult female:1
630-670 (646); 180-220 (190); 88-90 (89.3). Skull: Adult male:?
Condylo-basal length, 88.3-90.8 (89.5); palatal length, 49.4-52.7 (51);
postpalatal length, 34-35.2 (34.6); length of nasals, 36.4-38.4 (37.4);
zygomatic breadth, 60.2-60.4 (60.3); breadth across mastoids, 41.5—
41.7 (41.6); least interorbital breadth, 21-21.1; breadth of rostrum,
21.9-23.3 (22.6); maxillary tooth row, 19-19.8 (19.4). Adult female:
Condylo-basal length, 84.5-89.5 (87.7); palatal length, 48.4-50.3
(49.3); postpalatal length, 31.9-35.7 (34.2); length of nasals, 35.4-39
(37.5); zygomatic breadth, 59.4-60.6 (59.8); breadth across mas-
toids, 41-43.8 (42.7); least interorbital breadth, 20.4-22.3 (21.1);
breadth of rostrum, 22.3-22.4; maxillary tooth row, 20-21.

Remarks.—This is the darkest and one of the largest of the races
of flaviventris. In its dark colors and the absence of light face mark-
ings it somewhat resembles the monaz group. It is closely related to
warreni and probably intergrades with it, but material from southern
Colorado is too scanty to show clearly its relationships. A badly worn
skin without skull from ‘Fort Massachusetts’ [=mountains near]
has been in the National Museum collection for many years, but not
until 1903 and 1904, when Vernon Bailey collected a fine series of
adults in the Pecos River and Taos Mountains, N. Mex., was it possi-
ble to determine the characters of the species.

In a series of 12 specimens from Osier, Colo., in the San Juan
Range, about half of the individuals are considerably paler above
than in the typical form, being uniformly pinkish cinnamon grizzled
with white, without prominent brownish markings. A badly worn
specimen from Florida, Colo., is decidedly redder above, particularly
on the head and feet, indicating apparent intergradation with

1 Four specimens from New Mexico.

2 Two specimens from Wheeler Peak, N. Mex.
3 Three specimens from Wheeler Peak and Pecos Baldy, N. Mex.

1915.] MARMOTA FLAVIVENTRIS GROUP 55

warrent. Strangely, however, the skull of this specimen is not like
that of either warren or obscura, but agrees well with that of luteola.

Several lower jaws and fragments of crania, found in a cave on
the Manzano Mountains by Archibald Rea, and broken pieces of a
skull secured by Dr. Walter Hough from a cave on the Tularosa River
near Old Fort Tularosa (south slope of the Datil Range) indicate the
former occurrence of this species in those ranges. The jaws from
the Manzano Mountains agree essentially with recent material, but
the fragment from the Tularosa River is not specifically identifiable.

Of the habits of this marmot, Vernon Bailey in his field notes
says: *

They live entirely in or among rocks and prefer open country, either in parks or
above timber line. They often burrow under large bowlders in the parks and mead-

ows, but more often live in fathomless piles of broken rock piled along the base of
cliffs, or in seams and crevices of the cliffs themselves.

Specimens examined.—Total number, 24, as follows:

Colorado: Osier, San Juan Mountains (altitude, 9,625 feet), 12;1 Florida, La
Plata County (altitude, 7,200 feet), 1;? ‘‘ Fort Massachusetts” [probably from
Sierra Blanca Peak], 1

New Mexico: Pecos Baldy, 2; Truchas Peak, 2; Wheeler Peak, 4; Aqua Fria
Peak, 2.

Cranial Measurements of the Marmota flaviventris Group.

| |

es i ial =
| | | ia |S 5
| op d Pest =p ioe Beedle
S | a 3p oo ba =}
oO n fo) So
moles (oat lesa Sale leech asl ss
; S |S123/14/8 |eaigs! $] 8
No. Species and locality. See | Sols ISS Sel etal | Remarks:
3 =| 3 3 ae ok ie ES
> |SIZiSlais lj |sis
iB Samii e bein 1. [eis
ma }o |e /MIHIN|IE WH |Ala
Marmota flaviventris flaviventris. : .
100532-i- Denner, Cal. ....2.:-...-.-.----5 roy 97.1 |55 38. 2/42. 6 62. 3/46. 2/23 |22.3)20.9) Adult.
4758" |; Fort Crook, Cal. ...-....--.----.- 3 | 94 (54 |37.6/42.5)..../44.4)21.2/23 |20 Do.
203080 | Mount Hood, Oreg2 see Se ee [oe2i 59255; 15223) 2225 --- -(41.1)22 |23 119.5 Do.
80360 | Crater Lake, ’Oreg ~soc5Sddoesscos 87 |48.7/33. 7137 _|58. 4/40. 7\19. 2}19. 6)20 Do.
Poros Donner Calo i222. sec ss 5. - 25 86.2 |48. 7/34. 5/37. 2)... .)40. 4/17. 3/19. 4/21 Do.
311901 | Glen ees Springs, Cal......-.. Q | 84.1 148. 2/32.7/37 |57.6/39.2)19 (21. 2/20.5 Do.
23951 | Carson, Nev.........-..-- Eat Q 84.6 |48. 5/31. 8135. 455. 4/39. 6/18. 7/21. 4/22.1 Do.
Marmota flaviventris avara.
99759 | Okanogan, British Columbia. . rol 85.7 |47.9/34. 8/37. 6)56. 7/42. 2)19.5}20 |19.3) Adult.
4917 | Cascade, British Columbia...... S | 86.5 |48.8/34 (36.6155. 6/39. 1/17. 8119. 4/19 Do.
94343 | Okanogan, British Columbia... 2 78 43.3)31 33. 2/52. 7/37. 6|18 16.817.8 Do.
178842 |..... 0 (02 Ae eR OBC See aera Le 76.4 42. 6/30. 3/31. 552.5 38. 6|17. 117. 3)19.1 Do.
41107 | Midway, British Columbia...... 79.4 |44. 6/30. 8/33. le ..-[37.8|17.4|17 |19 Do.
Marmota flaviventris sierrae |
315157 pana Meadows, Tulare Coun- |
Wavalee osu case tae. (ste seas [S?] | 85.5 |47.8)34.3 36. 6/57. 3/40. 7)18. 5/20 (20.2) Old.
315165 Whitney Creek, Tulare County,
A ee te a ae Se Si 90.2 |50.5)35 (37 |58.3/43.6/20.3/20 |20.5| Adult.
42641 | Mount Whitney, Cal............ of 79.1 [44 |30.832.6)52 |37.5)17 |17.4/20.2; Subadult.
41108.j..... cr EI SE ee OE ae Se S | 84.5 |48 [82.439 [54.6/41.5/17.8)17 |20.2 Do.
42859 |_.... t CS Ga SSR SS ODE Oe abe ae aenaers Q 82.1 |46. 3/32. 237. 4/55. 2/39. 3)18.118.2/20 | Adult.
315163 | Whitney Meadow, Cal.......... Jee 77.3 |44.7/28.732 |52.7/37 |16.5)17.4/19 Old.
41950 | Head San Joaquin. River, Cal. . 81 46 131 (34. 41..--137.9|18. 7/18. 6119.5 Do.

1 Collection Colo. Mus. Nat. Hist. 3 Collection Mus. Vert. Zool., Univ. of California.
2 Collection Amer. Mus. Nat. Hist. 4 Collection Victoria Mem. Mus.

56

NORTH AMERICAN FAUNA.

[No. 37.

Cranial measurements of the Marmota flaviventris group—Continued.

| : ale (ae
be a) ..| S18 le yseiie
& bp 4 Siig 21S sere!
= | a DS ca yell ae lines
BS |e|/2)/ 4/5 /Egiss| 3] 8
No. Species and locality. 3 PlalA|’o |Sslas = | ~ | Remarks.
o) = 8 by 3 rie 25 q Be ‘
td 3 ~ ~ ic
|e |$lelelir isigia
He lslai es Bie lo [21s
) O;eH/A HINA AH [als
Marmota flaviventris parvula.
93689 | Arc Dome, Toyabe Range, Nev.. g 73 {41 +128. 6/80. 5/48. 6/34. 3/15. 5/17. 2|17. 7| Adult.
93690 | Jefferson, Toquima Range, Nev. 71.3 |39.4/27 |29.4/49 |34.2/15.6/17. 4|18.6] Adult; type.
Marmota flaviventris engelhardti.
157828 | Beaver Mountains, Utah........ [2?] | 80 |46.1/30.7/34.6/54 |37.5]17 |17.8]19 | Adult.
15 SO1 Sul ee eG a ae ae: das cients é 78.4 |44. 8/30. 2/33 |52.3/36.5/16.3/18 |19 Do.
158500 | Parawan Mountains, Utah. ..-. 76.8 |43.5|29.5/31 |52.8/37.7|18 |18 18.8 Do.
Marmota flaviventris nosophora.
168493 | Ross Fork, Mont..............-. S | 86.7 |47. 7/34. 2/35. 1/55. 7/41. 3/17. 8/19. 5/20. 1] Young adult.
156924 | Sawtooth National Forest, Idaho rol 90 |50. 3/35. 5/38. 5/59. 5)43.1)19 |19.6/19.5| Adult.
191363 | Conant Creek, Idaho..........-- 3 | 92.2 |52. 5/35. 2/39. 1/57.3/42 |21 |20.5/20.8 Young adult.
66709 | Pryor Mountains, Monica sao So | 94.2 {53 |37. 2/40. 4/63. 7/45 |22.2/20 (20.4) Adult.
168473 | Bitterroot Valley, Mont......-.. 83.8 |46. 2133. 5/34. 8/54. 5/41. 6119. 5/19. 3/20. 2 Do.
168472 | ..... dO: oe aeweccisaicisen Se So 78.2 |42. 5/31. 8/30. 8/54 37. 6/19. 2/18. 7/20.3 Do.
168494 | ..... ra Re ERA tert a ESM te 79 =|45.5/80 |32.8/53.2/40 |19.7/17 {19.8} Adult; type.
* Marmota flaviventris dacota.
186474 Custer, S. Dak ice. secu jee S | 89.4 |50.8)35 [388 |59.5/43 |21.8/22 |21.5) Subadult;
type.
65920 | Bear Lodge Mountains, Wyo...-| S& | 95.7 152. 2/38. 2/41. 1/61. 3/44. 7/23. 2/23. 3/21. 7; Adult.
25529 | Bridger Pass, Wyo..-..-..-..-.-..-- 90.5 |51 136. 2/38. 4/61. 7/41. 8]18. 2/20. 5/20. 9 Do.
191365 eustal; S. Dak Siverseic retina tte eiatctal ers 83.5 |47. 6/31. 8/34. 1/56. 7/40. 6/20. 7/20 |20.1 Do.
TOTO GA OS ae ingens ie cis woe aeeer 81.8 |45. 4/32. 5/31. 5/55. 3/39. 5/20 |20.5|20.3 Do.
168884 paver, S eDake area meee erises 84.5 |47.7/33 |85 155. 7/40. 7/19. 6/20. 4/20. 5 Do.
Marmota flaviventris luteola.
186520 | Medicine Bow Mountains, Wyo.} & | 87 |48.4/83.5)36 |55.7/40.6]17.3/19 |21.4 gir adult
ype.
25523 | Laramie Mountains, Wyo...-... rol 87.8 |49.5)/35.5/37.3/60 |43. 4/20. 2/19. 6)19.9) Adult.
1175 | Mount Lincoln, Colo...........- [?] | 92.5 |51.1/36.5]41 |59.4/43 |21. 8/20. 8/21 Old.
2 2279 | Steamboat Springs, Colotessset 85 147. 6/33.3/34 }54.9/40. 5117. 4/17. 7/21. 4) Adult.
2/3998 | Lakevohn;'Colo2 ..- 22s. sees 81.3 |45.3/31. 7/33 [54 |39 |16.4/17.7/19.3 Do.
139082 | Coulter, Cole. isk Site ae 78.3 |43.7/30 |34 |52.3)/38. 5/16. 5/17. 3)19.5 Do.
Marmota flaviventris warreni.
202937 cacsied Butte, Colo.:: ss 2c. 89.3 |49. 3/36. 1139. 2159 44. 7/21. 2/20. 3)16.7} Adult; type
DBE Geos GOAN tive ee Gua teene sobinereee 83.3 |47. 4136. 9/36. 8157 |42.9)19 |19.9]14.9} Adult.
22512 | Mud Springs! Colossal eee 85.1 |47 |33.6/38.2/57 |48. 4/20. 3/18. 7/20. 5 Do.
Marmota flaviventris obscura
135504 AU hedler Peak, .NAMexes soo 52 3 3S | 90.8 |52.7/34 138. 4/60. 4/41. 7/21 |23.3)19 | Adult.
ASS50GUE 2. SAO SRS 28a SEs oe ae rol 88.3 |49. 4135. 2/36. 4/60. 2/41. 5/21. 1/21. 9/19. 8 Do.
128750 Bee Baldy, NoMex. Saocsee- 89.5 |50.3/85 89 {59. 4/43. 8/20. 4/22. 4/21 Do.
133505 | Wheeler Peak, N. Mex...-.-..-..- 84.5 |48. 4131. 9/35. 4159. 4/41 120. 5)22. 3/20 Do.
133507 |..... Ota ie Ope Sik con setae 88.1 |49. 3/35. 7/38. 2/60. 6/43. 3|22. 3/22. 3/20 Do.

1 Collection Mus. Comp. Zool. 2 Collection E. R. Warren, Colorado Springs, Colo.

Marmota caligata Group.

The caligata group includes three species: IM. caligata, M. olympus,
and M. vancouverensis.

External characters.—Size large; tail long (about 27 to 33 per cent
of total length); ears relatively small (actually smaller than those of

1 Weight of 6 specimens as follows: 9 lbs. (2 9, cascadensis); 11 lbs. (¢, cascadensis); 113 lbs. (2, olympus);
153 lbs. (3, orytona); 17 lbs. (old 9, cascadensis).

1915.] MARMOTA CALIGATA GROUP. 57

M. monaz); posterior pads on sole of hind foot subcircular and situ-
ated near edges of sole (see Pl. III, fig. 1); mammae:! P. 2; A. 2;

=10; colors mainly black and white, shaded with cinnamon-buff
on hinder parts, or upperparts of solid colors—brownish drab, russet,
or vandyke brown.

Cranial characters.—Skull with superior outline nearly straight (as
in the other American groups); interorbital region and postorbital
processes much as in the flavwentris group;? nasals narrowed posteri-
orly, usually about same width at posterior end as premaxillae or
slightly narrower (wider in olympus); temporal ridges uniting in old
age to form a pronounced sagittal crest; anterior portion of floor of
basi-occipital nearly flat, bounded posteriorly by two low processes
which unite at about middle of basi-occipital, continuing as a pro-
nounced ridge to the foramen magnum; anterior portion often with two
rather pronounced depressions on either side of the median ridge;
palate beveled at posterior border (as in flavwentris group); interptery-
goid fossa relatively narrow (compared with monaz); palatal foramina
variable in shape; molar teeth similar to those of monaz; maxillary
tooth rows divergent anteriorly; anterior face of incisors ivory yellow
to orange-buff.

Geographic distribution.—From the Endicott Range, Alaska—the
most northerly range in the Rocky Mountains—and the Alaska
Peninsula south to the Olympic Mountains and Mount Rainier, Wash.,
and the Bitterroot and Salmon River Mountains in central Idaho;
also on Vancouver Island. Confined entirely to mountain sides at
and above timberline except in the north, where the animals live in
open meadows and descend to tide water. (See fig. 3.)

Remarks.—The members of this group may readily be distinguished
by their greater size and their peculiar coloraticn—either mixed
black and white or solid brownish. All of the races of Marmota cali-
gata are colored much alike, differing mainly in relative proportions
of black and white and in skull characters. MM. olympus, isolated on
the Olympic Peninsula, has developed a brownish drab coloration
with relatively little black or white, and UM. vancouverensis, dwelling
on Vancouver Island, has lost all of the black and white colors and
attained a dark seal brown pelage.

MARMOTA CALIGATA (EscHscHOLTz).
[Synonymy under subspecies.]

External characters.—(See under Marmota caligata group, excepting
color. )

Cranial characters.—(See under Marriot caligata group. )

Color.—Fore part of back mixed black and white in varying pro-
portions, sometimes with a buffy or brownish tinge; hinder back
similar, but usually strongly suffused with cinnamon-buff or pinkish

1 Asin M. flaviventra. 2 Excepting M. olympus, which is relatively wider interorbitally.

58

Zz
nai

M caligata
93 ae
39
9
9?
1%
mJ

a
2.
J.
4h.
ire).
6.
Lie ms
8. °° olympus
9

NORTH AMERICAN FAUNA.

4
‘a 4
a,

i b
PNY 4
| i
p \
\
f]

LAs
EO
QOeX

caligata : fo;
sheldom ZB
" oxytona —:

okanagana
nivaria
cascadensis
vigilis

"™ wvancouverensis

‘many places theoretica

[No. 37. :

Fic. 3.—Distribution of the Marmota caligata group. Boundaries of ranges of subspecies are in
1.

1915.] MARMOTA CALIGATA GROUP. 59

cinnamon, sometimes tinged with hazel or vandyke brown; under-
fur dark hair-brown, bone brown, clove brown, bister, mouse gray,
fuscous, or fuscous-black; sides of head and neck cinnamon, ochrace-
ous-cinnamon, vandyke brown or blackish brown, more or less mixed
with white or buffy white; borders of nose, lips, and chin soiled whit-
ish; top of head and face black or blackish brown; face in front of
and between eyes more or less extensively marked with white; legs
usually same color as back; feet black or blackish brown, the front feet
often with white markings, the hind feet mixed with cinnamon; tail
above, cinnamon, pinkish cinnamon, cinnamon-buff, clay color,
chestnut-brown, or bay, often mixed with hazel or vandyke brown;
beneath, chestnut-brown, clove brown, blackish brown, or hessian
brown, mixed with hazel, bay, or ochraceous-tawny; underparts
white, grayish white, or clay celor, varied with cinnamon, cinnamon-
buff, or blackish brown.

Geographic disiribution.—From the Endicott Range and the Alaska
Peninsula, Alaska, south 1o Mount Rainier, Wash., and the Bitter-
root and Salmon River Mountains, Idaho.

MARMOTA CALIGATA CALIGATA (EscuscHoLtz).
NorTHERN Hoary Marmot.
(Pl. EV, fig: 6; PI.-TX, fie: 4; Pl. XV, fig. 3.)

Arctomys pruinosa Richardson, Zool. Jour., III, 1828, p. 518. (Not of Gmelin.*)

Arctomys caligata Eschscholtz, Zool. Atlas, Part II, 1829, p. 1, Plate VI.

Arctomys pruinosus Richardson, Fauna Boreali-Amer., I, 1829, p. 150; Baird, Mamm.
N. Am., 1857, p. 345; Allen, Mon. N. Am. Rodentia, 1877, p.924. (Not ofGmelin.

Arctomys caligatus Tyrrell, Proc. Canadian Inst., 3rd Ser., VI, 1888, p. 88.

Marmotta caligata Allen, Bul. Amer. Mus. Nat. Hist., XIX, 1903, p. 539.

Marmota caligata MacFarlane, Proc. U.S. Nat. Mus., XXVIII, 1905, p. 751.

Type locality.—Bristol Bay, Alaska.

Disiribution.—Alaska and Yukon, from the Portland Canal north
on the coast to Bristol Bay, and in the interior to the Endicott
Range and the mountains lying westward of Fort Good Hope,
Mackenzie.

Characters.—Size medium (for the group); colors moderately white;
skull relatively short and broad.

Color.—General tone of upperparts white, moderately tipped
with black, becoming cinnamon-buff on hinder back and rump;
underfur at base fuscous or dark mouse gray (shading on hinder
back to bister), succeeded on fore back by a-broad area of white
which gradually shades to cinnamon-buff on hinder back (in some

1 Arctomys pruinosa, Gmelin, Syst. Nat., I, 1788, p. 144—based on the hoary marmot of Pennant. A

careful comparison of Pennant’s description with specimens of the several American species leads to the
conclusion that this species is unidentifiable; the name pruinosa is therefore rejected.

60 NORTH AMERICAN FAUNA. [No. 37.

specimens underfur between shoulders white to roots); top’ of head
and face black, with a white patch in front of eyes often covering
the whole face; black of crown usually extending from ear back
over shoulders in the form of two divergent (more or less indistinct)
stripes; sides of face mixed brown and white (often nearly white) ;
fore legs white or buffy white, hind legs pale cinnamon-buff; fore and
hind feet black or blackish brown; tail above, cinnamon-buff, tipped
with blackish brown or bay, the bases of the hairs extensively
dark chestnut-brown (or sometimes natal brown); beneath, blackish
brown or chestnut-brown, sometimes tinged with hazel or ochra-
ceous-tawny; underparts soiled whitish, sometimes mixed with black
or blackish brown.

Skull.—Relatively short and broad with short, broad rostrum;
zygomata broadly expanded posteriorly; nasals long, extending
beyond ends of premaxillae; postorbital constriction rather broad.

Measurements.—Adult male: Total length, 710-715; tail verte-
brae, 210-218; hind foot, 91-105. Adult female:? 675; 190; 95.
Skull: Adult male:* Condylo-basal length, 96.5-100.6 (average98. 7);
palatal length, 55.7-57.4 (56.4); postpalatal length, 36.5-38.2 (37.4);
length of nasals, 38.4-42.3 (40.7); zygomatic breadth, 64.5-68 (66.3);
breadth across mastoids, 44-46.2 (44.4); least interorbital breadth,
24.3-27.6 (25.8); breadth of rostrum, 23.7-26.1 (24.6); maxillary
tooth row, 22-23.8 (22.6). Adult female:+ Condylo-basal length,
92.8-96.4 (94.2); palatal length, 53-54.8 (53.8); postpalatal length,
35-38 (36.3); length of nasals, 38-39.7 (38.8); zygomatic breadth,
61.6—64.1 (62.7); breadth across mastoids, 40—43.3 (41.9); least inter-
orbital breadth, 23-25 (23.9); breadth of rostrum, 22.1-24.5 (23.2);
maxillary tooth row, 20.6-22.5 (21.7).

Remarks.—This race has an extensive distribution in Alaska and
Yukon and shows little variation over its whole range. In the
southern part it is confined to the region near the coast but in the
north it occupies the interior mountain ranges as far east as the
main Rocky Mountains in northwestern Mackenzie. In specimens
from the Kenai Peninsula the nasals average somewhat shorter than
in typical specimens, rarely extending back of the posterior ends of
the premaxillae, but the skulls show no other differences. Speci-
mens from the southern coast of Alaska as far south as the Portland
Canal (with the exception of the vigilis series from Glacier Bay) are
typical, but intergradation with oxrytona occurs a short distance from
the coast in northern British Columbia, the series from Cheonee
Mountains being distinctly intermediate in character. The form
occupying the northern Rockies in eastern Yukon and western

1 Two specimens from head of Coal Creek, Yukon.
2 One specimen from same locality.

3 Five specimens from Becharof Lake, Alaska.
4 Seven specimens: 6 from Alaska Peninsula, 1 from Coal Creek, Yukon.

1915.] MARMOTA CALIGATA GROUP. 61

Mackenzie is provisionally referred to this race, no skulls from this
entire region being available.
Specimens examined.—Total number, 102, as follows:

Alaska: Aleknagik Lake, 2; Becharof Lake, 12; Cape Elizabeth, 8; Charlie
River (near head), 1; Chickamin River (Behm Canal), 2;! Cordova Bay, 3;}
Disenchantment Bay, 1;! Fort Yukon, 1;? Hinchinbrook Island, 4; Juneau,
4; Kanatak, Portage Bay, 1; Kenai Mountains, 10;} Kenai Peninsula, 4;
Mount McKinley, 1; Portage Bay, 5; Port Snettisham, 2;) Seldovia, 7;3
Toklat River (near head), 1; Valdez Narrows, 1;! White Pass, 4; Yakutat
Bay, 2; no specific locality, 2.

British Columbia: Bennett, 2; Cheonee Mountains, 15.3

Mackenzie: Fort Good Hope (mountains west of), 1.

Yukon: Coal Creek (head) 4; Kalzas Creek, Pelly River, 2.

MARMOTA CALIGATA VIGILIS HeEtuer.
GLaciER Marmot.
Marmota vigilis Heller, Univ. of California Pub. Zool., V, 1909, p. 248.

Type locality—_ West shore of Glacier Bay, Alaska.

Distribution.—Known only from type locality.

Characters.—Similar in size and skull characters to caligata;
variable in color, but constantly darker than caligata, with a strong
tendency to run to blacks and browns; top of head and hind feet always
brown (except in purely melanistic individuals); underparts darker.

Color.—Normal fresh pelage (June 12): Upperparts soiled whitish
with a buffy tinge; underfur clove brown at roots, sparingly mixed
with dull brown or black; hinder back shading to cinnamon-buff,
darkest on rump and hind legs, extensively tipped with blackish
brown; underparts dull clay color mixed with grayish white; nose,
top of head, and feet blackish brown; sides of face mixed cinnamon-
buff and blackish brown; tail above, mixed clay color and chestnut-
brown; beneath, blackish chestnut-brown shading to hessian brown
in anal region. Variation: The above description is based on the
lightest and apparently most normally colored individual in the
series of topotypes. Between this phase and the pure black phase
are found numerous intermediate specimens, some as white beneath
as caligata and as dark above as orytona, while others are solid black-
ish brown very similar to M. vancouverensis, but slightly blacker; the
hind feet and usually the tail are blackish brown; fore feet black or
very dark brown.

Skull.—Not appreciably different from that of caligata.

Measurements—Adult male:4 Total length, 630-745 (average, 676);
tail vertebrae, 197-210 (205); hind foot, 92-102 (97). Adult female: ®
620-680 (652); 170-212 (191); 90-95 (92). Skull: Adult male:* Con-
dylo-basal length, 97.2-98.4 (97.6); palatal length, 54.7-55.8 (55.3);

1 Collection Mus. Vert. Zool., Univ. of California. - 4 Five specimens from type locality.

2 Collection Mus. Comp, Zool. 5 Three specimens from type locality.
3 Collection Amer. Mus. Nat. Hist,

62 NORTH AMERICAN FAUNA. [No. 37.

postpalatal length, 36.7-38 (87.5); length of nasals, 38.9-41.5 (40.6);
zygomatic breadth, 62-67.7 (64.8); breadth across mastoids,43.2-44.6
(43.8); least interorbital breadth, 22.6-26 (24.3); breadth of rostrum,
21.1-23.5 (22.2); maxillary tooth row, 21.8-22.5 (22.2). Adult
female: 1 Condylo-basal length, 93.7; palatal length, 51.6; postpalatal
length, 37.7; length of nasals, 39.6; zygomatic breadth, 61.8; breadth
across mastoids, 42.3; least interorbital breadth, 24.5; breadth of
rostrum, 21.3; maxillary tooth row, 22.

Remarks.—This form is apparently an incipient species or race in
which the characters have not as yet become fixed. It shows the
same tendency to become brown that appears so strikingly in M. van-
couverensis, and to a lesser degree in M. olympus. Its range is appar-
ently limited to the region about Glacier Bay (since specimens from
Yakutat Bay, White Pass, and Juneau are referable to calgata) but
whether it is actually isolated is not known.

Specumens examined.—Ten, from type locality.?

MARMOTA CALIGATA SHELDONI Howe tt.
MontacueE Is~tanp Marmor.
Marmota caligata sheldoni Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 18.

Type localty.—Montague Island, Alaska.

Distribution.—Known only from type locality.

Characters.—Similar to caligata, but size smaller and nasals shorter.

Color.—-Upperparts mixed white and black, whitest on fore back,
shading to cinnamon-buff on rump; underfur fuscous, shading on
hinder back to natal brown; top of head black; a large white patch
on face in front of eyes; sides of face mixed blackish brown and cinna-
mon; tail deep cmnamon-buff or ochraceous-buff, mixed with black,
the bases of hairs chestnut-brown; underparts white, tinged with
cinnamon.

Skull.—Similar to that of caligata, but decidedly smaller; nasals
shorter and broader posteriorly, terminating usually about on a line
with ends of premaxillae; premaxillae narrower.

Measuremenis.—Adult male topotype: Total length, 670; tail ver-
tebrae,-185; hind foot, 94. Adult female topotype: 640; 180; 90.
Skull: Adult male: * Condylo-basal length, 89.5-96.6 (average, 93.6) ;
palatal length, 51.2-55.4 (53.3); postpalatal length, 34.3-37 (35.5);
length of nasals, 35-39 (37.3); zygomatic breadth, 61.5-63.4 (62.4);
breadth across mastoids, 41.3-42.4 (41.9); least interorbital breadth,
22.5-23.8 (23.3); breadth of rostrum, 20.5-21.8 (21.2); maxillary
tooth row, 22.2-23.1 (22.5). Adult female:1 Condylo-basal length,
88.7; palatal length, 50.2; postpalatal length, 34; length of nasals, 37;
zygomatic breadth, 59.4; breadth across mastoids, 39.7; least

1 One specimen from type locality. 2 Four specimens from type locality.
2 Seven in collection Mus. Vert. Zool., Univ. of California,

1915.] MARMOTA CALIGATA GROUP. 63

interorbital breadth, 22.2; breadth of rostrum, 20.5; maxillary tooth
row, 21.8.

Remarks.—This race is a small form of caligata, confined to Mon-
tague Island, and differing from the typical race in size and skull
characters. On Hinchinbrook Island, separated from Montague
Island by only a narrow channel, the typical form is found.

Specimens excamined.—Seven, from type locality.’

MARMOTA CALIGATA OXYTONA HOottister.
Rosson Hoary Marmot,
Gertie te Ie We hea PIO tip! 2: Pl XxX fig. 4.)

Marmota sibila Hollister, Smiths. Misc. Coll., Vol. 56, No. 35, 1912, p.l. (Not Arctomys
sibila Wolf.)

Marmota oxytona Hollister, Science, N. S., XX XIX, No. 998, Feb. 18, 1914, p. 251
(new name for Marmota sibila Hollister).

Type locality—Head of Moose Pass branch of Smoky River,
Alberta (altitude, 7,200 feet).

Distribution.—Interior of northern British Columbia, southwestern
Mackenzie (?),and southern Yukon, from Teslin Lake and Liard River _
south to Barkerville, British Columbia, and the Mount Robson region,
British Columbia and Alberta.

Characters.—Colors much blacker and tail darker than in caligata;
skull larger and relatively narrower; males but little larger than
females.

Color.—Upperparts as in caligata, but more extensively tipped with
black, the underfur fuscous, shading to deep mouse gray or fussous-
black; hinder back dark cinnamon-buff, tinged with hazel and
heavily mixed with black; top of head black with rather small white
patches; cheeks ochraceous-cinnamon, varying to buffy white; tail
above, dark cinnamon-buff mixed with bay, the bases of the hairs
chestnut-brown or blackish brown; beneath, blackish brown, tinged
with bay; underparts soiled whitish mixed with dull cinnamon, the
bases of the hairs blackish brown.

Skull.—Much longer and relatively narrower than that of caligata,
with long rostrum; zygomata less widely expanded posteriorly; zygo-
matic arch longer and distance from squamosal arm to tip of post-
orbital process greater; nasals terminating on a line with ends of
premaxillae or slightly posterior; interpterygoid fossa relatively
narrower.

Measurements —Adult male: ? Total length, 720-775 (average, 747);
tail vertebrae, 210-235 (221); hind foot, 100-110 (105.6). Adult
female: * 720-740 (730); 210; 95-105 (100). Skull: Adult male:

1 Five in collection Mus. Vert. Zool., Univ. of California.

2 Five specimens from British Columbia (Mount Robson to Thudade Lake).

3 Two specimens from Sustut Mountains, British Columbia; and head of Smoky River, Alberta.
+ Seven specimens from northern British Columbia.

64 NORTH AMERICAN FAUNA. [No. 37.

Condylo-basal length, 101—107.4 (103.6); palatal length, 57-62.5 (58.9) ;
postpalatal length, 38.3-41.7 (40.1); length of nasals, 41.5-45.3
(43.2); zygomatic breadth, 62.8-67.4 (65.8); breadth across mastoids,
44,2-48.3 (46.6); least interorbital breadth, 24-25.8 (25.3); breadth
of rostrum, 22-24.8 (23.1); maxillary tooth row, 22-23.5 (22.8).
Adult female:1 Condylo-basal length, 101.8-104.6 (1038.2); palatal
length, 56.5-61.2 (58.9); postpalatal length, 36.4-41.1 (38.8); length
of nasals, 41—-43.6 (42.3); zygomatic breadth, 65.3-66.2 (65.8); breadth
across mastoids, 45.9-47.3 (46.6); least interorbital breadth, 25.4-26.7
(26); breadth of rostrum, 23-23.3; maxillary tooth row, 22.7.

Remarks.—This is the darkest and one of the largest of the races
of cahgata. It intergrades with caligata in northern British Columbia
and southern Yukon, and with okanagana in southern British Colum-
bia, but the material at present available is not sufficient to show the
exact limits of its range. Specimens from the Liard River (Fort
Halkett and Fort Liard) are provisionally referred to this race, no
skulls from this region being available and the skins being rather
indeterminate in characters. Additional material from the northern
Rockies may extend the known range of this form farther northward
in the interior.

A series from near Teslin Lake, Yukon, is intermediate between
caligata and oxytona; the skulls are smaller than those of typical
oxytuna, some of them even smaller than those of caligata, but rel-
atively narrower. The nasals, though actually longer than in cali-
gata, do not extend so far back of the ends of the premaxillae, in this
character agreeing with oxytona.

Specumens examined.—Total number, 63, as follows:

Alberta: Smoky River (near Moose Pass), 2.

British Columbia: Babine (mountains near), 3; Barkerville, 5; Finlay River
(mountains near head), 1; Fort Halkett, 1;? Klappan River (mountains
near), 3; Laurier Pass, 1; Level Mountain, 1;? McConnell Creek (near Sus-
tut Mountains), 1; Moose Pass, 2; Moose River (north fork), 2; Sheslay
River, 2;? Stuart Lake, 2; Sustut Mountains, 4; Thudade Lake, 2.

Mackenzie: Fort Liard, 2.
Yukon: Teslin Lake (mountains near), 29.*

MARMOTA CALIGATA OKANAGANA (Krne).
OKANAGAN Hoary MARMOT.
(Pl. X, fig. 3.)

Arctomys okanaganus ® King, Narr. Journ. to Shores of Arctic Ocean, II, 1836, p. 236.

Arctomys pruinosus, Audubon and Bachman, Quad. N. Am., III, 1854, p. 17, Plate
CIII. (Not of Gmelin.)

[ Marmota] okanagana Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 17
(type locality fixed).

1 Two specimens from head of Smoky River, Alberta.
2 Collection Univ. of Michigan.

3 Collection Amer. Mus. Nat. Hist.

4 Collection Victoria Mem. Mus.; approaching caligata.
5 “Arctomys okanaganii’’ on plate.

1915.] MARMOTA CALIGATA GROUP. 65

Type locality —Gold Range, British Columbia.

Distribution —Gold and Selkirk Ranges, British Columbia, and
probably main range of the Rocky Mountains in Alberta from Banff
to Henry House; exact limits unknown.

Characters —Similar in color to ozytona, but averaging a little
whiter; skuil similar to that of caligata.

Color—Upperparts much as in ozytona, but averaging whiter
(some specimens almost as pale as caligata, but tail darker); under-
fur fuscous, shading to clove brown; fore back white, hinder back
cinnamon-buff, more or less overlaid with black: sides of face cinna-
mon mixed with white; feet blackish brown, the hind feet grizzled
with cinnamon; tail above, deep cinnamon (sometimes mixed with
hazel) bordered with blackish brown, the bases of the hairs light
chestnut-brown; beneath, blackish brown or dark chestnut-brown;
underparts soiled whitish mixed with dull cinnamon.

Skull—Similar to that of caligata, but averaging slightly larger,
with narrower rostrum and postorbital constriction; nasals shorter,
usually terminating little if any posterior to ends of premaxillae. Com-
pared with oxytona: Decidedly shorter and relatively broader; brain-
ease less elongated and distance from tip of postorbital process to
squamosal arm of zygoma much less.

Measurements.—Adult male:* Total length, 670-754 (average, 694) ;
tail vertebrae, 202-218 (212); hind foot, 91-106 (98). Adult female:?
659-735 (695); 202-224 (212); 93-103 (94). Skull: Adult male:?
Condylo-basal length, 95-100 (97.7); palatal length, 53.4-57.4 (55.5);
postpalatal length, 37-38.1 (37.8); length of nasals, 37-40.5 (38.9);
zygomatic breadth, 64.8-67 (66.2); breadth across mastoids, 45.2-46
(45.6); least interorbital breadth, 24.9-27 (25.7); breadth of rostrum,
23—25.2 (24.2); maxillary tooth row, 21.9-22.7 (22.4). Adultfemale: 4
Condylo-basal length, 94.4-97.6 (96.4); palatal length, 53.5-55.3
(54.5); postpalatal length, 36-38.7 (37.3); length of nasals, 38—40.3
(39); zygomatic breadth, 62.2-66.3 (64.4); breadth across mastoids,
42.2-45.5 (43.7); least interorbital breadth, 21.9-24.9 (23.7); breadth
of rostrum, 21—23.3 (21.9); maxillary tooth row, 21.4—22.4 (22).

Remarks.—This race apparently has a rather limited distribution,
but its characters are well marked. Jt is not in any way inter-
mediate between oxytona and nivaria, as might be expected from its
geographic position, for, while it agrees with the former in color, its
skull is much smaller than in either and more nearly resembles that
of caligata. It is very much darker in color than nivaria, the differ-
ences being especially noticeable in the young.

i Four specimens from Selkirk Range, British Columbia.
2 Six specimens from same localities.
3 Three specimens from Glacier and Nelson, British Columbia.
4 Five specimens from same localities.
69168°—15——5

66 NORTH AMERICAN FAUNA. [No. 37.

King, in naming the species in 1836, gave an excellent description
and figure of the animal, based on two living individuals which had
been brought from the Okanogan region to Norway House, Canada.
These were later presented by King to the Zoological Gardens in
London, where they were seen by Audubon, and after the death of
the animals the skins served as the basis of the figure of the hoary
marmot in his “‘Quadrupeds of North America.” The type specimen,
as | am informed by Oldfield Thomas, is still in the British Museum
collection (No. 55.12.24.126) and agrees in every detail with the
original description. The skull probably (not certainly) belonging to
the skin is so diseased by menagerie life as to be of no use for com-
parison.

The original description is so complete and agrees so well with the
form occurring in the Selkirks that I have no hesitation in fixing the
type locality in the Gold Range—the first range to the eastward of
Shuswap Lake—where it is likely the type was secured.!

Specimens examined.—Total number, 18, as follows:

Alberta: Henry House (mountains 15 miles south), 2.
British Columbia: Field, 2;? Glacier, 7; Spillimacheen River, 3;3 Toad Moun-
tain (6 miles south of Nelson), 4.

MARMOTA CALIGATA NIVARIA Howe tn.
Montana Hoary Marmot.
(PISX, figs 2: Plo ol fies 4s)
Marmota caligava nivaria Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 17.

Type locality—Mountains near Upper St. Marys Lake, Mont.
(altitude, 6,100 feet).

Distribution.—Upper slopes (at and above timberline) of high
mountains of northwestern Montana and of Bitterroot and Salmon
River Mountains, Idaho (limits of range imperfectly known).

Characters—Whitest member of the group, being very much
whiter than either okanagana or oxytona; similar in size and skull
characters to oxytona.

Color.—Adults: Fore part of back (to middle) snowy white, spar-
ingly grizzled with black, the underfur dark hair-brown; hinder back
pinkish cinnamon or cinnamon-buff mixed with black and white,
the underfur bone brown; top of head black, much mixed with white
and with a large white patch across face in front of eyes; sides of face
brownish, mixed with white and cinnamon-buff; fore feet black with
; 1 King defines the type region as follows: ‘‘In a small tract of country, on the borders of the Rocky Moun-
tains, lying between the Columbia and Fraser Rivers, these animals are found in abundance, supplying
with food and clothing the Okanagan Indians, whose territory is bounded to the north by the Seechwap
Lake, and to the south by the Spokane River * * *,” (King, R. Narr. Journ. to Shores of Arctic
Ocean, II, 1836, p. 241.)

2 Collection Victoria Mem. Mus.
3 Collection Amer. Mus. Nat. Hist.

arf

1915.] MARMOTA CALIGATA GROUP. 67

white patches; hind feet black, more or less mixed with cimnamon,;
underparts white, sparingly mixed on abdomen with cinnamon-bufl;
tail above, mixed pinkish cinnamon and chestnut-brown; beneath
clove brown or blackish brown. Young (specimen from Elk Summit,
Idaho): Nearly pure white above, shading to cinnamon-buff on hinder
back; the underfur dark mouse gray; tail cinnamon-buff fringed at
tip with blackish brown, the bases of hairs hair-brown; feet and top
of head fuscous-black.

Skull— (Known only from females.) Closely similar to that of
oxytona, possibly averaging a little shorter; much larger and relatively
narrower than that of okanagana, with broader rostrum.

Measurements — Young adult male from type locality: Total length,
755; tail vertebrae, 250; hind foot, 110. Adult female:* 700-820
(average, 751); 200-245 (224); 95-113 (105). Skull: Adult female:
Condylo-basal length, 99.5-106.5 (102); palatal length, 58-61.4
(59.4); postpalatal length, 37.5-39.6 (38.2); length of nasals, 40.6-
43.9 (42.2); zygomatic breadth, 64-66.6 (65.7); breadth across
mastoids, 44.8-45.7 (45.5); least interorbital breadth, 23.3-27.2
(25.5); breadth of rostrum, 21.6-25 (23); maxillary tooth row,
22.2-24.4 (22.9).

Remarks.—This race widely differs in color from its nearest rela-
tives, okanagana and oxytona, being even whiter than caligata. The
characters are strikingly shown by young specimens which are
almost pure white except on the hinder back and tail. Additional
material is needed to determine the exact range of the form and the
cranial characters of the males.

Specimens examined.—Total number, 14, as follows:

Idaho: Bitterroot Mountains (headwaters of Clearwater River), 3; Elk Summit,

Salmon River Mountains, 2.
Montana: Upper St. Marys Lake (mountains near), 9.

MARMOTA CALIGATA CASCADENSIS Howe tn.
CascaDE Hoary Marmot.
(Pl. X, fig. 1; Pl. XV, fig. 4.)
Marmota caligata cascadensis Howell, Proc. Biol. Soc. Washington, XX VII, 1914, p. 17.

Type locality—Mount Rainier, Wash. (altitude, 6,000 feet).

Distribution Cascade Range (at and above timberline) from
Mount Rainier, Wash., north to southern British Columbia.

Characters.—Size large (equaling olympus and ozytona; larger than
caligata); color similar to that of caligata, but head and feet usually
browner and underparts darker; skull similar to that of oxytona
but relatively broader, much larger than that of either caligata or

_ okanagana.

1 Four specimens from type locality.

68 NORTH AMERICAN FAUNA. [No. 37.

Color.—Fore part of back white or creamy white, sparingly tipped
with vandyke brown, the bases of the hairs with a broad area of the
latter color; hinder back more extensively shaded with brown and
sometimes tinged with cinnamon-buff; head and face vandyke
brown or blackish brown, often with a white patch in front
of eyes; sides of face and neck brownish, tinged with cinnamon
or buffy white; feet blackish brown, often mixed with cinnamon:
tail cmnamon-buff mixed with vandyke brown, becoming (in some
specimens) dark chestnut-brown below; underparts mixed grayish
white and blackish brown in varying proportions, faintly tinged with
pale cinnamon-buff.

Skull—Similar to that of orytona, but relatively broader across
zygomata and interorbital region; decidedly larger than that of its
nearest neighbor, okanagana, with broader postorbital constriction;
compared with nwaria it is shorter, with the zygomata more widely
expanded and the premaxillae narrower.

Measuremenis.—Adult male:1 Total length, 710-785 (average, —
749); tail vertebrae, 205-252 (232); hind foot, 98-112 (102). Adult
female: ? 680-765 (714); 195-247 (219); 94-107 (99). Skull: Adult
male: * Condylo-basal length, 106.2-107 (106.6); palatal length,
61.6-62.7 (62.2); postpalatal length, 39-40 (389.5); length of nasals,
42-44.5 (43.3); zygomatic breadth, 69.2-69.8 (69.5); breadth across
mastoids, 48.5—49.2 (48.9); least interorbital breadth, 27.1-29 (28);
breadth of rostrum, 24.5-24.7 (24.6); maxillary tooth row, 22.38-24.1
(23.2). Adult female:* Condylo-basal length, 95.4-102.5 (98.8);
palatal length, 56.3-59 (57.7); postpalatal length, 34.6-39 (37.7);
length of nasals, 37.8—44 (41.7); zygomatic breadth, 64.2-68.4 (65.8);
breadth across mastoids, 44.9-49.4 (46.7); least interorbital breadth,
22.5-24.6 (23.8); breadth of rostrum, 19.3-22.7 (21.2); maxillary
tooth row, 21-23.2 (22.1).

Remarks.—The Cascade hoary marmot differs from both okanagana
and oxytona in whiter and browner (less black) coloration above; from
nivaria in much more dusky colors, both above and below; and from
caligata in much larger size and darker coloration. It is widely dif-
ferent from olympus, both in color and cranial characters. It is appar-
ently isolated from all the other forms of the group, and no absolute
intergrades have been examined, but the characters separating it
from the forms of caligata are so slight that it seems best to.regard it
as a subspecies of the latter.

1 Four specimens from Mount Rainier and Cascade River, Wash.
4 Six specimens from same localities.

8 Two specimens from Cascade River, Wash., and Mount Baker Range, British Columbia.
4 Seven specimens from Cascade Range, Wash., and Mount Baker Range, British Columbia.

* 1915.) - MARMOTA CALIGATA GROUP. 69

Specimens examined.—Total number, 50, as follows:

British Columbia: Chilliwack (mountains near), 1;! Hope, 1;? Howe Sound
(near head), 3; Mount Baker Range (near United States boundary), 8;*
Skagit River (mountains near head), 6; Spences Bridge, 1;+ Tammi Hy
Mountain, 2.1

Washington: Camp Chiloweyuck, 2; Cascade River (near head), 8; Easton
(mountains near), 1; Mount Rainier, 7; ‘‘Northwest boundary survey”
(probably Mount Baker Range), 10.

MARMOTA OLYMPUS (Merriam).
Oxtympic Marmot.
CPln Tie hes 1. Pex fies 1) Pl. XV) fies 2b.)

Arctomys olympus Merriam, Proc. Acad. Nat. Sci., Philadelphia, 1898, p. 352.
[ Marmota] olympus Trouessart, Cat, Mamm., Suppl., 1904, p. 344.

Type locality—Head of Soleduck River, Olympic Mountains,
Wash. (at timberline).

Distribution—Upper slopes of the Olympic Mountains, Wash.,
above timberline (from about 4,000 feet altitude to near summits of
peaks).

Characters.—Size large (about equaling M. caligata cascadensis);
color in fresh pelage brownish drab mixed with white; feet brown;
skull with broad rostrum and interorbital region.

Color—Fresh pelage:* General tone brownish drab, more or less
mixed with white hairs; underfur between hair-brown and benzo
brown, becoming pale drab-gray at tips; long hairs glossy blackish
brown mixed with more or less pure white ones; top and sides of
head blackish brown, with a broad, white patch in front of eyes;
sides of nose, lips, and chin white; legs brownish drab shading
to blackish brown on the feet; tail clove brown, tipped with light
pinkish cinnamon; underparts brownish drab mixed with white (or
solid soiled whitish). Worn summer pelage: General tone of upper-
parts pinkish buff, varied with russet, the bases of hairs bister; feet
chestnut-brown; tail above, pinkish buff or cinnamon-buff, mixed with
snuff brown; beneath, snuff brown to chestnut-brown. Yowng (half-
erown August specimens): General tone of upperparts grayish
brown, becoming cinnamon on rump; underfur hair-brown at base,
tipped on fore back with white, and on hinder back and rump with
cinnamon or cinnamon-bufi; top and sides of head and fore legs bister;
feet dark clove brown; underparts light clove brown mixed with white
or pale buff.

1 Collection Victoria Mem. Mus.

2Collection Mus. Comp. Zool.

3 Including five in Mus. Comp. Zool.

4 Collection Amer. Mus. Nat. Hist.

5 No specimens in full winter pelage have been seen: description from August specimens just beginnings
to acquire the fall pelage.

70 NORTH AMERICAN FAUNA. _[No. 87. -

Skull—Similar to that of cascadensis but relatively narrower
across zygomata and broader between orbits and across rostrum;
postorbital constriction narrower; nasal branches of premaxillae
narrower than nasals at posterior end.

Measurements—Adult male:1 Total length, 720-750 (average,
740); tail vertebrae, 210-237 (219); hind foot, 100-110 (106). Adult
female:? 670-690 (680); 180-192 (186); 91-100 (95.5). Skull:
Adult male:* Condylo-basal length, 105-109.8 (106.1); palatal
length, 60-63 (61.1); postpalatal length, 39.6-42 (40.6); length of
nasals, 40.6-46 (43.7); zygomatic breadth, 64.8-67.1 (65.6); breadth
across mastoids, 46.2-47.3 (47); least interorbital breadth, 27.5-
31.2 (29.8); breadth of rostrum, 24.5-27.7 (25.8); maxillary tooth
row, 21.5-24.2 (23.1). Adult female:* Condylo-basal length, 99.5;
palatal length, 58; postpalatal length, 37.5; length of nasals, 41.5;
zygomatic breadth, 64.3; breadth across mastoids, 43.7; least inter-
orbital breadth, 26.5; breadth of rostrum, 24.6; maxillary tooth
row, 22.5.

Remarks—The Olympic marmot is one of the largest members of
the caligata group, about equaling in external measurements casca-
densis and oxytona. Its skull averages about the size of that of
cascadensis, but the largest male skull of olympus is longer than any
other American marmot skull examined. Im color the species is
decidedly browner than any of the forms of caligata, though not
nearly so brown as vancouverensis. The black colors of caligata have
almost entirely disappeared, but some of the white hairs remain.

This species is confined to the high mountains of the Olympic
Peninsula and is geographically isolated from its nearest relative,
cascadensis.

Specimens examined.—Total number, 17, as follows:

Washington: Happy Lake, Olympic Mountains, 7;° Mount Ellinor, 3; Mount
Steel, 4; Soleduck River (near head), Olympic Mountains, 3.

MARMOTA VANCOUVERENSIS Swarts.
VANCOUVER ISLAND MaARMotT.
(Pl. EX, fig.3: Pl. XV, fies Ze)

Marmota vancouverensis Swarth, Univ. of California Pub. Zool., VII, 1911, p. 201;
X, 1912, p. 89.

Type locality—Mount Douglas, Vancouver Island, British Colum-
bia (altitude, 4,200 feet).

1 Three specimens from Olympic Mountains, Wash.

2 Two specimens from same locality.

3 Five specimens from same locality.

4 One specimen from same locality. =
5 Collection Field Mus. Nat. Hist.

1915.] MARMOTA CALIGATA GROUP. al

Distribution.—V ancouver Island, British Columbia; apparently not
generally distributed, and known at present only from “the moun-
tains at the head of China Creek, some 20 miles south of Alberni, in
the Golden Eagle Basin, and King Solomon Basin, and on the sur-
rounding slopes and ridges.”’ }

Characters.—Size of M. caligata cascadensis; color uniformly dark
brown; skull relatively narrow with peculiarly shaped nasals.

Color.—Entire body, legs, and tail dark vandyke brown, , the
underfur being of the same color, the long hairs more blackish and
glossy; sides of nose and chin soiled whitish; underparts sometimes
irregularly blotched with white, and back rarely with a few scat-
tering white hairs; feet glossy blackish brown. In worn pelage the
upperparts and tail fade to sayal brown or clay color.

Skull.—Similar to that of cascadensis, but smaller and relatively
narrower; zygomata less widely expanded; premaxillae relatively
wider; nasals deeply emarginate posteriorly, the frontals projecting
forward and forming a V-shaped notch between nasals; incisors
white or yellowish white.

feasurements.—Adult male:? Total length, 660-710 (average,
684); tail vertebrae, 200-300 (222); hind foot, 90-102 (98.4). Adult
female: ? 680-720 (700); 210-240 (225); 100-105 (102.5). Skull:
Adult male: Condylo-basal length, 98-103.2 (99.9); palatal length,
7-59.5 (57.8); postpalatal length, 36.5-39.5 (387.8); length of nasals,
40-42.7 (41.4); zygomatic breadth, 64.2-65.5 (64.7); breadth across
mastoids, 45.7-47 (46.1); least interorbital breadth, 23.2-24.4 (24);
breadth of rostrum, 22-24 (22.9); maxillary tooth row, 23.6-24.7
(24.1). Adult female:* Condylo-basal length, 93.7-96.6 (95.2);
palatal length, 54—-54.3 (54.2); postpalatal length, 35.2-37.5 (36.3);
length of nasals, 39-39.6 (39.3); zygomatic breadth, 61.6-62.6 (62.1);
breadth across mastoids, 44; least interorbital breadth, 21.7-23
(22.3); breadth of rostrum, 21 521.6: maxillary tooth row, 23.1-23.7
(23.4).

Remarks.—This peculiar marmot, although clearly related to the
mainland species (caligata) has, through isolation, developed striking
characters, both external and cranial. The tendency of isolated
coastal forms in this group to become brown (shown in a lesser
degree by MM. calhigata vigilis and M. olympus) has reached the greatest
extreme in this species, the black colors of the mainland forms being
entirely lacking and the white reduced to scattering hairs.

1 Swarth, H.S. Loc. cit., X, 1912, p. 89.
2 Five specimens from vicinity of type locality.

3 Two specimens from vicinity of type locality.
¢ Four specimens from vicinity of type locality.

72 NORTH AMERICAN FAUNA. [No. 37.

After a season’s exploration of the southern part of Vancouver
Island, Swarth came to the conclusion that the species is probably
confined to a small area in the vicinity of Mount Douglas. He states:

We found them in the mountains at the head of China Creek, some 20 miles south
of Alberni, in the Golden Eagle Basin, and King Solomon Basin, and on the sur-
rounding slopes and ridges. They were most abundant on Mount Douglas, the peak
to the west of King Solomon’s Basin. Wherever the ground was bare of timber, or
but sparsely covered, as is the case over extensive areas at this point, the marmots
had established themselves, burrowing under the rocks, and apparently never wander-
ing very far from home. * * * ‘Their extreme wariness is correlated with con-
spicuousness, for the dark brown pelage shows in marked contrast against either gray
rocks or green grass. * * * They whistled but seldom, only one or two being
heard during the three weeks we spent in their territory.

None was found in apparently suitable situations on Mount Arrow-
smith, and certain timber cruisers who had explored the wilder parts
of the island stated that they had never seen a marmot.

Specimens examined.—Total number, 11,’ as follows:*

British Columbia: Golden Eagle Basin, 1; King Solomon Basin, 3; Mount
Douglas, 7.

Cranial Measurements of the Marmota caligata Group.

ie |

= tae ae age
~ = :
ef aE acca ee] ©
& Bo | | Sy Omen li ates
+ q fas} oD |B m +, »
So |S$(2/13)} 8 leuss1 aie
No. Species and locality. 8 we) | a a S3.8| "|S | Remarks.
Be |S) 2/3 | ss) joel Sips
A ue! is is wi oils 3 a |‘
K g = D =| &) | o o ro) tal
oO So S io) o mB |e u Ss
7) O JARIAIHIN IA WR |Aala
Marmota caligata caligata.
131440/ Becharof Lake, Alaska .......-.. [$2] | 99 |56.8)36. 8/38. 4/68 |46. 2/27.6)26.1)22 | Adult.
131441)..... dO....------ +22 - +222 eee ee eee [S?] | 96. 5/55. 7/36. 5/41. 4/64. 7/44 |24.3]24 22.3 Do.
191449 ee Os een Aveiara eaten Me po stece ears [of ?] | 100. 6/57. 4/38. 2/42. 3/67.8/44 |25.8)24. 1123.8 Do.
98154) White Pass, Alaska......-...-.- S| 100 [57.7/37.8 43 |65. 2/44. 2/23. 4/23. 6/23. 4 Do.
131444] Becharof Lake, Alaska.......... [22] | 92.8/53 /35.8/38 163. 4/43. 2/24. 3/22. 9121.9 Do.
128069}..... GO laszece cent eae =P ssc ale be. 96. 4/53.9/38 |39.1/64.1/43 23. 9/24. 5122.5 Do.
98153] White Pass, Alaska.......-...-- 2 88.9/51 |33.5)40.5/57.540 |22.1/20 |22 Do.
135161| Head of Coal Creek, Yukon...... 2 94 |53.9)86 (88 |62.3/48.3)24 122.1/21.7 Do.
Marmota caligata vigilis.
4418) Glacier Bay, Alaska...........-- $ | 97.2/55.8'36.738.9/62 |43.5)22. 6/21. 1/22.3 Do.
BADE eae Ons ee seaeee: Sere sisarsters Sissel 3 97. 3|54. 7/37. 8/41. 4]64. 8/43. 2/24, 2/22 {21.8 Do.
97952)... - - dO....--2- +--+ +--+ 222-2 eee g 98.4/55.5,38 |41. 5/67. 7/44.6)26 |23. 5122. 5 Do.
97953)... GO. ooo 2 ee owe wie wel os 2 93. 7/51. 6:37. 7/39. 6161. 8/42. 3/24. 5/21. 3/22 Do.
Marmota caligata sheldoni, |
137319} Montague Island, Alaska........ S | 89,5/51.2'34.3/35 61. 5/41, 3/22. 5/20. 5]22. 5 Do.
4962)... -- OM eee rie eetiein cette ee 3 96.6/55.4/36 |39 |63 |42.3/23.5/21.6/23.1 Do.
4961|..... GO eee case emastie mane [S?]| 96 [54.5137 |38. 8/63. 4/42. 4/23. 8121. 8/22. 4 Do.
4969|..... GOs acest cd civ oui ee et eaen [S?] | 92.3|52. 2/34. 8/36. 4/61. 5/41. 5/23. 2121 122.2 Do.
4960I..... GN ee ree eects Q 88. 7/50.2134 [37 159, 4139. 7/22. 2120, 5121.8 Do.

1Swarth, H,S. Univ. of California Publ. Zool., X, 1912, pp. 89-90.

2 Allin collection Mus. Vert. Zool., Univ. of California,

3 All about 20 miles south of Alberni, British Columbia, within a radius of 10 miles.
4 Collection Mus. Vert. Zool., Univ. of California.

1915.] MARMOTA CALIGATA GROUP. 13

Cranial measurements of the Marmota caligata group—Continued.

i: ie ls le
a : s ah e |
eile ele @ \e|2
© wmi4/el2 io | 8 | 2
= . oP s > |S o rl ~~ wri
| Gio = ee BS
Ss ee) a B22 =P S 5 .
No. Species and locality. 3 BO Merl le Ie Be Helse, 3 += | Remarks.
¢{/2/5|/°} 8 ia" ae a|&
We Ss) gle> lan |S
Calebais Sp Oe is | Scie
ra 0) Oils
ee eae | Sauls a a ea i
Marmota caligata oxyiona. |
175565| Moose Forks, British Columbia...) ¢ 101 157 |40 |41.5/62.8/44.2/25 23 |22 | Adult.
202789] Sustut Mountains, Brit. Col.....; ¢ 102 |58. 6139. 5/42. 7/66. 8/46. 5/25. 8/22. 2/23 Do.
202791; Lake Thudade, Brit. Co!.......- é 107. 4/62. 5|40. 2145. 3167. 4/48. 3/25. 7/23. 6/23. 3 Do
1 20766] Level Mountain, Brit. Col....... 3 105 159 {41 |42.3/66.5)47 125. 7|24. 8/22 Do
174503) Head of Smoky River, Alberta... 101. 8/56. 5}41. 1143, 6/65. 3)45. 9125. 4123 |22.7 Do
174502|..... LO a oe SOS ciate | 104. 6/61. 2/36. 4/41 (66. 2/47. 3126. 7/23. 3/22. 7 Do

67073| Glacier, British Columbia......- Peete 95 |53.4/37 (37 |66.8/45.5/24.9/23 (22.7 Do.
G07G\222 22 GUO eae iu rol 100 (5%. 4)37.3/40.5/67 |46 |27 125.2)22.5 Do
66695| Nelson, British Columbia......- rol 98. 1/55. 7/38. 1/39. 3/64. 8/45. 2/25. 3/24. 4/21.9 Do.
67072| Glacier, British Columbia....... 96 154.4/87 140.3164 |43.7/23.5/21 |22.1 Do.
C1074 eee (OU) a Si Ie 97. 6|55. 3/38. 7/38. 6/66. 3/45. 5/21. 9/21. 9 21. 4 Do.
67075|..... LOS ene ence patac sie Sane « | 94. 4/53.3)36 [38 (65 |42.9/24,6/23.3,22.3 Do
Marmota caligata nivaria. |
72222| St. Marys Lake, Mont........-... | 106. 5/61. 4/39. 6/41. 2.66. 6/45. 7/27.2'25 |22.8 Do
12223 eee (SD) SEES Ae Neer ee 101 [58 |37. 8/43. 9:66. 3/45. 6/26. 1/23. 4 22.2 Do
(2225\eeece COREE ei ryt terate Vale claiels eave ote LOL (59 (37.8/43 |65.8/44.8/25.4.21.624.4 Do
(2B esos MORSE rae Na see ek. 99.5)59 137. 5/40.6.64 |45.7/23.3)22 (22.3 Do
Marmota caligata cascadensis. |
6871) Washington - British Columbia é 107 |62.7/40 |44.5 69, 2/48. 5|27, 1124. 7/24. 1 De.
boundary.
42793) Head of Cascade River, Wash...) g | 106.2/61.6)39 |42 |69.8/49.2/29 (24. sloo, 3 Do.
2 6840] Mount Baker Range, Brit. Col...| @ | 101 [59 (37. 7/43. 2 65. 6/47. 6/23. 3 21.8 21 Do.
42792} Head of Cascade River, Wash... 102.5)58.7,.39 44 /68. 4/49. 4/23. 1 22.7:23.2 Do,
90133} Mount Rainier, Wash.......-.... 98.5|56.3 38 41.666. 2/46. 7/24. 3 20. 7/22.1 Do.
90134|..... CORE aha Satintis eulsiste dis hte Q 95. 4/56. 3,38. 4/43. 9,66. 7/47. 8/22.5.21 |22.3 Do.
Marmota olympus. | |
67611} Olympic Mountains, Wash...... ref 105 60. 2/40.2/44 |64.8)47. 3/27. 5 24. 6)22.9 Do.
PAGS pte SAO se eases he oe 2 seins ge ete = S | 105.3/61.8/39.6)43 |65. 1/47. 3/31. 2 26. 3/21. 5 Do.
2'6230|--.- CGO) sees lel ea SEES ee Nota 3 109.8/63 |42 (46 |67.1/47.3/31.1 27. 7/23.7 Do.
67612)..... CO CS Ute ea BN nsec ea 2 | 99.5)58 137. 5)41. 5/64. 3/43. 7/26, 5 24. 6122.5 Do
Marmota vancouverensis.
4 12099] Vancouver Island, Brit. Col..... 3 98 (57 137 (405/255-1447) 4o32. 124 123.7 Do.
£12092)..... CORRE aes Cesena lear é 98. 2157. 2/36. 5/40 164. 2/45. 7128.2.22 (24.3 Do.
#412090)... .- GO een tase eae Sa ee rol 100. 4)57. 7/38. 2/42. 4/64. 5/46 124.4 22. 6/23.6 Do.
412091)..... MOWER se este layne aitle ace cof 103. 2)59. 5/39. 5)42. 7/65. 5/45. 8/24. 3 23, 2/24. 7 Do.
ROA RCL Olas nae tioeiee Sas cya eee So Su areas g 93.7)54 |85.2/39 |61.6)44 (21. 7/21. 5/23.7 Do.
#12100)..... (DS UC Ale NL aS A 96. 6|54. 3:37. 5]39. 6/62. 6/44 ie re pay. U Do.

1 Collection Amer. Mus. Nat. Hist.

2 Collection Mus. Comp. Zool.

3 Collection Field Mus. Nat. Hist.

4Collection Mus. Vert. Zooi., Univ. of California.

Fie. 1.

Bre.

EXPLANATION OF PLATES.
PLATE II.
{Much reduced.]

Marmota olympus, & adult, Olympic Mountains, Wash., (altitude 5,000 feet),
Aug. 28, 1897. (No. 92768, U. S. Nat. Mus., Biological Survey collection.)

. Marmota flaviventris obscura, § adult, Wheeler Peak, N. Mex. (altitude 12,400

feet), July 27, 1904. (No. 133506, U. S. Nat. Mus., Biological Survey
collection.)

PLATE III.

[Natural size.]

Marmota caligata oxytona. Klappan Mountain, British Columbia. (No. 170-
683, U. S. Nat. Mus., Biological Survey collection.)

Marmota flaviventris flaviventris. Summer Lake, Oreg. (No. 89311, U. S.
Nat. Mus., Biological Survey collection.)

Marmota monax preblorum. Woburn, Mass. (No. 78358, U. S. Nat. Mus.,
Biological Survey collection.)

PLATE IV.

[Three-fourths natural size.]

. Marmota flaviventris dacota, ¢ adult, Bridger Pass, Wyo. (No. 25529, U. 8.

Nat. Mus., Biological Survey collection.)

. Marmota monax monaz, S adult, Peaks of Otter, Va. (No. 143962, U. 8.

Nat. Mus.)

. Marmota caligata oxytona, 3 adult, Lake Thudade, British Columbia. (No.

202791, U. 8. Nat. Mus., Biological Survey collection.)

. Marmota flaviventris sierrae, 2 adult, head of San Joaquin River, Cal. (No.

41950, U. S. Nat. Mus., Biological Survey collection.)

Marmota monax rufescens, 3 adult, Lake George, N. Y. (No. 67695, U. 8.
Nat. Mus., Biological Survey collection.)

Marmoia caligata caligata, [% ?] adult, Becharof Lake, Alaska. (No. 131437
U.S. Nat. Mus., Biological Survey collection.)

b)

PLaTe V.

[Three-fourths natural size.)

. Marmota monax monax, 3g adult, Peaks of Otter, Va. (No. 143962, U. S.

Nat. Mus.)

. Marmota monax rufescens, 3 adult (type), Elk River, Minn. (No. 186521,

U.S. Nat Mus., Merriam collection.)

. Marmota monaz preblorum, 3 adult, Wilmington, Mass. (No. 78355, U. S.

Nat. Mus., Biological Survey collection.)

. Marmota monaz ignava, $ adult, L’ Anse au Loup, Labrador. (No. 8871, Mus.
Comp. Zool.)
4

1915.]

Fic. 1.

Fig. 1.

Fig. 1.

~~

or

EXPLANATION OF PLATES.

PLATE VI.

[Three-fourths natural size.]

. Marmota monax canadensis, 3 adult, Murray Bay, Quebec. (No. 7603, Field

Mus. Nat. Hist.)

Marmota monaz petrensis, $ adult (type), Revelstoke, British Columbia.
(No. 203532, U. 8. Nat. Mus., Biological Survey collection.)

Marmota monaz ochracea, [3 ?]subadult, Babine Mountains, British Columbiz.
(No. 202785, U.S. Nat. Mus., Biological Survey collection.)

Marmota flaviventris warreni, 2 adult (type), Crested Butte, Colo. (No. 202987,
U.S. Nat. Mus., Biological Survey collection.)

Pruate VII.
(Three-fourths natural size.]

Marmota flaviventris flaviventris, § adult, Donner, Cal. (No. 100532, U. &.
Nat. Mus., Biological Survey collection.)

Marmota flaviventris parvula, 2 adult (type), Jefferson, Nev. (No. 93690, U. &
Nat. Mus., Biological Survey collection.)

Marmota flaviventris avara, 3 adult, Okanogan, British Columbia, (No. 9975,
U.S. Nat. Mus., Biological Survey collection.)

. Marmota flaviventris engelhardti, Q adult, Beaver Mountains, Utah. (No.

158978, U. 8. Nat. Mus., Biological Survey collection).

Prate VIII.

[Three-fourths natura! size.]

Marmota flaviveniris nosophora, & adult, Sawtooth National Forest, Idaha
(No, 156924, U. 8. Nat. Mus., Biological Survey collection.)

- Marmota flaviventris luteola, [gf ?] adult, Mount Lincoln, Colo. (No. 175, Mus.

Comp. Zool.)

. Marmota flaviventris dacota, ¢ adult, Bear Lodge Mountains, Wyo. (Ne.

65920, U. S. Nat. Mus., Biological Survey collection.)

. Marmota flaviventris obscura, f adult, Wheeler Peak, N. Mex. (No. 135504,

U.S. Nat. Mus., Biological Survey collection.)

PLATE IX.
[Three-fourths natural size.]

Marmota olympus, 3 adult, Olympic Mountains, Wash. (No. 6325, Fieid
Mus. Nat. Hist.)

Marmota caligata oxytona, 3 adult, Lake Thudade, British Columbia. (Ne
202791, U.S. Nat. Mus., Biological Survey collection.)

. Marmota vancouverensis, § adult, Vancouver Island, British Columbia. (Ne.

12091, Mus. Vert. Zool., Univ. of California.)

. Marmota caligata caligata, [3 ?] adult, near Portage Bay, Alaska. (Ne.

131440, U.S. Nat. Mus., Biological Survey collection.)

76

Fia.

Fie.

Oo

pS

NORTH AMERICAN FAUNA, [No. 37.
PLATE XxX

(Three-fourths natural size.)

Marmota caligata cascadensis, § adult, head of Cascade River, Wash. (No.
42793, U.S. Nat. Mus., Biological Survey collection.)

. Marmota caligata nivaria, 2 adult, St. Marys Lake, Mont. (No. 72222,

U.S. Nat. Mus., Biological Survey collection.)

. Marmota caligata okanagana, 3 aduit, Glacier, British Columbia. (No. 67073,

U.S. Nat. Mus., Biological Survey collection.)

. Marmota caligata oxyiona, 2 adult (type), Head of Smoky River, Alberta.

No. 174503,U. S. Nat. Mus.)

PLATE: 26h
[fhree-fourths natural size.]

Marmota monax monaz, $ adult, Peaks of Otter, Va. (No. 143962, U. S
Nat. Mus.)

Marmota monax rufescens, ¢ adult (type), Elk River, Minn. (No. 186521,
U.S. Nat. Mus., Merriam collection.)

. Marmota monax preblorum, 3 adult, Wilmington, Mass. (No. 78355, U.S.

Nat. Mus., Biological Survey collection.)

. Marmota monaz canadensis, § adult, Murray Bay, Quebec. (No. 7603, Field

Mus. Nat. Hist.)

PLATE XII.
{Three-fourths natural size.}

Marmota monaz ignava, adult, L’ Anse au Loup, Labrador. (No. 8871, Mus.
Comp. Zool.)

Marmota flaviventris warrent, 2 adult (type), Crested Butte, Colo. (No. 202937,
U.S. Nat. Mus., Biological Survey collection.)

Marmota monaz ochracea, [3 ?|subadult, Babine Mountains, British Columbia.
(No. 202785, U.S. Nat. Mus., Biological Survey collection.)

. Marmota caligata nivaria, Q adult, St. Marys Lake, Mont. (No. 72222,

U.S. Nat. Mus., Biological Survey collection.)

PLATE XIII.
(Three-fourths natural size.)

Marmota flaviventris flaviventris, § adult, Donner, Cal. (No. 100532, U.S.
Nat. Mus., Biological Survey collection.)

Marmota flaviventris parvula, 2 adult (type), Jefferson, Nev. (No. 93690,
U.S. Nat. Mus., Biological Survey collection.)

. Marmota Maen avara, adult, Okanogan, British Columbia. (No. 99759,

U.S. Nat. Mus. Cpoigncal Survey colleetion.)

. Marmota Heer ee engelhardii, Q adult, Beaver Mountains, Utah. (No.

158978, U. 8S. Nat. Mus., Biological Survey collection.)

1915.]

| 2 Cereal I

EXPLANATION OF PLATES.
PLATE XIV.
[Three-fourths natural size.}
(No. 156924, U. S. Nat. Mus., Biological Survey collection.)

65920, U.S. Nat. Mus., Biological Survey collection.)

. Marmeta flaviventris luteola, [f ?] adult, Mount Lincoln, Colo.

Mus. Comp. Zool.)

U.S. Nat. Mus., Biological Survey collection.)

PLATE XV.

[Three-fourths natura! size.]}

. Marmota flaviventris dacota, § adult, Bear Lodge Mountains, Wyo.

77

. Marmota flaviveniris nosophora, ¢ adult, Sawtooth National Forest, idaho.

(No.

(No. 175,

. Marnota flaviventris obscura, 3 adult, Wheeler Peak, N.Mex. (No. 135504,

Marmoia olympus, $ adult, Olympic Mountains, Wash. (No. 6325, Fieid

Mus. Nat. Hist.)

Marmota vancouverensis, § adult, Vancouver Island, British Columbia.

(No. 12091, Mus. Vert. Zool., Univ. of California.)
Marmota caligata caligaia, | 3 ?| adult, near Portage Bay, Alaska.
U.S. Nat. Mus., Biological Survey collection.)

. Marmota caligata cascadensis, § adult, head of Cascade River,

42793, U.S. Nat. Mus., Biological Survey collection.)

(No. 131440;

Jash.

(No.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE II.

SKINS OF (1) MARMOTA OLYMPUS AND (2) M. FLAVIVENTRIS OBSCURA, SHOWING MOLT.

PEATE III

| Survey

iologica

B

Dept. Agr.

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37,

ican Fauna No

North Amer

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PLATE IV.

| Survey,

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Biolo

North American Fauna No. 37, U. S. Dept. Agr.

BIBAT[BI VIBSI[BO ‘IA (9) ‘OVIIOIS SLUIJUOATABE “YL 'F “XBUOUL XBUOUL “J °G

SUdDSOJLI XBUOUT "J “CG ‘BUOJAXO VIBSI[BVO "PL “G “VIOOVP SITPUDATARY "JL “LE

VLOWHVI JO SAISIGNVI GNV VINVYO

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE V.

SKULLS OF MARMOTA.

1. M. monax monax, 3, M. monax preblorum.,
2. M. monax rufescens. 4, M. monax ignava.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE VI;

SKULLS OF MARMOTA.

1. M. monax canadensis. 3. M. monax ochracea.
2. M. monax petrensis. 4, M. flavyiventris warreni.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE VII

SKULLS OF MARMOTA.

1. M. flaviventris flaviventris. 3. M. flaviventris avara.
2. M. flaviventris parvula. 4, M. flaviventris engelhardati.

North American Fauna No. 37, U. S. Dept. Agr. Biological Survey.

PLATE VIII.

-
~3—
Or bso

‘
4
:
‘

esau

SKULLS OF MARMOTA.

. faviventris nosophora.

1. M 3. M. flaviventris dacota.
2. M. flaviventris luteola.

4, M. flaviventris obscura.

North American Fauna No. 37, U. S. Dept. Agr. Biological Survey. PEATE De

SKULLS OF MARMOTA.

1. M. olympus. 3. M. yancouverensis.
2. M. caligata oxytona. 4. M. caligata caligata.

North American Fauna No. 37, U. S. Dept. Agr. Biological Survey. PLATE X.

SKULLS OF MARMOTA.

1. M. ealigata cascadensis. 3. M. caligata okanagana.
2. M. caligata nivaria. 4, M. caligata oxytona.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. LATE XI.

SKULLS OF MARMOTA.

1. M. monax monax. 3. M. monax preblorum.
2. M. monaxrufescens. 4. M. monax canadensis.

h American Fauna No. 37, U. S. Dept. Agr. Biological Survey. PLATE XIl.

SKULLS OF MARMOTA.

3. M. monax ochracea.
4, M. caligata nivaria.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE XIII.

SKULLS OF MARMOTA.

1. M. flaviventris flaviventris. 3. M. flaviventris avara.
2. M. flaviventris parvula. 4, M. flaviventris engelhardti.

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE XIV.

SKULLS OF MARMOTA.

M. flaviventris nosophora. 38. M. flaviventris luteola.
M. flaviventris dacota. 4. M. flaviventris obscura.

por

North American Fauna No. 37, U.S. Dept. Agr. Biological Survey. PLATE XV.

SKULLS OF MARMOTA.

1. M. olympus. 3. M. caligata caligata.
2. M. yancouveren 4. M. caligata cascadensis.

INDEX.

(New names in bold=face type; synonyms in italics.)

A.
alpina, Marmota, 18.
Arctomys, genus, 18,
avarus, 41.
brachyurus, 40.
caligata, 59.
caligatus, 59.
canadensis, 30.
dacota, 49.
empetra, 30.
flaviventer, 39, 49, 50.
ignavus, 29.
melanopus, 30.
mona, 22,
okanaganii, 64.
okanaganus, 64.
olympus, 69.
pruinosa, 59.
pruinosus, 59, 64.
sibila, 30.
avara, Marmota flaviventris, 41-42.
avarus, Arctomys flaviventer , 41.

B.
brachyurus, Arctomys, 40.

Cc.

caligata, Arctomys, 59.
Marmota, 57.
Marmota caligata, 59-61.
Marmotia, 59.
caligatus, Arctomys, 59.
canadensis, Arctomys, 30.
Glis, 30.
Marmota monax, 30-32.
cascadensis, Marmota caligata, 67-69.

D.

dacota, Arctomys, 49.
Marmota flaviventris, 49-50.

Distribution maps, caligata group, 68.

fiaviventris group, 38.
monax group, 23.

E.

engelhardti, Marmota, 50.
Marmota flaviventris, 45-46.
empetra, Arctomys, 30.
Mus, 30.

¥F.

flaviventer, Arctomys, 39, 49, 50.
Marmota, 39, 50.

flaviventris, Marmota, 36.
Marmota flaviventris, 39-41.

G.
Glis, genus, 18.

I.

ignava, Marmota monax, 29-30.
ignavus, Arctomys, 29.
Was

luteola, Marmota flaviventris, 50-52.

M.

Marmot, Black Hills, 49-50.
Cascade hoary, 67-69.
dusky, 53-55.
Engelhardt, 45-46.
glacier, 61-62.
golden-mantled, 46-49.
Montague Island, 62-63.
Montana hoary, 66-67.
Nevada, 44.
northern hoary, 59-61.
Okanagan hoary, 64-66.
Olympic, 69-70.
pallid yellow-bellied, 41-42.
park, 50-52.

Robson hoary, 63-64.
southern Sierra, 43-44.
Vancouver Island, 70-72.
Warren’s, 52-53.
yellow-bellied, 39-41,

Marmota, genus, 18.
alpina, 18.
avara, 41-42.
caligata, 57, 59-61.
canadensis, 30-32,
cascadensis, 67-69.
dacota, 49-50.
engelhardti, 45-46,
engelhardti, 50.
flaviventer, 39, 50.
fiaviventris, 36, 39-41.
ignava, 29-30.
luteola, 50-52.
marmota, 19.
monax, 21, 22-25.
nivaria, 66-67.
nosophora, 46-49.
obscura, 53-55.
ochracea, 34-35.
okanagana, 64-66,
olympus, 69-70.
oxytona, 63-64.
parvula, 4445.
petrensis, 33.
preblorum, 27-29.
rufescens, 25-27.

79

80 NORTH AMERICAN FAUNA.

Marmota sheldoni, 62-63. P,
sibila, 63.
sierrae, 43-44.
vancouverensis, 70-72.
vigilis, 61-62.
watreni, 52-53.
Marmotta caligata, 59.
Measurements, cranial, explanation of, 18. R.
caligata group, 72.
flaviventris group, 55.

parvula, Marmota flaviventris, 44-45,
petrensis, Marmota monax, 33.
preblorum, Marmota monax, 27-29,
pruinosa, Arctomys, 59.

pruinosus, Arctomys, 59, 64.

rufescens, Marmota monax, 25-27.

monax group, 35. s
melanopus, Arctomys, 30. :
monarx, Arctomys, 22. sheldoni, Marmota caligata, 62-63.
Glis, 22. sibila, Arctomys, 30.
Marmota, 21. Marmota, 63.
Marmota monax, 22-25. sierrae, Marmota flaviventris, 43-44,
Mus, 18, 22.
N. Vv.

vancouverensis, Marmota, 70-72.

nivaria, Marmoia caligata, 66-67. icilis. M ta caligata. 61-62
viguis, armota Ca. 1iga ay, Use

nosophora, Marmota. flaviventris, 46-49.

QO. Ww.

obscura, Marmota fiaviventris, 53-55. warreni, Marmota flaviventris, 52-53.
ochracea, Marmota monax, 34-35. Woodchuck, British Columbia, 33.
okanagana, Marmota caligata, 64-66. Canada, 30-32.
okanaganii, Arctomys, 64. Labrador, 29-30.
okanaganus, Arctomys, 64. New England, 27-29.
olympus, Arctomys, 69. ochraceous, 34-35.

Marmota, 69-70. rufescent, 25-27.
oxytona, Marmota caligata, 63-64. southern, 22-25.

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At
se WOVEIO ET
OA.

is

eee
ges

__'U.S. DEPARTMENT OF AGRICULTURE
a - BUREAU OF BIOLOGICAL SURVEY
; +9 a HENRY W. HENSHAW, Chief

| NORTH AMERICAN FAUNA —
| / No: 88

[Actual date of publication, September 30, 1915]

it ge
ae
| ig

=

A REVIEW OF THE AMERICAN MOLES

BY

HARTLEY H. T. JACKSON

x _ ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY

= -

be : : Qn
ins

\
WASHINGTON
GOVERNMENT PRINTING OFFIOE
ae 1915 -

U.S. DEPARTMENT OF AGRICULTURE

BUREAU OF BIOLOGICAL SURVEY
HENRY W. HENSHAW, Chief

NORTH AMERICAN FAUNA

No. 38

[Actual date of publication, September 30, 1915]

A REVIEW OF THE AMERICAN MOLES

BY

HARTLEY H. T. JACKSON

ASSISTANT BIOLOGIST, BIOLOGICAL SURVEY

aa eau
he
coo

WASHINGTON
GOVERNMENT PRINTING OFFICE
1915

LETTER <OF TRANSMIT

Unirep States DEPARTMENT OF AGRICULTURE,
Bureau oF BroLocicaL SURVEY,
Washington, D. C., March 26, 1918.

Sm: I have the honor to transmit herewith, for publication as
North American Fauna No. 38, a review of the American moles
by Hartley H. T. Jackson, assistant biologist, Biological Survey.
Widely distributed throughout a large part of North America and
very numerous in many places, moles have in some localities proved
"injurious to agriculture, though they are generally beneficial through
their destruction of insects, which form much of their food. Their
economic status has not yet been fully determined but is now being
carefully investigated by this bureau, and for this work the present
paper will serve as a basis.

Respectfully,
Ernry W. HEnsHaw,
Chief, Brological Survey.

Hon. Davi F. Hovston, ;

Secretary of Agriculture.
2

ADDITIONAL COPIES

OF THIS PUBLICATION MAY BE PROCURED FROM
THE SUPERINTENDENT OF DOCUMENTS
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AT

20 CENTS PER COPY
Vv

: CONTENTS.

Characteristics and development of the young........----.-.---.--.---.--
Petron ara MOLES Ee cs eis aso es Se. lawns oe
eerie VATIALION. 2.3 tas Sa eee eee rk. ae eked
Peng HOM tRVATIAbON. Seo ite ee er oe Sa eek oes eee
Semualvariation.. 0.2.2... ee Ae Ee os Ae en ens
SPM OM als ios eR ee een ne ol dele eke Sees Been.
eae Mm ION cia) Satara Eugen ne ink ee hk ea ge 03 ieee ad
beapiaeaenollet seria s 5! 1s Se SLE Jal SO kk SUN Re
MicmorranecuG ackuowledoments: 22 0 uid eee eS a

Demarini ce ce tie ee Se aie ie See oe

SULLS TBST CES eee ae nr a A es ese Sees ee ire 2 Re
Rare ee CNS See SG Sake ced hOlkig wee hd
List of generic names that have been used for American moles...........
Sele a MOnpAe ENCTA‘Ol MOLES. ois occ ese ns Voc oe oh eS ek ee eet
List of American genera, species, and subspecies, with type localities ....

Bee BD ete 5 Sai oe Seesctiais a. Ree aes = kum dint 2 ok bee oe

iMevs ro spectes and subspeciese-s .< 122. esse Sed sete kee ee oss 3
Peserpiions ol species and subspecies 2.2.02 jist ss. 522s se ee eee ee

ee REE IS ee aegis) i else e eM. AIG ote Sees

Key to species and subspecies. ...--- GP pees. Wy. Nei carole a8
Wexzeraptious OL species and subspecies... .2. iste. 1-52 seks osl dee eke t

Dee RE PELOSI ODS ens Gola, sarees ici atte bin ss Se ee ate eee tL ae kee ets
eet UNE sm Mane Nee ote eS Sse One ma te cele ae sacle oS
aS CH OLICH Sea ae aoe le oe oe see hageie SEIS Se ok Haren io Geico

HaeN LOIS SPCCICS ee ceo. We cs oe ok eatin Measles or He see ie ee eeaew
Deseriptions of species'and subspecies... ..... 4.2.2 220422... 22-22 ee:

[LLUSTRATIONS:

PLATES.

[Following page 98.]

PiatE I. Molting process of Scalopus aquaticus aquaticus; young of S. a. howelit.

II. Skulls (dorsal view) of Scalopus aquaticus aquaticus, S. a. howelli, S. a.
australis, S. a. anastase, S. a. parvus, S. a. machrinus, S. a. machrinoides,

S. a. pulcher, S. a. caryt, S. a. texanus, S. inflatus, and S. xreus

TIT. Skulls (ventral and lateral views) of Scalopus aquaticus aquaticus, S. a.
howelli, S. a. australis, S.a. parvus, 8. a. texanus, S. inflatus, and S. xreus.

3

f

ILLUSTRATIONS.

Puate IV. Skulls (dorsal view) of Scapanus townsendit, S. orarius orarius, S.o0. schefferi,
S. latimanus latimanus, 8. 1. occultus, S. 1. grinnelli, 8.1. sericatus, S. 1.
minusculus, S. 1. dilatus, S. anthonyi, Parascalops breweri, and Condylura

Fia.

cristata.

V. Skulls (ventral and lateral views) of Scapanus townsendii, S. orarius orarius,
S. latimanus latimanus, S. anthonyi, Parascalops breweri, and Condylura

cristata.

VI. Skulls (dorsal, ventral, and lateral views) of Netirotrichus gibbsii gibbsti
and NV. g. hyacinthinus. Mandibles (lateral view) of Scalopus aquaticus

“IO OT BR GC bd Fe

co oO

. Skull of young of Scalopus aquaticus machrinoides
. Geographic range of species and subspecies of the genus Scalopus.. .

. Tail of Scalopus aquaticus aquaticus...._.2 32550 2
. Snout of Scalopus aquatieus aquaticus...-2.55..-5 2 ee
. Fore foot of Scalopus aquaticus aquaticus..-... 0... ee
. Hind foot of Scalopus aquaticus aquaticus_..22.22.2...
. Geographic range of Scapanus townsendir, S. latimanus and subspecies,

aquaticus, S.a. howelli, S. a. australis, S. a. machrinus, S. a. texanus, S.
inflatus, S. xreus, Scapanus townsendu, S. orarius orarius, S. 0. schefferi,

S. latimanus latimanus, S. 1. occultus, S.1. grinnelli, 8.1. sericatus, S. 1.
minusculus, S. 1. dilatus, S. anthonyi, Parascalops breweri, Condylura

ertstata, Netirotrichus gibbsi gibbsu, and N. g. hyacinthinus.

TEXT FIGURES.

and 8. ANthONY toc. 55 sajeiste ore et ieee

. Tail-of Scapanus latimanus latimanus. 2222-22 22 en
- Snout of Scapanus latumanus laiimanus... 5.2 ee
10.
11.
Ae
13.
14.
15.
16.
Ly
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.

Fore foot of Scapanus latimanus latimanus?: 2.22.22 2 ee
Hind foot of Scapanus latemanus latimanus.. icc eee eee
Geographic range of Scapanus orarius and mubepeces Ee yas, oS tS
Geographic range of Parascalops brewert../--- 2-2 = = =) ee
Tail of Parascalops brewert.....22055022 2522) eee
Snout of Parascalops brewert.......----- Bt yee
Fore foot of Parascalops brewert..23 222552242422. 2 ee
Hind foot of Parascalops brewert....22-+:-92.222.- 2
Geographic range of Condylura cristata:..252*_-.- = 2a
Tail of Condylura cristata....0.055252 02-00)
Snout of Condyluracristata...2-220 9-2) 0 2k
Fore foot of Condylura, cristata.....:.12222 5212-2. eee
Hind foot.of Condylura cristata’ =2 222 22 = eee eat ae
Geographic range of Netirotrichus gibbsit and subspecies.....-.--.-----
Tail of Netirotrichus.gibbsit hyacnthinus.. 2. 5-4-5. --- eee eee
Snout of Neurotrichus gubbsi gibbsis. <2... 2-22. - eee
Fore foot of Netirotrichus gibbsit hyacintlinus. .._-22222- 25 ssn eee
Hind foot of Netirotrichus gibbsit hyacinthinus.......-----.-.---------

No, 88, NORTH AMERICAN FAUNA. Sept., 1915,

A REVIEW OF THE AMERICAN MOLES.

By Hartiey H. T. JACKSON.

INTRODUCTION.

On account of their subterranean and secretive habits, American
moles apparently were not observed by the earlier American ex-
plorers. As the settlement of the United States progressed, however,
these animals became well known, and in many cases proved injurious
to agriculture. Moles occur rather generally in eastern North
America along the Atlantic and Gulf coasts from Labrador to Florida
and in northeastern Tamaulipas, Mexico, and range westward to
Manitoba and northeastern Colorado. Within this area are found
three genera, Scalopus, Parascalops, and Condylura. All three genera
occur in some localities, but no one covers the entire region occupied
by the group. West of the area mentioned no moles are found until
the Pacific coast region is reached. There, two other genera, Scapa-
nus and Neiirotrichus, occur, their ranges being confined mainly to the
humid and semihumid region west of the Cascade Range and the
Sierra Nevada, from southern British Columbia to northern Lower
California.

In general, the several genera have the following distribution:

Scalopus, comprising the common naked-tailed moles of eastern
United States, is the most widely distributed and best-known genus
of the family. It ranges from southern Massachusetts, southern
Ontario, central Minnesota, and northeastern Colorado south to the
Gulf States and northeastern Tamaulipas, Mexico. It is confined
almost entirely to the Upper and Lower Austral Zones.

Parascalops, comprising only a single species—Brewer’s, or the
hairy-tailed mole—is much more restricted in range than Scalopus,
occurring from southern New Brunswick to northeastern Ohio,
southern Pennsylvania, and south along the Appalachian Moun-
tains to North Carolina. Although living in the midst of the range
of Scalopus, in reality the two occur together at comparatively few

5

6 NORTH AMERICAN FAUNA. [No. 38

places, since Parascalops is confined exclusively to the Transition and
Canadian Zones, while Scalopus inhabits chiefly the Austral Zones.

Condylura, also comprising only a single species—the star-nosed
mole—is widely distributed, but apparently not very abundant. It
ranges from southern Labrador and southeastern Manitoba south to
northern Illinois, along the Appalachian Mountains to western North
Carolina, and on the Atlantic coast to Georgia. Apparently not lim-
ited in its distribution by zones, it occurs from the Boreal to the Lower
Austral.

Scapanus fills the place on the Pacific coast occupied by Scalopus
on the Atlantic slope, being abundant and widely distributed from
British Columbia south to northern -Lower California. It ranges
through all the zones from Boreal to Lower Sonoran.

Neiirotrichus has a limited distribution on the Pacific coast west
of the mountains from southwestern British Columbia south to
Monterey County, California, occurring in the Boreal and Transition
Zones.

HABITS AND ECONOMIC STATUS OF MOLES.

While all American moles have certain general habits in common,
as, for example, spending most of their lives underground and feeding
largely upon insects, their habits vary in details. The genera Sca-
lopus, Scapanus, and Parascalops usually inhabit drier soils, burrow
deeper, and confine themselves more to subterranean runways than
do Condylura and Neiirotrichus. All moles prefer loamy or sandy
soil where burrowing is easier, and consequently they are scarce or
absent in heavy clay, stony, or gravelly soils. Insufficient food is
often the cause of their scarcity in excessively sandy soils.

The common mole (Scalopus) is found in almost any area where
soil.and food are suitable; it is most plentiful in meadows, gardens,
and similar habitats, but is by no means confined to them, and fre-
quently is found in open woodland, along the banks of streams, and
in other environments. It dwells in a series of subterranean tunnels
10 to 18 inches beneath the surface, and from these it forces.to the
outside small piles of earth, scarcely large enough to be worthy the
name ‘‘molehills.’”’ A second series of tunnels is made just beneath
the surface of the soil and appears as a series of small ridges, usually
more or less branching and at times ramifying in all directions. This
second series seems to be made chiefly during the animal’s hunt for
food and may be occupied. but once; generally, however, the main
surface tunnels are used for a considerable time. During dry weather
the mole works deeper and practically deserts the surface ridges. Its
change of habit is due in part to the increased hardness of the surface
soil, but undoubtedly is more the result of its pursuing worms and

1915.) INTRODUCTION. 7

insects into moister regions. Essentially the same condition is pro-
duced during winter, when the surface soil is frozen. The common
mole seldom leaves its tunnels. Its nest is about 5 or 6 inches in
diameter and usually 12 to 18 inches beneath the surface; most fre-
quently it is placed under roots of shrubs or pasture grass and is
made of grass and rootlets, but occasionally partly of leaves. In the
northern half of its range the young are born during March or April;
‘in the southern part they appear earlier in spring. The number of
young in a litter varies from two to five, the usual number being four;
and there is probably only one litter produced each year.

The habits of Scapanus are much like those of Scalopus. The
ridges formed by Scapanus upon the surface are usually more con-
spicuous than those of Scalopus, and its ‘‘molehills,” thrown up at
frequent intervals, much larger. The hills made by Scapanus often
contain a half bushel or more of dirt and resemble those made by the
pocket gopher. The mounds made by Scapanus, however, show no
trace of an opening, while those of the pocket gopher do. The
mounds of Scapanus in most cases are more nearly circular than those
of the pocket gopher. So far as known, the breeding habits of sub-
species of Scapanus latimanus do not differ essentially from those
of Scalopus; the time of breeding and the number of young in a litter
are about thesame. With Scapanus townsendu, however, the breed-
ing time is later, the young usually being born during May or June.
The number in a litter is less than with S. latimanus, there being usually
only two or three, seldom four, and not infrequently only one.

The habits of Parascalops are not well known, but in general they
appear to be much the same as those of Scalopus.

Star-nosed moles, genus Condylura, prefer to make their homes in
wet meadows or marshes, though occasionally they: may choose the
same habitat as Scalopus, or even occupy the same tunnels with
them. The surface ridges made by Condylura are more irregular
and broken than those of Scalopus and usually smaller and more
crooked; the burrows seem to be deep for a short distance, then
appear as surface ridges, shortly to disappear again. Besides the
subterranean tunnels, the star-nosed mole uses surface runways
under and through the grass in marshes and meadows. Unlike
other moles, Condylura frequently leaves its tunnels in winter and
burrows in the snow, or even runs on top of it. Little is known con-
cerning the nesting and breeding habits of this genus. A family of
five young about one-third grown, collected May 22, 1888, by Morris
M. Green, and now in the Biological Survey collection, was found in
a nest under a log on the flats of the Potomac River a short distance
north of Georgetown, D.C. Two of the young from this family are
slightly more developed than the others, but it is not known whether

8 NORTH AMERICAN FAUNA. [No. 88.

this is due to difference in time of birth or to subsequent conditions.
Another nest, containing young, is described by Bishop as follows:

On May 22, 1890, while having some apple trees planted, I had the good luck to
find a nest containing four young. :

The locality where the nest was found was two miles south of Kentville in Kings
County, Nova Scotia. The land had been cleared of small forest trees several years
before and had grown up with grass and was mowed every year.

The particular spot where the nest was found was a little hillock or cradlehill which.
had been formed apparently by a tree having been blown down. When the roots
had rotted away, a small dry mound of soft black sedimentary earth was formed,
and in this the nest was built. This mound was high enough to be out of reach of
storm water during wet weather.

The elevation containing the nest was ten inches below the surface, and was made
in circular form, seven inches in diameter. The nest was built of old dry grass, and
was very compact and neatly made. Although the mound contained a complete
network of roadways, no earth was thrown to the surface within ten feet of the nest.!

The little. Netirotrichus prefers a damp habitat and is seldom
found far from swamps, marshes, or streams. In the extreme
southern part of its range it is most frequently found in swampy
places overgrown with sedges or shrubs. Farther north its habitat
is less confined, and it is found along streams or even in moist dense
woods. Its tunnels are more like those of Condylura than other
moles, and it seems to spend no small part of its time in surface
runways or under logs. In fact, the tunnels are often open above
for some distance, and in this respect resemble the tunnels of the
eastern pine mouse (Pitymys pinetorum). The nesting and breeding
habits of Netirotrichus are unknown.

The economic status of American moles has been the subject of
much dispute, authorities differmg as to whether the animals are
beneficial or harmful. The genera Parascalops, Condylura, and
Netirotrichus are local in distribution, seldom abundant in any
locality, and most frequently inhabit waste or uncultivated lands;
they can not, therefore, have extensive economic importance. Such
is not the case, however, with Scalopus and Scapanus, which are
more generally distributed throughout their ranges, and usually
abundant in lawns, fields, and gardens. The food of moles consists
in large measure of insects, and in this they are beneficial. However,
moles destroy large numbers of earthworms, make unsightly ridges
in lawns, and dig tunnels which permit incursions of rodents injurious
to roots, tubers, and planted seeds, and which frequently are directly
responsible for the weathering away of humus deposits and super-
soils on hillsides; in these activities moles are harmful. The economic
status of the mole is being extensively investigated by Theodore H.
Scheffer, under the direction of the Biological Survey; the results of
his studies will be published later.

1 Bishop, W. L., Trans. Nova Scotia Inst. Sci., vol. 10. pp. 348-349, October 1, 1902.

1915.] INTRODUCTION. g
CHARACTERISTICS AND DEVELOPMENT OF THE YOUNG.

The young of Scalopus (Pl. I, fig. 3) are born hairless. Vibrissz
very soon appear on the lips, but hair does not show until the animal
is at least a week or ten days old. The fresh, first pelage remains
short and grows little until the animal is nearly one-third grown;
it is exceedingly fine and silky, and lies close to the body, giving
the animal a smooth, sleek appearance. Two young, probably
about a week old, from Jackson, N.C., have the general proportions
of the body much as in adults. The feet, both fore and hind, have
much the same shape as in adults, and are relatively about the
same size, though, on account of the bones of the arm thickening
rather than lengthening during growth, the A
fore feet of the young, being relatively more
projected from the body, appear at first glance to
be relatively very large. The hind feet are rela-
tively alittle wider than those of adults. The claws
of the fore feet are soft and weak, though relatively
thick and broad; those of the hind feet are very
soft and only slightly developed. The external ear
appears as a thickening of the dermis into a flat
papilla 1.5 mm. in diameter. The center of this
is penetrated by a minute auditory opening that i
seems to be closed by the contact of its sides; as aes
an auditory organ its function is probably exceed- ie. 1.—skutl of young of
ingly limited. The rudimentary eye appears as a = S*#/opus aquaticus mach-

; Fi rinoides (X 3). [Recon-
small pigmented spot covered by dermis; aminute, structed and drawn by
imperfect opening passes through the dermis to the author from speci-
the eye proper, and may be sufficiently penetrable Haak ike Nae
for the animal to perceive light from darkness; it Tom Greenway, Ark.]
seems improbable, however, that the eye is sufficiently developed for
form perception, and it is probable that with advanced age the sense
of light perception becomes less acute.

Skulls of young (fig. 1) a week or ten days old distinctly show the
sutures between the principal cranial bones; the sutures between
the frontals and parietals, however, are now nearly closed. In
comparison with skulls of adults, the mastoid region is high and
swollen and the zygomata are heavy. ‘The auditory meatus is
distinct and more pronounced than in adults, the bulle small, but
relatively high and rounded and not so much flattened. The nasals
are short and broad, becoming wider posteriorly; the premaxille are
triangular, adjoining the anterior two-fifths to one-half of the nasals.
During the development of the skull the premaxille push forward

1 Alcoholic, No. 7250, U. S. Nat. Mus.

10 NORTH AMERICAN FAUNA. [No. 38.

and gradually enclose the nasals anteriorly. Most of the cranial
bones are anastomosed and the sutures closed before permanent
dentition is acquired. The suture betwe@én the interparietal and
the parietals is the last to close and is the only one which may remain
open and distinct in extreme old age. The sutures between the
premaxille and the maxille and those between the nasals and the
maxille usually remain indistinctly visible until permanent dentition
is acquired.

The writer has examined no very young moles of genera other
than Scalopus, but it would seem that the general relationships of
characteristics of young to those of adults would be essentially the
same in the different genera. Bishop, writing about Condylura, states:
‘‘The young were probably ten days old, the fur just beginning to start,
which gave the skin a dark brown colour.” The most remarkable
difference between young and adults of Condylura is in the snout.
The character of that organ in the young has been well described
by Ayres, as follows:

At birth the star-nosed mole is nearly destitute of visible hair and the tactile
bristles of the facial region have not made their appearance at the surface. The
snout of the young Condylura lacks all the distinctive characteristics of the adult,
and the entire body resembles that of Talpa much more than it does its parent. On
a close examination of the distal end of the snout of such a new-born animal one can
distinguish a tract of skin which covers four-fifths of the circumference of the organ
(the part not specially marked off is the median ventral fifth). This dermal tract
extends for three millimeters toward the base of the snout and is marked off from the
remaining surface by a series of furrows running parallel to the long axis of the body.
A series of parallel ridges is thus formed, each ridge being bounded on either side by a
furrow. At their anterior and posterior ends these ridges pass gradually into the
neighboring smooth surface. By a gradual ingrowth of the bottoms of the furrows
each groove is deepened and each ridge suffers a correspondingly increased definition
of form, while at the same time the posterior end of each groove grows toward its
neighbor on either side. When the grooves have all united there is formed by their
union a common groove which nearly encircles the snout and separates the tactile.
from the remaining surface of that organ.

Commencing at the posterior margin of the tactile surface and advancing toward
the tip of the snout, the grooves deepen and grow toward each other in their bottom
portions until they finally coalesce underneath the ridges. The result of this process
is the production of free, finger-shaped processes composed exclusively of ectoderm,
attached to the anterior end of the snout in the manner already described for the
adult.?

The pelage of individuals of Scapanus, Parascalops, and Condylura,
one-third grown, has much the same compact, sleek appearance as
that of Scalopus, indicating a slow and probably entirely postnatal
srowth.

1 Bishop, W. L., Trans. Nova Scotia Inst. Sci., vol. 10, p. 349, Oct. 1, 1902.
2 Ayres, H., Biol. Centralb., band 4, pp. 358-359, 1885.

1915.] INTRODUCTION. Ua

PELAGES AND MOLTS.

The hair of all American moles is fine and silky, producing a soft
and velvetlike pelage. In Scalopus, Scapanus, and Parascalops the
hairs are nearly equal in length and there is no distinct underfur.
In Condylura some of the hairs are distinctly longer and coarser than
the major portion, the latter forming an underfur, and the whole
producing a pelage much less velvetlike than that of any other
genus. In Netirotrichus the condition of the pelage is somewhat as
in Condylura; the fur is shorter, however, and the underfur difficult
to detect.

The basal pelage reveals a series of transverse vermiculations, most
pronounced in Scalopus and Scapanus, least in Netirotrichus; in all
genera these markings are more noticeable in the fur on the back, less
on the ventral parts. Microscopic examination shows that these
vermiculations are due to structural as well as chromatic differences.
Each hair consists of normally pigmented, gray, cylindrical sections,
1 to 2 mm. long, alternated with finer flat sections, 0.2 to 0.5 mm.
long and unpigmented, or with the pigment reduced to a small amount
of yellow. Each one of these fine, flat sections acts as a hinge upon
which the hair bends; this in part produces the velvetlike texture of
the pelage and permits the hair to be rubbed either forward or back-
ward with little friction—a distinct advantage to a subterranean
mammal. The vermiculations usually show more clearly in worn
pelage than in fresh. The young of Scalopus, Scapanus, and Para-
scalops, in their first winter pelage, are more grayish than adults; of
Condylura, paler and more brownish. Winter pelage of adults of all
moles is usually darker than that of summer; worn pelage is faded
and frequently more brownish than the fresh. The color of speci-
mens retained in a cabinet or storage case for a few years usually
fades, becoming more brownish than that of recently killed animals.

TIME OF MOLTING.

There are two molts annually in Scalopus, one in spring and the
other in fall. Throughout most of the range of the genus the
spring molt is usually completed by the last of May or the first of
June; the fall molt, in the northern half of the range, by the first
week in October; in the southern part of the range the fall molt
naturally occurs later in the year, and the spring molt earlier than
farther north. The winter pelage of breeding females becomes worn
early in the season, but, as has been suggested by True,! the actual
molting in such individuals may frequently be delayed. Examina-
tion of a large series of specimens from Washington, D. C., shows that

1 True, F. W., Proc. U. 8. Nat. Mus., vol. 19, p. 37, 1896.

12 NORTH AMERICAN FAUNA. [No. 38.

the spring molt in that region occurs usually during the months of
April and May, though most of the animals are probably in full sum-
mer pelage by the middle of May. A large series of specimens from
the same locality, taken during October and the first part of Novem-
ber, are all in full winter pelage; the fall molt seems to take place
there during the latter part of August and the first half of September.
Specimens of Scalopus aquaticus machrinus from northern Illinois
show molting during the first half of May; one taken June 15, 1907,
at Joliet, Hl., is in fresh summer pelage, except for a small place on
the face; in a topotype of S. a. machrinus collected September 5, 1898,
the ventral parts are in short, fresh, winter pelage, the dorsal parts
in old, summer pelage. Specimens of S.a. pulcher, taken at Delight,
Ark., the middle of September, are in nearly complete winter fur; an
adult male taken April 30, 1910, at Lake City, Ark., is in fresh sum-
mer pelage except on the nose; specimens taken at Sour Lake, Tex.,
the latter part of March and first of April, 1905, show the beginning
of molt on the underparts. The spring molt of S. a. australis seems
to occur from about the middle of March to the middle of April. A
topotype of S. a. texanus, collected February 9, 1893, has fresh
summer pelage on the abdomen.

The time of molting seems to be more irregular in Scapanus than
in Scalopus, and the period of seasonal molt during which numbers
of individuals may be found molting is more prolonged than with
Scalopus; it is possible that this difference may be correlated with
climatic differences. The spring molt in Scapanus latimanus lati-
manus May commence any time from the middle of January to the
last of April; specimens in which the spring molt has just begun were
collected at Santa Cruz, Cal., January 14 and February 28, and at
Red Bluff, Cal., April 24; one collected at Lower Lake, Cal., April 12,
still retains the worn and faded winter pelage; a specimen collected
March 18, at Santa Cruz, is in fresh summer pelage, and the majority
of specimens show this condition by the middle of April. The
autumnal molt of S. 1. lattimanus may begin from the middle of August
to the last of November; in a male collected August 14, at Santa
Cruz, the molt is well begun; a specimen taken November 30 at
Fort Bragg, Cal., has fresh pelage on the head only, and another
taken the same day at King City, Cal., has obtained about one-third
of its new pelage; in most cases, however, the winter pelage is com-
plete by the last of November. The autumnal molt in 8S. J. dilatus is
usually completed a month earlier than in S. 1. latvmanus, though one
specimen collected at Chico, Cal., December 20, has just started to
molt. A female collected August 4, at Round Mountain, Cal., is
about two-thirds covered with fresh pelage, and is probably in a
delayed spring molt. The specimens of 8S. 7. occultus examined indi-
cate that the spring molt in that form may begin as early as the first

1915.] INTRODUCTION. 13

of January or as late as the latter half of June; a specimen in which
the molt is just begun was collected January 2 at Somerset, Cal.,
and a male in similar condition of molt was taken at Alhambra, near
by, on June 21; spring molt in S. l. occultus is usually completed by
the first of April and the autumnal molt about the first of December.
The material available does not show many specimens of S. townsendit
in process of spring molting; two collected April 7, 1914, at Puyallup,
Wash., are in nearly complete summer pelage, while in three others
the molt is fully complete. The majority of specimens show full
summer pelage the latter part of April. The fall molt of S. townsendit
occurs most frequently during October; specimens occasionally show
molting later in the year, as, for example, one collected December 20,
1912, at Ferndale, Cal., which shows the last remnants of summer
fur; on the other hand, the fall molt may begin as early as the latter
part of August; several females collected during July and the first
half of August, near Portland, Oreg., are in various stages of molting
from the beginning to half complete, which may be a delayed spring
molt. A specimen of S. orarius orarius collected March 11, 1913, at
Ferndale, Cal., has begun to molt; one of S. o. schefferi collected at
Lester, Wash., May 14, 1914, shows only a trace of old pelage on the
back and underparts; others secured the middle of May are in full
summer pelage. The change to winter pelage in S. o. orarius seems
to occur mostly during October, though a male taken at Eureka,
Cal., August 17, 1910, shows the beginning of the autumnal molt.

Material examined is insufficient for a satisfactory determination
of the time of molting of Parascalops; in a male from Magnetic City,
N. C., collected March 26, 1894, the ventral parts and the rump
are in fresh summer pelage, the rest of the fur being worn winter
pelage; other specimens in various stages of molt from the same
locality were taken between April 24 and July 5. Many specimens
collected at various localities durmg August and September show
a much-worn pelage; it seems probable that winter fur appears
during October and November; in fact a specimen collected October
12, 1910, in Wetzel County, W. Va., has traces of, a new pelage
under the old on the breast. An adult male from Magnetic City,
N. C., collected August 21, 1893, shows molting and has the entire
ventral parts in fresh pelage; it is impossible to determine whether
this is an early fall molt or a delayed spring molt.

The spring molt in moles of the genus Condylura occurs late, the
molting process being at its height durmg June and the first half
of July, though it is usually completed by the middle of the latter
month; very rarely molting may begin during the last few days of
May, and, equally as rarely, traces of winter pelage may remain
well into August. The autumnal molt of Condylura takes place

te NORTH AMERICAN FAUNA. [No. 88.

during October, though occasionally it begins the last of September;
the full winter pelage is usually obtained before the last of October.

In a large series of Netirotrichus from Sumas, British Columbia,
the spring molt appears to be at its height during the latter half of
May and the first week of June, though one or two specimens indicate
a beginning of the molt as early as the latter part of April; most
individuals are in full summer pelage by the first of July; none of
these specimens show autumnal molt. Farther south, in Washington,
Oregon, and northern California, spring molt is earlier, and specimens
taken there during the first part and middle of June are in summer
pelage. The available material has been insufficient to determine
the time of the fall molt, though apparently it occurs during October.
A specimen from Goldbeach, Oreg., collected September 21, 1901,
has a trace of new pelage under the old on the breast; two from
Steilacoom, Wash., October 9 and 12, 1891, and one from Palo
Alto, Cal., October 17, 1897, have the molt well advanced; a male
taken October 18, 1891, at Tenino, Wash., is in nearly complete
winter pelage.
MANNER OF MOLTING.

Molting in Scalopus occurs more or less regularly in definite
sequence on the different parts of the body, and the same order is
followed in both the vernal and autumnal molts. The fresh pelage
first appears on the breast and abdomen (PI. IJ, fig. 1) and gradually
replaces the old until the entire underparts, except the chin and
throat, have molted; at this stage there is a sharp lateral line of
demarcation between the new and the old fur (PI. I, fig. 2); the fresh
pelage gradually extends up over the back, generally encroaching
upon the posterior part first and working forward toward the nose.
The chin and throat in most individuals retain the old pelage for
several days after all the rest of the molt is complete. There are,
of course, exceptions to this general order of molting but most of
these occur in animals which are molting either earlier or later
than normally, and the writer is inclined to believe that these varia-
tions are either due to retarded or stimulated physiological processes,
or else result from injuries to the animal. Two specimens show
distinctly that the molting process has been retarded on account
of injuries; one? is in full winter pelage except a very small place
on the throat and a small patch directly posterior to a flesh wound
on the left side of the posterior part of the back; the other? is in
complete winter pelage except a circular patch, about 20 mm. in
diameter on the occiput, which is also mostly posterior to a flesh

1No. 180769, U. S. Nat. Mus., Biological Survey collection; ¢ adult, collected at Washington, D.C.,

October 11, 1912.
2 No. 6190, Acad. Nat. Sci. Philadelphia; 9 adult, collected at Audubon, N. J., October 28, 1908.

1915.] INTRODUCTION. 15

wound and partly encloses it; winter pelage surrounding the patches
is in both specimens long and well ‘developed, in marked contrast
with the short fur of delayed growth. A male’ from Connecticut, in
somewhat premature molt, has the entire dorsal parts in fresh
pelage except a patch about 30 mm. in diameter on the right shoul-
der; the anterior three-fifths of the ventral parts are in old pelage,
and this extends well up the sides. A postbreeding female, ? the
molting of which has already been mentioned by True, ? is in retarded,
much-worn winter pelage, except the posterior half of the back,
anterior to the rump, and a very small portion of the abdomen,
which are in rather incomplete spring pelage.

The sequence of molting in Scapanus is less definite than in
Scalopus. ‘The differences of color, texture, and length of hair be-
tween the old and new pelages of Scapanus are usually slight; often
the line of demarcation separating the two pelages is scarcely dis-
tinguishable, and seldom sharp as in Scalopus. The sequence of
molting on the various parts of the body appears in a few cases to be
not unlike that of Scalopus, the underparts molting first, followed
consecutively by the sides and back. More frequently, however, the
new pelage appears first on the head and throat, then works down
over the nape and back, encroaching last upon the abdomen; or, as
is shown most beautifully in a specimen‘ of Scapanus latimanus
latimanus from Petrolia, Cal., the new pelage may appear simul-
taneously in separate patches upon head, back, and rump.

The material has been inadequate to show anything definite re-

garding the method and sequence of molting in Parascalops, but the
specimens examined seem to indicate that these processes are not
unlike corresponding processes in Scalopus.
. The new fur of Condylura generally appears first on the posterior
part of the flanks, but the body sequence is inconstant;,the molt on
the flanks usually spreads forward and ventrally, while at the same
time on the back fresh pelage replaces the old, which sloughs off in
irregular blotches. Probably in most cases the ventral parts are in
fresh pelage before the major portion of the back has molted; a small
posterior rump patch is almost invariably the last to molt. The
contrast between new and old pelages during the spring molt is
marked; the autumnal molt, however, is often difficult to detect.

In the genus Netirotrichus, new pelage ordinarily first replaces the
old on top of the head; this is soon followed by the molting of the
posterior part of the back almost simultaneously with the beginning

1 No. 4276, Mus. Comp. Zool., Bangs collection; ¢ adult, collected Februrary, 21, 1896, at Liberty Hill,
Conn.
2 No. 22858, U. 8. Nat. Mus., Biological Survey collection; collected July 1, 1888, at Washington, D. C.
3 True, F. W., Proc. U. S. Nat. Mus., vol. 19, p. 37, 1896.
* No. 140706, U. 8. Nat. Mus., Biological Survey collection; f adult, collected Nov. 6, 1905.

v

16 NORTH AMERICAN FAUNA. [No. 38.

of the molting of the ventral parts on the throat and breast; the
molting areas on the back increase in size and finally enclose each
other; the area on the breast works posteriorly, then dorsally; the
flanks are the last to molt. The molting process in Neiirotrichus,
once well begun, seems to be very rapid, and this may account for
the sparsity of material of this genus showing molt.

GEOGRAPHIC VARIATION.

The maximum of geographic variation occurs in Scalopus and Scapa-
nus. In Scalopus it consists in general of a decrease in size toward
the south and an increase in pallor toward the west. The maximum
size of individuals is found in the north-central part of the range of
the genus; the size decreases gradually both to the east and west,
and rather abruptly toward the south. The rostrum tends to shorten
in the western and southwestern portions of the range. The palest
members of the genus are found in the region of western Nebraska;
the color darkens toward the east and south, becoming somewhat
ochraceous in the southwestern part of the range.

The size of Scapanus (except in S. townsend and S. orarws orarius),
like that of Scalopus, decreases toward the south. In the coast
region the pallor increases toward the south, and reaches its maxi-
mum in specimens of Scapanus latimanus occultus from San Diego
County, Cal.; farther south, however, in the San Pedro Martir moun-
tain region of Lower California, the color of Scapanus seems to be-
come darker again. In the interior, more mountainous regions, the
color of Scapanus becomes darker toward the south, the darkest
specimens being from Yosemite Valley, Cal., while the palest are
from Crater Lake, Oreg., and the Mount Shasta region, Cal.

Geographic variation in Netirotrichus manifests itself in a slight in-
crease in size and a very slight darkening of color in the southern
part of the range. A tendency appears also for the more pronounced
development of an anterior cusplike process on the cingulum of the
second upper premolar in specimens from the southern half of the
range of the genus.

Geographic variation in Parascalops and Condylura is negligible,
though a slightly decreasing size southward is noted in Condylura.

INDIVIDUAL VARIATION.

The general size, shape, and proportions of skulls of moles, con-
specific and of corresponding maturity, from any given locality are
usually constant, variation seldom exceeding more than 4 per cent.
On the other hand, the shape of individual cranial bones is noticeably
variable, especially that of the interparietal. In a series of specimens
of Scalopus from a given locality the posterior border of the palate

1915.] INTRODUCTION. heey , LY

may be simple, spined, or notched; in Scapanus, simple, but usually
notched; in Condylura usually simple, but sometimes spined; and in
Parascalops and Neiirotrichus, almost invariably simple. There is a
marked tendency in Scalopus and Scapanus toward abnormal denti-
tion; this appears not only in the production of accessory cusplike
processes on the cingula of the premolars, but also in the increase or
reduction of the number of premolars ordinarily present. The
tendency in Scalopus is toward extra premolars; in Scapanus:it is
toward the suppression of premolars, though in each of at least seven
skulls examined there is a supernumerary premolar.

The general color of specimens from a given locality is constant,
except for seasonal variations. Individual abnormalities, however,
occur frequently in Scalopus and Scapanus. The common type of
chromatic variation in Scalopus is the occurrence of white, cream,
orange, or ochraceous spots or blotches, appearing usually either on
the face or on the ventral parts. Scheffer ' has remarked upon the
prevalence of this form of variation in Scalopus at Manhattan, Kans.,
where he finds that certain regions of the body of a large majority
of the moles taken are washed with a tinge of orange, or that on the
head or abdomen are distinct patches of this color. Variations simi-
lar to these, however, crop out in many localities. They are not
confined to any particular species or subspecies, but appear most
frequently in S.a.machrinus and S.a.machrinoides. Inmanyspecimens
this form of variation is, apparently, partial albinism, the patches
being nearly white and the slight creamy or brown tinge probably
being due to glandular or other stains. Blotches in other specimens
range from cream color to shades of buff, orange, and brown, and show
clearly that they are due to mutations of color. A unique variation
occurs in a specimen from Warsaw, Ill. (No. 5429, American Museum
of Natural History). The entire underparts of this animal, except
the vent and right hind foot, are a beautiful capucine buff, as are
also the nose and right side of the face; a narrow band of this color
extends for a short distance up the posterior part of each flank, and
anteriorly another band passes over the shoulders and across the nape,
forming a complete collar; the remaining dorsal parts are much as in
normal specimens, but perhaps slightly more vinaceous. Another
peculiar variation occurs in a specimen from Madison, Ind. (No.
112008, United States National Museum). ‘The entire dorsal parts
of this individual are normally colored; the ventral parts are pale
ochraceous-salmon to ochraceous-buff, irregularly spotted and
blotched especially laterally with grayish fuscous, the whole effect
reminding one in certain respects of the ventral parts of some of the
spotted tiger-cats or ocelots.

1 Scheffer, T. H., Kansas State Agr. College Exp. Sta. Bul. 168, p. 4, 1910.
90549°—15——2

18 ; ; NORTH AMERICAN FAUNA. [No. 88.

That these chromatic mutations are inherited in Mendelian ratio
among the offspring is strongly suggested in the specimens examined
from Point Pelee, Ontario. From the small and somewhat isolated
colony of Scalopus inhabiting this locality 25 specimens collected
between October 18, 1908, and June 2, 1913, have been examined.
Of these specimens 18 have ochraceous patches on the face, while
the remaining 7 are normally colored. On purely unsubstantiated
evidence, it would appear that ‘‘lack of gray-producing pigment”’ is
here a dominant character, and that the hereditary tendency is
probably toward pattern development rather than toward the primi-
tive mammalian monochromatic grayish coloration.

Mutations of color seem to occur less frequently in Scapanus than
in Scalopus; they are most marked in specimens of Scapanus orarius
scheffer. from Walla Walla and Wenatchee, Wash. Abnormalities
in, color in the genera Parascalops, Condylura, and Netirotrichus, are
confined to albinistic spots which occur only rarely and usually
appear to have been caused by injuries.

The writer has seen no melanistic specimens of American moles.

SEXUAL VARIATION.

The only marked sexual variation in American Talpide is in size of
individuals. This is very noticeable in Scalopus and Scapanus, in
which genera males are considerably larger than females. The same
condition probably holds in other genera; in the specimens exam-
ined of Parascalops, Condylura, and Neiirotrichus there is no apparent
sexual variation, but this may be due to incorrect determinations
of sex.

AGE VARIATION.

The most characteristic change in American Talpide occurring
with increasing maturity is a flattening of the skull and a broad-
ening of the base of the rostrum. These tendencies prevail in all
genera. Broadening of the base of the rostrum is due to lateral
thickening of the maxille. In immature specimens the external
roots of the molars are not infrequently exposed in places through
the maxille. In all genera except Condylura the upper premolars
tend to become less cuspidate with increasing age, due not alone to
wear of the teeth, but apparently to absorption and physiological
processes. :

The genus Condylura furnishes an anomalous and unique age vari-
ation; old adults possess a distinct median longitudinal crest on the
upper posterior half of the rostrum. Another peculiar variation
accompanying age occurs in Parascalops in which the hair on the
nose and tail frequently turns white in old adults.

1915.] INTRODUCTION. 19

SEASONAL VARIATION.

Winter pelage of most moles is slightly darker than that of summer.
No other seasonal variations occur except in Condylura: The tail of
Condylura during the summer months is narrow and elongate; during
the winter months it becomes much thickened, abruptly constricted
at the base, and tapered apically. Both males and females have
_ been found with tails in both conditions. The variation probably is
due to assimilation of fat. Enlargement of the tail may begin as
early as August, but usually it does not begin until September, or
possibly later; the maximum size is reached in most cases by the
latter part of November and is retained until March or April, when the
tail gradually diminishes to its minimum summer size by the last of
May or the middle of June.

EXPLANATIONS.
EXTERNAL MEASUREMENTS.

External measurements of moles are in millimeters and, unless
otherwise stated, are those made by the collector from the animal
in the flesh. The following have been used:

Total length.—Tip of nose to end of terminal tail vertebra.

Tail vertebre.—Base of tail at upper surface to end of terminal tail
vertebra.

Hind foot.—Heel to end of longest claw.

CRANIAL MEASUREMENTS.

Cranial measurements were made by the author with a vernier
caliper. The following have been employed:

Greatest length—Antero-posterior diameter of skull from anterior
median point between bases of first upper incisors to most posterior
point of supraoccipital in median line.

Palatilar length.—Antero-posterior diameter of palate from poste-
rior median point between bases of first incisors to posterior median
border of palate.

Mastoidal breadth.—Lateral diameter of skull measured through
mastoids. Greatest lateral diameter of skull.

Interorbital breadth.—Lateral diameter of cranium measured at
coronal suture.

Maxillary tooth row.—Antero-posterior diameter of upper molar-
premolar row measured at alveolar border. ;

Mandibular molar-premolar row.—Antero-posterior diameter of
lower molar-premolar row measured at alveolar border.

The bone immediately anterior to the lambdoidal crest and just
posterior to the parietals is here called the ‘‘interparietal.” This
bone in moles has been considered by some writers part of the supra-
occipital.

20 NORTH AMERICAN FAUNA. [No. 38,
COLORS.

The names of colors used throughout the text are those of Ridg-
way.’ On account of iridescence and reflection of light from the fur
the color of any single animal may appear to vary considerably when
viewed from different angles. For this reason the following method
of making comparisons and observations of colors was used consist-
ently: Diffused daylight from a window was allowed to strike the
animal at an angle of 30° to 45° posterior to a plane perpendicular to
the median longitudinal line of the animal. The mole was then
viewed from its anterior end at an angle of about 60° from the light
rays and in the same plane with them. In spite of this care colors
have sometimes been exceedingly difficult to determine and describe.

MATERIAL AND ACKNOWLEDGMENTS.

The present revision recognizes 28 forms of 10 species of Amer-
ican moles and is based upon a study of 2100 specimens, mostly skins
accompanied by skulls. Of this number, Scalopus comprised 945,
Scapanus 604, Parascalops 129, Condylura 218, and Neiirotrichus 204.
While type specimens or essential topotypes of all described forms
have been examined, the material has been inadequate for an
entirely satisfactory understanding of the group, and this is partic-
ularly true of the genus Scapanus, and of Scalopus from some of the
southern and western parts of its range. Unreliable determinations
of sex also have been a serious handicap.

Although the study has been based primarily upon specimens in
the Biological Survey and other collections in the United States
National Museum, including the Merriam collection, it became evi-
dent early in the investigation that this material would be insufficient
for a proper elucidation of the group. Accordingly specimens were
borrowed from various museums and private collections until nearly
all that were available in the United States and Canada were exam-
ined. For the loan of specimens and for various other courtesies I
wish to express my appreciation to the following: Dr. J. A. Allen, of
the American Museum of Natural History; Mr. Samuel Henshaw
and Mr. Outram Bangs, of the Museum of Comparative Zoology,
Harvard College; Dr. Joseph Grinnell, of the Museum of Vertebrate
Zoology of the University of California; Mr. Charles B. Cory and
Mr. W. H. Osgood, of the Field Museum of Natural History; Dr. Wit-
mer Stone, of the Academy of Natural Sciences of Philadelphia;
Dr. W. J. Holland and Mr. W. E. Clyde Todd, of the Carnegie
Museum; Mr. H. L. Ward, of the Public Museum of the City of Mil-
waukee; Dr. A. G. Ruthven, of the University of Michigan Museum

1 Ridgway, R., Color Standard. »3 d Color Nomenclature, 1912.

1915.] ‘THE FAMILY TALPIDA. 91

of Zoology; Mr. P. A. Taverner, of the Victoria Memorial Museum,
Ottawa, Ontario; Mr. Stanley G. Jewett, of the Oregon State Game
Commission; Dr. Charles H. Gilbert, of Stanford University;
Mr. Myron H. Swenk, of the University of Nebraska; Mr. W. H. Over,
of the University of South Dakota; Mr. H. E. Anthony, Mr. D. E.
Kent, Mr. G. L. Kirk, Mr. W. E. Saunders, and Mr. Thaddeus Surber.
The writer is also indebted to Mr. Gerritt S. Miller, jr., and Mr. N.
Hollister, of the United States National Museum, and Dr. M. W.
Lyon, of the George Washington University, for many courtesies and
suggestions. The text drawings, with the exception of figure 1, are
the work of Miss Ruth Gibson Collette.

THE FAMILY TALPIDZ.

The family Talpide, exclusive of fossil forms, is composed of
thirteen genera peculiar to the temperate regions of the Northern
Hemisphere.t Five of these (Scalopus, Scapanus, Parascalops,
Oondylura, and Neiirotrichus) are indigenous to North America.
Scapanus and Neiirotrichus are found only on the Pacific slope;
Scalopus, Parascalops, and Condylura, only in the Atlantic drainage.
The family is a rather heterogenous group which, in certain genera,
has characters in common with other families of Insectivora. The
characters taken in combination, however, are diagnostic. The
nearest relationships of the family are with the Soricidz, which are
approached most closely in the Thibetan genus Uropsilus. The
American Talpide, however, are distinct from any of the American
Soricidz and may be distinguished by the following characters:

AMERICAN TALPIDA (MOLES). AMERICAN SORICIDH (SHREWS).
Ear-conch absent. Ear-conch present (small and incon-
spicuous in Blarina and Cryptotis).
Clavicle short and broad. Clavicle long and slender.
Humerus short and broad (length less Humerus relatively long and slender
than twice the width). (length more than twice the width).
Pelvis relatively narrow (length more Pelvis relatively broad (length less
than thrice the width). than thrice the width).
Os falciforme present on the fore foot No os falciforme on the fore foot.
(rudimentary and indistinct in Neitiro-
trichus).
Terminal phalanges of fore feet bifur- Terminal phalanges of fore feet simple,
cate. not bifurcate.
Zygomata present. Zygomata absent.

Audital bulle present, complete or in- Audital bulle absent.
complete.
Exterior pterygoid region rotund and Exterior pterygoid region angular and

much inflated. not inflated.
First upper incisor flat, without elon- First upper incisor not flat, with very
gated crown. elongated crown.

1 A closely related family, Chrysochlorida, is found in central and southern Africa.

ad

99 NORTH AMERICAN FAUNA, [No. 38,

SUBFAMILIES.

Several mammalogists have undertaken to divide the Talpidee
into groups or subfamilies. The most recent classification of this
sort is that of Thomas,’ who divides the family into five subfamilies,
namely, Desmanine; Talpine, Scalopine, Condylurine, and Urop-
siline. The American genera Scalopus, Scapanus, Parascalops, and
Netiroirichus would fall in the subfamily Scalopine, according to this
arrangement, while the Condylurine would be represented only in
America by Condylura. Such a system of classification, however, is
superficial and unnatural. The anatomical differences, other than
dentition, between Parascalops or Neiirotrichus and each of the
other genera of American moles are as great as those between Con-
dylura and each of the other genera or even greater. In other words,
in order to recognize fundamental and consistent subfamilies it
would be necessary to raise nearly every genus to the rank of a sub-
family. The convenience of such a classification is not apparent,
and accordingly subfamilies have been disregarded here.

HISTORY.

No reference appears in literature to a mole inhabiting America
until Seba,? in 1734, described and Abieineted two mammals which he
called ‘ Tatne, Vien. mger’ and “Talpa, rubra, Americana.’’
Seba, apparently erroneously, gives ‘‘ America” as the habitat of each
of these animals. His figure and description of Talpa virginianus
niger clearly indicate that Talpa europzxa Linneus was the animal in
mind. It is impossible definitely to determine the current species
to which his name Talpa rubra americana refers; probably he had
reference to a specimen of the African genus Chrysochloris, but it is
possible that he based his account upon a verbal description of some
species of the American genus Geomys. The essentials of his de-
scription of Talpa rubra americana refer to the red color, the short,
white, scantily haired tail, and the ‘‘tridactyle” fore feet. Never-
theless, his account shows that the presence of moles in America
probably was suspected at that time. The first definite knowledge
of their accurrence seems to have been obtained by Kalm,’? who,
October 28, 1748, saw burrows and runways of moles near Phila-
delphia, Pa. He captured one of the animals, and remarked upon
its strength and ferocity.

Linnzeus,* in 1758, described two American moles under the names
Sorex aquaticus and Sorex cristatus, basing his accounts largely upon
Kalm’s work. During the century following Linneus’s descriptions

1 Thomas, Oldfield, Ann. & Mag. Nat. Hist., series 8, vol. 10, p. 397, October, 1912.
2 Seba, A., Locupletissimi Rerum Naturalium Thesauri, vol. 1, p. 51, pl. 32, 1734.

8 Kalm, P., Beschreibung der Reise nach dem noérdlichen America, vol. 1, pp. 190-191, 1769.
4 Linnseus, Systema Nature, ed. 10, p. 53, 1758.

1915.] THE FAMILY TALPIDZ. 23

of these two forms, many papers on the habits, anatomy, and
taxonomy of American moles appeared—the result of an interest
aroused largely by the odd habits, specialized anatomy, and unknown
relationships of the group. To discuss here these almost innumer-
able writings would be impracticable, and only the more important
taxonomic revisions and synopses and the first usage of current
generic names will be mentioned. All other generic, specific, and
subspecific names will be discussed in their proper sequence.

The American Talpidse were confused by early zoologists with the
genus Sorex and the Kuropean genus Talpa until the year 1811, when
Illiger based the genus Condylura! upon Sorex cristatus Linneeus, and
the genus Scalops* upon Sorex aquaticus Linneus. The generic name
Scalops had been used previously by Cuvier * who, however, gave no
description or type species. The name Scalops was generally used
for the common mole of eastern United States until 1904, when
Palmer* revived the accepted name, Scalopus Geoffroy,> which ap-
parently had never been employed since proposed in 1803.

One of the first important treatments of a genus of American
moles, other than accounts in books on general natural history, is
given by Bachman,® who revised the genus Scalops, including therein
the genera now recognized as Scalopus, Scapanus, and Parascalops.
He remarked upon the morphology and distribution of Scalops
aquaticus and Scalops townsend, and described two new species
under the names Scalops brewert and Scalops latumanus. All of these
species are recognized in the present revision.

The genus Scapanus was proposed by Pomel,’ in 1848, to include
Bachman’s two species, Scalops townsendw and Scalops brewert. The
type of the genus Scapanus automatically became Scalops townsendii
Bachman when True,’ nearly fifty years after Pomel’s publication,
described the genus Parascalops and designated ee brewerr
Bachman as its type.

The genus Scalops was restored to the genus Tolga by Le Conte ®
in 1854, but his classification was not adopted by zoologists. He
divided Yalpa into three groups: under group i he included the
European genus known to-day as Talpa; under group 2 the modern
genera Scapanus and Parascalops; and under group 3 the genus now
known as Scalopus. In his revision he described two new forms,
one of which, Talpa reposta, is a synonym of Parascalops brewer

1 Tlliger, C., Prod. Syst. Mamm. et Avium, p. 125, 1811.

2 Tiliger, loc. cit., p. 126, 1811.

3 Cuvier, G., Lecons d’Anat. Comp., vol. 1, tab. 1, 1800.

4 Palmer, T. S., Index Generum Mamm., N. Am. Fauna No. 23, p. 620, Jan. 23 1904.
5 Geoffroy Saint Hilaire, E., Cat. Mamm. Mus. Nat. Hist. Nat., p. 77, 1803.

6 Bachman, J., Boston Journ. Nat. Hist., vol. 4, pp. 26-35, January, 1842.

7 Pomel, A., Archiv. Sci. Phys. et Nat., vol. 9, p. 247, November, 1848.

8 True, F. W., Proc. U.S. Nat. Mus., vol.17, p. 242, April 26, 1894.
9 Le Conte, Joseph, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, pp. 326-327, 1854.

94 NORTH AMERICAN FAUNA. [No. 38.

(Bachman); the other, Talpa teniata, is synonymous with Scapanus
townsendii (Bachman).

The same year that Le Conte’s revision appeared, a mole related
to the Japanese genus Urotrichus was collected by George Gibbs in
the Cascade Mountains of western Washington. This specimen later
became the type of Urotrichus gibbsic Baird,' a species which subse-
quently became the type of the genus Netirotrichus Ginther.?

In 1875 Gill? presented a synopsis of the Insectivora, in which he
classified families, subfamilies, and genera, dividing the family
Talpide into two subfamilies, Talpinee and Myogaline. The Talpinze
were further subdivided into three sections, Talpze, Condylure, and
Scalopes; the Myogaline into the sections Mygale and Urotrichi.
The American genus Condylura represented the section Condylure;
Scalops and Scapanus (including also the form now known as Para-
scalops) represented the section Scalopes; Urotrichus (including also
the present Netirotrichus) and the shrewlike Uropsilus formed the
section Urotrichi.

Two years after Gill’s synopsis was published there appeared a
preliminary revision of the American Insectivora by Coues,* in
which he listed six species of moles belonging to four genera.

Dobson,° in an extensive revision of the Insectivora published in
1883, gave detailed accounts of the taxonomic characters and anatomy
of moles, recognizing five American species which he placed in four
genera.

However, it was not until True’s monumental revision ® of the
American moles, in 1896, that the distribution and taxonomic rela-
tionships of the group in America began to be understood. ‘True
recognized eleven forms, distributed among eight species of five
genera, and described one new species, Scapanus orarwus.”

LIST OF GENERIC NAMES THAT HAVE BEEN USED FOR AMERICAN
MOLES.

Astromycter Harris, Amer. Journ. Sci. and Arts, vol. 9, p. 400, June, 1825 (from
Machias, Me., ‘‘Star” [newspaper]). A synonym of Condylura Iliger. Based
on an abnormally colored specimen of Condylura cristata with the tail in the
enlarged winter condition.

Astromyctes Gray, List Spec. Mamm. Brit. Mus., p. 76, 1843. Misprint for Astro-
mycter Harris (=Condylura Illiger).

Astromydes Blyth, Cat. Mamm. Asiat. Soc. Mus., p. 87, 1863. Misprint or emenda-
tion for Astromycter Harris (=Condylura Iliger).

1 Baird, S. F., Reports Explor. & Surv. Pacific Railroad, vol. 8, part 1, p. 76, 1857.

2 Ginther, A., Proc. Zool. Soc. London, 1880, p. 441, October, 1880.

3 Gill, T., Bul. U.S. Geol. & Geog. Surv. Terr., no. 2, series 2, pp. 91-120, 1875.

4Coues, E., Bul. U. 8. Geol. & Geog. Surv. Terr., vol. 3, no. 3, pp. 631-653, May 15, 1877.

6 Dobson, G. E., Monograph of the Insectivora, systematic and anatomical. Part 2, including the
families Potamogalidxz, Chrysochloride, and Talpidez, 1883.

6 True, F. W., Proc. U. 8. Nat. Mus., vol. 19, pp. 1-112, 1896.

7 True, F. W., loc. cit., p. 52.

&

1915.] THE FAMILY TALPID&. 25

Condylura Illiger, Prod. Syst. Mamm. et Avium, p. 125, 1811. Earliest available
name for the genus of which the type is Sorex cristatus Linneus. ‘‘ We owe the
name Condylura to the faulty figure of the animal given by De La Faille, in
which the tail is represented as constricted at intervals, the whole resembling
astring of beads. From this Illiger was led to include in his diagnosis the expres-
sion ‘cauda mediocris nodosa,’ and to bestow an inappropriate name.’’? Illiger
includes two species under the genus: Sorex cristatus Linneus and Talpa longi-
caudata Erxleben.

Condylurus Blainville, Ann. Francais et Etrangéres d’Anat. et de Physiol., vol. 2,
p. 219, 1838. Emendation for Condylura, used subgenerically ad Talpa cristata.

Condylus Van Hyning, Science, n. s., vol. 38, p. 243, August 15, 1913. Misprint
for Condylura.

Condytura Todd, Cyclopedia Anat. and Physiol., vol. 2, p. 994, 1839. Used erro-
neously, but consistently, for Condylura Illiger.

Neourotrichus Rye, Zool. Record, vol. 17, index, p. 8, 1881. EZmendation for Netiro-
trichus Giinther.

Neiirotrichus Giinther, Proc. Zool. Soc. London, 1880, p. 441, October, 1880. Har-
liest name for the genus of which Urotrichus gibbsit Baird is the type. Some-
times used without the dizresis (Neurotrichus Forbes, Zool. Record, vol. 17,
Mammalia, p. 14, 1881).

Néurotrichus Giinther, Proc. Zool. Soc. London, 1880, plate 42, October, 1880.
Misprint for Netrotrichus.

Parascalops True, Proc. U. 8. Nat. Mus., vol. 27, p. 242, April 26, 1894. The first
available name for the genus of which Scalops brewert Bachman is the type.
Perascalops Beddard, Cambridge Nat. Hist., vol. 10, Mammalia, p. 518, 1902.

Misprint for Parascalops.

Rhinaster Wagler, Nat. Syst. Amphib., p. 14, 1830. Name proposed by Wagler to
replace Condylura Illiger which he considered misleading. Under this he
mentions two species: Sorex cristatus Linnezeus and Condylura prasinata Harris.

Scalops Cuvier, Lecons d’Anatomie Comparée, vol. 1, Premier Tableau Général des
Classes des Animaux, 1800. Nomen nudum. The generic name Scalops later
was based upon Sorex aquaticus Linnzeus by Iliger (Prod. Syst. Mamm. et Avium,
p. 126, 1811), but is preoccupied by Scalopus Geoffroy.

Scalopus Geoffroy, Cat. Mamm. Mus. Nat. Hist. Nat., p. 77, 1803. Earliest available
name for the genus of which Sorex aquaticus Linneeus in the type.®

Scalpos Brooks, Rept. W. Va. State Board Agric., Quarter ending December 30, 1910,
p. 28, 1911. Misprint for Scalops.

Scapanus Pomel, Archives Sci. Physiques et Nat., vol. 9, p. 247, November, 1848.
Tenable name for the genus of which Scalops townsendii Bachman is the type.
Pomel’s description ° is faulty in that it states that the nostrils are lateral, a
condition which is not found in Scapanus townsendit.

1 True, F. W., Proc. U. S. Nat. Mus., vol. 19, p. 78, 1896.

2 “Wie bekannt, ist der Schwanz dieses Thieres vollkommen eben.”” (Wagler, loc. cit.)

3 Type fixed by Palmer, T. S. (Index Gen. Mamm., N. Am. Fauna No. 23, p. 621, 1904). The original
generic description reads: ‘‘Caract. nat. Deux incisives & la m&choire supérieure, quatre 4 l’inférieure, les
intermédiaires fort petites; un boutoir, une queue assez longue; pieds pentadactyles, doigts des pAtes
antérieures réunis jusqu’aux ongles seulement; ces ongles assez longs, plats, et dirigés un peu en arriére;
corps couvert de poils.’”’ (Geoffroy, 1803, loc. cit.)

4 Type fixed by Elliot, D. G. (Field Columb. Mus., publ. 45, zool. series, vol. 2, p. 391, 1901).

5 “Je type, LEPTORHINIENS, trompe gréle aigué, narines ouvertes prés de l’extrémité.

“‘ Genres Hyporyssus?, Scalops, Scapanus.

““Nota.—Ce troisiéme genre différe des scalops par la position latérale et non supérieure del’ouverture des
narines, et par la formule dentaire comprenant une intermédiaire supérieure et trois inférieures de plus.
Les espéces sont: Scapanus Towsendii (sic) et Breweri(Scal. Towsendii(sic) et Breweri Bachm.).”’ (Pomel,
A., 1848, loc. cit., p. 247).

26 NORTH AMERICAN FAUNA. [No. 38.

Scapasius Beddard, Cambridge Nat. Hist., vol. 10, Mammalia, p. 518, 1902. Mis.
print for Scapanus.

Scaphanus Herrick, Mamm. Minnesota, Geol. and Nat. Hist. Surv. Minnesota, Bul.
7, p. 55, 1892. Misprint or emendation for Scapanus.

Sorex Linnzeus, Syst. Nat., ed. 10, vol. 1, p. 53, 1758. The American moles Con-
dylura cristata and Scalopus aquaticus were placed in the genus Sorex by Linneeus.
Shaw (Gen. Zool., Mamm., vol. 1, p. 531, 1800) described Sorex radiatus (=Con-
dylura cristata) from a description and faulty figure by De La Faille.

Talpa Linneus, Syst. Nat., ed. 10, vol. 1, p. 52,1758. Type species: Talpa europea
Linneus. A Palearctic genus with which some or all American moies were
considered congeneric by certain early writers. Last used for American Talpide
by Le Conte (Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p.326, 1854), who referred
to it the genera now known as Scalopus, Scapanus, and Parascalops.

Talpasorex Schinz, Cuvier’s Thierreich, vol. 1, p. 191, 1821. Substitution for
Condylura Mliger.

Talpasorex Lesson, Manuel de Mammalogie, p. 124, 1827. Synonym of Scalopus
Geoffroy. Based upon Scalops pennyslvanica Harlan (=Sorex aquaticus Linnzeus),
in the original description of which the dental formula was erroneously stated.
Nec Talpasorex Schinz, by which it is preoccupied.

Urotrichus Temminck, Van der Heeven’s Tijdschr. Nat. Geschied. Physiol., vol. 5,
p. 286, 1838-1839. A Japanese genus with which WNeiirotrichus was included
until 1880.

Keys to the Genera of Moles.
[Based on external characters. ]
a. Length of tail vertebree less than one-fourth the total length; width of palm equal-
ing or exceeding length of palm.

6.1 Tail naked or but scantily haired; nostrils superior.
c!, Tail somewhat slender, essentially naked; fore toes webbed; geographic range

east of Rocky Mountains::..22 222125. 2) ee ee Scalopus (p. 27).

c?, Tailsomewhat fleshy, scantily haired; fore toes not webbed; geographic range
west of Rocky Mountains 2. -2: 80522225 22 S32 eee ee Scapanus (p. 64).

b.2 Tail densely covered with hair; nostrils lateral........... Parascalops (p. 77).

a.2 Length of tail vertebre more than one-fourth the total length; width of palm
less than length of palm.
6.1 Anterior end of snout surrounded by fringe of processes; nostrils circular to
oval, anterior; geographic range east of Rocky Mountains . .. Condylura (p. 82).
b?, Anterior end of snout not surrounded by fringe of processes; nostrils crescentic,
lateral; geographic range west of Rocky Mountains: . .. Netrotrichus (p. 92).
[Based on cranial and dental characters.]

a. Audital bulle complete; interior basal projection of upper molars narrow, simple.

b'. Mastoids relatively heavy; interparietal small; no functional lower canine;

lower incisors two; geographic range east of Rocky Mountains. Scalopus (p. 27).

b?, Mastoids relatively weak; interparietal large; lower canine present; lower

incisors three; geographic range west of Rocky Mountains. .-Scapanus (p. 54).

a?, Audital bulls incomplete; interior basal projection of upper molars relatively
broad, lobed.

b'. Interior basal projection of first and second upper molars trilobed; premolars, 4.

c!, Braincase relatively broad and flat; anterior nares directed forward; first

upper incisors broad, directed downward, with an exterior basal acces-

BOTY: ;CUSD). teseis-arcisseais oe Sa ASE Ee ae ee eee Parascalops (p. 77).

1915.] GENUS SCALOPUS. 27

c?. Braincase relatively narrow and high; anterior nares directed obliquely
upward; first upper incisors spatulate, directed somewhat forward, without

BISA ACCESSOFYPCUSP rast. .2 coke UE Condylura (p. 82).
b?. Interior basal projection of first and second upper molars bilobed; premo-
1 LTTE, i ies i aI ces nl ge Netiroirichus (p. 92).

List of American Genera, Species, and Subspecies, with Type Localities.
Scalopus aquaticus aquaticus (Linnezus)... Philadelphia, Pa. (p. 32).

aquaticus howell: Jackson...........-- Autaugaville, Ala. (p. 36).

aquaticus australis (Chapman)..-..... Gainesville, Fla. (p. 38).

aquaticus anastase (Bangs)......-.-- Point Romo, Anastasia Island, Fla. (Pp. 39).

aquaticus parvus (Rhoads).........-.-- Tarpon Sees Hla W(peAl)e

aquaticus machrinus (Rafinesque).-.-- Lexington, Ky. (p. 42).

aquaticus machrinoides Jackson......- Manhattan, Kans. (p. 45).

aquaticus pulcher Jackson.........--- . Delight, Ark. (p. 46).

aquaticus caryi Jackson...........---- Neligh, Nebr. (p. 48).

aquaticus intermedius (Elliot). .....-- Alva, Okla. (p. 49).

aquaticus texanus (Allen).....-..-.-..- Rockport, Tex. (p. 50).

SEOEIIGS (I BT PEED cae Alpe ae Stilwell, Okla. (p. 52).

POU SIACKSOM 05 koh o2 ees 5 os 45 miles from Brownsville, Tex., in

Tamaulipas, Mexico (p. 53).

Scapanus townsendvi (Bachman).......-.-- Vicinity of Vancouver, Wash. (p. 58).

Oranies oranus Wruie. 2.20. 22 2. Shoalwater Bay, Wash. (p. 61).

orarius scheffert nobis.....-.-----.---- Walla Walla, Wash. (p. 63).

latimanus latumanus Caen re ee Santa Clara, Cal. (p. 64).

latimanus occultus Grinnell & Swarth. Santa Ana Canyon, Cal. (p. 68).

latimanus grinnelli Jackson........--- Independence, Cal. (p. 69).

latimanus sericatus Jackson.........-- Yosemite, Cal. (p. 71).

latimanus minusculus Bangs........ Fyffe, Cal. (p. 72).

lavimanus dilatus True. .2 5.0.22... - Fort Klamath, Oreg. (p. 72).

latimanus alpinus Merriam ........... Crater Lake, Oreg. (p. 75).

Oheraniewilen. 2. on eee eo ee San Pedro Martir Mountains, Lower

California. (p. 75).

Parascalops brewert (Bachman).........-- Marthas Vineyard, Mass. (p. 79).
Condylura cristata (Linneeus)........---- Eastern Pennsylvania (p. 86).
Netirotrichus gibbsit gibbsit (Baird)........ White River Pass, Wash. (p. 94).

gibbsir hyacinthinus Bangs............ Nicasio, Cal. (p. 97).

Genus SCALOPUS Geoffroy.

Scalops Cuvier, Legons d’Anat. Comp., tome 1, tabl. 1, 1860. (Nomen nudum.)

Scalopus Geoffroy, Cat. Mamm. Mus. Nat. Hist. Nat., p. 77, 1803.

Scalops Illiger, Prod. Syst. Mamm. et Avium, p. 126, 1811.

Talpasorex Lesson, Manuel de Mamm., p. 124, 1827. Based upon Scalops pennsyl-
vanica Harlan. Nec Talpasorex Schinz.

Scalpos Brooks, Rept. W. Va. State Board Agric. for Quarter ending December 30
SIO. p: 28, 1911.

Type species.—Sorex aquaticus Linnzus.
Geographic range.—Hastern and central Massachusetts, south-

eastern New York, southern Pennsylvania, extreme southern
Ontario (Point Pelee), southern Michigan, northern Illinois, western

28 NORTH AMERICAN FAUNA. [No. 38.

Wisconsin, central Minnesota, extreme southeastern South Dakota,
northern Nebraska, extreme northeastern Colorado, south and east
to northeastern Tamaulipas, Mexico (45 miles from Brownsville,
Texas), to the Gulf of Mexico, and in Florida to Tampa Bay and
Lemon City. (See fig. 2).

External characters.—Body robust, depressed; tail short, round,
indistinctly annulated, very scantily haired (in appearance, essen-

B2008-103
Fic. 2.—Geographic range of the species and subspecies of Scalopus.

1. S. aquaticus aquaticus. 6. S.a. machrinus. 10. S. a. intermedius.
2. S.a. howelli. 7. S.a. machrinoides. ll. S.a. tezanus.

3. S. a. australis. 8. S.a. pulcher. 12. S. xereus.

4, S. a. anastase. 9. S.a. caryi. 13. S. inflatus.

5. S. a. parvus.

tially naked) (fig. 3). Head conoidal, depressed. Nose elongated
into a distinct snout, apical portion naked to line of anterior edge of
nasals; nostrils superior, crescentic, with concavities turned in later-
ally (fig. 4). Eyes mimute, concealed in fur. Auricular orifice small.
Legs short and stout. Feet large, fleshy, scantily haired above,
naked below, without tubercles. Fore feet handlike, the palms

1916.]

GENUS SCALOPUS. 29

broader than long (fig. 5). Fore toes and hind toes webbed. Claws
of fore feet broad, flat, and heavy; those of hind feet relatively short

B2009-103
Fie. 3.—Tail of
Scalopus aquati-

cus aqguaticus .

(X14).No.203190;
U.S. Nat. Mus.,
Biological Sur-
vey collection;
from Woodside,
Montgomery
County, Md.

and weak (fig.6). Fur
dense, soft, silky, the
hairs nearly equal in
length, producing a vel-
vetlike pelage. Mamme
6: latero-pectoral, 1-1; any
B2010-103
latero-abdominal, 1-1; Fre. 4.—Snout of S.a.aquaticus (X 13). In-
inguinal, ae: dividual referred to in fig. 2.
Skeletal characters.—Clavicle short and heavy, about
two-thirds as broad as long, penetrated antero-posteri-
orly through the center by a small circular foramen;
humerus heavy, about three-fourths as broad as long.
Pelvis narrow, bones of the opposite sides adjunct un-
der acetabula; two osseous bridges connect sacral ver-
tebree with ischium and produce four large foramina

HY
Pi \,

or openings, one in each of the angles formed by median lines of
acetabula and sacral vertebre. Superior surface of last sacral ver-

tebra with a very
small, low, late-
rally flattened,
longitudinal proc-
ess. Os falciforme
long, reaching to
proximal end of
terminal phal-
ange of first digit,

wider and rather ye B2011-103 °
sharply incurved FG. 5.—Forefoot of S. a. aquaticus (X14). Individual referred to in fig. 2.
at base, narrower and gradually tapering distally.

Skull conoidal, flat, with relatively broad braincase, and consider-

ably constricted interorbitally. Mastoids

» NSA A A relatively heavy and prominent. Inter-
PY Sa AAW ° °
4, ¥ £2) parietal short ‘and narrow, somewhat ir-
fZ rs! i e @

e # regular in outline, but usually narrower

anteriorly. Frontal region flat, not much
sloping; frontal sinuses swollen. Rostrum
relatively short; anterior ends of pre-
7 maxille much thickened and extending
beyond nasals, forming an acute notch
anterior to nasals. Anterior nares open-

—yYy
Se ee ieee!

B2012-103

Fic. 6.—Hind foot of S. a. aquatius ing forward. Zygomata moderately long

(X14). Individual referred toin fig. 2.

and heavy, not much curved, descending

slightly and slanting inward anteriorly, the posterior end well back
onsquamosal. Foramen magnum oval, of moderatesize. Infraorbital

30 NORTH AMERICAN FAUNA. [No. 88.

foramen relatively small, the plate forming its outer wall moder-
ately broad (slightly narrower than diameter of foramen). Au-
dital bullz complete, depressed (relatively higher than in Scapanus) ;
auditory meatus short, scarcely developed. External pterygoid
region much inflated posteriorly, slightly inflated anteriorly. Mes-
opterygoid space broad, the sides gently concave. Palate mod-
erately elongate, terminating at a distance posterior to last molar -
about equal to diameter of that tooth; posterior border of pal-
ate truncate, frequently with a median notch or spine. Anterior
palatine foramina small, oval to elliptical; first (anterior) pair of
posterior palatine foramina small (smaller than anterior palatine
foramina), round to oval; second (posterior) pair minute. Hori-
zontal ramus of mandible heavy, curved upward both at posterior
and anterior ends; coronoid process moderately elongate, triangular,
somewhat acute, directed shghtly posteriorly; angle of mandible rela-
tively short, broad (broader than coronoid), and truncate; inferior
mandibular notch moderate, subcircular, relatively narrow and deep.

Dental characters.—First upper incisor long and broad, convex in
front, flat posteriorly; second and third upper incisors lateral, simple,
conical, minute. Upper canine about two-thirds as large as first
incisor, simple. Upper premolars indistinctly cuspidate, the second
about twice the size of the first and about equal in size to canine (but
broader), the third about half as large agam as the second. Upper
molars W-shaped in transverse section, with an antero-internal
V-shaped cusplike shelf (not lobed); first and second molars subequal,
the third much smaller.

First lower incisor small, elongate, conical; second lower incisor
about twice the size of the first, elongate, caninelike, with an infero-
internal longitudinal groove; no persistent lower canine. Premolars
successively increasing in size posteriorly, somewhat simple, with a
slight tendency toward the development of a cingulum posteriorly.
Lower molars M-shaped in transverse section, the antero-internal cusp
bilobed; molars successively decreasing in size posteriorly. Func-
tional dentition: 1. 3; c. 4; pm. 3; m. 3; total 36.1

Keys to Species and Subspecies of Scalopus.

I. KEY TO ADULT MALES.

a.! Geographic range east of Mississippi River.
b.1 Total length more than 153 mm.; greatest length of skull more than 33 mm.;
breadth of skull across mastoids usually more than 17 mm.
c.! Total length less than 190 mm.; length of hind foot less than 21.5 mm.; greatest
length of skull less than 37 mm.; breadth of skull across mastoids less than
VO Wags 2.2.32 ogee en 2s se eee eee Scalopus aquaticus aquaticus (p. 32).

1 The actual dentition of Scalopus is: i. 3; c. 4; pm. 3; m. $; total, 40. The third lower incisor and the
lower canine are nonpersistent and disappear before the animal is mature.

1915.] GENUS SCALOPUS. ae

c.2 Total length more than 190 mm.; length of hind foot more than 21.5 mm.;
greatest length of skull more than 37 mm.; breadth of skull across mastoids
more than 19 mm...................-Scalopus aquaticus machrinus (p. 42).
6.2 Total length less than 153 mm.; greatest length of skull less than 33 mm.;
breadth of skull across mastoids usually less than 17 mm.
c.1 Length of hind foot usually less than 17 mm.; greatest length of skull 31.2 mm.
or less; breadth of skull across mastoids less than 16 mm.,

Scalopus aquaticus parvus (p. 41).
c.2 Length of hind foot usually more than 17 mm.; greatest length of skull more

than 31.2 mm.; breadth of skull across mastoids more than 16 mm.
d.1 Color golden sepia; face, chin, and wrists bright zinc orange; geographic
range Anastasia Island, Fla........-- Scalopus aquaticus anastasx (p. 39).

d.? Color not golden sepia; face, chin, and wrists not bright zinc orange.
e.! Color paler; greatest length of skull usually more than 32 mm.; geo-
. graphic range North and South Carolina, northern and western Georgia,
west to mouth of Mississippi River. ...Scalopus aquaticus howelli (p. 36).
e.2 Color darker; greatest length of skull usually less than 32 mm.; geo-
graphic range southeastern Georgia and eastern Florida,
Scalopus aquaticus australis (p. 38).
a.2 Geographic range west of Mississippi River.
6.1 Breadth of skull across mastoids 19.3 mm. or more.

c.1 Breadth of skull across mastoids more than 20 mm.; palatilar length more than

GMM «2... Br aeiatelaye ata) 8 at wale Siete cieisre 2 Scalopus. aquaticus machrinus (p. 42).
c.2 Breadth of skull across mastoids less than 20 mm.; palatilar length less than
MUGEN ee a8 ciaiciese'sia cles ais isin clole-w Scalopus aquaticus machrinoides (p. 45).

6.2 Breadth of skull across mastoids less than 19.3 mm.
c.1 Greatest length of skull more than 33 mm.; interorbital constriction more than
7.3 mm.
@ Ooler coppery snudl, DrOWD: 22.5... sce. vob bee seine: - Scalopus xreus (p. 52).
d.2 Color not coppery snuff brown. .
e.! Color dark; geographic range east of 95th meridian,
Scalopus aquaticus pulcher (p. 46).
e.2 Color pale; geographic range west of 95th meridian.
f. Nose and ankles tinged with ochraceous,
| Scalopus aquaticus intermedius (p. 49).
Jf. Nose and ankles not tinged with ochraceous, but usually with ivory

eM OMise sodas crs seid: cee te a als Scalopus aquaticus cary? (p. 48).

c.2 Greatest length of skull less than 33 mm.; interorbital constriction less than
7.3 mm.

d.1 Prelachrymal region much inflated; rostrum broad and truncate; geo-

graphic range Tamaulipas, Mexico.............- Scalopus inflatus (p. 53).

d.? Prelachrymal region not much inflated; rostrum not broad and truncate;

geographic range Texas. ............... Scalopus aquaticus texanus (p. 50).

4 II. KEY TO ADULT FEMALES.

a.1 Geographic range east of Mississippi River.
b.1 Total length usually more than 148 mm.; greatest length of skull more than
32 mm.; breadth of skull across mastoids usually more than 16.5 mm.

c.1 Total length less than 173 mm.; length of hind foot less than 21 mm.; greatest
length of skull less than 35.2 mm.; breadth of skull across mastoids less than

Dey ee eh. ee Scalopus aquaticus aquaticus (p. 32).

c.? Total length more than 173 mm.; length of hind foot more than 21 mm.; great-

est length of skull more than 35.2 mm.; breadth of skull across mastoids

MMOre tial US. (MM oe ee. ws Scalopus aquaticus machrinus (p. 42).

32 NORTH AMERICAN FAUNA. [No. 38.

6.2 Total length usually less than 148 mm.; greatest length of skull 32 mm. or less;
breadth of skull across mastoids usually less than 16.5 mm.
c.! Total length less than 130 mm.; length of tail less than 18 mm.; greatest length
of skull’29'3 mm“ or less: 7232 s2ace see Scalopus aquaticus parvus (p. 41).
c.2 Total length more than 130 mm.; length of tail more than 18 mm.; greatest
length of skull more than 29.3 mm.
d.' Color golden sepia; face, chin, and wrists bright zinc orange; geographic
range Anastasia Island, Fla.......... Scalopus aquaticus anastasz (p. 39).
d.? Color not golden sepia; face, chin, and wrists not bright zinc orange.
e.! Color paler; greatest length of skull usually more than 31 mm.; geographic
range North and South Carolina, northern and western Georgia, west to
mouth of Mississippi River.-.......- Scalopus aquaticus howelli (p. 36).
e.2 Color darker; greatest length of skull usually less than 31 mm.; geographic
range southeastern Georgia and eastern Florida,
Scalopus aquaticus australis (p. 38).
a.? Geographic range west of Mississippi River.
b.! Total length more than 160 mm.; breadth of skull across mastoids more than

18.4 mm.
c.! Breadth of skull across mastoids more than 19 mm.; palatilar length more than
d Bao 06) 40 Pear bis Ae ae es MS Scalopus aquaticus machrinus (p. 42).
c.2 Breadth of skull across mastoids less than 19 mm.; palatilar length less than
BSS 000 00 REM e pa a OL i calaye Aaa Soa ba Scalopus aquaticus machrinotides (p. 45).
b.? Total length less than 160 mm.; breadth of skull across mastoids less than 18.4
mm.
c.! Greatest length of skull more than 32 mm.; interorbital constriction more than
7.2 mom.
d* Color coppety snuli brown 222) ee Scalopus xreus (p. 52).

d.* Color not coppery snuff brown.
e.' Color dark; geographic range east of 95th meridian,
Scalopus aquaticus n (p. 46).
e.” Color pale; geographic range west of 95th meridian.
jf.) Nose and ankles tinged with ochraceous,
Scalopus aquaticus intermedius (p. 49).
J.? Nose and ankles not tinged with ochraceous, but usually with ivory

OVO We SU US UN eatin ae Scalopus aquaticus caryt (p. 48).

c.2 Greatest length of skull less than 32 mm.; interorbital constriction less than
7.2 Tom.

d.' Prelachrymal region much inflated; rostrum broad and truncate; geo-

graphic range Tamaulipas, Mexico............. Scalopus inflatus (p. 58).

d.? Prelachrymal region not much inflated; rostrum not broad and truncate;

geographic range Texas...........--.. Scalopus aquaticus texanus (p. 50).

Descriptions of Species and Subspecies of Scalopus.
SCALOPUS AQUATICUS AQUATICUS (Linnzus).
EASTERN MOLE.
(Pl. I, figs. 1, 2; Pl. II, fig. 1; Pl. TU, figs. 1, 1a; Pl. Vi, fig. 32)

Sorex aquaticus Linneus, Syst. Nat., ed. 10, p. 53, 1758.

Talpa europexa flavescens Erxleben, Syst. Reg. Anim., p.118,1777. Based on ‘“‘yellow
mole” of Pennant; type locality, New York(?).

Talpa Flava Zimmermann, Specimen Zool. Geog., p. 496, 1777. Based on ‘‘yellow
mole” of Pennant; type locality, New York(?).

1915.] SCALOPUS AQUATICUS AQUATICUS. 33

Talpa fusca Zimmermann, Specimen Zool. Geog., p. 497, 1777. Based on ‘‘ Brown
mole” of Pennant; type locality, New York(?).

[ Talpa} [europxa] flava Gmelin, Linn. Syst. Nat., ed 13, p. 110, 1788.

Scalopus virginianus Geoffroy, Cat. Mamm. Mus. Nat. Hist. Nat., p. 78, 1803. Type
locality, Virginia(?).

Talpa cupreaia Rafinesque, Précis des découv. et travaux somiologiques, p. 14, 1814.
Type locality, ‘‘Atlantic States’’ (see Atl. Journ. 1832, p. 61).

Scalops canadensis Desmarest, Mammalogie, 1'° partie, p. 155, 1820. New name for
Sorex aquaticus Linneus.

Scalops pennsylvanica Harlan, Fauna Amer., p. 33, 1825. Type locality unknown,
probably Pennsylvania.

Talpasorex pensylvanica Lesson, Manuel de Mamm., p. 124, 1827.

Sclalops| aquaticus Fischer, Synop. Mamm., p. 249, 1829.

Talpa virginiana Blainville (nec Talpa eee Breen), Annales Franc. et Etrang.
d’Anat et de Physiol., tome 2, p. 219, 1838.

Talpa sorex pensylvanicus Blainville, Annales Frang. et Etrang. d’Anat. et de Physiol.,
tome 2, p. 219, 1838. (In synonymy.)

Talpa (Scalops) Virginiana Blainville (nec Talpa virginiana Brisson), Ostéographie,
Atlas 1, table des planches, p. 4; Insectivores, pl. 5 (skull), pl. 9 (teeth), 1839-
1864.

T{alpa] aquatica Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327,
1854.

Scalops aquaticus aquaticus True, Proc. U. 8. Nat. Mus., vol. 7, 1884, p. 606, 1885.

Scalopus aquaticus Oberholser, Mammals and Summer Birds of Western North Caro-
lina (publ. by Biltmore Forest School, Biltmore, N. C.), p. 3, June 30, 1905.

Scalopus acquaticus (sic) Hahn, Proc. U. 8. Nat. Mus., vol. 32, p. 464, 1907.

Scalpos (sic) aquaticus Brooks, Rept. W. Va. Board Agric., 1910, p. 28, 1911.

Scalopus aquaticus aguaticus Miller, U. 8. Nat. Mus., Bul. 79, p. 7, December 31, 1912.

Type locality.— Philadelphia, Pennsylvania.

Geographie range.—Eastern United States from eastern and south-
ern Massachusetts, southeastern New York, and southeastern Penn-
sylvania, south through Virginia, and in the Appalachian Mountains
south through western North Carolina and eastern Tennessee.

General characters.—Size medium; color dark; skull high, heavy,
and angular; rostrum truncate; dentition moderate.

Color.—F resh winter pelage: Back fuscous, fuscous-black, or blackish
brown, becoming paler (drab-gray) on nose and ankles; fur at base
of hairs dark neutral gray; underparts slightly paler than back,
usually showing more neutral gray of base of hairs and frequently
tinged ventrally with mummy brown or fuscous. Summer pelage:
Slightly paler than in winter, usually more brownish; back sepia,
fuscous, or fuscous-black; underparts paler than back, more grayish
and usually tinged on chest with mummy brown or sepia. Young:
More grayish than adults and seldom, if ever, tinged ventrally with
brown.

Skull.—Medium in size, angular, much constricted interorbitally
and usually somewhat depressed postorbitally, slightly swollen
supraorbitally; mastoids moderately heavy; interparietal narrow;

90549°—15—_3

34 NORTH AMERICAN FAUNA, © [No. 38.

coronoid process and angle of mandible broad and heavy, the former
usually with a small mammiform process on posterior border; teeth
medium. :
Measurements.—Average of 2 adult males from type locality:
Total length, 182.5 (180-185); tail vertebree, 29.5 (28-31); hind foot,
21 (21-21). Average of 2 adult females from type locality: 154.5
(154-155); 23 (28-23); 19.5 (19-20). Average of 15 adult males
from Washington, D.C.: 163.4 (154-175); 26.5 (22-29); 19.8 (18-21).
Average of 8 adult females from Washington, D. C.: 152.6 (146-168);
26 (21-28); 19 (18-20). Average of 6 adult males from Wareham,
Mass.: 159.8 (154-167); 26.3 (21.5-30); 19.9 (19.5-21). Skull:
Average of 3 skulls of adult males from type locality: Greatest length, .
35.3 (34.6-35.7); palatilar length, 14.8 (14.7-15); mastoidal breadth, _
18 (17.8-18.2); interorbital breadth, 7.4 (7.3-7.5); maxillary tooth
row, 10.9 (10.9-11); mandibular molar-premolar row, 10.7 (10.6-
10.8). Average of 21 skulls of adult males from Washington, D. C.,
and vicinity: Greatest length, 34.3 (33.2-35.6); palatilar length,
14.7 (14.3-15.2); mastoidal breadth, 17.7 (17-18.3); interorbital
breadth, 7.4 (7.2-7.8); maxillary tooth row, 10.8 (10.4-11.3); man-
dibular molar-premolar row, 10.4 (10.1-10.8). Average of 15 skulls
of adult females from Washington, D. C., and vicinity: Greatest
length, 32.9 (32.3-34.2); palatilar length, 13.9 (13.5-14.7); mastoidal
breadth, 17.1 (16.3-17.5); interorbital breadth, 7.4 (7-7.6); maxil-
lary tooth row, 10.4 (10.1-10.8); mandibular molar-premolar row,
10.2 (9.8-10.4). Average of 5 skulls of adult males from Wareham,
Mass.: Greatest length, 34.3 (83.8-34.8); palatilar length, 14.8
(14.5-15); mastoidal breadth, 18.1 (17.8-18.4); interorbital breadth,
7.7 (7.6-7.7); maxillary tooth row, 11 (10.8-11.2); mandibular
molar-premolar row, 10.7 (10.6-10.8). Average of 2 skulls of adult
males from Dismal Swamp (Wallaceton), Va.: Greatest length, 35.2
(35.1-35.2); palatilar length, 14.8 (14.6-15); mastoidal breadth, 18.1
(18-18.2); interorbital breadth, 7.5 (7.4-7.6); maxillary tooth row,
11.1 (11-11.2); mandibular molar-premolar row, 10.7 (10.6-10.7).
Skull of adult male from Roan Mountain, N. C.: Greatest length, 36.1;
palatilar length, 15.5; mastoidal breadth, 18.4; interorbital breadth,
7.8; maxillary tooth row, 11.38; mandibular molar-premolar row, 11.
Remarks.—The first specific description of the common mole of
eastern, United States is that under Sorex aquaticus Linneus;! Lin-
neus places in its synonymy “ Talpa, Virgimanus, noger’ of Seba; but
the description and figure? given by Seba seem to characterize the
European rather than the American mole; the Linnean description,
however, clearly refers to the common mole of eastern America; as an
1 Linneeus, Syst. Nat., ed. 10, p. 53, 1758.
2 Seba, A., Thesaurus, vol. 1, p. 51, pl. 32,1734. Seba’s description of Talpa virginianus niger became the

basis for Shaw’s diagnosis of Talpa purpurascens (Shaw, G., Gen. Zool., Mamm., vol1, p. 521, 1800), which
is therefore a synonym of Talpa europza Linneus.

1915.] SCALOPUS AQUATICUS AQUATICUS. 35

authority for the habitat of the species, Linnzeus cites Kalm, who saw
mole runways along the Schuylkill River near Philadelphia.

In 1771 Pennant‘ described two abnormally colored or faded
specimens of this species under the names of “Yellow Mole” and
“Brown Mole’’; the yellow mole became Talpa europea flavescens at
the hands of Erxleben? in 1777, and the same year Zimmermann ®
named the yellow species Talpa flava and the brown one Talpa fusca.
Geoffroy * described Scalopus virgimianus in 1803, placing in synonymy
under it Talpa virgunianus mger Seba and Sorex aquaticus Linneeus.

The description given by Rafinesque * of Talpa cupreata is unsatis-
factory, but seems to apply to the subspecies S. a. aquaticus. The
synonymy of this form was still further encumbered when Desmarest ®
proposed the name Scalops canadensis in 1820, apparently basing the
name upon “Le Scalope du Canada”’ of Cuvier and others. The last
specific name added to the synonymy of this form is Scalops pennsyl-

vanica Harlan;’? Harlan believed the structure of the molars in the
specimen he described to be different from that of S. aquaticus, but
his description applies very accurately to S. aquaticus except as to the
number of teeth, which he states to be forty; elsewhere, however, he
writes that ‘‘this species corresponds in the number and arrangement
of its teeth with the genus Scalops of F. Cuvier.”’ ®

Scalopus aquaticus aquaticus, the common mole of northeastern
United States, in full winter pelage is the darkest of the genus. Itis
subject to slight local variations in size and color, and even in cranial
characters, which, were they constant over any considerable geo-
graphic range, might be considered differential enough for subspecific
recognition. Specimens from Marthas Vineyard and Nantucket
Islands and from the mainland of southeastern Massachusetts aver-
age very slightly paler and smaller than typical aquaticus, and their
skulls are slightly shorter and relatively broader than those from
the type locality. Specimens from Liberty Hill, Conn., average
large; eastern New York specimens are much like those from the
vicinity of Philadelphia, Pa., while those from Long Island are very
slightly smaller and possibly more grayish. Toward the south aqua-
ticus gradually decreases in size, intergrading with S. a. howella in
northern and western North Carolina. A large series from the
District of Columbia averages smaller than specimens from the
type region, but in all essential characters except size they are

1 Pennant, T., Quadrupeds, 1771. The writer has not had access to this publication, but presumes the
descriptions are about the same as those in Pennant’s History of Quadrupeds, ed. 3, vol. 2, pp. 230, 232, 1793.

2 Erxleben, J.C. P., Syst. Reg. Anim., p. 118, 1777.

3 Zimmermann, E. A. W., Spec. Zool. Geog., pp. 496-497, 1777.

4 Geoffroy Saint Hilaire, E., Cat. Mamm. Mus. Nat. Hist. Nat., p. 78, 1803.

5 Rafinesque, C. S., Précis des découv. et trav. somiolog., p. 14, 1814.

6 Desmarest, A. G., Mammalogie, part 1, p. 155, 1820.

7 Harlan, R., Fauna Amer., p. 33, 1825.
8 Harlan, loc. cit., p. 34, 1825.

36 NORTH AMERICAN FAUNA. [No. 38.

typical of aguaticus. Dismal Swamp, Va., produces a mole which
is as large as the typical form and has a little more tendency toward
a mummy brown shade con the back. Material from the Appa-
lachian Mountain region is entirely too scarce to determine the exact
relationship and area of intergradation of aquaticus with S. a. machri-
nus; a specimen from Roan Mountain, N. C., shows a very slight
approach toward machrinus in size. A skin without skull from
Walden Ridge, Tenn., is here rather arbitrarily referred to aquaticus
on account of its size and color.
Specimens examined.—Total number, 322, as follows:

Connecticut: Cos Cob, 3; East Hartford, 2; Liberty Hill, 6.1

District of Columbia: Washington, 72.

Maryland: Anne Arundel County, 2; Baltimore, 2; Berwyn, 2; Branchville, 5;
Cabin John, 1; Capitol View, 1; Chesapeake Beach, 1; Chevy Chase, 1; High-
land, 1; Howard County, 1; Landover, 1; Laurel, 11; Mount Rainier, 1;
Plummer’s Island, 1; Rockville, 2; Seven Locks, 1; Siiver Spring, 1; Wood-
side, 5.

Massachusetts: Holyoke, 1;1 Middleboro, 1; Nantucket, 6;! Wareham, 32;!
West Tisbury, 4.1

New Jersey: Afton, 1; Audubon, 6;? Englewood, 1;3 Fairhaven, 1;! Haddon-
field, 3.?

New York: Brooklyn, 1; Cold Spring Harbor, 1;° Cypress Hills, 1;? Dobb’s
Ferry, 1; Dunwoodie, 1;3 Hastings, Westchester County, 6;3)4 Lake Grove,
16; Locust Valley, 1;3 Long Island, 1; Mount Sinai, 2;3 New Rochelle, 1;3
New York City, 17;3+ Pelham Manor, 1;? Oyster Bay, 1; Piermont, 1;3
Rye, 1;} Sing Sing, 15; Southhampton, 1; Tarrytown Heights, 2.3

North Carolina: Asheville, 1;3 Buncombe County, 2;! Highlands, 1; Magnetic
City, 1; Roan Mountain (altitude 2500 feet), 1; Weaverville, 8.354

Pennsylvania: Carlisle, 6; Collingdale, 2;? Delaware County, 1; Marple, 1;?
Mechanicsville, 1;? Media, 2;? Mifflintown, 1; Philadelphia (type locality), 6.?

Tennessee: Blount County, 1; Walden Ridge (near Soddy), 1.

Virginia: Alexandria County, 3; Arlington, 3; Bristow, 2; Clark County, 1;
Dismal Swamp, 9; Eastville, 1; Falls Church, 9; Fort Myer, 6; Springvale, 1.

West Virginia: Berkeley Springs, 1.

SCALOPUS AQUATICUS HOWELLI Jackson.
Howe .v’s MOLe.
(PL. 1, fig. 3;, PL. II, fie. 25° Pl: ITI, fies. 2, 2a; Pl Vibes

Scalopus aquaticus howelli Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 19, Feb-
ruary 2, 1914.

Type locality.— Autaugaville, Autauga County, Alabama.

Type specimen.—No. 177931, U. S. Nat. Mus., Biological Survey
collection; ¢ adult, skm and skull; collected January 4, 1912, by
L. S. Golsan.

Geographic range.—North Carolina (except in Appalachian Moun-
tains), South Carolina, northern Georgia, thence southwest across

1 Collection Mus. Comp. Zool., Harvard College. 4 Collection Amer. Mus. Nat. Hist.
2 Collection Acad. Nat. Sci. Philadelphia. 4 Collection Field Mus, Nat. Hist,

1915.) SCALOPUS AQUATICUS HOWELLI. a

central Alabama and southern Mississippi to Pensacola Bay and the
Mississippi River.

General characters.—Intermediate in size between S. a. aquaticus
and S. a. australis; usually paler than either aquaticus or australis;
skull flat, less angular than in aquaticus; rostrum long and narrow.

Color.— Full winter pelage: Back dark drab, hair-brown, or fus-
cous, becoming in most specimens bufiy brown on head; nose and
wrists usually slightly suffused with ochraceous-tawny or cinna-
mon; underparts more grayish than back, usually much tinged
with Saccardo’s umber or tawny-olive. Fresh summer pelage:
Back dark cinnamon-drab or drab, sometimes with a very slight
coppery sheen, becoming cinnamon-bufi or ochraceous-tawny on
face; underparts hair-brown to neutral gray, grading into sepia on—
chest.

Skull.—Size medium (smaller than that of S. a. aquaticus, larger
than that of S. a. australis) flat; rostrum long and narrow; man-
dible weak; coronoid process and angle of mandible weak and very
narrow, incurved toward condyle, making superior and inferior
notches small and relatively deep; teeth small.

Measuremenis.—Type (adult male): Total length, 152; tail verte-
bre, 20; hind foot, 18. Skull: Average of 5 skulls of adult males
ee ae eeskt Greatest length, 32.3 (81.8-32.7); palatiler
length, 13 (12.7-13.3); mastoidal breadth, 16.7 (16.3-17.1); inter-
orbital breadth, 7.2 (7-7.4); maxillary tooth row, 10.1 (10-10.3);
mandibular molar-premolar row, 10.1 (9.9-10.2).

Remarks.—Howell’s mole differs from both S. a. aquaticus and
S. a. australis in cranial characters and is not strictly intermediate
between the two. It shows slight local variations which on account
of scarcity of material are not fully understood. Skulls from Bay
St. Louis and Washington, Miss., and from New Orleans and St.
Tammany Parish, La., have braincases higher and narrower than
typical specimens, and the rostra taper more gradually; none of
the skulls, however, is from a fully adult animal, and it is possible
that age might change this condition. Skulls from Biloxi, Cedar-
bluff, and Kemper County, Miss., are slightly heavier and more angu-
lar than those from the type region. In northern North Carolina
S. a. howell, begins to approach aquaticus in size. A series of speci-
mens from Young Harris, Ga., shows in color a tendency toward
aquaticus and has skulls which in angularity and height of braincase
are like those of aquaticus, but which in size and breadth of rostrum
are hike those of howell. A skin without skull from Pensacola, Fla.,
is indeterminable, but is provisionally referred to howelli.

Specimens examined.—Total number, 106, as follows:

Alabama: Ardell, 1; Auburn, 1; Autaugaville (type locality), 15; Castleberry,

2; Cottondale, 1; Eutaw, 1; Greensboro, 3; Huntsville, 1; Sand Mountain
(near Carpenter), 3.

38 NORTH AMERICAN FAUNA. [No. 28.

Florida: Pensacola, 1.1

Georgia: Columbus, 2; Crawfordsville, 1; Griffin, 1; Young Harris, 12.

Louisiana: New Orleans, 2; St. Francisville, 1; St. Tammany Parish, 1.?

Mississippi: Bay St. Louis, 3; Biloxi, 2; Cedarbluff, 1; Kemper County, 1;
Washington, 1.

North Carolina: Apex, 1;* Bertie County, 1; Jackson, 2; Kinston, 3; Moran,
Chowan County, 1; Raleigh, 23; Wilkesboro, 1.

South Carolina: Abbeville, 1;+ Beaufort County, 4; Calhoun Falls,1;* Catawba,
2;% Charleston, 1; Frogmore, 4; Georgetown, 1; Oakley, 2; Society Hill, 1.

SCALOPUS AQUATICUS AUSTRALIS (Chapman).
Fiormwa Mote.
(P1. II, fig. 3; Pl. TIL, fies. 3, 3a: PL VEL fezs

Scalops aquaticus australis Chapman, Bul. Amer. Mus. Nat. Hist., vol. 5, p. 339,
December 22, 1893.

Scalopus aquoticus australis Elliot, Field Columb. Mus., publ. 105, zool. series, vol. 6,
p- 470, 1905.

Type locality—Gainesville, Alachua County, Florida.

Type specuomen.—No. 334%, Amer. Mus. Nat. Hist.; young adult,
sex unknown, skin and skull; collected May 4, 1891, by F. M. Chap-
man.

Geographic range.—Southeastern Georgia and the eastern portion
of peninsular Florida south to Lemon City.

General characters.—Size small (smaller than howelli); feet rela-
tively large; tail relatively short; skull short and broad (relatively
broader and higher than that of howell); teeth small.

Color—Full winter pelage: Back clove brown, mummy brown, or
dark fuscous, paler on wrists, slightly tinged with ochraceous-buff on
nose; underparts similar to back but showing more dark neutral gray
at base of hairs. Worn winter pelage: Variable; back clove brown or
mummy brown to vinaceous-buff, drab, or drab-gray with or without
pinkish buff or ochraceous-buff on nose; beneath much as above,
more grayish.

Skull.—Small, not angular, less constricted interorbitally than in
S. a. aquaticus, and not much depressed postorbitally; mastoids not
heavy; coronoid process and angle of mandible narrow and weak, the
former generally without mammiform process on posterior border;
horizontal ramus of mandible weak; teeth small.

Measurements.—Average of 6 adult males from type locality: Total
length, 145 (141-150); tail vertebre, 24.9 (22-28); hind foot, 18 (17-
19). Average of 10 adult females from type locality: 138.8 (131-148);
22.6 (21-26); 17.3 (16-18). Skull: Average of 6 skulls of adult males
from type locality: Greatest length, 31.7 (31.3-32); palatilar length,
12.9 (12.4-13.3); mastoidal breadth, 16.6 (16-16.8); interorbital

1 Collection Mus. Comp. Zool., Harvard College. 2 Collection Field Mus. Nat. Hist.
2 Collection Amer. Mus. Nat. Hist. 4 Collection Univ. Michigan Mus.

1915.] SCALOPUS AQUATICUS ANASTASA., 39

breadth, 7.2 (7.1-7.3); maxillary tooth row, 9.9 (9.6-10.3); mandib-
ular molar-premolar row, 9.6 (9.4-9.8). Average of 11 skulls of
adult females from type locality: Greatest length, 30.4 (29.5-30.8) ;
palatilar length, 12.4 (12-12.7); mastoidal breadth, 16.1 (15.6—16.4);
interorbital breadth, 7 (6.7-7.3); maxillary tooth row, 9.6 (9.4-9.7);
mandibular molar-premolar row, 9.4 (9.29.6).

Remarks.—Thesmall size of S.a. australis, together with its relatively
short, broad, and high skull, readily distinguish it from S. a. howell,
its nearest relative toward the north. Its characters are retained very
constantly throughout most of its geographic range. Skulls of speci-
mens from St. Catherines Island, Ga., are actually and relatively longer
than those of typical australis, and in this respect approach howellv;
in other characters the skulls do not differ essentially from those of
typical australis; in color these specimens are inseparable from aus-
tralis. A similar though less marked tendency toward howelli is
noticeable in specimens from Ossabaw Island and Barrington, Ga.
South of the type locality australis decreases slightly in size; speci-
mens from Oak Lodge and Lemon City, Fla., are of minimum size
and average very slightly smaller and darker than those from the type
region.

Specimens examined.—Total number, 115, as follows:

Florida: Canaveral, 1; East Micco, 2;1 Enterprise, 6;!)? Eustis, 1; Gaines-
ville (type locality), 25; Georgiana, 1; Indian River, 1; Jacksonville, 3; Lake
Harney, 3; Lake Worth, 1; Lemon City, 1; Lynne, 5; New Berlin, 6; 2)
Micanopy, 2; Oak Lodge (East Peninsula opposite Micco), 21;3 Orange Ham-
mock, Kissimmee River, 1; Saint Augustine, 1; Saint Charles Creek, 1;?
West Jupiter, 1.? ;

Georgia: Barrington, 2; Cumberland Island, 2; Hursmans Lake, 10;3 Mont-
gomery, 3;3 Nashville, 1; Ossabaw Island, 4;3 Pinetucky, 3;3 Saint Cath- .
erines Island, 5;? Saint Simons Island, 1; Sterling, 1.3

SCALOPUS AQUATICUS ANASTAS (Bangs).
ANASTASIA ISLAND MOLE.
(Plt, hes 4.)

Scalops anastase Bangs, Proc. Boston Soc. Nat. Hist., vol. 28, p. 212, 1898.
Scalops anastasiz (sic) Elliot, Field Columb. Mus., publ. 45, zool. series, vol. 2, p. 391,
1901.

Scalopus anastase Cory, Field Mus. Nat. Hist., publ. 153, zool. series, vol. 11, p. 438,
June, 1912.

Type locality.—Point Romo, Anastasia Island, Florida.

Type specvmen.—No. 7192, Mus. Comp. Zool., Harvard College,
Bangs collection; ¢ adult, skin and skull; collected February 16,
1897, by Outram Bangs.

Geographic range.—Anastasia Island, Fla.

1 Collection Amer. Mus. Nat. Hist. 2 Collection Field Mus. Nat. Hist. 3 Collection Mus. Comp. Zool.

40 NORTH AMERICAN FAUNA. [No. 38.

General characters.—Size of S. a. australis: Fore feetrelatively large,
nails long and heavy; skull short, massive; mastoids, heavy; rostrum
short.

Color.—Late winter pelage: Back golden sepia; face, chin, and
wrists bright zine orange; underparts Sudan brown. Worn winter
pelage: Paler than in late winter pelage with less golden and zinc
orange, and showing more light neutral gray of base of hairs.

Skull.—Small, short, and heavy, narrow through mastoids; mas-
toids heavy; rostrum short; mandible heavy; coronoid process and
angle of mandible short ; ascending ramus wide; superior notch shallow.

Measurements.—Average of 2 adult males from type locality: Total
length, 137.5 (134-141); tail vertebre, 21.8 (21-22.5); hind foot, 17.8
(17.5-18). Skull: Average of 2 skulls of adult males from type
locality: Greatest length, 31.5 (31.4-31.6); palatilar length, 12.8
(12.6-12.9) ; mastoidal breadth, 16.5 (16.4-16.6); interorbital breadth,
7.1 (7-7.1); maxillary tooth row, 10 (9.9-10); mandibular molar-
premolar row, 9.7 (9.6-9.8).

Remarks.—¥our of the eight moles ecarined from Anastasia Island
are topotypes in the Bangs collection and were collected by Outram
Bangs, February 12-16, 1895; the other four are in the Field Museum
and were collected by Thaddeus Surber at Espanita, Anastasia Island,
January 25-29, 1901. The two series are very unlike. The speci-
mens collected at Espanita have no orange or golden suffusfons and
can not be separated by skin characters from S. a. australis of the
mainland; the skulls, however, are shorter than those of typical
australis and have shorter rostra and slightly heavier mastoids, and
in these characters are more like those of S. a. anastase. In a letter,
in regard to these specimens, dated August 19, 1913, Surber states:

‘““Eispanita” was the home of a Mr. Middleton (a Georgian) located well toward the
southern end of the island about 15 miles south of St. Augustine, on the Matanzas
River side of island. Most of my work was done in the vicinity of his house, near
which I was camped. * * * In the flesh I could never detect any difference
between these island moles and those from the mainland. * * * There were no
towns nor villages on the island during my visit in 1901, so that I was forced to use this
designation (Espanita) for the locality. Mr. Bangs’s specimens came from the lower
end of the island, I believe, but two or three miles from where these were secured.

Bangs, in a letter to the writer October 23, 1913, referring to the
locality where Surber collected, states:

Tam quite sure * * * that that part of the island is quite different from where
I was. He got Peromyscus floridanus where he was. In the parts of the island I
worked I never saw it at all though I trapped hundreds of small mammals. In fact,
there was no country suitable toit. Where I caught my moles they were not common.
They were in the salt flats and low sandy stretches, where their long tunnels extended
about in the white sand and through the flats. There was very little vegetation
here except, of course, ‘‘sea oats.”’

1915.] SCALOPUS AQUATICUS PARVUS. 4]

All the specimens from Anastasia Island are here provisionally
called anastase, although additional material may result in some
change of decision. Were the specimens from Espanita older indi-
viduals, their skulls might be inseparable from those from Point
Romo. The topotype series was taken about two weeks later in the
year than that from Espanita. Two weeks is a short time for changes
in the pelage, yet such changes are apparently rapid in Scalopus, and
it would not be entirely impossible for the full winter pelage of the
mole at Espanita to develop before molting into a pelage similar to
that of the topotype series.

Specimens examined.—Total number, 8, as follows:

Florida: Espanita, Anastasia Island, 4;! Point Romo (type locality), Anastasia
Island, 4.?

SCALOPUS AQUATICUS PARVUS (Rhoads).
LirrLte Mote.
(Pl. II, fig. 5; Pl. III, figs. 4, 4a.)

Scalops parvus Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1894, p. 157, 1894.
[Scalops] [aquaticus] parvus Elliot, Field Columb. Mus., publ. 45, zool. series, vol. 2,
p. 390, 1901.

Type locality.—Tarpon Springs, Hillsboro County, Florida.

Type specimen.—No. 8468, Acad. Nat. Sci. Philadelphia (No. 1468,
Rhoads collection); ? adult, skin and skull; collected December 24,
1893, by W. S. Dickinson.

Geographic range.—Region north of Tampa Bay, in Hillsboro and
Pasco Counties, Fla.

General characters—Smallest of the genus; tail relatively short,
shorter than that of S. a. australis; color much like australis; skull
small, weak; rostrum very narrow (narrower than that of australis).

Color.—Full winter pelage: Much like corresponding pelage of S. a.
australis; back sepia or dark sepia, becoming pinkish cinnamon or
ochraceous-buff on nose; underparts slightly paler than back and
showing dark neutral gray of base of hairs, sometimes tinged with
cimnamon-brown.

Skuil.—Like that of S. a. australis but smaller, weaker, and with
rostrum actually and relatively narrower; teeth small.

Measurements.—Average of 3 adult males from Port Tampa City,
Fla.: Total length, 134.3 (131-136); tail vertebre, 19.7 (19-20.5);
hind foot, 16.8 (16.5-17). Skull: Average of 3 skulls of adult males
from Port Tampa City, Fla.: Greatest length, 30.6 (30.2-31.2);
palatilar length, 12.7 (12.5-12.9); mastoidal breadth, 15.7 (15.5-
15.8); interorbital breadth, 7.1 (7.1-7.3); maxillary tooth tow, 9.8

1 Collection Field Mus. Nat. Hist. 2 Collection Mus. Comp. Zool, Harvard College.

49, NORTH AMERICAN FAUNA. [No. 38.

(9.6-10); mandibular molar-premolar row, 9.7 (9.5-10). Skull of
type (probably a female, though sexed ‘‘male’”’ by collector): Great-
est length, 29.3; palatilar length, 11.9; mastoidal breadth, 15.1; inter-
orbital breadth, 6.6; maxillary tooth row, 9.2; mandibular molar-
premolar row, 9.2.

Remarks.—The original description of this form was based upon a
single specimen which was unique in collections for several years.
True,’ on account of insufficient material, placed parvus in question-
able synonymy under S. a. australis, where it has since remained.
The number of specimens now available from the type region is far
from satisfactory, but they show characters sufficient to warrant
their separation from australis. The type of S. a. parvus is in worn
pelage; True ? was inclined to believe on account of the appearance
of the skin and the peculiarly worn state of the teeth that the mole
had been kept in confinement; there appears no reason for such a
conclusion, however, the wear in the teeth and pelage of the type
being nothing which might not occur in the normal habitat of the
mole. Three males from Port Tampa City, Fla., while distinctly dif-
ferent from australis, do not appear to be the extreme type of parvus;
unfortunately, however, the only specimens from the immediate
vicinity of the type locality, except the type, are young.

Specimens examined.—Total number, 9, as follows:

Florida: Belleair, 2; Port Richey, 1; Seven Oaks, 1; 4 Port Tampa City, 3;°
Tarpon Springs (type locality), 2.8

SCALOPUS AQUATICUS MACHRINUS (Rafinesque).

PRAIRIE MOLE.
(BL. Ty eG PIS ie. 62)

Talpa machrina Rafinesque, Atlantic Journ., vol. 1, p. 61, 1832.

Talpa sericea Rafinesque, Atlantic Journ., vol. 1, p. 61, 1832. Type locality near
Nicholasville and Harrodsburg, Ky.

Scalops argentatus Audubon & Bachman, Journ. Acad. Nat. Sci. Philadelphia, vol. 8,
p. 292, 1842. Type locality southern Michigan.

Talpa Pennantii Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327,
1854. Type locality unknown.

Scalops aquaticus argentatus Coues, Bul. U. 8. Geol. & Geog. Surv. Terr., vol. 3, p.
633, 1877.

Scalops (aquaticus var.) argentatus Herrick, Mamm. of Minnesota, Geol. & Nat.
Hist. Surv. Minnesota. Bul. 7, p. 54, 1892.

Scalops aquaticus machrinus True, Proc. U. 8. Nat. Mus., vol. 19, p. 20, December 21,
1896.

Scalopus aquaticus machrinus Elliot, Field Columb. Mus., publ. 105, zool. series, vol.
6, p. 470, 1905.

1 True, F. W., Proc. U.S. Nat. Mus., vol. 19, p. 21, 1896. 4 Collection of Hartley H. T. Jackson.
2 True, loc. cit., p. 34, 1896. 5 Collection Mus. Comp. Zool, Harvard College.
3 Collection Acad. Nat. Sci. Philadelphia.

1915.] SCALOPUS AQUATICUS MACHRINUS. 43

Type locality — Lexington, Fayette County, Kentucky.

Type specimen.—None known to exist.

Geographic range-——EKastern Iowa, and east of the Mississippi
River west of the Appalachian Mountains from western Wisconsin,
northern Illinois, southern Michigan, southwestern Ontario (Pomt
Pelee), and northern Ohio, south to central Tennessee.

General characters.—Largest of the genus; tail relatively long;
color paler than that of S. a. aquaticus and usually more reddish
brown; skull flat, broad, heavy, and angular; rostrum massive;
teeth very large.

Oolor.—Full winter pelage: Above, sepia, mummy brown, or hair-
brown, occasionally showing pinkish buff or cinnamon-buff on nose;
underparts more grayish than back and usually tinged with Prout’s
brown or cinnamon-brown. Summer pelage: Slightly paler than
winter pelage and usually more grayish.

Skull.—Large, broad, angular, and massive; mastoids very
massive; rostrum large; coronoid process and angle of mandible
heavy, the former frequently with a distinct secondary process on
posterior margin; dentition very heavy.

Measuremenis.—Adult male from type locality: Total length, 208;
tail vertebre, 38; hind foot, 24. Adult male from Midway, Ky.:
190; 29; 22.5. Adult female from Midway, Ky.,175;27;22. Average
of 3 adult males from Warsaw, Ill., 199 (194-206), 35.7 (81-38).
Skull: Adult male from Midway, Ky.: Greatest length, 39.1;
palatilar length, 16.7; mastoidal breadth, 20.7; interorbital breadth,
8.2; maxillary tooth row, 12.2; mandibular molar-premolar row,
11.6. Skull of adult female from Midway, Ky.: Greatest length,
37.7; palatilar length, 16.2; mastoidal breadth, 19.2; interorbital
breadth, 7.7; maxillary tooth row, 12.1; mandibular molar-premolar
row, 11.7. Average of 5 skulls of adult males from Warsaw, IL:
Greatest length, 39.2 (88.8-39.5); palatilar length, 16.8 (16.6-17);
mastoidal breadth, 20.5 (20.3-20.7); interorbital breadth, 7.9
(7.7-8.1); maxillary tooth row, 12.4 (11.9-12.6); mandibular molar-
premolar row, 12 (11.5-12.2).

Remarks.—The name Talpa machrina Rafinesque ! and the earliest
description of this form were apparently lost to science from shortly
after they were published until True? resurrected them in 1896;
Rafinesque very accurately describes the form and distinctly indi-
cates “‘near Lexington’’ as the locality where: the species occurred.
In the same publication Rafinesque describes another mole, ‘‘found in
woods near Nicholasville and Harrodsburg,” under the name Talpa
sericea;* his second species is clearly the young of his Talpa machrina.
Audubon and Bachman‘ described a mole from southern Michigan,

1 Rafinesque, C. S., Atlantic Journ., vol. 1, p. 61, 1832.

2 True, F. W., Proc. U.S. Nat. Mus., vol. 19, p. 20, 1896.

3 Rafinesque, loc. cit., p. 62, 1832.

4 Audubon and Bachman, Jour. Acad. Nat. Sci. Philadelphia, vol. 8, p. 292, 1842.

44 NORTH AMERICAN FAUNA. [No. 38.

in 1842, under the name Scalops argentatus; although Michigan speci-
mens of Scalopus average slightly smaller than typical S. a. machrinus,
and possibly also slightly paler, the difference is insufficient for sub-
specific recognition. Le Conte! in 1854 described a mole under the
name of Talpa pennantir which he seemed to believe was the yellow
mole of Pennant; Pennant’s yellow mole, however, recetved Latin
binomial designation as early as 1777? and is purely a synonym of
S. a. aquaticus. Le Conte states in his description of T. pennantii
that it ‘differs from S. aquaticus in being much larger (equal in size
to Talpa europea) and in having much larger anterior feet.’”* It is
evident, therefore, that the specimen he had in hand was probably
S. a. machrinus.

This large mole retains its characters with remarkable regularity
throughout its range. Specimens from the Mississippi Basin in
western Wisconsin, as far north as Prescott, Pierce County, are
as large as specimens from the type region, if not slightly larger;
specimens from eastern lowa are indistinguishable from specimens
from Kentucky. The color tends to become more grayish in northern
and western Illinois and in western Wisconsin, possibly indicating an
approach toward S. a. machrinoides. Moles from Big Sandy and
Nashville, Tenn., have somewhat weaker dentition than typical
machrinus but do not differ essentially in other respects.

Specumens examined.—Total number, 159, as follows:

Dlinois: Alton, 2; Belleville, 1; Calhoun, 3;* Chicago, 9;° Fremont, 1; Hamilton,
1; Joliet, 1;° Olive Branch, 1;° Olney, 5; Ozark, 1;° Parkersburg, 2; Ravens-
wood, 1;® Richland County, 7; Riehl Station, 1; Rosiclare, 1;° ‘‘Southern
Tilinois,’’ 3; Union County, 2; Warsaw, 18.
Indiana: Denver, 2;+ Effner, Newton County, 1; Fort Wayne, 1; Lake Maxin-
kuckee, 3; Madison, 1; New Lebanon, 1; Waterloo, 2; West Baden, 2; Wheat-
land, 2.
Iowa: Hillsboro, 1; Knoxville, 1.°
Kentucky: Eubanks, 1; Lexington (type locality), 1; Midway, 3
Michigan: Ann Arbor, 2;+ Chelsea, 1;+ Denton, 2;* Flushing, 3;’ Greenfield, 1;8
Hoiland, 1;° Lansing, 1; Manchester, 1; Milan, 1;* Portage Lake, 11;
Saline, 1.
Ohio: Cleveland, 1; Fairfield County, 1; Madisonville, 3; Salem, 1.
Ontario: Point Pelee, 25.%, °
Tennessee: Big Sandy, 1; Clarksville, 1; Nashville, 3;° Tennessee River, 1.
Wisconsin: Camp Douglas, 2; Durand, 1; Fountain City, 4;° Maiden Rock, 3;°¢
Prescott, 1;° Wyalusing, ih 6
1 Le Conte, Joseph, Proc. Acad. Nat. Sci., Philadelphia, vol. 6, 1853, p. 327, 1854.
2 Talpa européa flavescens Erxleben (Syst. Reg. Anim., p. 118 )and Talpa flava Zimmermann (Spec. Zool.
Geog. p. 497).
3 Le Conte, loc. cit.
4 Collection Univ. Michigan Mus.
6 Collection Field Mus. Nat. Hist.
6 Collection Milwaukee Public Mus.
7 Collection of Hartley H. T. Jackson.

8 Collection Victoria Mem. Mus.
3 Collection of W. E. Saunders, London, Ontario.

15... SCALOPUS AQUATICUS MACHRINOIDES. A5

SCALOPUS AQUATICUS MACHRINOIDES Jackson.
Missourrt VALLEY Mote.
GBIETT figs: 7.)

Scalopus aquaticus machrinoides Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 19,
February 2, 1914.

Type locality —Manhattan, Riley County, Kansas.

Type specimen.—No. 169717, U. S. Nat. Mus., Biological Survey
collection; ¢ adult, skin and skull; collected June 1, 1910, by
W. E. Berg.

Geographic renge.—West of the Mississippi River, except eastern
Iowa, from central Minnesota, southeastern South Dakota, and the
eastern border of Nebraska, south through northeastern Kansas to
extreme northern Arkansas.

General characters.—Size large, exceeded only by S. a. machrinus;
color more grayish than machrinus; skull heavy, angular, smaller
than that of machrinus, with a shorter rostrum, and relatively smaller
inferior mandibular notch.

Color.— Winier pelage: Upperparts ranging from bister to clove
brown, becoming paler on face and wrists; underparts slightly paler
than back, and usually showing more slate color of base of hairs,
washed ventrally with raw umber or mummy brown. Summer pelage:
Upperparts light drab, drab, or wood brown, paler on face, nose, and
ankles; beneath slightly paler than back, more grayish.

Skull_—Most nearly like that of S. a. machrinus but smaller, with
a relatively shorter rostrum; ascending ramus of mandible not so
heavy as in machrinus, and inferior mandibular notch smaller;
rostrum short and broad; molariform dentition very heavy.

Measurements.—Average of 3 adult males from Elk River, Minn.:
Total length, 172 (168-178); tail vertebre, 30 (27-32); hind foot,
22.2 (22-22.5). Average of 3 adult females from Fort Leavenworth,
Kans., Bismarck, Mo., and Council Blufis, Iowa: 181 (180-182); 32
(31-33); 22.3 (22-23). Skull: Average of 3 skulls of adult males from
type locality: Greatest length, 37.1 (86.2-37.7); palatilar length, 15.3
(15.2-15.5); mastoidal breadth, 19.4 (19.3-19.5) ; interorbital breadth,
8 (7.9-8.1); maxillary tooth row, 12 (11.8-12.1); mandibular molar-
premolar row, 11.8 (11.5-12).

Remarks.—The Mississippi River separates the range of S. a.
machrinus from that of S. a. machrinoides except for a short distance
_ where the former extends into lowa. Specimens from St. Louis, Mo.,
are somewhat intermediate between the two forms, being larger than
typical machrinoides and having relatively longer rostra. Toward
the north (Elk River, Minn.), machrinoides decreases slightly in size;
along the western border of its range it intergrades with S. a. caryi,

46 NORTH AMERICAN FAUNA. [No. 38.

specimens from Lincoln, Nebr., being smaller and paler than typical
machrinoides, while those from Vermilion, 8. Dak., though large, are
distinctly paler than the typical form and have higher, more rounded
skulls. ‘Two specimens from Carthage, Mo., and one from Winslow,
Ark., are smaller than specimens from the type region, and have
smaller teeth and narrower rostra; the heavy mastoids and the
massive skulls are much as in machrinoides. A series from Greenway,
Ark., shows a very slight approach toward S. a. pulcher in color and
width of rostrum, but is easily referable to machrinoides.
Specimens examined.—Total number, 77, as follows:
Arkansas: Greenway, 7;1 Winslow, 1.
Iowa: Council Blufis, 1.
Kansas: Burlington, 1; Fort Leavenworth, 4; Manhattan (type locality), 6;
Neosho Falls, 2; Onaga, 2; Stillwater Creek, 1.
Minnesota: Elk River, 4; Fort Snelling, 2.
Missouri: Bismarck, 1; Carthage, 2;? Charleston, 1; Columbia, 11; Independ-
ence, 1; Marble Hill, 1; St. Louis, 7; Stotesbury, 9.3
Nebraska: Everett, 1; Fort Crook, 2;* Lancaster County, 1;° Lincoln, 5;°

Perch, 1.+*
South Dakota: Big Sioux River (at mouth), 1; Vermillion, 2.6

SCALOPUS AQUATICUS PULCHER Jackson.
ARKANSAS MOLE.
(Pl. TF, fie.(8:)

Scalopus aquaticus pulcher Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 19, Feb-
ruary 2, 1914.

Type localhity.— Delight, Pike County, Arkansas.

Type specomen.—No. 170698, U.S. Nat. Mus., Biological Survey
collection; ¢ adult, skin and skull; coliected January 20, 1911, by
W. G. Savage.

Geographic range—Humid lowland region of southern and eastern
Arkansas, southeastern Oklahoma, northwestern and central Louisi-
ana, and eastern Texas.

General characters About the size of S. a. aquaticus; hind foot
larger; skull larger than that of aquaticus, flatter, less swollen supra-
orbitally, wider interorbitally; interparietal wider than in aquaticus;
skull narrower through mastoids than that of S. a. machrinoides,
with narrower rostrum and smaller teeth. Slightly larger than Scalo-
pus ereus; skull relatively wider interorbitally, through mastoids
and through rostrum, flatter and more angular.

Color.—Full winter pelage: Back dark fuscous with many hairs
tipped with pearl gray, producing in places a slightly frosted appear-
ance; top of head mummy brown; nose cinnamon-brown; underparts

1 Collection Field Mus. Nat. Hist. 4 Collection Amer. Mus. Nat. Hist.

2 Collection of Hartley H. T. Jackson. 5 Collection Univ. Nebraska.
3 Collection Mus. Comp. Zool., Harvard College. 6 Collection Univ. South Dakota.

1915.] SCALOPUS AQUATICUS PULCHER. 47

sepia, showing much blackish plumbeous of base of hairs. Worn
winter pelage: Above fuscous or olive-brown, usually tinged on
head with mummy brown or Saccardo’s umber; underparts dark
neutral gray, becoming paler anteriorly, tinged with bister. Tresh
summer pelage: Upperparts olive-brown shading into coppery seal
brown on head and face; nose and wrists slightly tinged with ochra-
ceous-orange; beneath, bright cinnamon, becoming grayish poste-
riorly.

Skull—Somewhat similar to that of S. a. aquaticus, but slightly
larger, less angular, with flatter braincase, and averaging wider inter-
orbitally and through mastoids; postorbital region less depressed
than in aquaticus, zygomata heavier posteriorly, and horizontal! ramus
of mandible more arched ventrally; no distinct secondary process on
posterior margin of coronoid process, though central portion of pos-
terior border of ascending ramus is sometimes slightly expanded
posteriorly.

Measuremenis.—Average of 6 adult males from type locality:
Total length, 155.9 (153-170); tail vertebree, 25 (23-29); hind foot,
22 (21-23). Average of 3 adult females from type locality: 149.3 .
(146-156); 23.3 (20-25);°21 (20-22). Skull: Average of 7 skulls of
adult males from type locality: Greatest length, 35.6 (34.7-37.4);
palatilar length, 14.7 (14.2-15.4); mastoidal breadth, 18.2 (17.6-19.2);
interorbital breadth, 7.8 (7.5-8.2); maxillary tooth row, 11.3
(11-11.7); mandibular molar-premolar row, 11 (10.8-11.5). Aver-
age of 3 skulls of adult females from type locality: Greatest length,
34.3 (34.1-34.6); palatilar length, 14.38 (14.2-14.4); mastoidal
breadth, 18 (17.5-18.3); interorbital breadth, 7.7 (7.4-8); maxil-
lary tooth row, 10.9 (10.7-11); mandibular molar-premolar row,
10.7 (10.6-10.8).

Remarks.—Like other members of the genus, this beautiful mole is
subject to slight local variations throughout its range. Specimens
from Grand Coteau and Clarks, La., are somewhat smaller than the
typical form; one from Clarks is unusually grayish for the form, but
‘the much-worn fur and the partial molt may account for the color.
The very few not immature in a series from Mer Rouge, La., are
much alike in color, but are not so richly colored as most specimens
of S.a. pulcher; skulls of males, however, show the greatest individual
variation observed in any series from a single locality; two skulls
are indistinguishable from topotype skulls of pulcher; two others
are long, narrow, high, and rotund, and have narrow rostra; another
is short, broad, and flat, and has a broad rostrum; age may possibly
account for some of the variation, and incorrect sex determinations
may also be partly responsible. An old male from Lake City, Ark.,
is In some respects intermediate between pulcher and S. a. machri-
nodes; the teeth are large; the mastoids almost as heavy as those

48 NORTH AMERICAN FAUNA, : [No. 38.

of machrvnowdes; and the rostrum is slightly heavier than that of
typical pulcher, but in size and other characters it is very similar to
skulls from the type locality. ‘The specimen is in very fresh spring
pelage, a little of the winter fur remaining on nose and rump, and in
color it seems to be more nearly like machrinoides than pulcher. A
specimen from Fort Smith, Ark., and one from Wister, Okla., neither
adult, are referable to pulcher.
Specumens examined.—Total number, 67, as follows:
Arkansas: Camden, 1; Delight (type locality), 15; Lake City, 1; Fort Smith, 1;
Wilmot, 1.
Louisiana: Clarks, 1; Columbia, 1;! Grand Coteau, 1; Mer Rouge, 23; Natchi-
toches, 1; Shreveport, 3.

Oklahoma: Wister, 1. ?
Texas: Joaquin, 1; Sour Lake, 16.

SCALOPUS AQUATICUS CARYI Jackson.
NorTHERN Puatins MOLE.
(Pl. II, fig. 9.)

Scalopus aquaticus caryi Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 20, Feb-
ruary 2, 1914.

Type locality—Neligh, Antelope County, Nebraska.

Type specumen.—No. 116799, U.S. Nat. Mus., Biological Survey col-
lection; ¢ young adult, skin and skull; collected September 18, 1901,
by Merritt Cary.

Geographic range.—Arid and semiarid plains region of central and
western Nebraska, northeastern Colorado, and northwestern Kansas.

General characters.—Size medium; tail long; color palest of the
genus; most nearly like S. a. intermedius but much paler and lacking
ochraceous suffusions on nose and wrists; skull slightly shorter than
that of antermedius and relatively broader interorbitally.

Color.—Autumn pelage: Back light drab slightly tending toward
avellaneous, becoming paler on head and shading in some cases into
ivory yellow on nose; underparts much the same color as back, more
mixed with neutral gray and occasionally washed with Saccardo’s -
umber or cinnamon-brown.

Skuill.—Size moderate, short, broad, and rotund; rostrum short;
mandible heavy; dentition relatively heavy. Very similar to the
skull of S. a. intermedius but slightly shorter and relatively broader
interorbitally. Compared with the skull of S. a. pulcher that of
_ S. a. cary? is shorter and higher, wider interorbitally, and has weaker
mastoids, shorter, wider rostrum, and larger teeth; it is larger, much
higher, and more massive than the skull of S. greus, and is much
smaller, higher, and less angular than that of S. a. machrinoides,

1 Collection Field Mus. Nat. Hist. 2 Collection Carnegie Mus.

1915.] SCALOPUS AQUATICUS INTERMEDIUS. 49

though in extreme old age it tends to become angular and the brain-
case flattens.

Measuremenits.—Average of 3 males from type locality: Total
length, 159 (157-160); tail vertebrae, 32 (31.5-32.5); hind foot, 21.1
(20.3-22). Skull: Average of 3 skulls of males from type locality:
Greatest length, 34.2 (33.1-34.8); palatilar length, 14.2 (14.1-14.4);
mastoidal breadth, 17.8 (17-18.4); interorbital breadth, 8.1 (7.7-8.4) ;
maxillary tooth row, 11.2 (11.1-11.3); mandibular molar-premolar
row, 10.9 (10.8-11).

Remarks.—This subspecies can be distinguished from all other
moles by its pale color and lack of ochraceous suffusions. Its nearest
relationships are with intermedius with which it probably intergrades
in southern Kansas.

Specimens excamined.—Total number 16, as follows:

Colorado: Wray, 1.

Kansas: Long Island, 3.'

Nebraska: Kennedy, 1; Neligh (type locality), 4; Long Pine, 2;? Niobrara
River, 1;3 ‘‘Sandhills,’’ 1; Warbonnet Canyon, Sioux County, 3.°

SCALOPUS AQUATICUS INTERMEDIUS (Elliot).
SOUTHERN Puains Mote.

Scalops machrinus intermedius Elliot, Field Columb. Mus., publ. 37, zool. series,
vol. 1, p. 280, 1899.

[Scalops| [aquaticus] intermedius Elliot, Field Columb. Mus. publ. 45, zool. series,
vol. 2, p. 390, 1901.

Scalopus aquaticus intermedius Bailey, N. Am. Fauna No. 25, p. 207, October 24, 1905.

Type locality.— Alva, Woods County, Oklahoma.

Type specimen.—No. 6832, Field Mus. Nat. Hist.; ¢ adult, skin
and skull; collected February 23, 1899, by Thaddeus Surber.‘

Geographic range.-—Central and western Oklahoma and adjacent
parts of northern Texas.

General characters.—Size medium; tail relatively long; color pale
with ochraceous on nose and wrists, darker and more ochraceous
than caryi; skull relatively short, high and heavy, but not angular;
rostrum short; teeth large.

Color.— Winter pelage: Back light drab to drab tinged with buff-
pink; nose and wrists ochraceous-buff to zinc orange; underparts

1 Collection Amer. Mus. Nat. Hist.

2 Collection Stanford Univ.

3 Collection Univ. Nebraska.

4Mr. C. B. Cory, in a letter dated March 20, 1914, advises the writer that the specimen labeled as the
type of Scalops machrinus intermedius Elliot is No. 6829, Field Mus. Nat. Hist., collected at Alva, Okla.,
December 8, 1899, by Thaddeus Surber. In the original description Elliot designates no type specimen
by number, but describes an adult male collected at Alva, Okla., February 23, 1899, by Thaddeus Surber.
Since No. 6832, Field Mus. Nat. Hist. is the only specimen of the original series to which this description
can apply, it is necessary to consider it the type of intermedius.

90549°—15 4

50 NORTH AMERICAN FAUNA. [No. 38.

slightly paler than back, more silvery and showing less buff-pink;
base of hairs deep neutral gray. Young: More plumbeous than
adults, darker, less pink-buff tinge on back and little or no ochraceous-
buff on nose and wrists.

Skull.—Resembles that of S. a. caryi in nearly every particular,
but averages slightly larger, is usually slightly more depressed post-
orbitally, and has relatively heavier molariform dentition.

Measurements.—Average of 4 adult males from type locality:
Total length, 164.5 (160-169); tail vertebree, 28.8 (27-31); hind foot,
21.8 (21-22). Skull: Average of 4 skulls of adult males from type
locality: Greatest length, 35 (34.5-35.6); palatilar length, 14.7 (14.5-
14.9); mastoidal breadth, 18.3 (17.8-18.5); interorbital breadth, 8.1
(7.8-8.3); maxillary tooth row, 11.4 (10.9-11.7); mandibular molar-
premolar row, 11.1 (10.5-11.4).

Remarks.—The subspecies wntermedius can be distinguished from
its nearest congener, S. a. caryi, by its slightly darker color, and, in |
full adult pelage, by the ochraceous-buff or zinc orange on nose and
wrists; the skull of «ntermedius is generally longer than that of caryi.
Specimens from Mount Scott, Okla., are a little darker than typical
intermedius, and the skulls are slightly longer. Intergradation
between this subspecies and S. a. texanus apparently occurs in north-
central Texas. It seems probable that intermedius intergrades also
with S. a. pulcher in eastern Oklahoma, though specimens from Red
Fork, Okla., show no approach to pulcher. An alcoholic specimen
from Belknap, Tex., too young for positive identification, is provis-
ionally referred to intermedius.

Specumens examrned.—Total number, 22, as follows:

Oklahoma: Alva (type locality), 6; Dougherty, 1;! Fort Reno, 1; Mount Scott,
5; Red Fork, 3.
Texas: Belknap, 1; Lipscomb, 3; Mobeetie, 2.

SCALOPUS AQUATICUS TEXANUS (Allen).
Texas Mouse.
(PI. Il) fig. 10; Pl. LA, figs.;5,5a;; Pl. Vag hes)

Scalops argentatus texanus Allen, Bul. Amer. Mus. Nat. Hist., vol. 3, p. 221, April 29,
1891. :

Scalops tecanus Allen, Bul. Amer. Mus. Nat. Hist., vol. 5, p. 200, 1893.

Scalops aquaticus texanus True, Proc. U.S. Nat. Mus., vol. 19, p. 21, 1896.

[Scalops] [aquaticus] texensis (sic) Elliot, Field Columb. Mus., publ. 45, zool. series,
vol. 2, p. 390, 1901.

Scalopus aquaticus tecanus Bailey, N. Am. Fauna No. 25, p. 206, October 24, 1905.

Scalopus aquaticus texensis (sic) Elliot, Field Columb. Mus., publ. 105, zool. series,
vol. 6, p. 471, 1905.

1 Collection Field Mus. Nat. Hist.

1915.] SCALOPUS AQUATICUS TEXANUS. 51

Type locality—Rockport, Aransas County, Texas.

Type specimen.—No. 3438, Amer. Mus. Nat. Hist.; sex unknown;
skin and skull; collected September, 1887, by Wiliam Lloyd.

Geographic range.—Coast region of Texas from Matagorda Bay to
Cameron County, north in the interior to central and east-central
Texas.

General characters.—Size small; much smaller, darker, and more
brownish than intermedius; back generally much tinged with bronze,
and nose and wrists suffused with orange; skull small, flat, much
depressed postorbitally, and swollen supraorbitally; rostrum short;
teeth large. 3

Oolor.—Full winter pelage: Back Saccardo’s umber to mummy
brown with bronze tinge; nose and wrists zinc orange to xanthine
orange; underparts much like back, less bronze, occasionally tinged
with amber-brown; base of hairs dark mouse gray dorsally, becom-
ing slightly paler ventrally. Fresh summer pelage: Slightly paler
than winter pelage, less brown, with less orange on nose and wrists;
underparts distinctly more grayish than in winter pelage.

Skull.—Small, flat, constricted interorbitally, much depressed
postorbitally, and swollen supraorbitally; rostrum short; mandible
relatively heavy; teeth relatively large. The skull of S. a. texanus
is very much smaller than that of S. a. ontermedius or S. a. pulcher;
about the size of that of S. a. australis but differs from it in shape and
in having much heavier mandibles and dentition.

Measurements.—Average of 8 adult males from type locality:
Total length, 138.5 (128-152); tail vertebra, 23.8 (21-26); hind
foot, 17.2 (15.5-18). Average of 4 adult females from type locality:
133.8 (130-137); 22.3 (20-24); 16.3 (15-17). Skull: Average of 9
skulls of adult males from type locality: Greatest length, 31 (30-82);
palatilar length, 13 (12.7-13.7); mastoidal breadth, 16.6 (16.1-17);
interorbital breadth, 6.8 (6.6-7.1); maxillary tooth row, 10.3
(10-10.6); mandibular molar-premolar row, 10.1 (9.7-10.3). Aver-
age of 4 skulls of adult females from type locality: Greatest length,
29.8 (29.3-30.2); palatilar length, 12.6 (12.3-12.8); mastoidal
breadth, 16.2 (15.9-16.5); interorbital breadth, 6.9 (6.7-7.1); max-
lary tooth row, 9.8 (9.7-9.9); mandibular molar-premolar row,
9.7 (9.5-9.9).

Remarks.—Local variation in the genus seems to reach its maxi-
mum in S. a. texanus, there being no two localities from which speci-
mens of this form have been examined but exhibit differences either
in skin or cranial characters, or in both; there is, however, very little
individual variation in a series from any one locality. Unfortunately,
no considerable number of specimens are available except from Rock-
port, Tex., and until extensive series of specimens and careful habitat

BY NORTH AMERICAN FAUNA. . [No. 38.

studies are obtained from each of many localities it will be impossible
to determine the extent and meaning of these variations. The large
series from the type locality is remarkably uniform in characters
except for seasonal color variations. Skulls from Corpus Christi are
slightly larger, higher through the braincase, and broader through the
rostrum than typical tezanus, as are also two believed to have been
collected at Brownsville, Tex.; specimens from Padre Island are
very slightly longer than those from Rockport; one from Santa Rosa
is high and weak and has a rather short rostrum. The color of the
Santa Rosa specimen is much paler than that of typical tezanus due
probably to the peculiar condition of the pelage. Specimens from
Mason have large skulls and pale color and in these respects seem to
approach S. a. intermedius; the skulls, however, are flat and have
narrow rostra. A skin without skull from Waco is inclined toward
intermedius in color, but is much too small for that form and has
been provisionally called texanus; another skin, in very soiled pelage,
from Longpoint, Tex., does not differ from typical texanus in size or
color.
Specimens examrned.—Total number, 42, as follows:
Texas: Brownsville, 2; Corpus Christi, 2; Longpoint, 1; Mason, 3; Padre Island,
3; Rockport (type locality), 25; San Antonio, 4; Santa Rosa, 1; Waco, 1.!

SCALOPUS AREUS (Bangs).
CoprpERY MOLE.
(PL. T1, fig. 12: Pl. IMM, fies.7,-72; Ble V Eos.)

Scalops texanus xreus Bangs, Proc. Biol. Soc. Washington, vol. 10, p. 138, December 28,
1896.

[Scalops] xreus Elliot, Field Columb. Mus., publ. 45, zool. series, vol. 2, p. 390, 1901.

Scalops aquaticus ereus Miller & Rehn, Proc. Boston Soc. Nat. Hist., vol. 30, p. 250,
December, 1901.

Scalopus xreus Elliot, Field Columb. Mus., publ. 105, zool. series, vol. 6, p. 471, 1905.

Scalopus aquaticus xreus Miller, U. 8. Nat. Mus., Bul. 79, p. 8, December 31, 1912.

Type locality.—Stilweli, Payne County, Oklahoma.

Type specomen.—No. 5475, Mus. Comp. Zool., Harvard College,
Bangs collection; @ adult, skin and skull; collected August 13,
1896, by Thaddeus Surber.

Geographic range.—Known only from type locality.

General characters.—Size medium; differs from all other forms of
Scalopus in its color, a rich coppery snuff brown; skull weak and
narrow.

Color.—Type (female), in late summer pelage: Upperparts rich cop-
pery snuff brown; underparts less coppery than above, more grayish.

1 Collection Acad. Nat. Sci. Philadelphia,

1915.] SCALOPUS INFLATUS. 58

Skull. Mauch smaller, weaker, and narrower than that of S. a.
machrinoides, S. a. pulcher, or S. a. wntermedius; larger than that
of S. a. texanus, relatively longer, less depressed postorbitally and less
swollen supraorbitally, with longer, narrower rostrum; teeth small.

Measuremenis.—Type (adult female): Total length, 154; tail
vertebra, 24; hind foot, 19. Skull (of type): Greatest length, 33.2;
palatilar leneth, 14; mastoidal breadth, 17; iterorbital breadth, 7.5;
maxillary tooth row, 10.6; mandibular molar-premolar row, 10.5.

Remarks.—The status and relationships of this form are unknown.
The type and only specimen in collections was taken August 13,
1896; the rich coppery color may be due in part to a deadening of
the summer fur preceding molt and if such proves to be the case
the coppery color will not be of diagnostic value. The skull is weak,
narrow, moderately high and rounded, and has a long, narrow
rostrum; the teeth are much worn, the sutures closed, and from all
appearances the animal was adult; it was sexed female by the
collector.

Specumen examined.—One, the type.

SCALOPUS INFLATUS Jackson.
TAMAULIPAS MOLE.
(Pl. Il, fig. 11; P1.-111, figs. 6, 6a; Pl. VI, fig. 8.)
Scalopus inflatus Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 21, February 2, 1914.

Type locality.—State of Tamaulipas, Mexico (45 miles from Browns-
ville, Texas).

Type specvmen.—No. 52709, U. S. Nat. Mus., Biological Survey
collection; young adult, sex unknown; skin and imperfect skull; col-
lected in 1892 by F. B. Armstrong.

Geographic range—Known only from type locality.

General characters.—Size small, larger than S. a. teranus; skull
broad, high, and arched, much inflated in prelachrymal region;
rostrum broad, truncate; zygomata heavy.

Color.—Back between wood brown and drab, becoming ochraceous-
buff on cheeks; general tone of underparts much as on back but
more mixed with mouse gray.

Skull.—Size medium, high and arched; prelachrymal region much
swollen; audital bulle high and well defined; zygomata heavy;
posterior edge of lachrymal foramen meets zygoma at nearly right
angle; rostrum broad; mandible heavy, horizontal ramus much
arched ventrally; outer groove in third upper premolar pronounced;
first lower premolar small and inconspicuous.

Measurements.—Type (from dry skin, foot relaxed): Tail vertebre,
18; hind foot, 16.5. Skull (of type): Palatilar length, 13.4; mastoidal
breadth, 17; interorbital breadth, 7.1; maxillary tooth row, 11.1;
mandibular molar-premolar row, 11.

54 NORTH AMERICAN FAUNA. [No. 38.

Remarks.—The only-known specimen of this form is imperfect; it
lacks complete data, the pelage is ragged and apparently faded,
and the posterior portion of the braincase is broken away. Aside
from its many other distinctive characters, Scalopus inflatus can easily
be separated from all other members of the genus by the peculiarly

3

B2013-103

Fig. 7.—Geographic range of the species and subspe-

cies of Scapanus except S. orarius (see fig. 12).

1. S. townsendii. 6. S. 1. minusculus.
2. S. latimanus latimanus. 7. S. 1. dilatus.

3. S. 1. occuiltus. 8. S. 1. alpinus.

4. 8.1. grinnelli. 9. S. anthonyi.

5. S. 1. sericatus.

inflated prelachrymal region of
theskull. It was at first thought
that this inflation was due to
parasites; but the bilateral sym-
metry of the inflations, the fact
that no apparent work of para-
sites has been noticed in this re-
gion of the skull in the many
hundred specimens of Scalopus
examined, and, further, the fact
that a very slight and inconspic-
uous tendency toward prelachry-
mal inflation is present in all
members of the genus, leads to
the belief that the inflation in
this species is an extreme accen-
tuation of normal development.

Specimen ecamined.—One, the

type.
Genus SCAPANUS Pomel.

Scapanus Pomel, Archives Sci. Phys-
iques et Nat., tome 9, p. 247, Novem-
ber, 1848.

Scaphanus Herrick, Geol. and Nat. Hist.
Surv., Minnesota, Bul. 7, p. 55, 1892.

Scapasius Beddard, Cambridge Nat.
Hist., vol. 10, p. 518, 1902.

Type species.—Scalops town-
sendw Bachman.

Geographic range. — South-
western British Columbia (Fraser
Riverregion), western and south-
ern Washington, western Ore-
gon, extreme western Nevada,

California (except the southeastern desert region), south to San Pedro
Martir Mountains, Lower California (figs. 7 and 12).

External characters —Body robust, depressed; tail short, round,
thick and fleshy, tapering apically and slightly constricted proxi-
mally, indistinctly annulated, scantily haired with coarse hairs

1915.] GENUS SCAPANUS. 55

(fig. 8). Head conoidal, depressed. Nose elongated into a snout
(shorter and less truncate than in Scalopus), the apical portion naked
to line of anterior edge of nasals; nostrils superior, .crescentic, with
concavities turned in laterally (fig. 9). Eyes minute, concealed in
fur. Auricular orifice small. Legs short and stout.
Feet large, fleshy, scantily haired above, naked below.
Fore feet handlike, the palms as broad as long (fig.
10). Soles of hind feet with one to three (usually
two) distinct tubercles. Fore toes and hind toes not
webbed. Claws of fore feet broad, flat, and heavy;
those of hind feet relatively short and weak (fig. 11).
Fur dense, soft, silky, the hairs nearly equal in length,
producing a velvetlike pelage. Mamma, 8: latero-
pectoral, 2-2; latero-abdominal, 1-1; mguinal, 1-1.

Skeletal characters.—Clavicle short and heavy, about
three-fourths as broad as long, distinctly notched on
the inferior border, not penetrated by a foramen.
Humerus heavy, about three-fourths as broad as long. eee
Pelvis narrow, bones of opposite sides scarcely touch- py¢. grail of Sew
ing under acetabula; two osseous bridges, each bifur- —-2#¥s latimanus Tat-

é c : s imanus (X 14). No.
cate posteriorly, connect sacral vertebre with ischium — jos958, U. 8. Nat.
and produce six foramina or openings, alarge one in =‘Mus.; from Colma,
each of the anterior angles formed by median lines of as Oe
acetabula and sacral vertebre and two smaller ones in each of the
posterior angles. Superior surface of last sacral vertebra without
longitudinal process. Os falciforme relatively broad and long, reach-
ing proximal end of terminal phalange of first digit, gently incurved,
not much tapering distally.

Skull conoidal, flat, with relatively broad braincase, and slightly
constricted interorbitally. Mas-
toids moderately heavy, not prom-
inent. Interpariectal large, some-
what rectangular, breadth (antero-
posterior diameter) about one-
third the length, slightly convex
anteriorly and concave posteriorly
with a posterior median projection.
Bate Frontal region flat, not much slop-
Fig. 9.—Snout of 8.1. latimanus (X13). Individ- 1 2 Bt

ual referred to in fig. 8. ing; frontal sinuses somewhat
swollen. Rostrum moderately long; anterior ends of premaxille
slightly thickened and extending beyond nasals, forming a notch
(acute to truncate) anterior to nasals. Anterior nares opening for-
ward. Zygomata moderately long and heavy, rather sharply
curved inward anteriorly, the posterior end attached about medially
on squamosal. Foramen magnum oval,ofmoderatesize. Infraorbital

56 NORTH AMERICAN FAUNA. [No. 38.

foramen relatively small (larger than in Scalopus), the plate forming
its outer wall moderately broad. . Audital bullae complete, depressed;
auditory meatus short (more developed than in Scalopus). External
pterygoid region moderately inflated both posteriorly and anteriorly.
Mesopterygoid space moderately broad and relatively long, the sides
nearly straight and slightly converging posteriorly. Palate mod-
erately elongate, termin-
ating at a distance pos-
terior to the last molar
about equal to half the
diameter of that tooth;
posterior border of palate
truncate or slightly emar-
ginate, usually with a
shght median notch. An-
terior palatine foramina
poie-io3. «Moderate, oval to ellip-
Fic. 10.—Fore foot of 8.1. latimanus (X 14). Individual referred tical-oval; posterior pala-
to in fig. 8. tine foramina small (the
first pair larger than the second), round to oval. Horizontal ramus of
mandible moderately heavy, somewhat curved upward at posterior end,
nearly straight at anterior end; coronoid moderately elongate, quad-
rate, truncate, directed almost perpendicular to median line of horizon-
tal ramus; angle of mandible large and heavy (relatively longer than
in Scalopus); inferior mandibular notch
large, subcircular, relatively broad and deep.
Dental characters —First upper incisor
long and broad, convex anteriorly, flat pos-
teriorly ; second and third upper incisors and
upper canine simple, conical, of moderate
size, subequal, about two-thirds as long as
first incisor. First, second, and third upper
premolars similar to second and third inci-
sors, the third premolar usually with a small
postero-basal cusp; fourth premolar much "iia ee ne
larger than third, more cuspidate, with a ferred toin fig. 8.
large posterior cusp, usually with an intero-basal cusplike process,
and frequently with a more or less developed antero-basal cusplike
process. Upper molars W-shaped in transverse section, with an
antero-internal V-shaped cusplike shelf (not lobed); first and second
molars subequal, the third much smaller.
First seven lower teeth (incisors, canine, and first, second, and
third premolars) small, conical, the first and third incisors smaller
than the other teeth, which are subequal; premolars each with a

1915.] GENUS SCAPANUS® 57

slight cusplike process posteriorly; fourth premolar much larger,
more cuspidate. Lower molars M-shaped in transverse section, the
antero-internal cusp bilobed; first molar with a postero-internal
basal accessory cusp; second molar with a postero- and an antero-
internal basal accessory cusp; third molar with an antero-internal
basal accessory cusp; first and second molars subequal, the third
much smaller. Dentition: 1. 3; c.+; pm. 4;! m. 3; total 44.

Key to Species and Subspecies of Scapanus.
[Based upon specimens of adult animals in fresh pelage.]

a!. Unicuspid teeth evenly spaced and not crowded; rostrum relatively long and nar-
row; color dark, almost black (except in Scapanus orartus schefferi).
b!. Total length more than 200 mm.; greatest length of skull more than 40 mm.;
sublachrymal-maxillary ridge distinct.......... Scapanus townsendit (p. 58).
b?, Total length less than 200 mm.; greatest length of skull less than 40 mm.; sub-
lachrymal-maxillary ridge not much developed.
c!, Color darker; rostrum narrow; interorbital breadth of skull 8.2 mm. or less,
Scapanus orarius orartus (p. 61).
e?. Color paler; rostrum relatively broad; interorbital breadth of skull 8.2 mm.
TE EROS Sa ge thee pepe ee ror Scapanus orarius scheffert (p. 63).
a?. Unicuspid teeth usually crowded and not evenly spaced; rostrum relatively short
and broad; color usually brown or gray, seldom almost black.
61. Total length usually less than 165 mm.; greatest length of skull 33.5 mm. or less,
c!, Greatest length of skull less than 31 mm.; premolars 3,
Scapanus anthonyi (p. 75).
c?, Greatest length of skull 31 mm. or more; premolars $.
d'. Interorbital constriction more than 7.6 mm.; breadth of skull across mas-
toids more than 16.2 mm............ Scapanus latimanus grinnelli (p. 69).
d?. Interorbital constriction 7.6 mm. or less; breadth of skull across mastoids
16.2 mm. or less.
e!. Inferior mandibular notch deep; geographic range south of latitude

S17 5a es Re a St ee Scapanus latimanus occultus (p. 68).

e?, Inferior mandibular notch shallow; geographic range north of latitude

RIN tec e cto. © : stan latimanus minusculus (p. 72).

b?. Total length Tene = more Siben 165. mm.; greatest length of skull more than
33.5 mm.

c!, Geographic range, California west of Sacramento and San Joaquin Valleys,

including also Klamath Canyon...... Scapanus latimanus latimanus (p. 64).

c?. Geographic range, Oregon and California east of Sacramento and San Joaquin
Valleys, except Klamath Canyon.
d'. Color dark; interorbital constriction less than 7.6 mm.,
Scapanus latimanus sericatus (p.71).
d?. Color pale; interorbital constriction more than 7.6 mm.
e!. Greatest length of skull less than 36 mm.,
Scapanus latimanus dilatus (p. 72).
e”. Greatest length of skull more than 36 mm.,
Scapanus latimanus alpinus (p. 75).

Pm. 3in the type and only specimen of Scapanus anthonyi.

58 NORPH AMERICAN FAUNA. [No. 38.

Descriptions of Species and Subspecies of Scapanus.

SCAPANUS TOWNSENDII (Bachman).
TOWNSEND’S MOLE.
(PLAIV, fig. 1;.PlV jp figs a, la; Pl. Vile)

Scalops canadensis Richardson, Fauna Boreali-Amer., part 1, p. 9, 1829. (Not of
Desmarest or Harlan.)

Scalops Townsendii Bachman, Journ. Acad. Nat. Sci., Philadelphia, vol. 8, part 1,
p. 58, 1839.

Scalops Townsendi Bachman, Boston Journ. Nat. Hist., vol. 4, p. 31, January, 1842.

Scapanus Towsendii (sic) Pomel. Archiv. Sci. Physiques et Nat., vol. 9, p. 247, 1848.

Scalops metallescens Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 242,
1854. (Nomen nudum.)

Scalops zneus Cassin, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 299, 1854.
Type locality, Oregon.

| Talpa] znea Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327, 1854.

Talpa Townsendii Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327,
1854.

Talpa teniata Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327, 1854.
Type locality, banks of Columbia River.

Sclapanus] Townsendi Peters, Monatsber. Konig. Preuss. Akad. Wissensch., Berlin,
1863, p. 656, 1864.

Scapanus Townsendti True, Proc. U. 8. Nat. Mus., vol. 7, p. 607, 1885.

Type locality—‘ Banks of the Columbia River.” Probably from’
the vicinity of Fort Vancouver, Clarke County, Washington, which
it seems well to consider the type locality.

Type specumen.—Cotype, No. 449, Acad. Nat. Sci. Philadelphia;
collected May 9, 1835,1 by J. K. Townsend. ;

Geographic range.—Extreme northwestern California, Oregon, and
Washington west of the Cascade. Mountains.

General characters—Size largest of the genus; color dark, almost
black; skull large, mastoids relatively heavy, rostrum long; uni-
cuspid teeth evenly spaced and not crowded. The young of S.
townsendia in superficial skin characters often resemble adults of S.
orarius, but are easily separated from the latter by their large fore
feet, with thick, heavy claws.

Color.— Winter pelage: Upperparts blackish brown, fuscous-black,
~ sooty black, to almost black, usually with a purplish sheen; under-
parts very slightly paler than the back and frequently stained with
brown. Summer pelage: Much like winter pelage, but very slightly
paler, with purplish sheen more pronounced.

Skull—tLarge (greatest length of smallest skulls more than 40
mm.), flat, and angular; mastoids angular and rather heavy; inter-
parietal relatively narrower antero-posteriorly than in S. latima-
nus; slight but distinct sagittal crest between anterior portions

1 The date is not on the labels now attached to the specimen, but Bachman (loc. cit.) states it was col-
lected May 9, 1835.

1915.] SCAPANUS TOWNSENDII. 59

of parietals in adults; zygomata heavy; rostrum long and rela-
tively narrow; sublachrymal-maxillary ridge well developed; den-
tition heavy; unicuspid teeth, both maxillary and mandibular,
evenly spaced and not crowded asin latumanus; mandible relatively
weaker than in latimanus.

Measurements.—Average of 3 young adult females from vicinity
of Portland, Oreg.: Total length, 206 (195-222); tail vertebre, 48.3
(45-51); hind foot, 26.3 (24-28). Average of 7 adult males from
Ferndale, Humboldt County, Cal.: 224.1 (217-237); 41.1 (84-50);
26.7 (26-27). Average of 4 adult females from Ferndale, Cal.: 208
(202-210); 41.8 (87-46); 26.8 (26-27). Skull: Average of 10 skulls
of adult males from Puyailup, Wash.: Greatest length, 43.6 (42.3-
44.6); palatilar length, 18.1 (17.4-18.8); mastoidal breadth, 20.8
(20.3-21.4); interorbital breadth, 9.2 (9.1-9.4); maxillary tooth row,
14.1 (13.7-14.4); mandibular molar-premolar row, 13.8 (13.5-14.4).
Average of 10 skulls of adult females from Puyallup, Wash.: Greatest
length, 42.3 (41.5-44.1); palatilar length, 17.6 (16.9-18.1); mastoidal
breadth, 19.9 (19.3-20.4); interorbital breadth, 8.9 (8.4-9.2); maxil-
lary tooth row, 13.9 (13.5-14.4); mandibular molar-premolar row,
13.6 (13.2-14). Average of 3 skulls of young adult females from the
vicinity of Portland, Oreg.: Greatest length, 41.8 (41.2-42.1); pala-
tilar length, 17.6 (17.3-18); mastoidal breadth, 19.8 (19.4-20); inter-
orbital breadth, 9 (8.7-9.1); maxillary tooth row, 13.4 (13.3-13.5);
mandibular molar-premolar row, 13.2 (13.1-13.3). Average of 7
skulls of adult males from Ferndale, Humboldt County, Cal.: Great-
est length, 42.9 (42.3-44.2); palatilar length, 18.3 (17.7-18.7); mas-
toidal breadth, 20.9 (20-21.8); mterorbital breadth, 9 (8.7-9.5);
maxillary tooth row, 13.9 (13.5-142); mandibular molar-premolar
row, 13.7 (13.2-14.1). Average of 4 skulls of adult females from
Ferndale, Cal.: Greatest length, 41.5 (41.2-41.9); palatilar length,
17.9 (17.8-18); mastoidal breadth, 20 (19.9-20.2); interorbital
~ breadth, 8.9 (8.7-9.1); maxillary tooth row, 13.7 (13.5-13.9); man-
dibular molar-premolar row, 13.3 (13.2-13.5).

Remarks.—The presence of moles in the Pacific northwest was
known. to some of the early explorers, but the first one described was
by Richardson, who, though very accurately describing the animal
now known as Scapanus townsendi, referred his specimens to the com-
mon mole of eastern United States, then known as Scalops canadensis
Desmarest. It was not until ten years later that the species was
named, when Bachman? published his description based upon two
specimens. One specimen was a normally colored individual re-
ceived from Nuttall from a locality not stated; the other was col-
lected by Townsend and according to Bachman (lec. cit.) was

1 Richardson, J., Fauna Boreali-Amer., part 1, pp. 9-12, 1829.
2 Bachman, J., Journ. Acad. Nat. Sci. Philadelphia, vol. 8, part 1, pp. 58-60, 1839.

60 NORTH AMERICAN FAUNA, [No. 88. ”

labeled ‘‘Banks of the Columbia River, May 9, 1835.” The latter
specimen is a partial albino, having a narrow, irregular white streak
extending from chin to abdomen, and another from forehead to snout;
this specimen (No. 449) is now in the Academy of Natural Sciences
of Philadelphia, where sometime in later years it was marked ‘‘Type
of Scalops townsendu.” It is not the type, smce Bachman’s descrip-
tion was based in part if not primarily on the specimen submitted
by Nuttall; it may well be considered a cotype, however, since
Bachman does not designate a type; that he considered the two
specimens one and the same species is evident in his remark: ‘‘I
subsequently received from Mr. Townsend another specimen, a little
larger in size, which I presume to be a mere variety, although very
singularly marked” (loc. cit., p. 58). This same abnormal specimen
became, in 1854, the type of Talpa teniata Le Conte. Cassin ex-
hibited and described a mole which he called “Scalops metallescens”’
before the Philadelphia Academy in 1853, but in the published
account! of his talk and exhibition, which appeared the next year,
the name “Scalops metallescens” occurs without any description.
Subsequently, however, Cassin? described the animal under the
name “Scalops xneus;” the type* is now in the United States
National Museum. Cassin states: ‘‘In its dentition and otherwise it
is a strict congener of Scalops townsend, but is much smaller and of
a different color. Its black claws are especially remarkable, and dis-
tinguish it from all other species of the genus” (loc. cit.). Unfor-
tunately the skull has been lost, but the skin seems to show that it
is of a rather young specimen of Scapanus townsendw which has been |
shrunk and discolored by some chemical, possibly corrosive subli-
mate. The general tone of coler of the back is between Brussels
brown and Prout’s brown; the underparts are mostly buckthorn
brown, and on chin and ankles is a suspicious tinge of sulphine
yellow. The claws are heavy as in townsendw and do not indicate
specific relationship with S. orarius; both claws and soles of the feet
are black, which might readily be accounted for by the presence of
mercuric sulphid from the combination of carbon bisulphid and
corrosive sublimate used in preserving specimens.

Townsend’s mole, though showing considerable individual vari-
ation in size and in proportions and shape of skull, is subject to very
little geographic variation. In a large series of skulls from Puyallup,
Wash., are four which have supernumerary premolars. Three of
these have each an extra premolar between the second and third
premolars of the right mandible; the other specimen has a super-
numerary tooth between the second and third premolars of the left

1 Cassin, J., Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 242, 1854.

2 Cassin, loc. cit., p. 299, 1854.
8 No. 3725 ,U. S. Nat. Mus., skin without skull; collected in Oregon, by ‘‘U. 8. Exploring Expedition.”

1915.] SCAPANUS ORARIUS ORARIUS. 61

mandible, and a minute extra tooth between the third and fourth

right lower premolars. This is interesting siace Scapanus normally

has the theoretically complete mammalian dentition of 44 teeth.
Specimens examined.—Total number, 203, as follows:

California: Cresent City, 6; Ferndale, 11;' Smith River, 2.

Oregon: Beaverton, 3; Coquille, 1; Drain, 1; Goldbeach, 2; Grants Pass, 1;
Netarts, 1;? Oregon City, 3; Portland, 24; 2,%.* Salem, 3; Seaton, 1; ‘‘U.S.
Exploring Expedition,’”’ 1; Wells, 1.

Washington: Columbia River (type locality), 1;° Hot Springs Trail, Olympic
Mountains, 1;° Lake Cushman, 2; La Push, 1; Puyallup, 108;7 Renton,
1;! Roy, 1; Sauk, 1; Seattle, 1; Skykomish, 1; South Bend, 5; Steilacoom,
3; Tacoma, 1;> Tenino, 3; Vanccuver, 11; Vancouver Barracks (probably
exact type locality), 1.

SCAPANUS ORARIUS ORARIUS True.
Coast MOLE.
GRIN fies 2 PloNi fies. 2, 2a; Pl. Vi, nes 11.)
Scapanus orarius True, Proc. U. 8. Nat. Mus., vol 19, p. 52, December 21, 1896.

Type locality —Shoalwater Bay, Pacific County, Washington.

Type specimen.—No. 3383;, U. S. Nat. Mus.; 9 young adult, skin
and skuli (posterior portion of braincase broken and incomplete);
collected August 30, 1855, by J. G. Cooper.

Geographic range-——Humid coast region of northern California
(north of Mendocino), Oregon, and Washington.

General characters.—Size medium; color dark. The subspecies
orarivus is somewhat smaller than S.J. latimanus, with relatively smaller
fore feet, and slenderer claws; darker than latimanus. Similar in
color to S. townsend, but very much smaller, with actually and rela-
tively smaller feet and claws. The skull of S. o. orarius can always
be easily distinguished from that of townsendit by its much smaller
size, without reference to any other characters; from that of latumanus
it differs in its evenly spaced and uncrowded unicuspid teeth, its very
narrow rostrum, undeveloped and indistinct sublachrymal-maxillary
ridge, and very weak mandible; teeth, particularly the first incisors,
smaller than in latimanus.

Color.—General. tone much the same as in S. townsendit. Winter
pelage: Upperparts fuscous-black, chetura black, blackish brown, to
nearly black; underparts slightly paler and more grayish. Summer
pelage: Much like winter pelage but usually more brownish.

Skull—sSize medium (average greatest length of skulls of adult
males about 34 to 35 mm.); mastoid region weak; sublachrymal-
maxulary ridge only slightly developed; rostrum very narrow;

1 Collection Mus. Vert. Zool., Univ. California. 5 Collection Acad. Nat. Sci. Philadelphia.
2 Collection Oregon State Game Comm. 6 Collection Field Mus. Nat. Hist.
3 Collection of H. E. Anthony, New York City. 7 One hundred, skulls only; two skeletons,

* Collection of 8, G. Jewett, Portland, Oreg.

62 NORTH AMERICAN FAUNA. [No. 38.

horizontal ramus of mandible narrow and weak; teeth, especially first
incisors, small.

Measurements.—Average of 3 adult males from HKureka, Humboldt
County, Cal.: Total length, 167 (163-175); tail vertebree, 33.7 (31-35);
hind foot, 20.7 (20-22). Skull: Average of 2 skulls of adult males
from Ferndale, Humboldt County, Cal.: Greatest length, 34.3 (33.5-
35); palatilar length, 13.5 (13.8-13.7); mastoidal breadth, 16.4
eR pS (16.3-16.4); interorbital breadth,
8.1 (8-8.1); maxillary tooth row,
10.7 (10.5-10.9); mandibular mo-
lar-premolar row, 10.4 (10.3-10.5).

Remarks.—Judged by the speci-
mens examined, Scapanus orarius
(including the subspecies scheffert)
has a more extensive geographic
range than S. townsend; it occurs
both farther north and _ farther
south, as well as much farther east
(fig. 12). The ecological relation-
ship of the two forms is unknown;
its solution would involve most
valuable and interesting research
for some student favorably sit-
uated.

The few specimens of S. 0. ora-
rus available from the region of
\|sothe type locality are mostly of |

B2018-103 young animals or have imperfect

Fic, 12.—Geographic range of the subspecies of skulls. However, there seems to

OC ae be very little geographic variation,

either chromatic or cranial, in

specimens taken throughout the whole coastal strip extending from
Juan de Fuca to Mendocino County, Cal.

Specimens examined.—Total number, 46, as follows:

California: Crescent City, 3; Cuddeback, 1;1 Eureka, 6;1 Ferndale, 2;! Mendo-
cino, 1;? Orick, 1; Smith River, 1.

Oregon: Astoria, 1;? Myrtle Point, 1; Newport, 1; Portland, 3;4 Yaquina Bay, 4.

Washington: Chehalis County, 1; King County, 1;° La Push, 1; Neah Bay, 2;

Pacific County, 1;° Port Townsend, 1; Puyallup, 6; Shoalwater Bay (type
locality), 2; South Bend, 5; Steilacoom, 1.

os = eT DD oe OR SSeS

—s os

1. S. 0. orarius. 2. S. 0. schefferi.

1 Collection Mus. Vert. Zool., Univ. California. 4 Collection of H. E. Anthony, New York City.
2Collection Field Mus. Nat. Hist. 5 Collection Milwaukee Public Mus.
3 Collection Univ. Michigan Mus. 6 Collection Amer. Mus. Nat. Hist.

1915.] SCAPANUS ORARIUS SCHEFFERI. : 68

SCAPANUS ORARIUS SCHEFFERI subsp. nov.!
ScHEFFER’S MOLE.
CPV, fie. 32) P1) ‘V1, fig: 12.)

Type locality —Walla Walla, Walla Walla County, Washington.

Type specimen.—No. 204997, U. S. Nat. Mus., Biological Survey
collection; ¢ adult, skin and skull; collected August 8, 1914, by
Theodore H. Sees

Geographic range.—Extreme southwestern British Columbia, nor cde
western Washington (east of Puget Sound and north of eeecdc 48°
N.), central and southern Washington from the west slopes of the
Cascade Mountains east to Walla Walla, and both slopes of the Cas-
cade Mountains in northern and east-central Oregon.

General characters.—Similar to S. 0. orarius but much paler, slightly
larger, with both hind and fore feet relatively larger, and claws
heavier. Skull about the size of that of orarius, with relatively
shorter and broader rostrum and greater interorbital breadth.

Color.—Fresh autumn pelage: Upperparts glossy deep mouse gray;
underparts slightly paler and more grayish. Worn summer pelage:
Upperparts hair-brown to pale fuscous; underparts mouse gray.

Skull—Compared with that of S. 0. orarius, the skull of scheffert is
broader interorbitally and has a relatively and actually shorter and
broader rostrum, more evident sublachrymal-maxillary ridge, heavier
mandible, and oe dentition.

Measurements—Type (adult male): Total length, 170; tail verte-
bre, 35; hind foot, 23. Average of 3 adult males @nondine type)
ame ae feeatity 168.7 (165-171); 34.7 (84-35); 23 (23-23).
Skull: Type (adult male): Greatest length, 34.7; palatilar length,
14.6; mastoidal breadth, 16.7; interorbital breadth, 8.4; maxillary
tooth row, 11.2; mandibular molar-premolar row, 10.9. Average
of 3 skulls of adult males Gncluding type) from the type locaiity:
Greatest length, 34.4 (84.2-34.7); palatilar length, 14.4 (14.2-14.6);
mastoidal breadth, 16.4 (16.2-16.7); interorbital breadth, 8.3
(8.2-8.4); maxillary tooth row, 11.1 (11-11.2); mandibular molar-
premolar row, 10.8 (10.8-10.9). Average of 2 skulls of adult females
from the type locality: Greatest length, 32.9 (32.8-39); palatilar
length, 13.7 (13.6-13.8); mastoidal breadth, 15.9 (15.8-15.9); inter-
orbital breadth, 8.2 (8.2-8.2); maxillary tooth row, 10.6 (10.6-10.6);
mandibular molar-premolar row, 10.4 (10.3-10.5).

Remarks.—Typical specimens of scheffert are easily distinguishable
from S. o. orarius, its most closely allied form. The type specimen
has fresh autumn pelage on the upperparts, the underparts still re-
taining the worn summer pelage. On the face of the type specimen

1 Named for Theodore H. Scheffer in recognition of his careful investigations of the habits and ecology
of American moles. :

64 NORTH AMERICAN FAUNA. "OT Ne. 38.

is a small spot of ochraceous-buff and a faint wash of the same color
on the throat. Similar chromatic abnormalities crop out in several
specimens from the type locality, and one specimen (No. 204998,
U.S. Nat. Mus.) has as a dental abnormality a supernumerary left
upper premolar.

Intergradation of this form with orarius is clearly indicated in
specimens from the west slope of the Cascade Mountains in Oregon
and Washington. Specimens from Vida, McKenzie Bridge, and
Three Sisters, in Oregon, approach orarwus in color and in cranial
characters, though distinctly referable to schefferi. Specimens from
Mount Vernon, Wash., are slightly darker in color than specimens
from Walla Walla, but the skulls are almost identical in essential
characters with the type series. Specimens from Skykomish, Wash.,
are nearly as dark as typical orarius, but are referred to schefferi
because of cranial characters which, although approaching orarius,
are much nearer scheffert.

Specumens examrned.—Total number, 59, as follows:

British Columbia: Chilliwack Valley, 2; Chiloweyuck Depot, 2; Fraser River
(near New Westminster), 1;1 Sumas, 6.

Oregon: McKenzie Bridge, 2; Three Sisters, 2; Vida, 4.

Washington: Easton, 4; Fort Walla Walia, 4; Lester, 4; Merritt, 5; Mount Ver-
non, 2; North Yakima, 5; Skykomish, 6; Walla Walla (type locality), 5;
Wenatchee, 5.

SCAPANUS LATIMANUS LATIMANUS (Bachman).
CALIFORNIA MOLE.
(Pl, IV,, fig..4; Pl. V,-figs..3, 3a; Pl. Vi; fies13%)

Scalops latimanus Bachman, Boston Journ. Nat. Hist., vol. 4, p. 34, 1842.

Scalops californicus Ayres, Proc. California Acad. Sci., vol. 1, p. 54, 1856. Type
locality, San Francisco, California.

Scapanus californicus True, Proc. U. 8. Nat. Mus., vol. 19, p. 52, December 21, 1896.

Scapanus latimanus Osgood, Proc. Biol. Soc. Washington, vol. 20, p. 52, April 18, 1907.

Scapanus latimanus latimanus Miller, U.S. Nat. Mus., Bul. 79, p. 9, December 31, 1912.

Type locality—Probably Santa Clara, Santa Clara County, Cali-
fornia.’

Type specimen.—Mounted specimen, with imperfect skull, in the
Berlin Museum; collected during October, 1834.

Geographic range.—Western California west of the San Jacinto and
Sacramento Valleys, from Santa Maria River north to Cape Mendo-
cino, thence northeasterly to Klamath Canyon, Siskiyou County.

General characters.—Size medium (length of adults usually about
170 to 180 mm.); color fuscous or drab to chetura black, most
frequently chetura drab; skull flat, rather massive in the maxillary

1 Collection Amer. Mus. Nat. Hist.
2 Vide Osgood, W. H., Proc. Biol. Soc. Washington, vol. 20, p. 52, 1907,

1915 ] SCAPANUS LATIMANUS LATIMANUS. 65

region; rostrum short and relatively heavy; unicuspid teeth irregular
and crowded.

Oolor.—Fresh winter pelage: Above, fuscous-black, chetura drab,
or chetura black; old and faded specimens more brownish, fuscous
to mummy brown; beneath, much as above but paler, usually
slightly more grayish, and frequently stained with brown mid-
ventrally. Summer pelage: Usually paler and shghtly more brown-
ish than winter pelage, drab, hair-brown, or chetura drab, paler
beneath. Young: Usually darker and more grayish than adults.

Skull._Size medium, flat; sublachrymal-maxillary ridge heavy
and well defined; rostrum short and broad; unicuspid teeth irregular
in size, and crowded; second lower incisor caninelike, larger, and
longer than either the first or the third incisor. The skull of S. 1.
latumanus is very-‘much smaller than that of S. townsend, the skulls
of large adult males of latimanus being smaller than the skulls of
small adult females of townsend; the rostrum of latimanus is rela-
tively shorter and broader than that of townsendii, and the unicuspid
teeth are more irregular in size and shape, and more crowded.
Compared with S. o. orarius the skull of latimanus is wider and much |
heavier, with less rounded and more truncate braincase, much shorter
and broader rostrum, much heavier sublachrymal-maxillary ridges,
larger teeth, and heavier horizontal rami of mandibles. The skull
of typical latimanus is larger than that of any other subspecies of
S. latimanus except S. 1. alpinus, from which it is indistinguishable.

Measurements —Adult male from Gilroy, Santa Clara County,
Cal.: Total length, 173; tail vertebrz, 35; hind foot, 23. Average of
4 young adult males from Santa Cruz, Cal.: 175.8 (173-181); 33.3
(32-38) ; 20.8 (20-21). Average of 4 young adult females from Santa
Cruz, Cal.: 171.5 (162-185); 32.5 (81-35); 20.8 (20-21). Skull:
Adult male from Gilroy, Santa Clara County, Cal.: Greatest length,
36.2; palatilar length, 14.4; mastoidal breadth, 17.3; interorbital
breadth, 7.8; maxillary tooth row, 11.2; mandibular molar-premolar
row, 10.9. Average of 4 skulls of young adult males from Santa
Cruz, Cal.: Greatest length, 36.1 (35.7-36.6); palatilar length, 14.2
(13.8-14.6); mastoidal breadth, 17 (16.9-17.3); interorbital breadth,
7.8 (7.5-8); maxillary tooth row, 11 (10.8-11.1); mandibular
molar-premolar row, 10.6 (10.5-10.8). Average of 7 skulls of
adult males from Nicasio, Cal.: Greatest length, 37.1 (36.6-37.4);
palatilar length, 14.5 (14.3-14.7); mastoidal breadth, 17.3 (17-178);
interorbital breadth, 7.8 (7.5-7.9); maxillary tooth row, 11.3
(11.1-11.6); mandibular molar-premolar row, 11 (10.7-11.4). Aver-
age of 4 skulls of young adult females from Santa Cruz, Cal.: Greatest
length, 34.7 (34.4-35); palatilar length, 13.8 (13.4-14.1); mastoidal
breadth, 16.8 (16.6-17); interorbital breadth, 7.8 (7.7-8); maxillary

90549°—15——5

66 NORTH AMERICAN FAUNA. [No. 38.

tooth row, 10.7 (10.4-10.9); mandibular molar-premolar row, 10.3
(10.1-10.5). Average of 5 skulls of adult females fromeNicasio, Cal.:
Greatest length, 36 (85.3-36.6); palatilar length, 14.1 (13.7-14.5);
mastoidal breadth, 16.9 (16.7-17.2); interorbital breadth, 7.7 (7.4—
8.1); maxillary tooth row, 11.1 (10.8-11.4); mandibular molar-
premolar row, 10.8 (10.3-11.1).

Remarks.—Bachman’s name latvmanus was placed in synonymy
under Scapanus townsend by Peters! where it remained until
Osgood showed that it did not apply to townsendu but to the mole of
west-central California, then known as Scapanus califormcus (Ayres).
Osgood writes:

As stated by Peters (loc. cit.), it [the type specimen] was transmitted by Deppe
from Monterey, California. It was collected in October, 1834, at Santa Clara, not a
Mexican locality,? as suggested by Peters, but doubtless the*town of that name in
California not very distant from Monterey. Only one species of mole is known to’
occur at this locality, and the specimen is typicalof this species. The hind foot to
end of claws measures 18.7 mm. The fragmentary skull, which Dr. Matschie caused
to be removed from the mounted specimen, presents the following measurements, all
decidedly smaller than S. townsendi: Length of upper tooth row from front of incisor

to back of last molar, 15.4; of lower tooth row, 13.7; outside width at second upper
molar, 10.2.3

The measurements given by Osgood are somewhat less than those
of skulls of adults from the vicinity of Santa Clara, Cal., but this may
possibly be due to immaturity of the type specimen. The animal is
certainly not townsendu, and it seems best to accept Osgood’s verdict
and place californicus in synonymy under latumanus.

A skeleton * of this species in the United States National Museum |
has been set aside as the type of Scalops californicus Ayres, and of it
Lyon and Osgood state: ‘“‘This skeleton is one of Dr. Ayres’s original
specimens, and probably the only one of them now in existence. It
seems well to treat it as a type, although it was not so indicated by
the original describer.” > However, there seems to be no good reason
for designating this specimen as the type of californicus; it was
entered in the museum catalogue February 14, 1857, but in the col-
lection are two other specimens (alcoholic) which Lyon and Osgood
apparently overlooked. One® of these, without the date of collec-
tion, was entered in the catalogue May 4, 1857; the other,’ collected
several weeks after Ayres had read his description before the Cali-
fornia Academy of Sciences, was entered in the museum catalogue

1 Peters, W., Monatsber. Konig. Preuss. Akad. Wissensch., Berlin, 1863, p. 656, 1864. :

2 At the time the specimen was collected California was part of Mexico. Peters, however, states ‘in Sta.
Clara (Sonora?) gesammelt worden.’’

2 Osgood, W. H., Proc. Biol. Soc. Washington, vol. 20, p. 52, 1907.

4No. 3111, U. 8. Nat. Mus., skeleton (lacking right manus and forearm, and left last upper molar); col-
lected at San Francisco, Cal., by Dr. W. O. Ayres. °

5 Lyon, M. W., and Osgood, W. H., U.S. Nat. Mus., Bul. 62, p. 234, 1909.

6 No. 2673, U.S. Nat. Mus., alcoholic; collected at San Francisco, Cal., by Dr. W. O. Ayres.

7 No. 1288, U. S. Nat. Mus., alcoholic; collected at San Francisco, Cal., September, 1855, by Dr. W. O.
Ayres.

1915.] SCAPANUS LATIMANUS LATIMANUS. 67

February 4, 1856. Nothing in the original description would indicate
that Ayres had any one particular specimen in mind;"in the only
place in his description where specimens are mentioned, he refers to
the ‘‘color of fur, in the specimens seen.” + Furthermore if Ayres set
aside any specimen as the type it was probably in the Museum of the
California Academy of Sciences, since preceding his original descrip-
tion the donation of five moles to the Academy was acknowledged
and the following statement made concerning them: ‘‘In connection
with these, Dr. Ayres presented the following description”’ (loc. cit.).
It seems, therefore, that the specimens in the United States National
Museum are not a part of Ayres’s original series.

Slight local variations in size and shape of skull of latwmanus
might be worthy of subspecific recognition were they constant over
any considerable geographic area. However, they are so slight and
inconstant, and crop out so frequently, that to recognize them by
subspecific appellation would only be confusing, and add nothing
to the knowledge of the relationships of the group. Specimens from
the coast rezion north of San Francisco Bay seem to average slightly
larger and darker than typical specimens, but the difference is not
sufficiently pronounced to warrant subspecific designation. Speci-
mens examined from Klamath Canyon show little approach in color
toward 8S. l. dilatus, but the skulls are shorter than in typical Jati-
manus, showing in this respect intergradation with dilatus. Speci-
mens from Lower Lake and Mount Sanhedrin are paler than typical
latimanus, probably indicating an approach toward dilatus. Inter-
gradation with S. 1. occultus occurs in the region between Santa
Margarita, in San Luis Obispo County, and Santa Barbara; a speci-
men from the former locality is clearly intermediate, but appears to
be nearer latumanus.

Specumens excamined.—Total number, 171, as follows:

California: Aptos, 1; Bells Station, 1; Berkeley, 8; Berryessa, 1; Beswick, 2;
Bodega, 1; Bolinas, 1; Boulder Creek, 1; Brentwood, 1; Briceland, 1; Cahto,
2; Carmel Point, 1;3 Cazadero, 1; Colma, 1; Cuddeback, 2;? Eldridge, 7; Fort
Bragg, 2; Freestone, 3;? Gilroy, 1; Gualala, 1; Guerneville, 3; Haywards, 5;
Hornbrook, 1; Inverness, 1; King City, 1; La Honda, 3;?,3 Lower Lake 2;+
Marin County, 1; Mendocino, 3;?»> Menlo Park, 8;? Monterey, 1;4 Mount
Sanhedrin, 1;* Napa, 1;? Nicasio, 54; Oakland, 4;° Pacific Grove, 1; Palo
Alto, 1;+ Petaluma, 2; Petrolia, 2; Piedmont, 1;? Point Arena, 1;2 Point
Reyes, 1; Red Bluff, 2;? Rockport, 2; San Francisco, 10; San Leandro, 3;?
Santa Cruz, 9; Santa Margarita, 1;? Santa Rosa, 1;° Scott River, Siskiyou

County, 1;? Snow Mountain, Colusa County, 1;* Stanford University, 1;?
Stevens Creek, San Mateo County, 1;° Walnut Creek, Contra Costa County, 2.?

1 Ayres, W.O., Proc. California Acad. Sci., vol.1, p.54,1856. 4 Collection Mus. Comp. Zool., Harvard College.
2 Collection Mus. Vert. Zool., Univ. California. 5 Collection Field Mus. Nat. Hist.
3 Collection Amer. Mus. Nat. Hist. 6 Collection Milwaukee Public Mus.

68 NORTH AMERICAN FAUNA. [No. 88.

SCAPANUS LATIMANUS OCCULTUS Grinnell & Swarth.
SOUTHERN CALIFORNIA MoLgE.
(Pl. IV, fig. 5; Pl. VI, fig. 14.)

Scapanus latimanus occulius Grinnell & Swarth, Univ. California Publ. Zool., vol.
10, p. 131, April 13, 1912.

Type locality—Santa Ana Canyon, west slope of north end of
Santa Ana Mountains, Orange County, California; altitude 400 feet.

Type specomen.—No. 2369, Mus. Vert. Zool., Univ. California; 9
young adult, skin and skull; collected September 20, 1908, by H. S.
Swarth. ©

Geographic range.—Southern California west of the deserts, from
Olancha, at the south end of Owens Lake, in Inyo County; Sanger,
in Fresno County; and Santa Barbara, in Santa Barbara County,
south to the San Diegan region.

General characters.—Size small (length of adults usually about
145 to 160 mm.); color in winter pelage paler and shghtly more
brownish than that of S. J. latumanus; color in summer pelage much
like that of latimanus; skull small, weak; rostrum usually relatively
longer and narrower than that of latumanus.

Color.— Winter pelage: In most specimens paler and more brown-
ish than corresponding pelage of S. 1. latumanus; upperparts cheetura
drab or fuscous, and occasionally, in much faded specimens, olive-
brown or mummy brown; underparts somewhat paler and more
grayish. Summer pelage: Very slightly paler than winter pelage;
essentially like summer pelage of latumanus. :

Skull_—Very similar to that of S. 1. latumanus in general shape
and proportions, but much smaller, and in most specimens with rela-
tively longer and narrower rostrum.

Measurements.—Average of 13 young adult males from Compton,
Los Angeles County, Cal.: Total length, 151.9 (140-165); tail vertebrae,
25.3 (22-29); hind foot (measured, from relaxed foot of dry skin,
by the writer), 18.3 (17.5-19.5). Type (young adult female): 150;
33; 18. Two young adult females from Olancha, Owens Lake,
Cal.: Total length, 148, 155; tail vertebra, 30, 36; hind foot, 18, 19.
Skull: Average of 13 skulls of young adult males from Compton, Los
Angeles County, Cal.: Greatest length, 31.6 (80.7-32.8); palatilar
length, 12.3 (12.1-12.8); mastoidal breadth, 15.4 (15-16); imter-
orbital breadth, 7.1 (6.9-7.4); maxillary tooth row, 10 (9.7—10.3);
mandibular molar-premolar row, 9.5 (9.3-9.9). Average of 4 skulls
of adult males from Witch Creek, Cal.: Greatest length, 32.9 (32.6-
33.5); palatilar length, 12.9 (12.7-13.2); mastoidal breadth, 15.8
(15.5-16.1); interorbital breadth, 7.3 (7.2-7.5); maxillary tooth row,
10.3 (10.1-10.5); mandibular molar-premolar row, 9.8 (9.6—10).
Skull of type (young adult female): Greatest length, 31.5; palatilar

1915.1 SCAPANUS LATIMANUS GRINNELLI. 69

length, 12.2; mastoidal breadth, 15.6; interorbital breadth, 7.1;
maxillary tooth row, 9.9; manidibular molar-premolar row, 9.4.
Skulls of 2 young adult females from Olancha, Cal.: Greatest length,
31, 31.5; palatilar length, 12.2, 12.6; steenottell breadth, 15.1, 15.6;
ii rorbital breadth, 7.2, 7.3; sane dl lore tooth row, 9.9, 10; snes li:
ular molar-premolar row, 9. 4, 9.6.

Remarks.—The mole a ouiann California j is constant in charac-
ters over its entire range. ‘The type of occultus is a young adult
female; it has abnormal mandibular teeth; the right first lower
premolar is absent, and the left first lower premolar is crowded back
and inward, close to the second premolar and in the same enlarged
socket with it. Specimens from Witch Creek are slightly larger
than those from the type region. Toward the northern border of
its range the subspecies shows a gradual increase in size until it
intergrades with S./. laivmanus in the region north of Santa Barbara
County. <A young adult male from Tehachapi and a specimen from
Santa Barbara approach latumanus in size. A male from Porter-
ville, Tulare County, is larger and slightly darker than typical
specimens of occultus, and has a relatively narrower skull, probably
showing in these respects a tendency towards S. l. sericatus. An
imperfect skeleton, with broken skull, from Sanger, Fresno County,
is provisionally referred to occultus; in dentition and in the size and
proportions of the mandible it is indistinguishable from specimens
from the type region. Two females from the south end of Owens
Lake are almost indistinguishable from typical occultus, their only
approach toward S. 1. grinnelli being in slightly higher braincase.
The relationships of occultus with Scapanus anthonyi are not clear,
but the differences in size and color alone are sufficient to separate
the two forms, even though it should ultimately be proven that the
dentition of the type of anthonyi is abnormal (p. 76).

Specimens examined.—Total number, 73, as follows:

California: Alhambra, 10; Canyada Laga, 4 miles north of Ventura, 1;! Comp-
ton, 21; Julian, 2;? Los Angeles, 3; Olancha, 2; Pasadena, 2;? Porterville, 1;
Riverside, 1; San Bernardino, 5; San Bernardino Peak, 1; San Diego County,
5; San Gabriela: Sanger, 1;? Sun Jacinto Mountains, 2;? Santa Ana Canyon
one locality), 1;? Santa Bei 1; Sierra Madre, os Angeles County, 1;?
Somerset, 4; Ten achapy 1; Ventura, 1: ? Warner Pass, San Diego County, 1:2
Witch Creek, 5.

SCAPANUS LATIMANUS GRINNELLI Jackson.
GRINNELL’s Mote.
(Pl. IV, fig. 6; Pl. VI, fig 15.)
—Scapanus latimanus grinnelli Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 56,
March 20, 1914.

Type locality. —Independence, Inyo County, California; altitude
3900 feet.

1 Collection Milwaukee Public Mus. 2 Collection Mus. Vert. Zool., Univ. California.

70 NORTH AMERICAN FAUNA. [No. 38.

Type specumen.—No. 17785, Mus. Vert. Zool., Univ. California; ¢
adult, skin and skull; collected May 8, 1912, by H. A. Carr.

Geographic range.—Known only from type locality.

General characters.—Size small; very slightly larger than average
specimens of S. l. occultus or S. l. minusculus; smaller than S. 1.
sericatus; color darkest of the latumanus group; skull short and high,
broad interorbitally and through mastoids; rostrum short and wide.

Color.—Type, in worn and somewhat faded winter pelage: Upper-
parts between fuscous and fuscous-black; underparts more grayish,
stained on the throat and chest with Dresden brown. Topotype,
in fresh summer pelage: Upperparts fuscous-black; underparts dark
mouse gray.

Skull.—Short and high, not much depressed postorbitally, broad
interorbitally and through the mastoids; posterior base of zygoma
with a smail inconspicuous process on outer lateral margin; rostrum
short and wide; angle of mandible short and heavy. The skull of
grinnelli is about the size of large skulls of S. 1. occulius, but differs
from them, in its higher braincase, its shorter and much broader
rostrum, and in being much wider interorbitally and through the
mastoids. It is much smaller and relatively shorter and wider than
the skull of either S. 7. latumanus or S. l. sericatus.

Measurements.—Type (adult male): Total length, 156; tail verte-
bre, 31; hind foot, 21. ‘Topotype (adult female): Total length, 158;
tail vertebre, 36; hind foot, 20.5 Skull: Type (adult male): Greatest
length, 33; palatilar length, 13.1; mastoidal breadth, 17; interorbital
breadth, 7.8; maxillary tooth row, 10.4; mandibular molar-premolar
row, 10. Skull of topotype (adult female): Greatest length, 32.2; ©
palatilar length, 12.8; mastoidal breadth, 16.4; interorbital breadth,
7.8; maxillary tooth row, 10.3; mandibular molar-premolar row, 9.9.

Remarks.—This form is a well-marked subspecies which can be
separated from S. l. occultus either by skin or skull characters. Inter-
eradation with occultus probably occurs in the region north or west
of Owens Lake; a very slight indication of such intergradation is
noticeable in specimens of occultus from Olancha. The topotype .
is peculiar in that the left upper tooth row contains a supernumerary
premolar, apparently the one immediately posterior to the canine
and probably derived from the anlage of the first premolar. As is
the case in some specimens of S. townsend, previously mentioned,
this is especially interesting, since it gives this individual one more
tooth than the theoretically complete mammalian dentition of 44
teeth.

Specumens examined.—Iwo, from type locality.

1 Collection Mus. Vert. Zool., Univ. California.

1915.] SCAPANUS LATIMANUS SERICATUS. U1

SCAPANUS LATIMANUS SERICATUS Jackson.
‘ Yosemite MOotLe.
(Pl. IV, fig. 7; Pl. V1, fig. 16.)

Scapanus latimanus sericatus Jackson, Proc. Biol. Soc. Washington, vol. 27, p. 90,
March 20, 1914.

Type locality.— Yosemite, Yosemite Valley, Mariposa County, Cali-
fornia. © .

Type specimen.—No. 109548, U. S. Nat. Mus., Biological Survey
collection; @ adult, skin and skull; collected August 20, 1901, by
W. K. Fisher. ,

Geographic range.— Yosemite region, Mariposa County, Cal.

General characters.—Smaller than S. 1. latimanus, darker and more
grayish in fresh pelage; larger and darker than S. l. occultus or S. l.
minusculus; claws, especially of fore feet, longer and more slender
than those of latimanus; skull relatively long, narrow, and flat.

Color.—Fresh summer pelage: Upperparts fuscous-black; under-
parts more grayish, dark mouse gray. Worn summer pelage: Upper-
parts glossy olive-brown, underparts slightly paler.

Skull—Relatively long, narrow, especially through mastoids;
smaller than that of S. 1. latimanus; larger than that of S. 1. occultus,
S. 1. minusculus, or S. 1. grinnelli; about equal im length to that of
S. 1. dilatus, but much narrower, and not so high through the brain-
case.

Measurements.—Two adult females, type and virtual topotype:
Total length, 165, 171; tail vertebra, 36, 34; hind foot, 21, 22. Skull:
Skulls of two adult females, type and virtual topotype: Greatest:
length, 34.3, 34; palatilar length, 13.3, 13.7; mastoidal breadth,
16.2, 15.9; mterorbital breadth, 7.2, 7.5; maxillary tooth row, 11,
10.8; mandibular molar-premolar row, 10.4, 10.4.

Remarks.—The skulls of the type and topotype! of sericatus show
minor differences, but in essential features they are remarkably alike.
They are very unlike the skull of S. 1. minusculus, being distinctly
larger, and with different proportions. The subspecies sericatus is
intermediate in size between S. 1. lattmanus and S. 1. occultus, and in
fresh pelage is darker than either. The type of sericatus is mostly in
worn summer pelage, fresh pelage appearing on the abdomen and
flanks; the topotype is in fresh summer pelage. An adult male from
Bower Cave, Mariposa County, is not typical of the subspecies; it
has a shorter tail than specimens from the Yosemite Valley, and
the skull is flatter and wider through the braincase than that of typical
servcatus.

Specimens examined.—Total number, 3, as follows:

California: Bower Cave, 1; Yosemite (type locality), 1; Yosemite Valley, 1.?

1 No. 12980, Mus. Vert. Zool., Univ. California. 2 Collection Mus. Vert. Zool., Univ. California.

42 NORTH AMERICAN FAUNA, [No. 38.

SCAPANUS LATIMANUS MINUSCULUS Bangs.
SreRRA MOLE.
(PITY, fie. 85 -Pl) Vij fice ag)

Scapanus californicus minusculus Bangs, Proc. New England Zool. Club, vol. 1, p. 70,
July 31, 1899.

Scapanus latimanus minusculus Miller, U. 8. Nat. Mus., Bul. 79, p. 10, December 31,
1912.

Type locality Fyffe, El Dorado County, California.

Type specimen.—No. 9189, Mus. Comp. Zool., Harvard College,
Bangs collection; ? young adult, skin and skull; collected July 15,
1897, by W. W. Price and E. M. Nutting.

Geographic range.—Known only from type locality.

General characters.—Similar in size and color to S. l. occultus, pos-
sibly very slightly larger; skull higher and narrower through brain-
case than that of occultus; inferior mandibular notch very shallow,
the angle of mandible being much shorter than in occultus.

Color.—Type, in summer pelage: Upperparts hair-brown:; under-
parts deep mouse gray.

Skull.—Much like that of S. 1. occultus, but braincase narrower,
higher, and more rotund; inferior mandibular notch very shallow,
much shallower than in occulius; angle of mandible short and weak.

Measurements.—Type (young adult female): Total length, 160; tail
vertebre, 31; hind foot, 21. Skull: Type (young adult female):
Greatest length, 31.9; palatilar length, 12.6; mastoidal breadth,
15.3; interorbital breadth, 7.1; maxillary tooth row, 10; mandib-
ular molar-premolar row, 9.7.

Remarks.—The type and only-known specimen of minusculus is a
female, barely adult, the status of which can not be determined until
more specimens are obtained from the type locality. The essential
differences from S. 1. occultus are in the skull, which is relatively
narrower and higher and with the inferior mandibular notch very
shallow; the hind foot is longer than that of occultus. This form,
however, seems to be isolated from occultus by the interception of
sericatus. In color the type of minusculus is much like some speci-
mens of S. l. dilatus.

Specimen examined.—One, the type.

.

SCAPANUS LATIMANUS DILATUS True.
KiamMAtTH MOLE.
(PL. IV, fig. 9; Pl. VI, fig. 18.)

Scapanus dilatus True, Proc. U. S. Nat. Mus., vol. 17, p. 242, April 26, 1894.

Scapanus truei Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 102, April 26, 1897.
Type locality, Lake City, Modoc County, California.

Scapanus fruti (sic) Elliot, Field Columb. Mus., publ. 105, zool. series, vol. 6, p. 469,
1905.

1915.] SCAPANUS LATIMANUS DILATUS. 78

Type locality—Fort Klamath, Klamath County, Oregon.

Type specimen.—No. 186628, U.S. Nat. Mus., Merriam collection;
adult, sex unknown, skeleton; collected in 1883 by Charles EK. Bendire.

Geographic range.—South-central Oregon and Upper Sonoran and
Transition Zones of northeastern California and adjacent parts of
Nevada.

General characters.—Similar to S. 1. latimanus, but much paler and
averaging slightly smaller; skull shorter, higher, and more rotund
than in latimanus.

Color.—Summer pelage: Upperparts mouse gray, light drab, or
drab, in some specimens becoming more brownish on the nose;
underparts neutral gray, mouse gray, or smoke gray, sometimes
tinged with light drab and occasionally stained in midventral line
with Dresden brown.

Skull.—Smaller than that of S. 1. latumanus or S. l. alpinus, rela-
tively shorter, higher through the braincase, and relatively wider
through the mastoids. About equal in length to the skull of S. 1.
sericatus, but relatively much wider and higher through the brain-
case.

Measurements.—Two adult males from McCloud, Cal.: Total
length, 178, 170; tail vertebre, 36, 42; hind foot, 21, 21. Skull:
Type (sex unknown): Greatest length, 34.3; palatilar length, 13.6;
mastoidal breadth, 16.8; interorbital breadth, 7.8; maxillary tooth
row, 10.4;1 mandibular molar premolar row, 10.3. Skulls of two
adult males from McCloud, Cal.: Greatest length, 34.7, 35.2; palatular
length, 13.2, 13.7; mastoidal breadth, 17.3, 17.3; 1aterorbital breadth,
7.8, 7.9; maxillary tooth row, 9.8, 10.3; mandibular molar premolar
row, 9. 5, 9.5.

Penarks. —True based his description of this fan entirely upon
the cranial characters of a specimen with abnormal dentition, there
being in the type only three maxillary premolars. Later he irae
the name in synonymy? under Scapanus californicus (=8. l. lati-
manus). The skull of the type, however, is relatively shorter and
broader than that of typical latumanus. Unfortunately the exact
color of the mole found at Fort Klamath, Oreg., is not known, there
being no skins available from that locality; two alcoholics from the
type locality, however, seem to indicate that the animal is pale,
like the mole of northeastern California; from a geographic view
point this is what one would anticipate; specimens from Ashland
and Fremont, Oreg., are pale, like specimens from northeastern
California, and those from Ashland have skulls inseparable from

the skull of the type of dilatus. The type of Scapanus truei Mer-

riam, which is now placed in synonymy under dilatus, is a rather

1 Upper tooth formula abnormal: i. 3, c. 1, pm. 3, m. 3.
2 True, F. W., Proc. U. S. Nat. Mus., vol. 19, p. 52, 1896.

14 NORTH AMERICAN FAUNA. [No. 38.

young adult collected at Lake City, Modoc County, Cal.; in the
original description Merriam mentions the following cranial and
dental characters:

Skull similar to that of S. californicus, but slightly smaller, with narrower palate

and decidedly narrower and more slender rostrum. Last upper premolar with a
strongly developed, trenchant inner cusp, not present in californicus.}

The narrowness of the palate and rostrum is probably due in
part to immaturity; the skull of a young adult female, which has
been removed from an alcoholic specimen? from Fort Klamath,
has the rostrum as narrow as that of the type of truer though it is
slightly shorter. The presence of an inner cusp on the last upper
premolar is not of diagnostic value. In any large series of Scapanus
from a single locality this cusplike process may be present in various
stages of development in about half the specimens; it appears more
frequently in skulls of young individuals and seems to be absorbed
with age. In the type of truei this accessory cusp has about reached
its maximum development, yet in a specimen ® collected as nearby
as Cedarville, Modoc County, there is a barely perceptible trace of
a cusp.

Considerable local geographic variations are observed in this
form, but to recognize them by subspecific names would only be con-
fusing. With a large series of specimens available from each of many
localities it would be possible to learn more of the extent and rela-
tionships of these variations. Specimens from the upper Sacra-
mento Valley show intergradation with S. l. latvmanus, the color
darkening and the skulls becoming larger and more elongate. A
specimen from Fremont, Oreg., has certain cranial characters
which suggest that dilatus may possibly imtergrade with S. o.
schefferi in central Oregon.

Specimens examined.—Total number, 44, as follows:

California: Auburn, 1;+ Baird, 1; Bald Mountain, 1; Cedarville, 1; ° Chico, 1;
Fort Crook, 1; Hayden Hill, 1; Lake City, 1; McCloud, 2; McCloud River,
1; Millford, 2; Mosquito, El Dorado County, 1;° Mount Shasta, 1; Nevada
City, 1; Parker Creek, Warner Mountains, 1;° Plumas County, 1;’ Pratt-
ville, 1; Quincy, 4; Red Point, 1;* Round Mountain, Shasta County, 1;
Sisson, 6;° Susanville, 4;+ Tower House, Shasta County, 1.°

Nevada: Holbrook, 1.
Oregon: Ashland, 3; Fort Klamath (type locality), 3; Fremont, 1.

1 Merriam, C. H., Proc. Biol. Soc. Washington, vol. 11, p. 102, 1897.
2 No. 186627, U. S. Nat. Mus., Merriam collection.

3 No. 16723, Mus. Vert. Zool., Univ. California.

4 Collection Stanford Univ.

6 Collection Mus. Vert. Zool., Univ. California.

6 Collection Amer. Mus. Nat. Hist.

7 Collection Carnegie Mus.

1915.] SCAPANUS ANTHONYI. 75

SCAPANUS LATIMANUS ALPINUS Merriam.
Mount Mazama Mote.

Scapanus alpinus Merriam, Proc. Biol. Soc. Washington, vol. 11, p. 102, April 26, 1897.

Type locality—Crater Lake, Mount Mazama, Klamath County,
Oregon; altitude about 7000 feet.

Type specumen.—No. 79967, U. S. Nat. Mus., Biological Survey
collection; ¢ adult, skin and skull; collected August 18, 1896, by
Vernon Bailey.

Geographic range—Known only from type locality.

General characters—About the size of large specimens of S. 1. lati-
manus, but much paler; hind foot large; indistinguishable in color
from S. 1. dilatus, but larger; skull like that of large specimens of
latimanus, and much larger, flatter, and relatively longer and narrower
than that of dilatus.

Color.— Worn summer pelage (type): Upperparts mouse gray;
underparts (much worn) deep mouse gray; faded unworn pelage on
throat and chest with a distinct buffy sheen.

Skull.—Similar to that of S. 2. datvmanus; indistinguishable from
skulls of large adult males of latimanus.

Measurements.—Type (adult male): Total length, 188; tail verte-
bre, 38; hind foot, 24.5 Skull: Type (adult male): Greatest length,
36.9; palatilar length, 14.6; mastoidal breadth, 17; interorbital
breadth, 7.9; maxillary tooth row, 11.4; mandibular molar-premolar
row, 11.3.

Remarks.—The subspecies alpinus is a poorly differentiated form
having the color of S. 1. dilatus and the size and skull proportions of
large specimens of S. 1. latimanus; it is not strictly intermediate
between these two forms, however, since specimens of latimanus
from southwest of Klamath Lake do not reach the maximum size.
The type is a very old male with much-worn teeth; the skull is flat-
tened by age and the cusps of the teeth are reduced through wear
and absorption.

Specomen examined.—One, the type.

SCAPANUS ANTHONYT Allen.
AnTHONY’S MOLE.
(BL. EV, fig 10; Pl. V, figs. 4, 443° PIOVYE, fig: 19.)
Scapanus anthonyi Allen, Bul. Amer. Mus. Nat. Hist., vol. 5, p. 200, August 18, 1893.

Type locality—San Pedro Martir Mountains, Lower California;
altitude 7000 feet.

Type specimen.—No. $313, ee Mus. Nat. High ¢ adult, skin
and skull; collected May 8, 1893, by A. W. (nen

6 NORTH AMERICAN FAUNA. id. 38. |

Geographic range-—Known only from type locality.

General characters.—Size, smallest of the genus; feet and hands
small; in superficial appearance much like S. 1. occultus, but smaller
and darker; skull smaller than that of occultus, flatter, and with
relatively shorter rostrum; premolars 3.

Color—Type: Upperparts between fuscous and. fuscous-black;
underparts hair-brown to fuscous.

Skull_—Smallest of the genus; in general appearance and propor-
tions similar to that of S. Ll. occultus, but smaller, flatter, and with
relatively shorter rostrum; each of the parietals of occultus has a
small posterior projection, extending between the interparietal and
the mastoid, which is not present on the parietals of the type of
anthonyi; premolars, 3.

Measurements.—Type (adult male): Total length, 135; tail verte-
bre, 26; hind foot (measured by the writer from relaxed foot of dry
skin),17. Skull: Type (adult male): Greatest length, 30.1; palatilar
length, 12; mastoidal breadth, 16.2; interorbital breadth, 7; maxil-
lary tooth row, 9.3; mandibular molar-premolar row, 9.

Remarks.—As previous descriptions of the type and only-known
specimen of anthonyi have been somewhat inaccurate, particular
care has been taken in the present description and measurements.
Allen! gave the breadth of the interorbital constriction as 7.6
mm., an error probably due to inaccurate measuring instruments.
Allen (loc. cit.) claimed that the fourth premolar on one side was
wanting and on the other was rudimentary; True? in a footnote states
that he finds “‘only three premolars on either side,’ but elsewhere
(loc. cit., p. 51) gives the premolar formula as 3. As a matter of —
fact, the premolar formula is 3; True has correctly indicated this in
his sketch (loc. cit., pl. 3, fig. 6) of the mandible. It is not the fourth
premolar that is lacking, but either the first or second, probably the
first. It seems possible that the dentition of the type is abnormal;
however, the reduction of the premolars by one occurs uniformly in
each tooth row. It will be impossible to know the exact status of
the form until more specimens are available from the region of the
type locality. Full specific rank is here given the form because of the
absence of evidence showing intergradation with the nearest geo-
graphic neighbor, S. 1. occultus. The type of anthonyi is a fully adult
male, and if it represents an average specimen of the form, its size
and color alone are sufficient characters by which to separate it from
occultus.

Specumen examined.—One, the type.

1 Allen, J. A., Bul. Amer. Mus. Nat. Hist., vol. 5, p. 200, 1893.
2True, F. W., Proc. U.S. Nat. Mus., vol. 19, p. 53, 1896.

1915.] GENUS PARASCALOPS. fix
Genus PARASCALOPS True.

Parascalops True, Proc. U. S. Nat. Mus., vol. 27, p. 242, April 26, 1894.
Perascalops Beddard, Cambridge Nat. Hist., vol. 10, p. 518, 1902. .

Type species.—Scalops brewert Bachman.

Geographic range.—Southeastern Canada and northeastern United
States from southern New Brunswick, southern Quebec, and eastern
Ontario, south to northeastern Ohio and southern Pennsylvania, and
in the Appalachian Mountains to western North Carolina (fig. 13).

External characters —Body robust, not much depressed; tail short
(relatively longer than in Scalopus), round, thick, and fleshy, slightly
constricted at bas¢) TEES ee
annulated, densely |
covered with rather
long, coarse hairs
(fig. 14). Head co-
noidal, depressed.
Nose developed into
a conical snout
(shorter than in Sca-
lopus or Scapanus),
with a superior lon-
gitudinal median
groove extending
the anterior half of
its length; nostrils
lateral, crescentic,
with concavities up-
ward (fig.15). Eyes
minute, concealed in
fur. Auricular ori-
fices relatively large. , 3 |
Beem og. Neel So Bo
large, fleshy, sparse- Fic. 13.—Geographic range of the species Parascalops breweri.
_ly haired above, naked below. Fore feet handlike, the palms as
broad as long (fig. 16). Soles of hind feet each with two tubercles and
a distinct heel-pad (fig. 17). Toes not webbed. Claws of fore feet
broad, flat, and heavy; those of hind feet relatively short and slender.
Fur dense, soft, and silky (coarser than in Scalopus or Scapanus), the
hairs nearly equal in length, producing a velvetlike pelage. Mamma,
8: latero-pectoral, 2-2; latero-abdominal, 1-1; inguinal, 1-1.

Skeletal characters.—Clavicle relatively longer and weaker than in
Scalopus or Scapanus, length about equal to breadth, penetrated
antero-posteriorly by a foramen near the inferior border. Humerus

SS

0

p

78 NORTH AMERICAN FAUNA, [No. 38.

heavy, about two-thirds as broad as long. Pelvis narrow, bones of
opposite sides not touching under acetabula; no osseous bridges con-
necting sacral vertebre with ischium. Superior surface of last sacral
vertebra with a small, cuneate, longitudinal process. Os falciforme
short, reaching proximal end of first metacarpal; broad and triangular
at base, abruptly tapering into a narrow apical process.

Skull conoidal (less so than in Scalopus), flat (less so
than in Scalopus or Scapanus), somewhat elongate, with
moderately broad braincase, slightly constricted inter-
orbitally. Mastoids moderately heavy. Interparietal
moderately large (smaller than m Scapanus, larger than
in Scalopus), somewhat rectangular, breadth about one-
third the length, emarginate posteriorly and usually
anteriorly also, with a posterior median projection.
Frontal region depressed; frontal smuses moderately
swollen. Rostrum moderately long; anterior ends of
premaxille shghtly thickened and extending beyond
nasals, forming a truncate notch anterior to nasals.
Anterior nares opening forward. Zygomata moder-
ately long, relatively heavy, considerably out-curved,

Hi)
!
MKT
Halh
yyleh
Bhs!
y INF yk
i 4
y }
riot
Siti
A ir
0 b
Ht
Ma
/

B2020-103

Fie. 14—Tail of
Parascalops brew-
eri (X14). No.
186618, U. S. Nat.

the posterior end attached about medially on squamosal.
Foramen magnum oval, of moderate size. Infraorbital
foramen relatively small (about as in Scapanus), the
plate formmg its outer wall moderately broad (rela-

Mus Merman tively broader than in Scalopus or Scapanus). Audital
collection; irom 2 .
Locust Grove, bulle mcomplete; no auditory meatus. External

N.Y. pterygoid region moderately mflated (less so than

in Scapanus) both posteriorly and anteriorly. Mesopterygoid space
relatively long and narrow, the sides nearly straight and slightly
converging posteriorly. Palate moderately elongate, relatively nar-
row, terminating opposite posterior edge of last molar; posterior
border of
palate emar-
ginate, with-
out spine or
notch. An-
terior pala-
tine fora-
mina mod-
erate (relatively larger than in Scapanus), oval; first pair of poste-
rior palatine foramina moderate, oval to elliptical; second parr,
minute or obsolete. Horizontalramus of mandible moderately heavy,
curved upward at posterior end and downward at anterior end; coro-
noid elongate, quadrate, truncate, directed slightly forward; angle of

B2021-103
Individual referred to in fig. 14.

Fira. 15.—Snout of P. breweri (X14).

1915.1 PARASCALOPS BREWERI. fi

mandible relatively weak (about size of coronoid); inferior mandibu-
lar notch broad and shallow.

Dental characters.—First upper incisor short, broad, and flat, with
a distinct small external accessory cusp; Eecond a third upper
incisors, upper canine, and first, second, and third upper premolars
simple, conical, of Hodesats s1ze, subeqtial (except canine which is
larger), about half as high as first incisor; fourth premolar much
larger than third, with anterior cusp, and with narrow interior basal
ledge slichtly bilobed. Upper molars W-shaped in transverse sec-
tion, with an internal basal shelf distinctly trilobed in first and sec-
ond premolars and indistinctly bilobed in the third; second molar
slightly smaller than first, the third much smaller than second.

First lower incisor small, simple, slightly flattened; second lower
incisor larger than first, conical, somewhat caninelike; third lower

B2022-103 B2023-103

Fic. 16.—Fore foot of P. breweri (X14). Individual referred Fig. 17.—Hind foot of P. breweri
to in fig. 14. (X14). Individual referred to in
fig. 14.

incisor small (about size of first), conical. Lower canine, and first,
second, and third lower premolars essentially similar to third incisor.
Fourth premolar about one-third larger than third, the posterior base
with a small cusplike shelf or heel. Lower molars M-shaped in trans-
verse section, both antero- and postero-internal cusps bilobed;
first and second molars each with a postero-internal basai accessory
cusp; second and third molars each with an antero-internal basal
accessory cusp; first and second molars subequal, the third much
smaller. Dentition: i. 3; c.+; pm. 4; m. 3; total 44.

PARASCALOPS BREWKRI (Bachman).
Harry-TarLtep MoLe.
(RE TY, fie. 11: PL. V; fies: 5, da; Plo VE, fie: 20:)

Talpa Europxa Harlan (nec Talpa europea Linneus, 1758), Fauna Amer., p. 48, 1825.

Talpa americana Harlan (Bartram ms.) (misidentified with Talpa europxa Linnzus,
1758), Fauna Amer., p. 43, 1825.

Scalops Breweri Bachman, Boston Journ. Nat. Hist., vol. 4, p. 32, 1842.

80 NORTH AMERICAN FAUNA. [No. 38. -

[Scapanus] breweri Pomel, Archiv. Sci. Physiques et Nat., tome 9, p. 247, 1848.

T\alpa] reposta Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327, 1854.
Type locality unknown.

T\alpa] Breweri Le Conte, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 1853, p. 327, 1854.

Scapanus americanus Coues (based erroneously on Talpa americana Harlan [Bartram
ms.], 1825, qui Talpa europxa Linnzus, 1758), Amer. Nat., vol. 13, p. 190, 1879.

Scaphanus (sic) brewert Herrick, Geol. & Nat. Hist. Surv. Minnesota, Bul. 7, p. 55,
1892.

Parascalops brewert True, Proc. U.S. Nat. Mus., vol. 17, p. 242, April 26, 1894.

Type locality—Marthas Vineyard, Massachusetts.

Type specymen.—None known to exist.

Geographic range.—That of the genus (see p. 77).

General characters—Size medium (length averaging about 155
mm.); nostrils lateral, crescentic, with concavity upward; tail short,
thick, densely covered with hair; color dark; usually fuscous-black
or chetura black; skull flat; audital bulle incomplete; rostrum
slender; first upper incisors with a distinct external accessory cusp.

Color.—General tone, fuscous-black, chetura black, or chetura
drab, slightly paler and more grayish on underparts; hairs on feet,
and usually on nose and tail, more brownish, often becoming white
in old adults; throat and underparts sometimes stained with Dresden
brown or Saccardo’s umber. The color in study skins soon fades and
becomes slightly brownish.

Skull_—Size medium (length about 32 mm.), flat, depressed post-
orbitally, especially in adults, slightly constricted interorbitally;
zygomata moderately heavy; pterygoids small; audital bulle incom-
plete; auditory meatus absent; rostrum narrow; dentition moderate;
first upper incisor with distinct outer secondary cusp; internal edge
of second upper molar trilobed.

Measurements.—Average of 8 adult males from Magnetic City,
N. C.: Total length, 149.5 (139-152); tail vertebra, 30 (23-36);
hind foot, 19.5 (18-20). Average of 2 adult females from Lunen-
burg, Mass.:°153 (153-153); 29.5 (27-32); 18.5 (18-19). Skull:
Average of 10 skulls of adult males from Magnetic City, N. C.: Great-
est length, 32.4 (31-33.8); palatilar length, 12.6 (12.1-13.1); mas-
toidal breadth, 14.5 (13.9-15); interorbital breadth, 7.3 (7.1-7.5);
maxillary tooth row, 9.9 (9.2-10.2); mandibular molar-premolar row,
9.5 (8.9-9.8). Skull of adult female from Lunenburg, Mass.: Greatest
length, 31.2; palatilar length, 12.2; mastoidal breadth, 14.3; imter-
orbital breadth, 7.2; maxillary tooth row, 9.6; mandibular molar-
premolar row, 9.3.

Remarks.—Bartram was apparently the first to recognize this form,
to which he gave the manuscript name Talpa americana. Harlan?
placed Yalpa americana (Bartram ms.) in synonymy under Talpa
europea Linneus, apparently failing to distinguish the two forms

1 Harlan, R., Fauna Amer., p. 43, 1825.

1915.] PARASCALOPS BREWERI. Sl

even with the Bartram manuscriptin hand. Bachman‘ described the
species in 1842 under the name Scalops breweri, basing his description
upon a specimen ‘“‘found by Dr. L. M. Yale, at Martha’s Vineyard, an
Island on the coast of New Engiand.”’ Coues,? however, in 1879 ap-
plied the name Scapanus americanus to this species on the ground that
Harlan’s description was ‘‘applicable neither to Scalops [=Scalopus]
nor to Talpa” and that “he [i. e. Harlan] really had in view an
American mole, which he recognized as distinct, both generically and
specifically, from our common Scalops aquaticus.” The name Talpa
americana Harlan (Bartram ms.), however, is untenable for several
reasons: Harlan placed it in synonymy under Talpa europea Lin-
neus (loc. cit.); Harlan actually misidentified the American animal
with Talpa europea Linneus, as is evident from his mentioning in
his introduction a species of mole as common to both continents,?
and, by process of eliminating those species which he distinctly refers
in his descriptions to an American distribution, it becomes evident
that the animal he meant was Talpa europea Linneus; furthermore,
Harlan’s description, as has been shown by True,‘ is a translation,
word for word (with a very few omissions) of Desmarest’s * descrip-
tion of Talpa europea Linneeus.

The description of Talpa reposta Le Conte is based upon a specimen
from an unknown locality and seems to refer to a specimen of Para-
scalops brewert with slightly abnormal teeth.

The hairy-tailed mole shows remarkably little geographic varia-
tion, and such as does occur is obliterated by individual variation.
This variation is manifest mostly in size, and, in adults, size variation
may reach about 5 per cent below or above the average at a given
locality. White spots and blotches on the ventral parts of a few
specimens of young, as well as of adults, indicate an occasional tend-
ency toward partial albinism. In extreme old age the skull flattens
and becomes much depressed postorbitally. A most peculiar change
associated with old age is that, im many specimens examined, the
hair on the nose and tail is white; in other specimens, somewhat
younger but distinctly adult, some of the hairs on the nose and tail
are normally colored, others are white. This peculiar senile variation
is not confined to any particular region, but seems to be more preva-
lent north of Pennsylvania.

The hairy-tailed mole is rather rare and local in distribution, and
difficult to trap. It is, therefore, quite probable that in the course
of time its known geographic range may be extended considerably.

1 Bachman, J., Boston Journ. Nat. Hist., vol. 4, p. 32, 1842.
2 Coues, E., Amer. Nat., vol. 13, pp. 189-190, 1879.
3 Harlan, R., Fauna Amer., p. vili, 1825.

4 True, F. W., Proc. U.S. Nat. Mus., vol. 19, p. 76, 1896.
5 Desmarest, A. G., Mammalogie, Ire partie, p. 160, 1820.

90549°—15——6

82 NORTH AMERICAN FAUNA. [No. 38.

Bachman * lists it from Georgia but does not mention any speci--
men or definite locality. The writer has seen only three specimens
from Canada: One from Quebec, Quebec, is in the United States
National Museum; another, a flat skin without skull, was collected
at Meaches Lake, Quebec, and is in the Victoria Memorial Museum
at Ottawa; the third is a skeleton collected at Saint Catharines,
Ontario, and is in the exhibition cases of the United States National
Museum. Chamberlain? records seemg a specimen from Charlotte
County, New Brunswick. Specimens have been collected in Ontario,
“near Ottawa,’’? and at Guelph‘ and Acton.’ Nash ® records speci-
mens from several counties in Ontario, but mentions no specific
localities.
Specumens examined.—Total number, 129, as follows:

Connecticut: West Winsted, 1.

Maine: Lake Umbagog, 1.

Massachusetts: Harvard, 1;’ Lunenburg, 3.

New Hampshire: Dublin, 1; Ossipee, 1; Webster, 2.8

New York: Elizabethtown, 1;* Lake George, 7; Lansing, 3;% !° Locust Grove, 16;
Oswego, 1;1° Plateau Mountain, Catskills, 1; Peterboro, 3; Waterville, 1.

North Carolina: Magnetic City, foot of Roan Mountain, 31; Roan Mountain
(altitude 3000 feet), 1.

Ohio: Cleveland, 1; EUsworth, 2; Ravenna, 3.1°

Ontario: St. Catherines, 1.

Pennsylvania: Allegheny County, 1; Brownsburg, 1;11 Carnot, 1;12 Erie, 1;1°
Leasuresville, 9; New Lexington, 1;}? Ohiopyle, 1;!* Pittsburgh, 4;!2 Warren,
1.12

Quebec: Meaches Lake, Wright County, 1;'* Quebec, 1.

Vermont: East Wallingford, 5.

Virginia: Mountain Lake, 1; Mount Rogers, 1.

West Virginia: Cranberry Glades, 1; Franklin, 1; Traveller’s Repose, 1; Wet-
zel County, 1; White Sulphur Springs, 15.8) 9 19 12

Genus CONDYLURA Illiger.

Condylura Tliger, Prod. Syst. Mamm. et Avium, p. 125, 1811.

Talpasorex Schinz, Cuvier’s Thierreich, vol.1, p.191, 1821. Nec Talpasorex Lesson, 1827.

Astromycter Harris, Amer. Journ. Sci. and Arts, vol. 9, p. 400, June, 1825 (from
Machias, Me., “Star’’ [newspaper]).

Rhinaster Wagler, Nat. Syst. Amphib., p. 14, 1830.

Condytura Todd, Cyclopedia Anat. and Physiol., vol. 2, p. 994, 1839.

1In White, G., Statistics of the State of Georgia, Fauna and Flora, p. 4, 1849.
2 Chamberlain, M., Bul. Nat. Hist. Soc. New Brunswick, no. 3, p. 39, 1884.

2 Whiteaves, J. F., Ann. Rept. Dept. Interior Canada for 1888, part 3 (Geol. Surv.), p. 36, 1889.
4 Goldie, A., Ontario Nat. Sci. Bul., No. 3, p. 40, 1907.

5 Moore, T. J., Ontario Nat. Sci. Bul., No. 3, p. 41, 1907.

6 Nash, C. W., Check List of Vertebrates of Ontario, p. 26, 1906.

7 Collection of Hartley H. T. Jackson.

8 Collection Mus. Comp. Zool., Harvard College.

® Collection Field Mus. Nat. Hist.

10 Collection Amer. Mus. Nat. Hist.

11 Collection Acad. Nat. Sci. Philadelphia.

12 Collection Carnegie Mus.

18 Collection Victoria Mem. Mus.

14 Collection of D. E. Kent, Rutland, Vermont.

1915.] GENUS CONDYLURA. 83

Astromycies Gray, List Spec. Mamm. Brit. Mus., p. 76, 1843.
Astromydes Blyth, Cat. Mamm. Asiat. Soc. Mus., p. 87, 1863.
Condylus Van Hyning, Science, n. 8., vol. 38, p. 243, August 15, 1913.
Type species.—Sorex cristatus Linneus.
Geographic range.—Southeastern Canada and northeastern United
States, from southern Labrador, central Quebec and Ontario, and

Fra. 18.—Geographic range of the species Condylura cristata.
southeastern Manitoba, south to northeastern Illinois and northern
Indiana and Ohio; in the Atlantic coast region south to Virginia
(Dismal Swamp) and Georgia (Marlow) ; and in the Appalachian Moun-
tains to western North Carolina (fig. 18).

External characters.—Body semirobust, not much depressed. Tail
relatively long (about equal in length to body without head),

84 NORTH AMERICAN FAUNA, [No. 38. °

distinctly annulated, scaled, and covered with coarse blackish hairs;
in summer the tail is slender, slightly depressed, gradually tapering
apically, and slightly constricted proximally; in winter it is greatly
veae thick and fleshy, gradually tapering apically, and abruptly
much constricted at the base (fig. 19). Head narrow,
not much depressed. Snout moderately elongate, ter-
minating in a naked disk surrounded on its margin by
a fringe of twenty-two nasal processes symmetrically
arranged, eleven on each side of a median line (fig. 20).
Nostrils nearly circular, situated in anterior surface of
nasal disk. Kyes small (relatively large for Talpide;
larger than in Scalopus, Scapanus, or Parascalops), con-
cealed in fur. Auricular orifice large. Legs short;
weaker than in Scalopus, Scapanus, or Parascalops.
Feet large, not especially fleshy, scaled, scarcely haired
above, naked below. Fore feet handlike, the palms as
broad as long, toes relatively long; first, second, third,
and fourth toes each with three flat, triangular pro-
cesses on outer inferior edge, the basal process on the
first toe being small and inconspicuous. Fore claws
relatively long, broad (but narrower than in Scalopus,
Scapanus, or Parascalops), depressed, the superior sur-
face turned obliquely outward (fig. 21). Hind feet rel-
atively long and narrow, broader apically than basally;
a large depressed triangular plantar tubercle lies near
the base of the first digit, and distally and inwardly on
the sole are five small tubercles. Hind claws relatively —
long, laterally compressed, the outer surface turned
obliquely upward (fig. 22). Toes not webbed. Fur
dense, long, and silky (relatively coarser than in Scalo-
pus, Scapanus, or Parascalops) ; some of the hairs longer
and coarser than the major portion, which forms a dis-
B2025-103 tinct underfur. Mamme, 8: latero-pectoral, 2-2;
Fa. 19—Tail of Latero-abdominal, 1-1; inguinal, 1-1.
Condylura crista- Skeletal characters.—Clavicle relatively long and nar-
i 1D Aw vow (for Talpide), length about twice the breadth,
umnal enlarge- . : :
mentscarcely bee Slightly concave superiorly and convex inferiorly, not
For he iw’ penetrated by a foramen. Humerus about two-thirds
from Egelston aS broad as long, much weaker medially than in Para-
township, Mus- gcglops. Pelvis narrow, bones of opposite sides not
ae en touching under acetabula; no osseous bridges connect-
1906. ing sacral vertebree with ischium. Superior surface of
last sacral vertebra with a distinct, flat, deltoid, longitudinal process.
Os falciforme small but distinct, short, reaching proximal end of first
metacarpal; moderately broad, not much tapering distally.
Skull long and narrow, not much flattened, with relatively high
and narrow braincase, not constricted interorbitally, tapering

RS ae Ss SS > e
= bee 3 &
= 2 ae =
teh Soe oe

1915.] GENUS CONDYLURA. 85

distally. Mastoidsweak. Interparietal large, broad, irregularly cres-
centic, anteriorly deeply emarginate medially, posteriorly broadly

B2026-103
Fic. 20.—Snout of C. cristata (X 13). Individual referred to in fig. 19.

emarginate with a posterior median projection. Frontal region slop-
ing ventrally anteriorly; frontal sinuses scarcely swollen. Rostrum
long and narrow, in
adults with a distinct
basal supero-median
erest; infero-anterior
ends of premaxille ex-
tending much beyond
nasals; supero-anterior
ends of premaxille bare-
ly reaching beyond na-
sals; nasals in adults Sasa
terminating anteriorly in Fig. 21.—Fore foot of C. cristata (X 13). Individual referred to in
an acute median point. He 10:
Anterior nares opening obliquely upward. Zygomata short, narrow,
straight, directed obliquely downward anteriorly, the posterior end
well forward on squamosal. Fora-
men magnum elliptical, large. In-
fraorbital foramen very large, the
plate forming its outer wall rela-
tively very narrow. Audital bulle
incomplete; no auditory meatus.
External pterygoid region scarcely
inflated. Mesopterygoid space long
and broad, the sides converging pos-
teriorly (pterygoids, however, slight-
ly diverging). Palate elongate, very
i narrow, terminating opposite an-
eee Bae terior border of last molar; posterior
vidual referred to in fig. 19. border of palate deeply emargi-
nate, frequently with a median spine. Anterior palatine foramina
‘minute; posterior palatine foramina very large and conspicuous,

86 NORTH AMERICAN FAUNA. [No. 38.

the first pair! reniform or elliptical-oval and about twice as large as
the second pair which are usually elliptical. Horizontal ramus of
mandible weak, curved upward posteriorly, straight anteriorly; coro-
noid moderately long, somewhat broad, but relatively weak, erect,
slightly acute; angle of mandible very long and slender (much longer
and narrower than coronoid), the inferior edge turned inward; in-
ferior mandibular notch large and angular.

Denial characters ——F¥irst upper incisor large, semiovate, curved
inward and directed anteriorly; second upper incisor linear, minute,
lying close under base of first incisor; third upper incisor lateral,
large, elongate, caninelike, with a small postero-basal tubercle dis-
appearing with age. Upper canine short and slender (less than half
dimensions of third incisor), conical. First, second, and third upper
premolars small, laterally compressed, successively increasing in size
posteriorly, each with an anterior and a posterior basal tubercle;
fourth upper premolar similar to third, much larger, with an interior
basal cusplike process. Upper molars W-shaped in transverse sec-
tion, with an interior basal shelf having an indistinctly tricuspidate
edge; first and second molars nearly subequal, the third much
smaller.

First lower incisor moderate in size, spatulate, flat, directed ante-
riorly; second incisor close and superior to first, similar in shape,
much smaller (about half as large), directed anteriorly; third upper
incisor minute, slightly flattened apically, curved downward, directed
anteriorly. Lower canine long, slender, curved posteriorly, with a
rather large posterior basal accessory cusp and a very small anterior
one. Lower premolars small (successively slightly increasing in size
posteriorly), compressed laterally, each with a well-developed poste-
rior cusphike heel and a small cusplike development (minute in the
first premolar) of anterior portion of cingulum. Lower molars M-
shaped in transverse section, laterally compressed; interior shelf
narrow, tricuspidate, the median cusp indistinctly bifid; first and
third molars subequal, the second slightly larger. Dentition: i. 3;
c.+; pm. 4; m. 3; total 44.

CONDYLURA CRISTATA (Linneeus).
Star-Nosep Mote.
(PE IV; fig. 12; PIV V, fies.6, 6a;. PIV ties 2)

Sorex cristatus Linnzeus, Syst. Nat., ed. 10, vol. 1, p. 53, 1758.

Talpa longicaudata Erxleben, Reg. Anim., p. 118, 1777. Based on Pennant’s long-
tailed mole; type locality, North America.

Talpa Cristata Zimmermann, Specimen Zool. Geog., p. 496, 1777.

1Jt may be that these large palatine vacuities are not functional foramina. They have the position,
however, of the first pair of posterior palatine foramina, and, whatever their function, they are of consid-
erable generic diagnostic importance.

1915.] CONDYLURA CRISTATA. 87

Talpa Caudata Zimmermann (nec Talpa caudata of Linneeus and others), Specimen
_ Zool. Geog., p. 497, 1777. Based on Pennant’s long-tailed mole; type locality,

North America.

?Talpa canadensis De La Faille, Naturgesch. des Maulwurtes, t. 1, p. 3, 1778. [Not
seen. |

Talpa radiata Shaw, Gen. Zool., Mamm., vol. 1, p. 523, 1800. New name for Sorex
cristatus Linnzeus.

Sorex radiatus Shaw, Gen. Zool., Mamm., vol. 1, p. 531, 1800. Based upon a igure
and description of De La Faille. Tne eer aiee. (nde.

Scalopus cristatus Geoffroy, Cat. Mamm. Mus. Nat. Hist. Nat., p. 77, 1803.

Scalops cristatus Fischer, Zoognosia, vol. 3, p. 156, 1814.

Condylura cristata Desmarest, Journ. de Physique, de Chimie, d’Hist. Nat. et des Arts,
vol. 89, p. 230, September 1819.

Condylura longicaudata Desmarest, Journ. de Physique, de Chimie, d’Hist. Nat. et
des Arts, vol. 89, p. 232, September, 1819.

Tal{pasorex] cristata Schinz, Cuvier’s Thierreich, vol. 1, p. 191, 1821.

Condylura fissipes Schinz, Cuvier’s Thierreich, vol. 1, p. 191, 1821. (In synonymy.)

Talpa flava Schinz (nec Talpa flava Zimmermann), Cuvier’s Thierreich, vol. 1, p. 191,
1821. (In synonymy.)

| Talpa] purpurascens Shinz(nec Talpa purpurascens Shaw qui Talpa europxa Linneus),
Cuvier’s Thierreich, vol. 1, p. 191, 1821. (In synonymy.)

Condylura macroura Harlan, Fauna Amer., p. 39,1825. Type locality, New Jersey.

Astromycter prasinatus Harris, Amer. Journ. Sci. and Arts, vol. 9, p. 400, June, 1825
(from Machias, Me., “Star’’ [newspaper]). Type locality, Maine.

Condylura prasinata Harris, Boston Journ. Philos, and Arts, vol. 2, p. 582, July, 1825.

Talpasor[ex] longicaudata Schinz, Cuvier’s Thierreich, vol. 4, p. 312, 1825.

?Talpasorer fissipes Minding, Geog. Vertheilung der Sidugethiere, p. 64, 1829.
(Nomen nudum.)

astromycler prarinatus (sic) Rafinesque, Atlantic Journ. and Friend of Knowledge, vol.
1, p. 61, Summer 1832.

Condytura cristata Todd, Cyclopedia Anat. and Physiol., vol. 2, p. 996, 1839.

Rh{inaster] cristatus Wagner, Suppl. Schreber’s Saiugethiere, vol. 2, p. 114, 1841.

Rh{inaster] macrurus Wagner, Suppl. Schreber’s Saiugethiere, vol. 2. p. 115, 1841.

Rh{inaster] longicaudatus Wagner, Suppl. Schreber’s Siugethiere, vol. 2, p. 115, 1841.

Rh{inaster] macroura Wagner, Suppl. Schreber’s Saiugethiere, vol. 2, p. 117, 1841.

Talpa (Condylurus) cristata Blainville, Osteographie, Atlas 1, tables des planches, p. 4;
fasc. 6, Insectivores, pl. 1 (skeleton), pl. 5 (skull), pl. 9 (teeth), 1839-1864.

Astromydes (sic) cristatus Blyth, Cat. Mamm. Asiat. Soc. Mus., p. 87, 1863.

Condylus (sic) cristata Van Hyning, Science, n. s., vol. 38, p. 243, August 15, 1913.

Type locality.—EKastern Pennsylvania.

Geographic range.—That of the genus (see p. 83).

General characters.—Size medium; color dark, blackish; tail long,
about equal in length to body (without head), in autumn and winter
much enlarged, slightly shortened, but constricted at base, scaly,
haired; nostrils anterior, in nasal disk surrounded by twenty-two
fleshy processes; skull relatively long and narrow; audital bulle incom-
plete; premaxillee much extended beyond nasals anteriorly; first upper
incisors large, incurved, and projecting anteriorly.

Color.—Fresh pelage: Upperparts blackish brown to nearly black;
underparts paler and more brownish, fuscous to fuscous-black; tail

88 NORTH AMERICAN FAUNA. [No. 38.

similar to back, sometimes indistinctly bicolored. Worn pelage:
Paler and more brownish than fresh pelage; upperparts fuscous to
fuscous-black; underparts fuscous to hair-brown; wrists frequently
with a narrow ring varying from pinkish buff to clay color. Nasal
disk and processes in live animals rose color. Young usually paler
and more brownish than adults.

Skull.—Size medium (length about 34 mm.), elongate, narrow
(breadth across mastoids about 13 mm.), not depressed postorbit-
ally ; braincase moderately high and arched; interparietal wide antero-
posteriorly; audital bulle incomplete; premaxille much extended
beyond nasals anteriorly; adults with distinct crest between posterior
halves of nasals; zygomata short, narrow, straight, directed obliquely
downward anteriorly; palate narrow; dentition weak; first upper
incisor broad, incurved, projecting anteriorly; second upper incisor
minute; third upper incisor long, narrow, caninelike, in young with
a small postero-lateral basal tubercle; upper molars with indistinctly
trilobed inner basal ledge.

Measurements.—Average of 10 adult males from Digby, Nova
Scotia: Total length, 202.2 (189-211); tail vertebrae (summer), 78.4
(71-83.5); hind foot, 28.1 (26-30). Average of 2 adult males from
Washington, D. C.: Total length, 184 (183-185); tail vertebree (win-
ter), 65.5 (65-66); hind foot, 28 (28-28). Skull: Adult (male ?)
from Holmesburg, Pa.: Greatest length, 34.1; palatilar length, 12.9;
mastoidal breadth, 13.1; interorbital breadth, 7; maxillary tooth
row, 11; mandibular molar-premolar row, 11.3. Average of 3 skulls
of adult males from Locust Grove, N. Y.: Greatest length, 34.6
(34.1-35.2); palatilar length, 13.2 (18-13.3); mastoidal breadth, 13.6
(13.5-13.7) ; interorbital breadth, 7.2 (7.1-7.2); maxillary tooth row,
11.5 (11.4-11.5); mandibular molar-premolar row, 11.6 (11.5-11.7).
Skull of adult female from Locust Grove, N. Y.: Greatest length,
35; palatilar length, 13.3; mastoidal breadth, 13.4; interorbital breadth,
7.3; maxillary tooth row, 11.5; mandibular molar-premolar row, 11.9.
Average of 10 skulls of adult males from Digby, Nova Scotia:
Greatest length, 33.9 (83.1-35) ; palatilar length, 13 (12.9-13.6) ; mas-
toidal breadth, 13.4 (13-14); interorbital breadth, 7.2 (7-7.4);
maxillary tooth row, 11.1 (10.6-11.5); mandibular molar-premolar
row, 11.3 (11-11.9). Average of 2 skulls of adult males from Wash-
ington, D. C.: Greatest length, 33.8 (83.6-33.9) ; palatilar length, 13.1
(12.7-13.5) ; mastoidal breadth, 12.7 (12.6-12.8) ; interorbital breadth,
6.8 (6.7-6.8); maxillary tooth row, 11.2 (11.1-11.2); mandibular
molar-premolar row, 11.2 (11.1-11.3).

Remarks.—Linneus’s description, in 1758, of the star-nosed mole
under the name Sorex cristatus, appears to be its first mention in
literature; on the authority of Kalm, Pennsylvania is given as its

1915.] CONDYLURA ORISTATA, 89

habitat. Pennant,1! in 1771, described two moles under the names
‘““Radiated Mole” and ‘‘Long-tailed Mole’’; the former he correctly
considered to be the Sorex cristatus of Linnseus; his long-tailed species
must also be referred to Condylura cristata since he describes it as a
mole with a radiated nose and a tail two inches long, a description
which applies to no other mammal. Pennant’s description of the
long-tailed mole was probably based upon either a young or a summer
specimen in which the diameter of the tail was minimum; as early as
1777 this became the basis for two Latin binomials, Talpa caudata
Zimmermann? and Talpa longicaudata Erxleben.*’ The writer has
been unable to verify the name Talpa canadensis of De La Faille.
Shaw * renamed Sorex cristatus Linneus, calling it Talpa radiata;
he also recognized Talpa longicaudata, but in his remarks under
Talpa radiata states: ‘It is, perhaps, in reality no other than a
variety of the former species (1. e. T. longicaudata), or a sexual
difference.” > Shaw ® again renamed the species when he confused
a figure and description of it, given by De La Faille, with the genus
Sorex, and called it Sorex radiatus. That Shaw should thus have
been misled is strange, since he writes:

One would be inclined to think that the remarkable moniliform appearance of the
tail in this animal, as exhibited in M. de la Faille’s figure, may be partly owing to the
contraction of the interstices of the joints in drying.

It is evidently allied to the radiated Mole, but if the figure given by M. de la Faille
be accurate, must surely be a very distinct species.’

Schinz,® when substituting the generic name Talpasorex for Con-
dylura, used the name Condylura jfissipes in synonymy under Talpa-
sorez cristatus; it seems probable that the name fissipes had been
used in literature previous to this, but the present writer has been
unable to find an earlier usage. Schinz (loc. cit.) also placed the
names Talpa flava and Talpa purpurascens in synonymy under
Talpasorex cristatus; this was probably purely an error on his part,
since there is nothing in the original descriptions of Talpa flava
Zimmermann and Talpa purpurascens Shaw, nor in subsequent
descriptions of these forms, which would lead one to confuse either
with Condylura Illiger; the former name is a synonym of Scalopus a.
aquaticus (Linneus), the latter of Talpa europea Linneus. In 1825,
two other names were proposed which apply to the star-nosed mole—

1 Pennant, T., Quadrupeds, 1771. The present writer has not seen this work but presumes the descrip-
tions are essentially the same as those in Pennani’s History of Quadrupeds, ed. 3, vol. 2, p. 232, pl. 90, 1793.

2Zimmermann, E. A. W., Spec. Zool. Geog., p. 497, 1777.

8 Erxleben, J. C. P., Syst. Reg. Anim., p. 118, 1777.

4 Shaw, George, Gen. Zool., Mamm., vol. 1, p. 523, 1800.

6 Shaw, loc. cit., p. 524.

6 Shaw, loc. cit., p. 531.

7 Shaw, loc. cit., p. 532.
8 Schinz, H. R., Cuvier’s Thierreich, vol. 1, p. 191, 1821.

90 NORTH AMERICAN FAUNA. [No. 38.

Condylura macroura Harlan: and Astromycter prasinaius Harris.?
Both Harlan and Harris based their descriptions upon animals whose
tails were in the enlarged winter condition. Harris was further
deceived by his specimen being discolored with a shade of green, and
proposed the new genus Asiromycter to include this animal, though
later ? he returned it to the genus Condylura.

It is to be remarked that Condylura cristata has comparatively
little geographic variation over a wide zonal range; specimens from
the Lower Austral Zone, in Georgia and Virginia, are subspecifically
inseparable from those from the Boreal Zone in Quebec and Labra-
dor. A very slight increase in size is noticeable toward the northern
part of the range of the species, but it is inconstant, and insufficient
for subspecific recognition. Occasional local variations appear, but
when a large series is examined from any one locality these prove
inconstant, or else crop out in remote localities. Thus, in a series
from Digby, Nova Scotia, the skulls have on the average very slightly
higher braincases and more abruptly sloping frontals, but the differ-
ence is not constant and can be matched perfectly by specimens from
New York, Massachusetts, Pennsylvania, and Maryland; the same
is true of a small postpalatal process present in the majority of skulls
in the series from Digby. A specimen from Marlow, Georgia, is an
alcoholic from which the broken and imperfect skull has been re-
moved for study; it offers no characters by which it can be separated
from typical specimens from Pennsylvania. Possibly when larger
series of adult specimens are available differences of diagnostic
value not now discernible will become apparent.

The limits of the geographic range of Condylura are not satisfac-
torily known. Many published records of the star-nosed mole have
certainly been based upon erroneous identifications. For example,
the species is included among the mammals supposed to occur in
Oklahoma, and Townsend® lists it from ‘the Territory of the
Oregon.”

Richardson © describes a specimen said to have been taken by
Douglas on the “‘banks of the Columbia.”’” Richardson’s description
clearly indicates that the animal he had in hand. was Condylura
cristata, but it is almost certain that it was not collected near the
banks of the Columbia River. The most westerly known point of
the range of the species is in Manitoba, where, according to Seton,’

1 Harlan. R., Fauna Amer., p. 39, 1825.
2 Harr, T. W., Amer. Journ. Sci. and Aris, vol. 9, p. 400, June, 1825,

8 Harris, T. W., Boston Journ. Philos. and Arts, vol. 2, p. 582, July, 1825.

4 Woodhouse, S. W., Report of an Expedition Down the Zuniand Colorado Rivers, by Capt. L. Sitgreaves;
Mammals, p. 43, 1854.

5 Under the names Condylura longicaudata and Condylura macroura. Townsend, J. K., Narrative of
Journey across the Rocky Mountains to the Columbia River, p. 313, 1839.

6 Under the name Condylura macroura. Richardson, J., Fauna Boreali-Amer., vol. 1, p. 284, 1829.
7 Seton, Ernest Thompson, Life-Histories of Northern Animals, vol. 2, p. 1137, 1909.

1915.] CONDYLURA CRISTATA, 91

specimens have been brought to the taxidermist shop of W. R. Hine
in Winnipeg. Cory! records a specimen from Warsaw, IU., and
Professor Frank Smith, of the University of Ilinois, found a dead
specimen in the vicinity of Urbana in the same State.’

Specimens examined.—Total number 218, as follows:

Connecticut: East Hartford, 1; Liberty Hill, 1;? Norfolk, 1.

District of Columbia: Washington, 10.

Georgia: Marlow, 1.

Labrador: Black Bay, 1;* Hamilton Inlet, 2; Paradise, 2; Saint Michael de
1; Sandwich Bay, 2.

Maine: Eastport, 1; Freeport, 1; Oakland, 1; Penobscot River (Hast Branch), 2;
Small Point, 2.

Maryland: Brookeville, 1; Cabin John, 1; Chevy Chase, 1; College Park, 2;
Glendale, 1; Laurel, 2; Marshall Hall, 1; Prince Georges County, 1; Williams-
port, 1; Woodside, 2.

Massachusetts: Belmont, 4;? Gardner, 1; Lunenburg, 4; Medway, 1; ? Middle-
boro, 2; New Bedford, 1; Newburyport, 2; Seehonk, 1;* Watertown, 2;3
metic own, ip Watrineton: 4,

Michigan: Ann hon 8;° Chelsea, 1;° Douglas Lake, 1;5 Geddes, 1;° Goge-
bie 1; © Hancock, 1; ° Ralantareo County, 1;° iinaheear Connie 1; Por-
cupine Mountains, 2;° Portage Lake, 6. °

Minnesota: Elk River, 1; Fort Ripley, 1; Margie, 1.

New Brunswick: Arthurette, i;’ Hampton, 1.

New Hampshire: Ossipee, 4; Webster.?

New Jersey: Lake Hopatcong, 5; * Tuckahoe, 1.4

New York: Cross River, 1; Essex County, 1; Geneva, 2; Highland Falls, 2;7
Ithaca, 1;°® Lake George, 2; Lansing, 2; ° Lockport, 1; Locust Grove, 11;
Lyons Falls, 1; New York, 2; Nichols, 1; Oswego, 1; Pelham, 1; Peterboro, 5;
Rockland, 1;7 Saint Lawrence County, 1; Sing Sing, 3

~North Carolina: Magnetic City, 2; Weaverville, 1.3

Nova Scotia: Barrington Passage, 1; !° Digby, 18; Halifax, 7; James River, 1
Newport, 2.’

Ohio: Cleveland, 1; Ellsworth, 1; Garrettsville, 3.

Ontario: Landsowne, 1;7 Middlesex County, 1;!° Moose Factory, 1; Muskoka,
1;7 Ottawa, 4.

Pennsylvania: Carlisle, 1; Lake Leigh, 1;4 Meadville, 2; New Lexington, 2;4,"
Philadelphia, 1;* Radnor, 1.4

Quebec: East Main River, 1; Godbout, 3; Lake Edward, 1;? Montreal, 1.

Vermont: East Wallingford, 4; 12 Mount Mansfield, 2; Rutland, 1.8

Virginia: Dismal Swamp, 4.

Wisconsin: Colby, 1; Medford, 2; ® Merrill, 1; ® Newald, Forest County, 1.°

’

1 Cory, C. B., Field Mus. Nat. Hist., publ. 153, zool. series 11, p. 444, 1912. In apersonal letter dated March
20, 1914, Mr. Cory writes in regard to this specimen: “On a number of occasions Mr. Charles K. Worthen
sent me boxes of birds and mammals, from which I could select specimens TI desired and send back the
others. One lot contained a specimen of Condylura cristata simply labeled ‘ Warsaw’ (in pencil) with no
other data. Thespecimen was returned to him.’’

2 Wood, F. E., Bul. Illinois State Lab. Nat. Hist., vol. 8, p. 588, 1910.

3 Collection Mus. Cemp. Zool., Harvard College.

4 Collection Acad. Nat. Sci. Philadelphia. 9 Collection Field Mus. Nat. Hist.

5 Collection Univ. Michigan Mus. -10 Collection Victoria Mem. Mus.

6 Collection Milwaukee Public Mus. . 11 Collection Carnegie Mus.

7 Collection Amer. Mus. Nat. Hist. 12 Collection of D. E. Kent, Rutland, Vt.

8 Collection of Hartley H. T. Jackson. 13 Collection of G. L. Kirk, Rutland, V+.

99° NORTH AMERICAN FAUNA. ENo. 38.

Genus NEUROTRICHUS Giinther.

Netirotrichus Giinther, Proc. Zool. Soc. London, 1880, p. 441, October, 1880.
Néurotrichus Giinther, Proc. Zool. Soc. London, 1880, plate 42, October, 1880.
Neurotrichus Forbes, Zool. Record, vol. 17, Mammalia, p. 14, 1881.
Neourotrichus Rye, Zool. Record, vol. 17, Index, p. 8, 1881.

Type species.— Urotrichus gibbsit Baird.

Geographic range.—Pacific coast region of North America west of
the Cascades and Sierra Nevada, from southwestern British Columbia

oy osnaesctaers 5 Skene Te , (fraser River region) south to
Fremont Peak, Monterey County,
California (fig. 23).

External characters.—Smallest of
the American Talpide; body some-
what robust, not much depressed.
Tail moderate in length (about half
as long as head and body), moder-
ately fleshy, constricted at base,
scaled, very distinctly annulated,
sparsely covered with coarse hairs
(fig. 24). Head conoidal, relatively
long, not much depressed. Snout
elongate, terminating in a naked
disk or pad, apical superior surface
naked to line of anterior edge of
nasals; nostrils lateral in terminal
pad, slightly crescentic, with an-
terior end of crescent enlarged and
its concavity upward (fig.25). Eyes
minute (relatively about asin Con- —
dylura), concealed in the fur. Au-
ricular opening large. Legs short,
eee) weak (relatively as in Condylura).

B2029-103 Heetlarge, notfleshy, scaled, sparse-
Fig. 23.— Geographic range ‘of the subspecies of “1— haired above, iakedtselwe more

eine ie ae Te fot hardly handlike, the palms
longer than broad, toes relatively long. Fore claws relatively
long, not broad, not depressed (fig. 26). Hind feet long and
narrow, broader distally than proximally; six tubercles (varying
slightly in position) on each hind foot, usually located one at
base of third digit, one interdigital between the third and fourth
digits, one between the fourth and fifth digits, one postero-
interdigital between the second and third digits, and two near the
center of the sole. Hind claws moderately long and slender, slightly
compressed laterally, acute (fig. 27). Toes not webbed. Pelage
similar in general to that of Condylura, but shorter, finer, and with

1915.] GENUS NEUROTRICHUS. 93

underfur less clossly defined. Mamme, 8: latero-pectoral, 2-2;
latero-abdominal, 1-1; inguinal, 1-1.

Skeletal ee Cleicle relatively long and narrow (for the
Talpide), length about twice the breadth; concave supeonys infe-
rior surface with a flat process projecting ogee ae
ally; not penetrated by a foramen. Humerus about
three-fifths as broad as long, weaker medially (as in
Condylura). Pelvis narrow, bones of opposite sides sep-
arated by considerable space (about 2 mm.) under aceta-
bula; no osseous bridges connecting sacral vertebree with
ischium. Superior surface of last sacral vertebra with-
out process. Os falciforme rudimentary and scarcely
perceptible.

Skull conoidal, moderately depressed, without promi-
nent processes and ridges, with moderately broad brain-
case, Be eealy constricted interorbitally. Mastoids weak.
Interparietal large, broad, irregularly semicircular, ante-
riorly deeply emarginate medially, posteriorly slightly
emarginate and usually without a posterior median pro-
jection. Frontal region scarcely sloping ventro-anteri- A
orly; frontal sinuses very slightly swollen. Rostrum ri. 24—rail of
moderately elongate; anterior ends of premanillze Ot) epee de Pt
much thickened, extending beyond nasals, forming a — jnus(x1!). No.
somewhat truncate notch anterior to the sane TaN eee ewes Se

- : Nat. Mus., Bi-
rior nares opening forward. Zygomata short, narrow, ological survey
slightly out-curved, directed slightly downward anteri- “ollection; from

- ss Aptos, Cal.
orly, the posterior end forward on squamosal (less so

than in Condylura). Foramen magnum elliptical, relatively large.
Infraorbital foramen large, the plate forming its outer wall relatively
broad and heavy. Audital bulle incomplete; no auditory meatus.
External pterygoid region scarcely inflated (relatively more than in
Condylura). Mesopterygoid space moder-
ately long and broad, the sides usually gent-
ly concave. Palate moderately elongate,
po relatively narrow, terminating opposite pos-

p2031-103 terior border of last molar; posterior border

Fic. 25—Snout of Neiirotrichus of palate slightly emarginate, without spine

gibbsit gibbsii (X13). No. 83468, é 3 c

U. §. Nat. Mus.: from British Ornotch. Anterior palatine foramina mod-

Columbia. erate, oval; first pair of posterior palatine
foramina moderate, oval to elliptical-oval; second pairminute. Hor-
izontal ramus of mandible slender, nearly straight; coronoid rela-
tively long, erect proximally, curving posteriorly distally into an
acute process; angle of mandible relatively short and narrow
(smaller than coronoid) ; inferior mandibular notch relatively shallow
and broad.

94 NORTH AMERICAN FAUNA. [No. 38. °

Dental characters.—First upper incisors very short, relatively broad;
second and third small (about half as large as first incisor), somewhat
laterally compressed. Upper canine larger than third incisor, trian-
gular, compressed, two-rooted, with a small postero-basal accessory
cusp. First upper premolar similar to canine,
slightly smaller; second upper premolar large,
triangular, with a postero-basal cusp, a pos-
tero-internal basal cusplike heel, and some-
times with an anterior cusplike process on the
cingulum, which tends to disappear with age.

et nlate Upper molars W-shaped in transverse section,

eee Nn ied with an interior basal shelf having a distinctly

(X13). Individual referredtoin bicuspidate edge; first and second molars sub-
Heo equal, the third much smaller.

First lower incisor small, spatulate, directed slightly antero-
interiorly; second and third lower incisors and lower canine similar
to first incisor, smaller, successively decreasing in size posteriorly,
directed obliquely anteriorly, the canine with a smaller posterior
basal tubercle. Lower premolars triangular,
each with a posterior cusplike heel and an
indistinct posterior median sulcus, the second
about twice as large as first and with a small
antero-basal tubercle. Lower molars M-
shaped in transverse section, shghtly com-
pressed laterally (less so than in Condylura) ;
interior shelf low and narrow, tricuspidate,
the median cusp not bifid; first molar with

je
B2033-103

; : Fic. 27.—Hind foot of NV. g. gibbsii
minute basal tubercle between posterior and — (x 13). Individual referred to in

median interior cusps; first and second mo-_ 48-4.
lars subequal, the third smaller. Dentition: 1. 3; c. +; pm. 3; m. 3;
total 36.

Key to Subspecies of Netirotrichus.

a.! Size smaller; color paler; length of skull less than 23.2 mm.,
Neiirotrichus gibbsit gibbstvi (p. 94).
a.* Size larger; color darker; length of skull more than 23.2 mm.,
Neiirotrichus gibbst hyacinthinus (p. 97).

Descriptions of species and subspecies of Netirotrichus.

NEUROTRICHUS GIBBSII GIBBSII (Baird).
GiBBs’s MOLE.
CEL VI, figs tas dio 225)

Urotrichus gibbsit Baird, Report Pacific R. R. Survey, vol. 8, part 1, Mammals, p. 76,
1857.

Urotrichus Gibsii Lord, Naturalist in Vancouver Island and British Columbia, p. 338,
1866.

———————

1915.] NEUROTRICHUS GIBBSII GIBBSII. 95

Néurotrichus (sic) gibbsit Giinther, Proc. Zool. Soc. London, 1880, pl. 42, October, 1880.

Neiirotrichus gibbsti True, Proc. U. 8. Nat. Mus., vol. 7, p. 607, 1885.

Neurotrichus gibbsit Bryant, Zoe, vol. 1, p. 359, February, 1891.

Neurotrichus gibbsi Merriam, Mazama, vol. 1, p. 228, 1897.

Neurotrichus gibbsi major Merriam, N. Am. Fauna No. 16, p. 88, October 28, 1899.
Type locality, Carberry Ranch, altitude 4100 feet, between Mount Shasta and
Mount Lassen, Shasta County, California.

Neiirotrichus gibbsit gibbsii Miller, U.S. Nat. Mus., Bul. 79, p. 11, December 31, 1912.

Netirotrichus gibbsti major Miller, U. 8S. Nat. Mus., Bul. 79, p. 11, December 31, 1912.

Type locality—White River Pass, north of Mount Rainier, Pierce
County, Washington.

Type specimen.—No. 3882;, U. 5. Nat. Mus.; immature, sex un-
known; poorly made skin and fragmentary skull; collected July 15,
1854, by George Gibbs.

Geographic range.—Extreme southwestern British Columbia, west-
ern Washington and Oregon west of the Cascade Mountains, south in
the coast region to Eureka, Humboldt County, Cal., and in the inte-
rior, west of the Sierra Nevada, to South Yolla Bolly Mountain, Cal.

General characters.—Size small (total length averaging less than
120 mm.); color dark, usually dark mouse gray; tail medium in size
(about one-third of total length), scaled in transverse annular rows,
covered with a few blackish, coarse hairs; skull small (greatest
leneth usually less than 23 mm.), flat, and not angular.

Color.—General tone dark mouse gray to blackish mouse gray,
occasionally dusky neutral gray; upperparts and underparts essen-
tially the same color, the underparts rarely shghtly paler than the
back; the longer hairs in full pelage frequently tipped with whitish,
producing a frosted appearance; recently killed animals and speci-
mens little handled generally show purplish and greenish iridescence.
The worn pelage is paler and more brownish than the fresh.

Skull—Small (greatest length usually less than 23 mm.), smooth,
flat, not much depressed postorbitally, not much constricted inter-
orbitally; frontal sinuses slightly swollen; zygomata short, weak;
pterygoids short, weak, laterally flattened; audital bulle incomplete;
rostrum moderate in length and width; dentition moderate; first
upper incisors flat and rodentlike; upper canine (third lateral tooth)
flattened laterally, much like first upper premolar; anterior portion
of cingulum of second upper premolar not usually developed into a
superior cusplike process.

Measurements.—Average of 9 adult males from Sumas, British
Columbia: Total length, 113 (107-117); tail vertebra, 37.1 (34-39);
hind foot, 16.6 (15.7-17). Average of 5 adult females from Sumas,
British Columbia: 116.6 (111-120); 36.4 (83-40); 17 (17-17). Av-
erage of 3 males from Carberry Ranch (type locality of N. g. major),
Shasta County, Cal.: 119.7 (118-121); 40.3 (89-42); 17 (17-17).
Skull: Average of 10 skulls of adult males from Sumas, British Co-
lumbia: Greatest length, 22.5 (22.1-23); palatilar length, 9.4 (9.1-9.6);

96 NORTH AMERICAN FAUNA, - [No. 38. °

mastoidal breadth, 10.3 (10.1-10.5); interorbital breadth, 5.3
(5.2-5.4); maxillary tooth row, 7.1 (7-7.2); mandibular molar-
premolar row, 7.3 (7.1-7.5). Average of 6 skulls of adult females
from Sumas, British Columbia: Greatest length, 22.4 (21.5-23);
palatilar length, 9.4 (9.2-9.5); mastoidal breadth, 10.2 (9.6-10.5);
interorbital breadth, 5.3 (5.1-5.5); maxillary tooth row, 7.1 (6.9-7.2);
mandibular molar-premolar row, 7.2 (7-7.3). Average of 2 skulls
of males from Carberry Ranch (type locality of major), Shasta County,
Cal.: Greatest length, 22.7 (22.4-23); palatilar length, 9.3 (9.3-9.3);
mastoidal breadth, 10.6 (10.5-10.7) ; interorbital breadth, 5.4 (5.45.4);
maxillary tooth row, 7 (6.9-7); mandibular molar-premolar row, 7
(6.9-7.1).

Remarks.—This little mole, the most shrewlike of the American
members of the family, shows comparatively little geographic varia-
tion throughout its rather extensive range; in fact, the local almost
obscures the geographic variation. Southward a tendency appears
toward an increase in size and toward the development of a
cusplike process upon the anterior portion of the cingulum of the
second upper premolar; this reaches the climax in JN. g. hyacinthinus.
The presence of this cusplike process on the second upper premolar,
however, is not strictly diagnostic since it occasionally crops out,
slightly developed, in specimens taken near the northern limit of the
range of N.g. gibbsw, or may be absent in specimens of hyacinthinus
taken near the southern border of itsrange. Thus, it occurs, weakly
developed, in a few specimens from Sumas, British Columbia; it is
absent in two specimens from Crescent City, Cal., but is present in |
five young adults from Eureka, Cal., and in one from Goldbeach,
Oreg.; some specimens of hyacinthinus from Cuddeback and Aptos,
Cal., have the process, while others lack it.

The form Neurotrichus gibbst major Merriam? is here placed in
synonymy under WN. g. gibbsu. The type of major was collected at
Carberry Ranch, altitude 4100 feet, between Mount Shasta and
Mount Lassen, Shasta County, Cal.; a careful comparison of the type,
topotypes, and other specimens from the Shasta region, with a large |
number of specimens from Washington and British Columbia fails to
show any differences between major and gibbsw worthy of subspecific
recognition; specimens of major average very slightly larger than
gibbsii and in some other respects appear to be intermediate in ©
characters between gibbsit and hyacinthinus, but in size and general
proportions of skull they are much nearer gibbsw. The presence of
an anterior ‘‘cusp” on the cingulum of the second upper premolar in
major is not of diagnostic value, as has been shown in the preceding

1 Merriam, C. Hart, N. Am. Fauna No. 16, p. 88, 1899,

1915. ] NEUROTRICHUS GIBBSII HYACINTHINUS. 97

paragraph, and at best can be considered only an approach toward
hyacinthinus; nor does the second lower premolar differ essentially
from that of gibbsii, since it can be matched almost perfectly in any
large series of the genus Netrotrichus from any locality.

Five specimens examined from Eureka, Cal., are hardly adult,
but are provisionally referred to gibbsti on account of color and
size of skull; when a larger series of specimens, with more adults, is
available from this region, a change of decision may be necessary.

Specimens examined.—Total number, 146, as follows:

British Columbia: Chilliwack Valley, 6; Douglas, 1; Howe Sound, 3; Lang-
ley, 1; Sumas, 72; Tammi Hy Creek (Chilliwack Valley), 1;' Thurstons, 2.

California: Arcata, 1;? Beswick, 1; Carberry Ranch, Shasta County, 3; Crescent
City, 2; Eureka, 5;3 Hoopa Valley, 1; Mount Shasta, 4; Salmon Mountains
(near Eina Mills), 1; South Yolia Bolly Mountain, 1;2 Tower House, Shasta
County, 2;? Trinidad, 3.?

Oregon: Anna Creek, Mount Mazama, 1; Astoria, 1; Crater Lake, 1; Elk Head, 1;
Eugene, 1; Fort Klamath, 4; Goldbeach, 1;3 McKenzie Bridge, 1; Mulinomah
Falls, 1; Salem, 1;* Seaside, 1; Siskiyou, 1; Vida, 2; Yaquina Bay, 3.

Washington: Kirkland, 1; Lake Cushman, 4; Mount Rainier, 2; Mount Vernon,
3; Neah Bay, 1; Seattle, 1; Steilacoom, 2; Tenino, 1; White River, Cascade
Mountains (type locality), 1.

NEUROTRICHUS GIBBSII HYACINTHINUS Bangs.
SouTtTHERN GiBBsS’s Mole.
(PE VE, fies. 2, 2a, 2b,-23.)

Netiroirichus gibbsi hyacinthinus Bangs, Amer. Nat., vol. 31, p. 240, March, 1897.
Neurotrichus gibbsii hyacinthinus Miller & Rehn, Proe. Boston Soc. Nat. Hist., vol.

30, p. 254, December, 1901. :
Neurotrichus gibbsi hyacinthinus Elliot, Field Columb. Mus., publ. 105, zool. series,

vol. 6, p. 467, 1905.

Neiirotrichus gibbsit hyacinthinus Miller, U. 8. Nat. Mus., Bul. 79, p. 11, December 31,
1912.

Type locality.—Nicasio, Marin County, California.

Type specimen.—No. 1240, Mus. Comp. Zool., Harvard College,
Bangs collection; @ adult, skin and skull; collected March 10, 1894,
by C. A. Allen.

Geographic range-—Coast region of California from Cuddeback,
Humboldt County, south to Fremont Peak, Monterey County.

General characters.—Larger than N. g. gibbsii; color usually slightly
darker; skull larger, relatively wider through braincase and rostrum,
with more angular mastoid region.

Color.—Much like that of N. g. gibbsii but averaging darker;
usually blackish mouse gray or dusky neutral gray; often almost
black, especially in fresh pelage; in full unworn pelage sometimes

1 Collection Victoria Mem. Mus. 8 Collection Field Mus. Nat. Hist.
3 Collection Mus. Vert. Zool., Univ. California. 4 Coliection Amer. Mus. Nat. Hist.

90349°—15——7

98 e NORTH AMERICAN FAUNA. [No. 38.

distinctly overcast with hoary, due to whitish tips of the longer hair;
worn pelage paler and more brownish; purplish and greenish irides-
cence shows in fresh specimens, as in gibbsii.

Skull.—Similar to that of N. g. gibbsii but larger (greatest length
usually over 23.6 mm.) and relatively wider through braincase,
interorbitally, and through rostrum; mastoid region usually heavier
and more angular than in gibbsvi; anterior portion of cingulum of
second upper premolar usually developed into a superior cusplike
process. ;

Measurements.—Average of 6 males from Aptos, Santa Cruz
County, Cal.: Total length, 121.5 (118-126); tail vertebre, 38.3
(37-39); hind foot, 16.9 (16.5-17). Skull: Young adult male from
type locality: Greatest length, 24.2; palatilar length, 9.8; mastoidal
breadth, 11; interorbital breadth, 5.7; maxillary tooth row, 7.4;
mandibular molar-premolar row, 7.3. Skull of young adult female
from type locatity: Greatest length, 23.5; palatilar length, 9.6;
mastoidal breadth, 10.6; interorbital breadth, 5.4; maxillary tooth
row, 7.3; mandibular molar-premolar row, 7.3. Average of 6 skulls
of males from Aptos, Santa Cruz County, Cal.: Greatest length,
23.8 (23-24); palatilar length, 9.9 (9.6-10); mastoidal breadth, 10.9
(10.7-11.1); mterorbital breadth, 5.6 (5.5-5.7); maxillary tooth row
7.6 (7.4-7.8); mandibular molar-premolar row, 7.6 (7.5—7.7).

Remarks.—The southern-coast form of Gibbs’s mole, NV. g. hyacin-
thinus, differs from the typical northern form, JN. g. gibbsii, chiefly
in its larger size and darker color, though frequently specimens
are no darker than typical gibbsu. Specimens from the south of San
Francisco Bay are not quite so intensely colored as those from the
type region, and, on the average, seem to have slightly shorter
rostra, but the differences are too trivial for subspecific designation.

Specimens examined.—Total number, 58, as follows:

California: Aplae, 10; Burlingame, 1; Cazadero, 1;! Cuddeback, 7;! Fairfax, 1; '
Freestone, 1;' Fremont Peak, 1; Gualala, 9; Guerneville, 3;* Inverness, 4;?

Mendocino, ot t Nicasio (type locality), 6; Palo Alto, 1;? Pent Arena, 4;1
San Gerouing, 1;! Santa Cruz, 7.

PLATE TI.
[Natural size.]

Fie. 1. Early stage in molting process of Scalopus aquaticus aquaticus (Linneeus);
? adult; Fort Myer, Va., April 15, 1897. (No. 83686, U. S. Nat. Mus.)
2. Middle stage in molting process of Scalopus aquaticus aquaticus (Linnzus);
? adult; Falls Church, Va.,. May 26,1907. (No. 144453, U. S. Nat. Mus.)
3. Nestling young of Scalopus aquaticus howelli Jackson; Jackson, N.C. (No.
7250, U. S. Nat. Mus.)

1 Collection Mus. Vert. Zool., Univ. California. ? Collection Field Mus, Nat. Hist.

North American Fauna No. 38, U. S. Dept. Agr. Biological Survey. PLATE |.

B2001-103

MOLTING AND YOUNG OF SCALOPUS.

1, 2. Scalopus aquaticus aquaticus.
3. Scalopus aquaticus howelli,

90549 °—15—_8

Hrge le

10.

dls

PLATE II.
[Natural size. ]

Scalopus aquaticus aquaticus (Linnzeus); essentially a topotype; ¢ adult;
near Media, Pa. (No. 9845, Acad. Nat. Sci. Philadelphia; 2845, Rhoads
collection. )

. Sealopus aquaticus howelli Jackson; type; ¢ adult; Autaugaville, Ala. (No.

177931, U. S. Nat. Mus., Biological Survey collection. )

. Scalopus aquaticus australis (Chapman); topotype; ¢ adult; Gainesville, Fla.

(No. 7783, Field Mus. Nat. Hist.)

. Scalopus aquaticus anastasx (Bangs); topotype; 3 adult; Point Romo, Anas-

tasia Island, Fla. (No. 7193, Mus. Comp. Zool., Harvard College, Bangs
collection. )

. Scalopus aquaticus parvus (Rhoads); ¢ adult; Port Tampa City, Fla. (No.

7202, Mus. Comp. Zool., Harvard College, Bangs collection.)

. Scalopus aquaticus machrinus (Rafinesque); ¢ adult; Oakwoods Cemetery,

Chicago, Ill. (No. 8155, Field Mus. Nat. Hist.)

. Scalopus aquaticus machrinoides Jackson; type; ¢ adult; Manhattan, ‘Kans.

(No. 169717, U. S. Nat. Mus., Biological Survey collection.)

. Scalopus aquaticus pulcher Jackson; type; % adult; Delight, Ark. (No.

170698, U. S. Nat. Mus., Biological Survey collection.)

. Scalopus aquaticus caryt Jackson; type; ¢ young adult; Neligh, Nebr. (No.

116799, U. S. Nat. Mus., Biological Survey collection.)

Scalopus aquaticus texanus (Allen); topotype; 3 adult; Rockport, Tex.
(No. 51885, U. S. Nat. Mus., Biological Survey collection.)

Scalopus inflatus Jackson; type; young adult, sex unknown; State of Tamau-
lipas, Mexico (45 miles from Brownsville, Tex.). (No. 52709, U. S. Nat.
Mus., Biological Survey collection.)

. Scalopus xreus (Bangs); type; @ adult; Stilwell, Okla. (No. 5475, Mus.

Comp. Zool., Harvard College, Bangs collection. )

North American Fauna No. 38, U.S. Dept. Agr. Biological Survey.

PLATE Il.

B2002-103

SKULLS OF SCALOPUS AQUATICUS (SUBSPP.), S. INFLATUS, AND S. AREUS.

S. a. aquaticus.
S. a. howelli.

. S. a. australis.
S. a. anastasee.

. S. a. parvus.
. S.a. machrinus.

S. a. machrinoides.,

. S. a. pulcher.

9. S. a. Caryl.
10. S. a. texanus.
11. S. inflatus.

. ereus.

Fies. 1, 1a:

PLATE III.
[Natural size. ]

Scalopus aquaticus aquaticus (Linnezeus); essentially a topotype; ¢ adult;
near Media, Pa. (No. 9845, Acad. Nat. Sci. Philadelphia; 2845,
Rhoads coliection.)

. Scalopus aquaticus howelli Jackson; type; ¢ adult; Autaugaville, Ala.

(No. 177931, U. S. Nat. Mus., Biological Survey collection.)

. Scalopus aquaticus australis (Chapman); topotype; ¢ adult; Gainesville,

Fla. (No. 7783, Field Mus. Nat. Hist.)

a. Scalopus aquaticus parvus (Rhoads); 3 adult; Port Tampa City, Fla.

(No. 7202, Mus. Comp. Zool., Harvard College, Bangs collection.)

a. Scalopus aquaticus texanus (Allen); topotype; ¢ adult; Rockport, Tex.

(No. 51385, U. 8S. Nat. Mus., Biological Survey collection.)

a. Scalopus inflatus Jackson; type; young adult, sex unknown; State of

Tamaulipas, Mexico (45 miles from Brownsville, Tex.). (No. 52709, |
U.S. Nat. Mus., Biological Survey collection.)

. Scalopus xreus (Bangs); type; Q adult; Stilwell, Okla. (No. 5475, Mus.

Comp. Zool., Harvard College, Bangs collection.)

North American Fauna No. 38, U. S. Dept. Agr. Biological Survey. PLATE III.

SKULLS OF SCALOPUS AQUATICUS (SUBSPP.), S. INFLATUS, AND S. AEREUS.

1, la. S. a. aquaticus. 3, 3a. S. a. australis. 6, 6a. S. inflatus.
dg é

2, 2a. S. a. howelli. 4, 4a. S. a. parvus. a. §. sereus.

5, da. S. a. texanus.

12 Cetera bs

10.

dle

PLATE IV.
[Natural size. ]

Scapanus townsend (Bachman); 3 adult; Ferndale, Humboldt County, Cal.
(No. 19141, Mus. Vert. Zool., Univ. California.)

. Scapanus orarius orarius True; % adult; Ferndale, Humboldt County, Cal.

(No. 19111, Mus. Vert. Zool., Univ. California.)

. Scapanus orarius schefferi nobis; type; 3% adult; Walla Walla, Wash. (No.

204997, U.S. Nat. Mus., Biological Survey collection.)

. Scapanus latimanus latimanus (Bachman); ¢ adult; Menlo Park, San Mateo

County, Cal. (No. 18779, Mus. Vert. Zool., Univ. California.)

. Scapanus latimanus occultus Grinnell & Swarth; ¢ adult; San Gabriel, Cal.

(No. 30310, U. 8S. Nat. Mus., Biological Survey collection.)

. Scapanus latimanus grinnelli Jackson; type; ¢ young adult; Independence,

Cal. (No. 17785, Mus. Vert. Zool.; Univ. California.)

. Scapanus latimanus sericatus Jackson; type; @ adult; Yosemite, Cal. (No.

(109548, U. S. Nat. Mus., Biological Survey collection.)

. Scapanus latimanus minusculus Bangs; type; 2 young adult; Fyffe, Cal.

(No. 9189, Mus. Comp. Zool., Harvard College, Bangs collection.)

. Scapanus latimanus dilatus True; type; young adult, sex unknown, prob-

ably ¢; Fort Klamath; Oreg. (No. 186628, U. 8. Nat. Mus., Merriam
collection.)

Scapanus anthonyi Allen; type; ¢ adult; San Pedro Martir Mountains, Lower
California. (No. 4947, Amer. Mus. Nat. Hist.)

Parascalops brewer 1(Bachman); ¢ young adult; Leasuresville, Pa. (No. 603,
Carnegie Mus.)

. Condylura cristata (Linneus); ¢ adult; Lunenburg, Mass. (No. 96075, U.S.

Nat. Mus., Biological Survey collection.)

North American Fauna No. 38, U. S. Dept. Agr. Biological Survey. PLATE IV.

B2004-103

SKULLS OF SCAPANUS, PARASCALOPS, AND CONDYLURA.

. S. townsendii. 5. S. 1. occultus. 9. 8.1, dilatus.
S. 0. orarius, 6. S. 1. grinnelli. 10. S. anthonyi.
5. oO. schefferi. . S. 1. sericatus. 11. P. breweri.
5. 1. latimanus. . S. 1. minusculus. 12. C. cristata.

PLATE V%
[Natural size.]

Fies. 1, la. Scapanus townsendii (Bachman); $ adult; Ferndale, Humboldt County

Cal. (No. 19141, Mus. Vert. Zool., Univ. California.)

2, 2a. Scapanus orarius orarius True; ¢ adult; Ferndale, Humboldt County,
Cal. (No. 19111, Mus. Vert. Zool., Univ. California.)

3, 3a. Scapanus latimanus latimanus (Bachman); ¢ adult; Menlo Park, San
Mateo County, Cal. (No. 18779, Mus. Vert. Zool., Univ. California.)

4, 4a. Scapanus anthonyi Allen; type; ¢ adult; San Pedro Martir Mountains,
Lower California. (No. 4947, Amer. Mus. Nat. Hist.)

5, 5a. Parascalops brewert (Bachman); ¢ young adult; Leasuresville, Pa. (No.
603, Carnegie Mus.)

6, 6a. Condyiura cristata (Linneus); ¢ adult; Lunenburg, Mass. (No. 96075,
U.S. Nat. Mus., Biological Survey collection.)

North American Fauna No. 38, U. S. Dept. Agr. Biological Survey.

B2005-103

SKULLS OF SCAPANUS, PARASCALOPS, AND CONDYLURA.

S. townsendii. 3, 3a. S. 1. latimanus. 5, 5a. P. breweri.
. 8. 0. orarius. 4, 4a. S. anthonyi. 6, 6a. C. cristata.

Fic.

20.

21.

PEALE: vale
[Natural size.]

la, 1b, 22. Netirotrichus gibbsu gibbsi (Baird); g adult; Sumas, British
Columbia. (No. 62952, U.S. Nat. Mus.)

, 2a, 2b, 23. Netirotrichus gibbswi hyacinthinus Bangs; $ young adult; Aptos,

Cal. (No. 160900, U.S. Nat. Mus., Biological Survey collection.)

. Scalopus aquaticus aquaticus (Linnzeus); essentially a topotype; 3 adult;

near Media, Pa. (No. 9845, Acad. Nat. Sci. Philadelphia; 2845, Rhoads
collection.)

. Scalopus aquaticus howelli Jackson; type; ¢ adult; Autaugaville, Ala. (No.

177931, U. 8S. Nat. Mus., Biological Survey collection.)

. Scalopus aquaticus australis (Chapman); topotype; 3 adult; Gainesville, Fla.,

(No. 7081, Mus. Comp. Zool., Harvard College, Bangs collection.)

. Scalopus aquaticus machrinus (Rafinesque); ¢ adult; Oakwoods Cemetery,

Chicago, Ill.. (No. 8155, Field Mus. Nat. Hist.)

. Scalopus aquaticus texanus (Allen); topotype; ¢ adult; Rockport, Tex. (No.

51885, U.S. Nat. Mus., Biological Survey collection.)

. Scalopus inflatus Jackson; type; young adult, sex unknown; State of Tamau-

lipas, Mexico (45 miles from Brownsville, Tex.). (No. 52709, U.S. Nat.
Mus., Biological Survey collection.)

. Scalopus xreus (Bangs); type; 2 adult; Stilwell, Okla. (No. 5475, Mus.

Comp. Zool., Harvard College, Bangs collection.)

. Scapanus townsendw (Bachman); ¢ adult; Ferndale, Humboldt County, Cal.

(No. 19141, Mus. Vert. Zool., Univ. California.)

. Scapanus orarius orarius True; & adult; Ferndale, Humboldt County, Cal.

(No. 19111, Mus. Vert. Zool., Univ. California.)

2. Scapanus orarius scheffert nobis; type; ¢ adult; Walla Walla, Wash. (No.

204997, U. S. Nat. Mus., Biological Survey collection.)

. Scapanus latimanus latimanus (Bachman); ¢ adult; Menlo Park, San Mateo ©

County, Cal. (No. 18779, Mus. Vert. Zool., Univ. California.)

. Scapanus latimanus occultus Grinnell & Swarth; g adult; Alhambra, Cal.

(No. 30299, U. S. Nat. Mus., Biological Survey collection.)

5. Scapanus latimanus grinnelli Jackson; type; ¢ young adult; Independence,

Cal. (No. 17785, Mus. Vert. Zool., Univ. California.)

. Scapanus latimanus sericatus Jackson; type; Q adult; Yosemite, Cal. (No.

109548, U. S. Nat. Mus., Biological Survey collection.)

- Scapanus latimanus minusculus Bangs; type; 2 young adult; Fyffe, Cal.

(No. 9189, Mus. Comp. Zool., Harvard College, Bangs collection.)

. Scapanus latimanus dilatus True; type; young adult, sex unknown, proba-

bly g; Fort Klamath, Oreg. (No. 186628, -U. S. Nat. Mus., Merriam
collection.)

. Scapanus anthonyi Allen; type; ¢ adult; San Pedro Martir Mountains,

Lower California. (No. 4947, Amer. Mus. Nat. Hist.)

Parascalops breweri (Bachman); ¢ young adult; Leasuresville, Pa. (No. 603,
Carnegie Mus.)

Condylura cristata (Linnzeus); ¢ adult; Lunenburg, Mass. (No. 96075, U.S.
Nat. Mus., Biological Survey collection.)

North American Fauna No. 38, U. S. Dept. Agr.

Biological Survey.

PLATE VI.

B2006-103
SKULLS OF NEUROTRICHUS, AND MANDIBLES OF AMERICAN TALPIDA.

1, la, 1b, 22. N. g. gibbsii. 8. Scalopus inflatus.

2, 2a, 2b, 23. N. g. hyacinthinus. . Scalopus ereus.

3. Scalopus a. aquaticus.
4. Scalopus a. howelli.

5. Scalopus a. australis.

). Scalopus a. machrinus.
. Scalopus a. texanus.

15. Scapanus l. grinnelli.
16. Scapanus 1. sericatus.
J. Scapanus townsendii. 17. Scapanus 1. minusculus.
. Scapanus o. orarius. 18. Scapanus 1, dilatus.
2. Scapanus o. schefferi. 19. Scapanus anthonyi.
3. Scapanus |. latimanus. 20. Parascalops breweri.
. Scapanus 1. cecultus. 21. Condylura cristata.

IN Dae

[New names in bold=face type; synonyms in italics.

acquaticus, Scalopus, 33.
znea, Talpa, 58.

zneus, Scalops, 58.
zreus, Scalops, 82.

Scalopus, 52-53.
alpinus, Scapanus, 75.
americana, Talpa, 79.
americanus, Scapanus, 80.
anastas#, Scalops, 39.

Scalopus, 39-41.
anastasi%, Scalops, 39.
anthonyi, Scapanus, 75-76.
aquatica, Talpa, 33.
aquaticus, Scalops, 33.

Scalopus, 32-36.

Scalpos, 33.

Sorez, 32.
argentatus, Scalops, 42.
Astromycter, 82.

prarinatus, 87.

prasinatus, 87.
Astromyctes, 83.
Astromydes, 83.

cristatus, 87.
australis, Scalops, 38.

Scalopus, 38-39.

breweri, Parascalops, 79-82.

Scalops, 79.
Scapanus, 80.
Scaphanus, 80.
Talpa, 80.

californicus, Scalops, 64.
Scapanus, 64.

canadensis, Scalops, 33, 58.
Talpa, 87.

caryi, Scalopus, 48-49.

caudata, Talpa, 87.

Condylura, 82-86.
cristata, 86-91.
fissipes, 87.
longicaudata, 87.
macroura, 87.
prasinata, 87.

Condylus. 83.
cristata, 87.

Condytura, 82.
cristata, 87.

cristata, Condylura, 86-91.
Condylus, 87.
Condytura, 87.
Talpa, 86, 87.
Talpasorez, 87.

cristatus, A stromydes, 87.
Rhinaster, 87.
Scalops, 87.
Scalopus, 87.
Sorer, 86.

cupreata, Talpa, 33.

dilatus, Scapanus, 72-75.
europea, Talpa, 79.

fissipes, Condylura, 87.
Talpasorer, 87.
flava, Talpa, 32, 33, 87.
flavescens, Talpa, 32.

fusca, Talpa, 33.

gibbsi, Neurotrichus, 95.

gibbsii, Neurotrichus, 95.
Netirotrichus, 94-97.
Néurotrichus, 95.
Urotrichus, 94.

gibsii, Urotrichus, 94.

grinnelli, Scapanus, 69-70.

howelli, Scalopus, 36-38.
hyacinthinus, Neurotrichus, 97.
Netirotrichus, 97-98.

inflatus, Scalopus, 53-54.
intermedius, Scalops, 49.
Scalopus, 49-50.

latimanus, Scalops, 64.
Scapanus, 64-67.
longicaudata, Condylura, 87.
Talpa, 86.
Talpasorez, 87.
longicaudatus, Rhinaster, 87.

machrina, Talpa, 42.

machrinoides, Scalopus, 45-46.

machrinus, Scalops, 42.
Scalopus, 42-44.

macroura, Condylura, 87
Rhinaster, 87.

macrurus, Rhinaster, 87.

major, Neurotrichus, 95.
Neitirotrichus, 95.

metallescens, Scalops, 58.

| minusculus, Scapanus, 72.

Neourotrichus, 92.
Neurotrichus, 92.
gibbsi, 95.
gibbsii, 95.
hyacinthinus, 97.
major, 95.
Néurotrichus, 92.
gibbsii, 95.
Netirotrichus, 92-94.
gibbsii, 94-97.
hyacinthinus, 97-98
major, 95.
occultus, Scapanus, 68-69.
orarius, Scapanus, 61-62.

Parascalops, 77-79.
breweri, 79-82.

99

100

parvus, Scalops, 41.
Scalopus, 41-42.
pennantti, Talpa, 42.
pennsylvanica, Scalops, 33.
pensylvanica, Talpasorex, 33.
pensylvanicus, Talpa, 33.
Perascalops, 77.
prarinatus, Astromycter, 87.
prasinata, Condylura, 87.
prasinatus, A stromycter, 87.
pulcher, Scalopus, 46-48.
purpurascens, Talpa, 87.

radiata, Talpa, 87.
radiatus, Sorex, 87.
reposta, Talpa, 80.
Rhinaster, 82.
cristatus, 87.
longicaudatus, 87.
macroura, 87.
macrurus, 87.

Scalops, 27.
eneus, 58.
xreus, 52.
anastas#, 39.
anastasiz, 39.
aquaticus, 33.
argentatus, 42.
australis, 38.
breweri, 79.
californicus, 64.
canadensis, 33, 58.
cristatus, 87.
intermedius, 49.
latimanus, 64.
machrinus, 42.
metallescens, 58.
parvus, 41.
pennsylvanica, 33.
texanus, 50.
texensis, 50.
townsendi, 58.
townsendii, 58.
Scalopus, 27-30.
acquaticus, 33.
ereus, 52-53.
anastase, 39-41.
aquaticus, 32-36.
australis, 38-39.
caryi, 48-49.
cristatus, 87.
howelli, 36-38.
inflatus, 53-54.
intermedius, 49-50.
machrinoides, 45-46.
machrinus, 42-44.
parvus, 41-42.
pulcher, 46-48.
texanus, 50-52.
tcexensis, 50.
virginianus, 33.
Scalpos, 27.
aquaticus, 33.
Scapanus, 54-57.
alpinus, 75.
americanus, 80.
anthonyi, 75-76.
breweri, 80.

NORTH AMERICAN FAUNA.

Scapanus californicus, 64.
dilatus, 72-75.
grinnelli, 69-70.
latimanus, 64-67.
minusculus, 72.
occultus, 68-69.
orarius, 61-62.
schefferi, 63-64.
sericatus, 71.
townsendi, 58.
townsendii, 58-61.
towsendii, 58.
truei, 72.
truii, 72.

Scapasius, 54.

Scaphanus, 54.
breweri, 80.

schefferi, Scapanus, 63-64.

sericatus, Scapanus, 71.

sericea, Talpa, 42.

Sorex aquaticus, 32.
cristatus, 86.
radiatus, 87.

teniata, Talpa, 58.
Talpa xnea, 58.
americana, 79.
aquatica, 33.
breweri, 80.
canadensis, 87.
caudata, 87.
cristata, 86, 87.
cupreata, 33.
europea, 79.
flava, 32, 33, 87.
flavescens, 32.
Jusca, 33.
longicaudata, 86.
machrina, 42.
pennantii, 42.
pensylvanicus, 33.
purpurascens, 8%.
radiata, 87.
reposta 80.
sericea, 42.
teniata, 58.
townsendii, 58.
virginiana, 33.
Talpasorex, 27, 82.
cristata, 87.
fissipes, 87.
longicaudata, S87.
pensylvanica, 33.
teranus, Scalops, 50.
Scalopus, 50-52.
terensis, Scalops, 50.
Scalopus, 50.
townsendi, Scalops, 58.
Scapanus, 58.
townsendii, Scalops, 58.
Scapanus, 58-61.
Talpa, 58.
towsendii, Scapanus, 58.
truei, Scapanus, 72.
truii, Scapanus, 72.

Urotrichus gibbsii, 94.
gibsii, 94.

virginiana, Talpa, 33.

virginianus, Scalopus, 33.

C)

[No. 38,

'S. DEPARTMENT OF AGRICULTURE ~
BUREAU OF BIOLOGICAL SURVEY | |
Ai 2 HENRY WwW. HENSHAW, Chief - 3 awe

;
{
;
=

[ORTH AMERICAN FAUNA —

[Actual date of publication, November 15, 1915]

3 EVISION OF THE POCKET GOPHERS OF —
_ * THE GENUS THOMOMYS — |

j BY
- CHIEF FIELD NATURALIST, BIOLOGICAL SURVEY.
# that te
eae WASHINGTON
GOVERNMENT PRINTING OFFICE
ros LG 2 1g16 : bee eee

Biological Survey. PLATE I.

North American Fauna No, 39, U. S. Dept. Agr.

W6LE

“Uo0jog *F, ysourg Aq ‘eqoyrueyy ‘Arioqiey utoIy Weeds B JO SUIMeLIP Po10[OD WIOI, ‘ozIs [eRanyeU SPIIY}-OM} JnoGy

(‘SNSOSSANY SAGIOd1V.L SANOWOH |) YSHdOH) LEMO0d ZIMIVYd

bes PEPAREMENT OF AGRICULTURE
BUREAU OF BIOLOGICAL SURVEY

HENRY W. HENSHAW, Chief

or eee ere tt

NORTH AMERICAN FAUNA
INSo.. 39

[Actual date of publication, November 15, 1915]

REVISION OF THE POCKET GOPHERS OF
THE GENUS THOMOMYS

BY
VERNON BAILEY

CHIEF FIELD NATURALIST, BIOLOGICAL SURVEY

WASHINGTON _
GOVERNMENT PRINTING OFFICE
1915

LETTER OF TRANSMITTAYT:

UNITED STATES DEPARTMENT OF AGRICULTURE,
BurEAv OF BIoLoGicaL SURVEY,
Washington, D. C., April 29, 1915.

Sir: I have the honor to transmit herewith for DAL ation as
North American Fauna No. 39, a revision of the pocket gophers of
the genus Thomomys, by “Tenner Bailey, chief field naturalist of the
Biological Survey.

Pocket gophers are burrowing rodents which in pursuit of food
and in running tunnels often damage crops, young trees, and irriga-
tion ditches. Moreover, the heaps of earth ejected from the tunnels
not only cover more vegetation than the animals eat, but also obstruct
the operation of mowing machines. Gopher burrows frequently
penetrate and cut through irrigation-ditch banks, and the escaping
water becomes a menace to agriculture and sometimes to human life.
The animals are thus of considerable economic importance.

The present paper completes the technical revision of the pocket |

gophers (family Geomyidee), the first part of which was prepared by
Dr. C. Hart Merriam and published in 1895 as North American
Fauna No. 8. The revision will supply definite information regarding
the status and geographic distribution of the several forms, which is
important in connection with attempts to control or exterminate

them.
Respectiully, —
| Henry W. HENsHaw,
Chief, Brological Survey.
Hon. Davin F. Houston,

Secretary of Agriculture.
2

ee

CONTENTS:

LEEDS eee {H@) CLROY OS ea hs Paes ene
“Tur iDAN@e Le] OFS Sa es Oa aN ee ice ae ng ea ee Pree
Ee IPM ARNE eee een Ae ees ke) ee ee AL
WOSMMOM Gc ke ee a ee af poe eae eam
aekemeopiersas 1000 - .. 22-3 ee ee ee eee bees eles ee
PeMULE MIRCHI CTMCs ae eo Nees eS et tee 8s LE ee Pa Sea cs ee
MecancinOlsOr POCKEL COPNCIS: 2.2222... =. 0-5-2 oi Seen ete See
SELES, GIRS ONS NRS al R FPSE eg e e
MaomictleehiaraCters.......-.. 2) 222.522 ee ST GRD IOS AAR IN ee Rea!
Mine ee ee ee ck oe

melkmowledements.-:-.2.2-- 2.2... soe pe. ig tee en Pp aa he,

SELES LEDUC CPS ok a he a ee a a

Bae Mem AChCIS hae ee ep aie lat ete ke ie eee tyes
Gromp relationships. ........-... ..: Pcs Re ELT, as SRE tes aS TANNA PORE
LE GEST TRESS TELL: 0) OS ee a P= er eT
SLC TG vB TCS AAT AG ea 0109 0s ae eee gg ra a
List of species and subspecies, with type localities................-.-----
HexaLomrouporor related species. .-..: =... 32 - sees Dees bee es
iMewaoriecics amd SUDSPECClES <0... 5... 5252 levee eo ee al se dk ele
Description and distribution of species and subspecies. .:....-----.------
A oMOUtY SHOE OIVOTUSIOTOUD s6e2 623 ssa sok se ood ee oe. Bn SR
MP OMOULUSLOWUMSCIGL STOMP 2.5 5 Nasa ete er on Del est ee ee
INO MUS TUOULE CL OUP RES: rr 52 2 tie Sepa. ta ae BS eee.
ME AMIOIIUSIAL PINUS PTOUDS)- <2: 22. es ei Plo
apna Se Der PAlldUs CTOUPs 2. 2-202 Seed 2s Sen ee bes eke eee eS
EO TELOTELIS. J RUMBLES Ser ROT ane et el ace as ok, oye ee
AOU S AEMUTULUSIOTOUP 22 oe oe oe Me Bet ee
WGIIO NISC! DOLGCS OTOUD a. 22 ie. ea ee, ea as
DRDO NUSHOSSOMOTOU Ps 8 aos Sat eee cent eee We ci eee ay
Ehomomys douglast croup... -2:--------.-- Pai ya ae saa eee ae
ERO S OR LICOLO STOMP: <2 eS et er ee Sak Seger te es a ea

Renee Sater er 2 er iE NE ES Se ee Be ee SE eS

PuATE I.
TI.

Il.

VI.

VII.

VIII.

Fre. 1.

ILLUSTRATIONS

PLATES.

[Plate I, frontispiece; Plates II-VIII, following nage 132.]

Prairie pocket gopher (Thomomys talpoides rufescens).

Skulls (ventral views) of Thomomys bulbivorus, T. townsend: townsendi, T.
bottx botix, T. perpallidus aureus, T. talpoides rufescens, T. t. clusius,
T. idahoensis, T.fossor, T. monticola monticola (and lateral view of monti-
cola), T. douglasi douglasi, T. bottx leucodon, T. ocius, and T. umbrinus
umobrinus.

Skulls (dorsal views) of Thomomys bulbivorus, T. townsendi townsend, T. t.
nevadensis, T. boits botte, T. b. leucodon, T. b. laticeps, T. b. anguluris,
T. b. pallescens, and T. altivallis.

. Skulls (dorsal views) of Thomomys bottx pascalis, T. b. minor, T. 6. navus,

T. fulvus texensis, T. f. pervagus, T. perpallidus apache, P. sinalox, T. b.
alticolus, and T’. b. russeolus.

. Skulls (dorsal views) of Thomomys perpallidus albatus, T. p. perpallidus,

T. p. perpes, T. operarius, T. p. aureus, T. p. canus, T. latirostris, T. caba-
zone, T. alpinus awahnee, T. bottze mewa, T. b. anitx, T. magdalenx, T. a.
alpinus, T. jacinteus, T. neglectus, and T. 6. nigricans.

Skulls (dorsal views) of Thomomys fulvus fuluus, T. cervinus, T. f. toltecus,
T. mearnsi, T. f. ntermedius, T. f. desertorum, T. baileyi, T. lachuguilla,
T. perditus, T. goldmani, T. nelsoni, T. sheldoni, T. umbrinus umbrinus,
T. u. peregrinus, T. u. orizabe, and T. atrovarius.

Skulls (dorsal views) of Thomomys talpoides rufescens, T. t. bullatus, T. t.
nebulosus, T. t. agrestis, T. t. clusius, T. t. caryi, T. t. pryort, T. quadratus
quadratus, T. ocius, T. idahoensis, T. pygmxus, T. columbianus, T. fossor,
T. winta, T. bridgeri, and T. quadratus fishert.

Skulls (dorsal views) of Thomomys monticola monticola, T. m. mazama, T.
m. pinetorum, T. m. nasicus, T. douglasi douglast, T. d. melanops, T.
niger, T. d. oregonus, T. fuscus loringi, T. hesperus, T. d. yelmensis, T.
limosus, T. f. fuscus (male and female), T. f. myops, and T. f. saturatus.

TEXT FIGURES.

Map showing distribution of the genus Thomomys...............-.--- 9

. Skins of adult females of Thomomys, showing number and position of

mamme in different groups and species: 1, Thomomys wmbrinus
umbrinus; 2, T. bottx bottx; 3, T. fossor; 4, T. talpoides rufescens. . . 12

. Skins of Thomomys, showing waves of pelage during molt: 1, Thomo-

mys talpoides rufescens; 2, 4, T. quadratus fisheri; 3, T. t. clusius... - 19

. Skins of Thomomys showing abnormal pelage conditions, due to tick

bites, mange, or skin disease: 1, Yhomomys fossor; 2, T. fuscus
ST LS SLES se Ee I ee et Sa a geo ee 21

. Map showing distribution of certain species and subspecies of Thomo-

mys: 1, fuscus; 2, loringi; 3, saturatus; 4, myops; 5, columbianus; 6,
bridgeri; 7, winta; 8, quadratus; 9, townsendi; 10 nevadensis; 11, canus;
12, monticola; 13, pinetorum; 14, mazama; 15, nasicus; 16, helleri;
17, bulbivorus; 18, hesperus; 19, niga 20, oregonus; 21, yelmensis;
22, melanops; 23, Timosue: ZA SNUG ET Ae eae Se tis en) erie a eee 23

6 ILLUSTRATIONS.

Page.
Fic. 6. Map showing distribution of certain species and subspecies of Thomo-
mys: 1, bottx; 2, minor; 3, laticeps; 4, leucodon; 5, navus; 6, mewa; 7,
pascalis; 8, angularis; 9, diaboli; 10, infrapallidus; 11, pallescens; 12,
alpinus; 13, awahnee; 14, neglectus; 15, altivallis; 16, jacinteus; 17,
cabezonx; 18, puertx; 19, perpallidus; 20, albatus; 21, perpes; 22,
operarvus; 23, fisher... - 2.23345 -404 2422s eee 24
7. Map showing distribution of certain species and subspecies of Thomo-
mys: 1, apache; 2, fulvus; 3, toltecus; 4, desertorum, 5, texensis; 6, baileyi;
7, mearnsi; 8, lachuguilla; 9, cervinus; 10, sinalox; 11, chrysonotus;
12, nigricans; 13, martirensis; 14, russeolus; 15, magdalenex; 16, anitx;
17, alticolus: ifiesiel ot. ee ee 25
8. Map showing distribution of certain species and subspecies of Thomo-
mys: 1, fossor; 2, agrestis; 3, pervagus; 4, desertorum; 5, aureus; 6,
latirostris; 7, chrysonotus; 8, cervinus; 9, intermedius..........-.--.- 26
9. Map showing distribution of certain species and subspecies of Thomo-
mys: 1, umbrinus; 2, orizabe; 3, peregrinus; 4, atrovarius; 5, sheldoni;
6, nelsoni; 7, goldmani, 8, perdituss: =. 22a) es ee 2
10. Map showing distribution of certain species and subspecies of Thomo-
mys: 1, talpoides; 2, rufescens; 3, bullatus; 4, nebulosus; 5, clusius; 6,
caryi; 7, pryori; 8, idahoensis; 9, pygmexus; 10, ocius...-.-.--------- 28

No. 39, NORTH AMERICAN FAUNA. Nov, 15, 1915,

REVISION OF THE POCKET GOPHERS OF THE
GENUS THOMOMYS.

By VERNON BAILey.

INTRODUCTION.

Pocket gophers, a group of rodents peculiar to North and Middle
America, belong to the family Geomyide, which includes the genera
Thomomys, Geomys, Pappogeomys, Cratogeomys, Platygeomys, Ortho-
geomys, Heterogeomys, Macrogeomys, and Zygogeomys. The present
revision deals only with the genus Thomomys,' which is distinguished
from the others by the practically ungrooved surface of the upper
incisors. The large fur-lined external cheek-pouches are a family
character shared by all genera and species, as are the general habits
of living in underground burrows and throwing up heaps of earth
along their lines of tunnels. The genus Thomomys includes 88
recognizable forms of 40 species. These vary in size from that of a
small mole to that of a large rat, and in color from light buff to yellow
and gray, and from many shades of brown and rusty to dusky and
even black. All are short-legged, robust, sturdy little animals,
with small ears and eyes, and short, smooth hair.

~ Gophers of the genus Thomomys are abundant over almost the entire
western half of the United States, in a great area of southwestern
Canada, and over much of Mexico (see fig. 1), covering considerably
more territory than all the other members of the family together.
The various forms do not overlap in range to any great extent, but
in the area occupied they are present in almost every locality, and
are especially abundant in fertile and valuable agricultural lands.
Their burrows are a constant menace to agricultural operations, irri-
gation ditches, and reservoirs. Their food is entirely of vegetable
matter, and of this roots, bulbs, and cultivated crops are preferred.
In destroying crops and fruit trees’they cause a heavy loss to western

1JIn 1895 Dr. C. Hart Merriam monographed the family Geomyide, exclusive of the genus Thomomys
(N. Am. Fauna No. 8), and also included a chapter on the anatomy and relationships of Thomomys and a
map of the genus without reference to its specife and subspeciic divisions. It was my privilege to assist

him in working over the material for his monographic revision and later I was assigned the task of com-
pleting the revision of the genus Thomomys.
7

red NORTH AMERICAN FAUNA. NO. 39.

farmers each year, and it is necessary in farming areas to use every
possible means of restraining their increase and checking their depre-
dations. In wild land, on the other hand, their presence is highly
beneficial, not only because they stir the soil and enrich it by burying
vegetation under their mounds, but also because they form a net-
work of underground channels which aid materially in retaining
water and carrying it deeper into the ground. Such of their beneficial
habits as have operated for ages in enriching the western soils are
far overbalanced in agricultural lands by their destructive habits.
On lands newly brought under cultivation it is often necessary to
wage an active campaign against gophers, in order to render the
crops fairly safe. (See footnote, p. 14.)

The present paper describes the recognizable species and sub-
species and defines their geographic distribution so far as at present
known.

GENERAL HABITS.

BURROWS.

Pocket gophers of the genus Thomomys are burrowing animals
which spend most of their lives underground. The several species
have practically the same habits except as modified by environment.
They are sturdy little miners, drifting their tunnels on and on in
search of food, throwing out little heaps of earth along the lines of
excavation, and then quickly closing the doorway. The tunnels vary
from about 6 inches to a foot below the surface, and from about 14
to 3 inches in diameter, according to the size of the gopher. The
length of the tunnels is practically interminable and their course ~
varies from a winding network to long direct lines.

The character of the burrows is determined by the food supply.
A row of potatoes may be followed across a field, or the spreading
roots of an apple tree may hold the attention of the gopher for some
time. If food is scarce the rows of mounds reach in direct lines for
long distances, but when it is abundant they wind back and forth
within a limited area. While the males move about more freely at
the beginning of the breeding season and there are indications of rare
migrations, pocket gophers are evidently the most restricted in their
habitat of any of our native mammals. Even moles, which live more
completely underground, burrow more rapidly and travel more
widely. The whole summer’s line of hills of one gopher can often
be seen at a glance. The animals rarely leave an alfalfa or potato
field, but hurry across waste places. Under ordinary circumstances,
however, a gopher often spends a year on a-single acre, and perhaps
the next year on an adjoining one.. Most gophers probably never
vet 40 rods from where they were born, but where environment is less
favorable and the gopher is more active, he may extend his tunnels
for half a mile or possibly a mile during his career.

INTRODUCTION.
y to 15 feet apart, mark the lines of the

B2042-122

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, the openings are securely

1915. ]

Little mounds of earth, thrown out as the burrows are excavated,

-and varying in bulk normally from a quart to a bushel and in dis-

tance from a close proximit

Fic. 1.—Map showing the distribution of the genus Thomomys (shaded area).

burrows.

After the loose soil is pushed out

packed with earth and closed from within as a fortification against

enemies.

The entrances are often closed so securely that even the

experienced trapper has difficulty in locating the burrow beneath.

10 NORTH AMERICAN FAUNA. [NO. 39.

Usually, however, the opening may be found by the depression
where the last load of earth was left, and the direction of its slope
downward determined by the position from which the mound of
earth was pushed out. This also serves to distinguish a gopher —
_ lull from a molehill, as the mole pushes up the earth from below
without making an opening to the daylight. Often the opening
is near some plant, as the gopher evidently recognizes a choice
species by the root and, following it to the surface, devours or
carries it away. : |

These mounds or “gopher hills,’ frequently cover low vegetation;
on wild land this serves to enrich the soil, but in fields and meadows
they often cover and destroy the growing grain and forage, and also
interfere with mowing and harvesting.

FOOD HABITS.

- Gophers of all species of Thomomys are strict vegetarians. Their
food consists for the most part of roots, bulbs, tubers, and the most
edible and nutritious underground parts of plants, but it includes
also much green vegetation from aboveground. As they extend
their tunnels gophers find wild onions, liliaceous bulbs, wild potatoes,
underground beans, grass tubers, and an endless number of edible
roots. These are cut in sections or, if of convenient shape and size,
tucked in the pockets whole, and when the pockets are well filled
are carried to the diming room -to be eaten at leisure or deposited
in the storeroom for future use. When gophers are trapped their
pockets usually contain articles of food, sometimes SO much that
they stand out on both sides of the head.

The list of plants found in the pockets is long and vanes Sarne
of the fleshy desert plants, as cactuses, yuccas, agaves, and sotol are
entered from below and the juicy centers eaten out. Wild clovers
and a great_many leguminous plants are favorite foods, while grass
stems and blades are eaten to some extent. Very small bushes are
sometimes cut down near the burrows and the pars eaten, but trees
are rarely injured aboveground.

Gophers apparently A not require tae and it is doubtful if.
they drink at all. Many desert species uccine for long periods,
sometimes a year or more, where they can not get water, but green
food and juicy roots evidently supply all the moisture they need.

DAMAGE TO CROPS.

Pocket gophers show a keen appreciation of man’s efforts in agri-
culture.and horticulture by resorting to cultivated areas. They
revel in potato fields, and one gopher will destroy a long row of the
tubers, taking one hill after another from the time they first appear
until they are harvested. They quickly attack newly planted areas,

1915.] INTRODUCTION. ifi

and where potatoes are stored in the field they often enter the pits
to eat the tubers or carry them away. All kinds of garden vegetables
and also most field crops are eagerly eaten, either below or above
the surface of the ground. Gophers sometimes burrow up into
pumpkins or squashes and eat out the insides without the mischief
being detected until the empty shells dry up. They will work all
summer in gram fields, cutting and eating the leaves, stems, and
erain heads and covering with their hills of earth even more grain
than they eat. In alfalfa and clover fields and in grass meadows they
destroy both roots and tops and pile their mounds over much of the
crop, and in addition, by leaving their mounds full of small stones,
they dull and break the knives of mowing machines. Their mischief
is perhaps most exasperating in orchards and nurseries, as they
often injure or kill choice fruit trees by their underground operations.
They are very fond of the roots of apple, pear, and fig trees.

In irrigation areas gophers have an annoying habit of burrowimg
through the ditch banks and letting the water cut its way cut, of
filling up the smaller ditches with their mounds, and of interfering
generally with the most efficient use of water. It is reported that
in the spring of 1915 their burrows caused two reservoir dams on
the Little Colorado River, near St. Johns, Ariz., to give way, causing
the death of eight persons by drowning, and damage to crops to the
extent of half a million dollars.

WINTER HABITS.

Apparently none of the pocket gophers hibernate, and in spring,
as the deep snows melt, their old burrow-molds are seen snakelike
over the surface of the ground as evidence of winter activity. They
work below the frost and instead of throwing out mounds they
tunnel under the snow along the surface of the ground, probably
gathering some green plants as they go, and then fill these snow
tunnels with earth from their burrows. This earth freezes and
hardens and often remains all summer as earth casts of gopher
burrows in the snow. The long safe winter under protection of
the snow has probably done much to conserve the species and facil-
itate their wide dispersal and abundance in high mountains and

northern areas.
BREEDING HABITS.

The little known of the breeding habits of Thomomys barely
suffices for a few general conclusions. In northern and more
elevated portions of the country female gophers usually contain
small embryos in June, and early in that month a male and female
are occasionally caught in the same burrow. This rarely happens
later in summer, and it is evident that only one litter of young
is regularly raised in a year. In warmer climates the young are

1G NORTH AMERICAN FAUNA. - ENO. 39.

born earlier, and in some regions two litters may be raised. Though
I have collected many specimens, I have never seen one younger
than the nearly half-grown stage, when they begin to move about,
nor have I found specimens of young in any of the museum collec-
tions. During July and August many half-grown gophers are

B2043-122 —
Fig. 2.—Skins of adult females of Thomomys, showing number and position of mamme in different
groups and species. (Exact location of nipple indicated by black dot. Relative position of posterior

pairs often distorted by method of sewing up skin.) 1, Thomomys uwmbrinus umbrinus; 2, T. bottzx
bott2; 3, T. fossor; 4, T. talpoides rufescens..

caught im traps, but then they are working in the parent burrow
or starting new lines for themselves. A little later old and young
are in independent burrows. Eee

The number of young in a litter varies in the different species, as
might be expected from the variation in the number of mamme

1915. ‘INTRODUCTION. 13

in the females (fig. 2). In the uwmbrinus group from Mexico, with
3 pairs of mamme, the number of embryos has been recorded as 2
and 3. In other species, with 4 pairs of mamme, the number of
embryos has been recorded as 4 and 5, while in the clusius and
ocvus groups, with 6 and 7 pairs of mamme, 6 and 7 embryos seem
to be the rule. Judging from nearly matured fetuses the young are
born helpless and naked, and must remain for some time in the
nest while cared for by the mother. Apparently nothing is known
of their nests or breeding quarters.

DISPOSITION.

Like all of the family, gophers of the genus Thomomys are pug-
nacious and fight man or dog as quickly as they would an animal |
of their own size. To the gopher everything that moves is an
enemy and is attacked with vicious wheezing and savage bites.
The toe of a boot or a stick is seized, and the heavy incisors make
deep cuts into wood or quickly cut through leather. The weasel
is probably the gophers only enemy of its own size that successfully
attacks and masters it in its burrows. The gopher’s vision is
extremely limited, and it seems not to see an object until very close,
when its instinct of self-preservation suggests prompt attack.

POCKET GOPHERS AS FOOD.

These gophers are not only strict vegetarians, but are in every
way cleanly and exemplary in their food habits. Their flesh is
tender and well flavored, and were the animals large enough they
might be made a valuable adjunct to our meat supply. They have
furnished me with many a palatable meal in the mountains when
other food was not available, and would have been of great value
to early explorers, who suffered from starvation, had a few gopher
traps been carried across the mountains and deserts. Two or three
gophers broiled over the coals of the camp fire furnish a fairly sub-
_ stantial and palatable meal, especially if accompanied by a stew of
wild onions and bulbs of the camas. Gophers are hardly large
enough to be worth catching for the market, however, but where large
numbers must be destroyed to protect crops, some may be utilized for
the table. The meat is rather dark and fine-grained, tender, and in
flavor not unlike that of the squirrel. Sometimes it is rather strongly
flavored with wild onions, but to some this is not unpleasant.

NATURAL ENEMIES.

~ While the underground life of pocket gophers would seem to render
them safe from attacks of enemies, yet they are preyed upon by a
great variety of animals. Hawks and owls pick up so many of them
at their doorways that gopher bones are among the most abundant

14 NORTH AMERICAN FAUNA. [No. 39.

in the pellets of these birds. Coyotes, foxes, and bobcats pounce
upon them as they push out their loads of earth, and badgers occa-
sionally dig them out. Weasels are probably their greatest enemies,
and when one gets into a gopher burrow there is no escape for the
occupant. Snakes also prey upon them to such an extent that one
common species is known as the gopher snake. The protection of
such natural enemies of the gopher as prove least harmful in other
ways, notably hawks, owls, weasels, and badgers, is of the utmost
importance as an adjunct in the control of what is proving in many

sections one of the worst of our rodent pests.

DESTRUCTION OF POCKET GOPHERS.

Pocket gophers are easily trapped and poisoned, and on limited
areas may be reduced to harmless numbers by systematic, intelligent
effort. In extensive fields and meadows and along systems of
irrigation ditches or reservoirs their control often presents problems
of considerable magnitude. The Biological Survey has made special
study of the many conditions: under which injuries from gophers
occur and of the most economical and efficient means of their control.
The results have appeared in several publications of the United
States Department of Agriculture. '

GENERAL CHARACTERS.

The short legs, strong frame, and powerful muscles of pocket
gophers peculiarly fit them for subterranean life. The long front
claws vary considerably in size with the different species, being largest
in 7. talpoides and T’. rufescens, which are medium-sized species, and
relatively smallest in 7’. bulbivorus, the largest of the genus; but in
all they are well adapted to rapid digging. The stout hind feet, with
wholly naked, plantigrade soles and stout nails are well adapted
to pushing, while the delicate naked soles of all the feet are doubtless
of great advantage as a substitute for eyes in an animal which spends
most of its life in utter darkness. . Fringes of stiff bristles bordering
the fingers and palms evidently assist in holding together the loose
earth as it is pushed out of the burrow. The very small eyes and
ears are keen at close range, but are of little use in the burrow where
most of the animal’s life is spent. The stubby and seemingly rather
useless tail is slightly tapering, thinly haired, and often naked near
the tip. It probably serves as a substitute for eyes in the animal’s
rapid backward progress through its burrow, as has been shown

1 Pocket Gophers of the United States, by Vernon Bailey. Bul. No. 5, Division Ornithology and Mam-
matlogy, 1895.

Directions for Destroying Pocket Gophers, by David E. Lantz. Circular No. 52, Biological Survey,
1908.

Harmful and Beneficial Mammals of the Arid Interior, by Vernon Bailey. Farmers’ Bulletin 335, 1908,

Pocket Gophers as Enemies of Trees, by David E. Lantz. Yearbook Separates No. 506, 1909.

4915.1 INTRODUCTION. 15

¢

by Dr. Merriam to be the case witk the very similar and closely
related genus Geomys.' The vibrisse are of just sufficient length
to span the diameter of the burrow and doubtless serve as a further
substitute for eyes.

The general pelage is fine and silky, becoming long, full, and soft in
winter in cold climates and again becoming thin and harsh in summer.
In the southern tropical or subtropical species the fur, if it may be
called fur, seems not to attain the full soft winter coat. The pelage
consists of a fine all-plumbeous underfur through which rises a
longer, coarser coat with plumbeous base and colored tips which give
the dominant shade characterizing the species. This usually varies
considerably with the season, in some species becoming much darker
in winter, in others much paler. The covering on the underparts is
usually much thinner, often allowing the skin to show through. The
tail, ears, and tops of feet are but thinly clothed with short, stiff hairs.
The small nose-pad is naked. :

The construction of the mouth is such that both upper and lower in-
cisors protrude from fur-incased mandibles, allowing perfect freedom
for cutting roots, digging in hard earth, or prying out stones without
danger of the earth entering the real mouth, which opens vertically
well back of the incisors and closes automatically as the incisors are
spread wide apart. The fur-bordered lips close in a vertical slit in
front of the real mouth or draw back and expose an ample cavity and
a thick, fleshy tongue.

The pockets, which give the animal unusual interest, have been the
cause of much speculation and misapprehension. Some of the speci-
mens first seen by naturalists were stuffed with the pockets wrong side

out, Say’s genus Pseudostoma being based on such a specimen, while
-Kuhl’s Saccophorus was based on a specimen showing the pockets in
their real position. The use of the pockets is still often misunderstood.
They are external invaginated folds of skin, fur lined, and opening
along each cheek parallel with the mouth and extending approximately
14 to 2 inches back under the skin of the shoulder. They are very
elastic and when empty might pass unnoticed, but when filled they
more than double the apparent size of the animal’s head. | have often
thrust two fingers half their length into the two pockets of a small-sized
gopher, reaching back almost to the middle of the animal’s body. In
large species the thumb fits nicely in the empty pocket. In removing
a gopher’s skin a thin band of muscle is discovered attached to the apex
of each pocket and by it they are drawn back and kept in place. The
pockets can not be fully everted without greatly stretching or possibly
breaking this muscular attachment, but they can be drawn out so that
a large fold of the lining will hang out of the pocket when the animal’s
muscles are relaxed. It is a common belief that the pockets are used

1N. Am. Fauna No. §, 16, 1895.

16 - NORTH AMERICAN FAUNA. ; [NO. 39.

in carrying earth from the burrows, but such is not the case as, in pre-
paring many hundreds of specimens, I have never found earth in the
pockets, except for minute particles from roots and bulbs. Other
naturalists have given the same negative evidence. On the other hand,
the pockets are commonly stuffed full of vegetation. I have often
watched the gopher come to the surface, opening a new doorway as he
came, and begin to draw down one plant after another, cuttmg and
tucking away the sections in his pockets as he drew them down. His
motions are quick as a flash, and that the hands are used in filling the
pockets is about all one can be sure of. Dr. Merriam found by watch-
‘ing a captive specimen of Geomys that the hands were used for both
fillmg and emptying the pockets.1

CRANIAL CHARACTERS.

In Dr. Merriam’s very full discussion of the morphology of the
skull in his Monographic Revision of the Geomyidx? numerous figures
and illustrations of the genus Thomomys are included. In the present
work it is therefore necessary only to mention some of the variations
and general characters useful in the separation of the various forms.

In all, the cranium is wide and low, with spreading zygomatic arches
for the accommodation of powerful muscles. ‘The mandibles are heavy
and irregular, and the incisors above and below are long, curved, deeply
embedded tools of labor and weapons of defense. The relative length
and breadth of the skull vary greatly in different species: shortening
of the total length generally implies a relatively wider skull; elongation
a relatively narrower. In some species, however, the rostrum alone is
elongated or shortened, in others the braincase varies in length and
breadth. The generally wedge-shaped nasals are extremely variable
in length and form, especially in form of termination at posterior tips.
The premaxillee in some terminate even with the posterior line of the
nasals, in others they extend well back into the frontals with pointed,
3 beveled or truncate tips.

The fiitonpenic a furnishes a convenient character for the recogni-
tion of many forms, varying from a minute oval to a large quadrate
bone covering nearly half the basal width of the skull. In some
species, however, it is obscured in old age until completely obliterated
in a high sagittal crest, in others advanced age produces no appre-
ciable change. The various forms of the line of junction between the
base of the zygomatic processes of the maxillze and the premaxille and
frontal and lachrymal bones furnish excellent specific as well as group
characters. In the umbrinus group the bases of these bones are forked
or concave; in most others convex, straight, or doubly curved. There
are often good characters in the symphysis of the zygomatic process

1N. Am. Fauna No. 8, 18, 1895. 2Tbid., 33 and 198, 1895.

1915.] INTRODUCTION. 17

of the maxilla and the jugal, but this changes considerably with age
and is not much used in diagnoses. The pterygoids have a wide
range of variation and furnish excellent characters in many species.
The large and concave pterygoids of 7. bulbworus have been used as
the main basis for a subgenus ( Megascapheus),! but while unique they
certainly do not carry more than specific distinction. The general
variation is in elevation or depression of the wings of the pterygoids,
wide or narrow, and sharply V-shaped or widely U-shaped fossa.
The audital bulle vary greatly in relative size and form, the largest
being smoothly rounded and globose, the smaller more elongated
and angular. The basioccipital varies greatly in relative length and
width, generally becoming narrower as the bulle increase in size and
crowd upon its margins. Many slight detailed differences can also be
used to advantage as characters. Many excellent characters in the
lower mandibles are not generally used in diagnoses, as in most forms
there are others more convenient.

DENTITION.

The dental armature of Thomomys is included and fully discussed
with the other genera in Dr. Merriam’s revision of the Geomyide.
Throughout the genus Thomomys there is surprising uniformity of
tooth form and enamel pattern in the molariform teeth; while
many minute variations furnish distinctive specific characters, the
necessity of studying them with a lens renders them inconvenient
for general use, and they rarely serve to separate closely related
forms. On the contrary, the incisors furnish conspicuous group as
well as specific characters. The angle of the upper incisors to the axis
of the skull varies from the very protruding incisors of T. bulbwworus,
T. leucodon, and T. umbrinus to the abruptly incurved incisors of T.
pygmexus, T. idahoensis, and 7. ocius. A narrow groove near the
inner edge of each upper incisor is deepest and most conspicuous in
the monticola group, but so slight and obscure in other groups as to
be detected only by critical examination with a lens; it is usually
most obscure in species with projecting upper incisors. In some
species the incisors are rather slender, and in others relatively stout.

SEXUAL VARIATION.

The male in most species is considerably larger than the female and
its skull is heavier, more angular, and more heavily ridged in old age.
This sexual difference is greatest in the bottz group and apparently
least in the monticoia and fuscus groups, but is noticeable in all.

The functional mamme vary in fully adult females of different
species from 3 to 7 pairs, usually arranged in 2 pairs of inguinal, 2 or

1 @lliot, Field Columb. Mus., zool. ser. III, 190, 1903. Type, Diplostoma bulbivorum Richardson.
2N. Am. Fauna No. 8, 69-108, 1895.

9812 1°—15——2

18 NORTH AMERICAN FAUNA, ‘ING. 39. -

~ none of abdominal, and 1, 2, or 3 of pectoral. The usual number of
embryos ranges from 3 to 7.
MOLT. _ e

In the genus Thomomys there seem to be normally two complete
changes of pelage during the year, from the winter to the summer coat
during spring or early summer, and from the summer to the winter
coat in fall. The time of these changes varies in different species and
often in the same species with different climatic conditions, and
apparently also with the varying physical condition of individuals,
as age and bodily vigor. In the northern and more elevated areas ©
the molt is more pronounced and its progress more rapid, owing to
greater extremes of climate; while in the equable climate of the
west-coast areas and in the Tropics of western Mexico very gradual
and slight change of pelage occurs. The progress of the molt 1s nor-
mally from the nose over the head and body to the base of the tail
along a crescentic line, a little faster on the back than on the sides.
On the belly the molt lags behind and is less reguiar. :

The spring molt is most conspicuous, as each succeeding wave of
hair is shorter than that displaced and leaves a dark line of plumbeous
basal color exposed on the old hair. The change is a‘creeping process.
New hair displaces old on the nose, apparently pushing out the old
and worn and taking its place along the line of advance backward
and downward. The line is normally a semicircle, but it often
becomes distorted by one part progressing more rapidly than another.
In some cases the change is complete as it progresses, all the old hair
being replaced, but in others the molt is only partial, a thinning
or filling-up process, but always progressing along the line in the
same way. In spring the first molt wave and often those up to the
fifth are but slight changes, a thinning. like the removal of one winter
shirt after another, but the final, perhaps fifth or sixth, change to the
thinnest summer clothing is a complete change as it progresses. The
number of molt waves is not easily determined from specimens, as
they often overtake and run into each other. The long winter coat
is retained on the rump often until late summer, and the successive
molt waves strike against this until the short summer coat pushes
‘it off at the base of the tail just before, or sometimes, in high or north-
ern localities, not until, the next winter coat has begun to make its
appearance on the nose. There are at least five waves in the spring —
molt, as this number appears on many specimens taken in May or
June. Those taken later in summer usually show a less number of
coats but with more deeply marked division lines due to the combi-
nation of several waves into one or more.

The fall molt is different. The winter coat is put on more rapidly
with the appearance of really cold weather, and apparently the full
change is accomplished in two molt waves that begin at the nose and

1915.) INTRODUCTION. 19

progress over the animal along a semicircular line. The line is not
so conspicuous, however, as the longer hair overlaps the short summer
coat which it is crowding out, and does not exhibit the plumbeous
color along its margin. Often before the first coat has reached the

| B2014-122

Fic. 3.—Skins of Thomomys (U.S. Nat. Mus. specimens) showing waves of pelage during molt.

1. Thomomys talpoides rufescens, male adult, No. 180486, Valley City, N. Dak., May 17, 1912, showing
five waves of pelage during spring molt.

2. Thomomys quadratus fisheri, male adult, No. 94411, Ruby Valley, Nev., June 23, 1898, showing four
waves of pelage during spring molt.

3. Thomomys talpoides clusius, female adult, No. 160396, Shirley Mountains, Wyo. (8,800 feet altitude)
Aug. 21, 1909, showing three waves of spring molt, with coat of previous winter on rump and two
waves of fall molt coming in on head: two winter coats at one time.

4. Thomomys quadratus fisheri, male adult, No. 80721, Cottonwood Range, Nev., Sept. 17, 1896, show-

| ing first wave of fall molt back to rump, second wave back to ears.

) base of the tail the second has begun on the nose and is following the

|. first (fig. 3). This seems not to replace the first but to fill up and
thicken the coat, while both may increase in length as the weather
becomes more severe.

20 - NORTH AMERICAN FAUNA. [NO. 39. -

Climatic conditions control the time and nature of 1nolt, but not
its general plan. Species from high in the mountains and from well
north wear heavy winter coats for a longer time: T. talpoides from
October to April; 7. fossor and T.fuscus from October to May; while
those from low hot valleys—T. perpallidus, T. albatus, and T. chryso-
notus—wear thinner winter coats from about the first of December
to the first of March. Species from the California coast show the
usual molts, but with slight contrast between summer and winter,
and with considerable individual variation in dates of change. The
one fully tropical species, 7. atrovarius, from western Mexico, is repre-
sented by specimens taken from April 5 to August 3, in most of which
one molt line appears across the back, and none shows more than one,
while there are several without any trace of molt. This condition is
rarely found in any other species in summer pelage and is probably
due to the tropical chmate. Specimens showing a molt line seem to
be changing from one thin, harsh pelage to another exactly like it,
and whether there is any change to a winter pelage in this species
remains to be shown by winter specimens.

When the young, which are evidently born naked, are old enough
to begin moving about, they have a fine, short coat of silky hair,
which a little later becomes longer and more lax as well as finer and
softer than in adults, and when they are about half grown the molt
begins on the nose and progresses slowly as in adults, with but one
molt to adult summer pelage. This change is not completed until
the animal is almost grown, and while it resembles the faded and
old worn pelage of adults, it can be recognized by its finer, softer
texture.

The number and sequence of molt waves can be satisfactorily deter-
mined only by a study of living animals as a supplement to the large
series of specimens available.

ABNORMAL PELAGE CONDITIONS.
ALBINISM.

Albinism is rare in the genus Thomomys, but a number of spec-
imens examined are fully albinistic and others partially so. One
albino of 7. bottx from San Francisco is in the U.S. National Museum
collection, and Townsend speaks of a ‘‘perfect albino’’ presented
to him at San Diego.’ Two specimens of T. fuscus in the Biologi-
cal Survey collection from Shuswap, British Columbia, are pure white
all over. A pure white specimen lacking all of the skull except the
front teeth was given me at Scottsburg, Oreg., but the species was not
identifiable; it was probably T. niger, a species otherwise known only
from pure black specimens. Specimensof T.awahnee from the Yosem-

1 Proc. U.S. Nat. Mus. X, 177, 1887.

1915.] INTRODUCTION. ol

ite, T. mewa from Fresno Flat, Cal., and 7. pascalis from Fresno, Cal.,
show large white patches on backs, arms, and breasts, and one of T.
yelmensis from Tenino, Wash., shows white patches on one side. The
white patches below, on chin, breast, or belly, are often constant and
furnish good specific or subspecific characters. Two specimens of
T. aureus from Shiprock, N. Mex., are light gray all over from a pre-
dominance of white or light gray hairs. One of 7. nevadensis from
Battle Mountain, Nev., is hoary over the head and back from a gén-
erous sprinkling of white hairs. A specimen of 7. fossor from the
Cochetopa Forest, Colo., is also gray over the head and back in the
same manner, but this may belong to another series of abnormal color
variation, caused by ticks or skin disease (fig. 4).

B2045-122

Fic. 4.—Skins of Thomomys (U.S. Nat. Mus. specimens), showing abnormal pelage conditions.
1. Thomomys fossor, No. 137661, female adult, Arapahoe Pass, Colo., July 5, 1905. The abnormal white
specking is probably due to tick bites.
2. Thomomys fuscus fuscus, No. 72949, female adult, Tobacco Plains, Mont., July 8, 1895. The white
on head may be due to ticks, mange, or skin disease.

' MELANISM.

~ Melanism is much more common than albinism, as one species, 7’.
mger, is known only from pure black individuals, and six other spe-
cies are more or less melanistic. In 7. townsendi, from southern
Idaho, there are 5 black specimens and 12 of the normal gray. The
Biological Survey series contains 36 normal yellow-gray, 18 black,
and 10 dark gray individuals. In 7. orizabx, of southern Mexico,
14 out of 16 specimens from the type locality on Mount Orizaba are
black. Of 8 specimens of. 7. wmbrinus from Boca del Monte 4 are
almost black and 4 are dark brown. In aseries of 11 specimens of 7.
atrovarvus, 3 are almost black. One specimen of 7’. peregrinus is fully
melanistic, but dichromatism may be rare in the species, as speci-
mens from many other localities are normal. In a large series of
specimens of 7. alticolus from southern Lower California 4 are so dark

99 NORTH AMERICAN FAUNA. [No. 39. .

as to be considered partially melanistic. There are two black skins of
T.talpoides from Red Deer, Alberta; one of T. rufescens from Fish Lake,
in the Turtle Mountains, N. Dak.; one of TJ. fulvus from Trumbull
Mountains, Ariz.; one of 7. toltecus from Fairbanks, Ariz.; and several
specimens of 7. botte from widely separated localities in California.
This is by no means an exhaustive list of melanistic specimens, but
merely serves to show how common and widely represented is this
variant phase.

Abnormal marking of another type is undoubtedly caused by -tick
bites. In many specimens of 7’. fossor from the mountains of Colo-
rado, Wyoming, and New Mexico small white specks occur scattered
irregularly over the top of head and shoulders, giving the animal a
speckled appearance similar to that of tick-bitten Texas horses. The
fact that these occur only on the parts of the gophers’ bodies not to
be reached by their teeth strongly suggests as their origin the pres-
ence of small ticks which engorge and remain attached to the skin
long enough to produce sores, which when healed are covered with
white instead of normal hair. Gophers with speckled heads are
represented from Colorado by specimens from 3 and 7 miles east of
Cochetopa Pass, from Hahns Peak, Rabbit Ear Mountains, Elk Head
Mountains, Longs Peak, Estes Park, Baxter Pass, and Pearl; from
Wyoming, by a specimen from Bridger Peak; from Utah, by speci-
mens from the Beaver Mountains; and from New Mexico, by specimens
from Costilla Pass, Costilla River (near source), Red River (near
source), Taos Mountains, and Twining.

A somewhat similar abnormal condition is found in a few speci-
mens of 7. fuscus from northwestern Montana. The top of the head
and shoulders becomes gray from a considerable mixture of white
hairs. The hair has a thin and unnatural appearance, the result, it
has been suggested, of mange or a similar skin disease.

VARIATION AND DISTRIBUTION.

The great variety of slight characters and lack of more trenchant
characters over a wide range, the abundance and almost continuous
dispersal of the genus in this range, and the evident uniform vigor
of the whole group indicate that the genus is at the height of its
development. It has pushed out from the other perhaps older, more
widely differentiated genera of the Geomyide and covered more ter-
ritory and produced more species and subspecies than all the rest of
the family. In 1895.Dr. Merriam recognized in the Geomyide, ex-
clusive of Thomomys, 8 genera embracing 37 species. The genus
Thomomys, without even a valid subgenus,! contains at least 88
recognizable forms of 40 species, and occupies about twice the area
of the rest of the family. (See figs. 5 to 10.)

1 See first footnote, p. 17.

1915.} INTRODUCTION. 23

Thomomys shows its vigor and adaptability by occupying the terri-
tory not inhabited by other more robust members of the family, which

a

al
5 (o)9 Vesti
); :
Sy)
Nai y

Aca

| B 2046-122
Fic. 5.—Map showing distribution of certain species and subspecies of Thomomys, and .ocalities at which
specimens were collected.

1. fuscus. | 6. bridgeri. 11. canus. | 16. helleri. 21. yelmensis
2. loringi. 7. uinta. 12. monticola. 17. bulbivorus. 22. melanops
3. saturatus. 8. quadratus. 13. pinetorum. 18. hesperus. 23. limosus.
4. myops. 9. townsendi. 14. mazama. 19. douglasi. 24. niger.

5. columbianus. 10. nevadensis. 15. nasicus. | 20. oregonensis.

hold the milder, more luxuriant valleys. It lives in almost every
desert valley and rugged desert range of mountains. as well as on the

94 NORTH AMERICAN FAUNA. : [No. 39.°

semiarid plains and stony slopes of the great mountain ranges, and
even up through the mountain forests to timber line and often far

— = = =

hi
\

Wi Hye:

i (((( Me \

: AQF
Zits

iS)

=

Dee Sy ere ic

B2047-122

Fic. 6.—Map showing distribution of certain species and subspecies of Thomomys, and localities at which
specimens were collected.

1.. bottx. 6. mewa. 11. pallescens. 16. jacinteus. | 21. perpes.
2. minor. 7. pascalis. 12. alpinus. 17. cabezonz. 22. operarius.
3. laticeps. 8. angularis. 13. awahnee. 18. puertz. 23. fisheri.

4. leucodon. 9. diaboli. 14. neglectus. 19. perpallidus.

5. navus. 10. infrapallidus. 15. altivallis. 20. albatus.

above on the peaks and crests of high ranges. That these gophers
however, prefer comfortable conditions and thrive on mellow

1915.1 INTRODUCTION. 25

soul and an abundant food supply is shown by the development of
more robust species under such favorable environment. All the
larger species inhabit rich valley bottoms. The smaller are mainly

ave San aca, |
B2048-122
Fic. 7.—Map showing distribution of certain species and subspecies of Thomomys, and localities at which
specimens were collected.

1. apache. 6. baileyi. 10. sinaloz. | 14. russeolus.
2. fulvus. 7. mearnsi. 11. chrysonotus. | 15. magdalenz.
3. desertorum. 8. lachuguilla. 12. nigricans. 16. anitz.

4. toltecus. 9. cervinus. 13. martirensis. 17. alticolus.
5. texensis.

from deserts where food is scarce and conditions for underground
life hard and unfavorable.

For unknown reasons there is practically no overlapping in range
of either genera or species in the family. This may be due to the

26 NORTH AMERICAN FAUNA. [No. 39. ,

ferocious dispositions of all members of thefamily.. For about 2,000
miles the ranges of Geomys and Thomomys meet across the Middle
West from Pembina, N. Dak., to the lower Rio Grande, apparently
without actual overlapping. The same conditions occur along the
ranges of large and small species of Thomomys, as at Fort Bridger,

Fic. 8.—Map showing distribution of certain species and subspecies of T’homomys, and localities at which
specimens were collected.

1. fossor. 4. desertorum. 7. chrysonotus.
2. agrestis. 5. aureus. 8. cervinus.
3. pervagus. 6. latirostris. 9. intermedius.

Wyo., where 7’. bridgerz is abundant in meadows and the little 7. ocius
in sagebrush only a few rods away; at Bear River, Wyo., where T.
pygmeus holds the same relation to T. bridgeri; in western Nevada,
where the small T. fisheri evidently keeps out in the sagebrush to avoid
the big T. nevadensis; and in Idaho, where T. townsend holds the rich
bottoms and the little 7. cdahoensis the dry sagebrush borders. Where

4915.) _ INTRODUCTION. D7

T.monticola meets the various forms of the bott# group along the west
slope of the Sierra Nevada, and other species along the east slope of
the range, the line of demarcation seems to be sharp, without actual
overlapping of the species, although one form may lap well past the

GUS af

= oe b> Ut eeeP aS ee
Ps . sO 70 5
: ~ Sree

2,
°
o
D

B n

by je

d b

f

B2050-122
Fic. 9.—Map showing distribution of certain species and subspecies of Thomomys, and localities at which
specimens were collected.

1. umbrinus. 4. atrovarius. | 7. goldmani.
2. orizabz. 5. sheldoni. | 8. perditus.
3. peregrinus. 6. nelsoni.

other in close juxtaposition, showing an apparent but not real over-
lapping of range. The two forms generally occupy different life
zones, or different types of ground, or else divide the territory on

28 NORTH AMERICAN FAUNA. [No. 39.

sonie other logical basis. This peculiarity of distribution is exempli-
fied by many species and seems to be universal between the slender-

B 2051-122

Frc. 10.—Map showing distribution of certain species and subspecies of Thomomys, and localities at which
specimens were collected.

1. talpoides. | 5. clusius. 9. pygmeus.
2. rufescens. 6. caryi. 10. ocius.

3. bullatus. 7. pryori.

4. nebulosus. 8. idahoensis.

and deep-rostrum groups. Further and more carefulstudy, however,
is needed to determine its cause.

1915. } INTRODUCTION. | 29

The more nearly related forms generally intergrade or coalesce
where their ranges join, some gradually, others abruptly, according
to the change of physiographic or climatic conditions.

In Dr. Merriam’s phylogenetic tree of the Geomyidex,! Thomomys is
given as one of the lower branches. The great numbers of genera
- and species of the family represented in southern Mexico and Central
America are spoken of as of the utmost interest in view of the time
and place of origin of the family to which they belong.’ If, as seems

probable, the family originated in tropical Mexico and Central Amer-
ica, Thomomys has pushed out to the greatest distance of any genus
in the family and now reaches from Boca del Monte (near Orizaba),
Vera Cruz, Mexico, north to Edmonton, Alberta, and from eastern
North Dakota to the Pacific coast.

Over this vast region the species occupy every life zone from the
Tropical to the Arctic Alpine, and the greatest extremes of heat and
cold, dryness and humidity, light and shade to which the area is sub-
ject. Although living underground and being partially nocturnal in
habits, the animals respond perfectly to environmental conditions and
vary in color according to light and shade, in claws and incisor teeth
according to soil texture, in length of fur according to temperature,
and in size and form of ears in accordance with length of fur. Thus
the great number of recognizable forms is a direct result of the wide
distribution and adaptation of a plastic group. Tae temptation to
theorize beyond the limits of actual knowledge in treating such a
group is difficult to resist.

The boundaries of ranges of the recognized forms follow closely
the border lines of known physiographic or climatic areas. Many
forms occupy definite divisions of a life zone, or, in some Cases,
as on a mountain slope or on several different mountain slopes,
extend regularly through a section of two or rarely three life zones,
maintaining their boundary lnes with surprising accuracy. On
some steep and narrow mountain peaks or ranges, however, a valley
species will extend up and down the sides through several climatic
zones without undergoing noteworthy change.

Permanent residence in a given region seems necessary for notice-
able change of characters, and even very restricted but very strongly
characterized areas have developed well-marked forms. Species with
the widest range are T. talpoides and T. rufescens of the great plains
and prairies of the Saskatchewan and Dakota regions, or T. fuscus and
T. fossor of the Canadian Zone areas of the Rocky Mountais. Some
interior valley forms also have extensive ranges, but to the south
and west, as the country becomes more broken and irregular, the
recognizable forms of Thomomys become more numerous and local.
California leads in the number of species, partly from its size and

IN. Am. Fauna No. 8, 24, 1895. 2 Tbid., 28.

30 NORTH AMERICAN FAUNA. [NO. 39.

great diversity of climatic and physical features and partly from the
fact that it has been the most thoroughly studied of the Western
States. The magnificent series of specimens in the Museum of
Vertebrate Zoology of the University of California has not only
added to the number of species known in the State, but has greatly
helped in defining the ranges of local species and correlating them
with the distribution areas.

MATERIAL EXAMINED.

In the present study of the genus Thomomys over 7,300 specimens
have been examined. Of these, 4,166 are in the Biological Survey
collection and 475 in the other collections of the National Museum.
Of the remaining specimens examined, 1,755 are in the Museum of
Vertebrate Zoology, 276 in the American Museum of Natural His-
tory, 234 in the Field Museum of Natural History, 177 in the Museum
of Comparative Zoology, 131 in the Stanford University collection,
90 in the Academy of Natural Sciences of Philadelphia collection, and
smaller numbers from the Victoria Memorial Museum, the Pro-
vincial Museum of British Columbia, the Kansas University collection,
the Oregon Fish and Game Commission collection at Reed College,
the California Academy of Sciences collection, the North Dakota
Agricultural College Museum collection, and private collections of
Mr. E. R. Warren, Mr. Stanley G. Jewett, Mr. W. D. Hollister, Dr.
William Bebb, and Mr. Ernest Thompson Seton. |

For the use of this material, which has in every case been most
cordially loaned, my appreciation is elsewhere expressed.

All of the types now in America (83 in number) have been ex-
amined, and through the courtesy of Mr. Oldfield Thomas a good
photograph of the skull of the type of T. umbrinus has been pro-
vided and also notes on the old types in the British Museum. Of
the eight types not examined, five (7. talpoides, T. umbrinus, T.
atrovarwus, T. alticola, and T. anit) are in the collection of the
British Museum. Good series of topotypes of all of these have been
available for study, and also topotypes of T. bulbivorus and T. douglast,
the types of which appear to be no longer in existence. Unfortu-
nately the type of T. talpoides is imperfect, lacking most of the skull,
but a good series of virtual topotypes have been available for study.

The type of T. botte in the Museum d’Histoire Naturelle, at Paris,
has not been seen, but large series of specimens from the type region
have been examined.

The study of this material shows many localities where a few speci-
mens would throw much light on the zonal distribution and relation-
ships of species, and brings up many problems that can be settled
only by careful field study and the collection of specimens at many

1915.1] INTRODUCTION. 31

localities. The distribution maps show many wide areas in which
no gophers have been collected and in which the resident species is
still unidentified.

The character of the specimens has generally been such as to
render the study most profitable. Well-made skins with cleaned
skulls, in series sufficient to show variation due to sex, age, and
abnormalities, have greatly simplified the task of classification.
The skulls of a few specimens collected on early expeditions have
been removed and cleaned and for the first time studied in relation
to ample material. In this way errors in the former use of names
have been corrected.

ACKNOWLEDGMENTS.

The greater part of the present revision was completed through a
study of specimens in the Biological Survey collection, but without
the additional material in other museums and private collections the
results would have been very imperfect. The friendly assistance and
cordial cooperation of the leading museums in the United States and
Canada and of private collectors have done much toward making
possible the results thus far obtained.

It is with great pleasure that I acknowledge my indebtedness to
Dr. Joseph Grinnell, director of the Museum of Vertebrate Zoology,
University of California; Mr. Gerrit 5. Miller, jr., curator of mam-
mals, United States National Museum; Dr. J. A. Allen, curator of
mammals and birds, American Museum of Natural History; Mr.
Charles B. Cory, curator of the Department of Zoology, Field Museum
of Natural History; Mr. Samuel Henshaw, curator of mammals,
Museum of Comparative Zoology; Dr. Witmer Stone, curator of
the Academy of Natural Sciences of Philadelphia; and Prof. J. C.
Branner, in charge of the biological collections at Stanford Univer-
sity; not only for the use of the large collections under their care,
but for courtesies and assistance in working on the collections at these
respective museums. For the use of specimens borrowed I am
indebted to Mr. Francis Kermode, curator of the Provincial Museum,
Victoria, British Columbia; Mr. P. A. Taverner, of the Victoria
Memorial Museum, Ottawa, Ontario; Prof. C. D. Bunker, in charge
of the collection of the Kansas University Museum; Prof. William B.
Bell, of the North Dakota Agricultural College; Dr. B. W. Evermann,
director of the Museum of the California Academy of Sciences;
Mr. William L. Finley, of the Oregon Fish and Game Commission; and
Prof. John F. Bovard, in charge of the University of Oregon Museum.
To private collectors my thanks are especially due to Mr. E. R.
Warren, Mr. Stanley G. Jewett, Dr. William Bebb, Mr. W. D.
Hollister, and Mr. Ernest Thompson Seton.

32, NORTH AMERICAN FAUNA. [NO. 39. .

Genus THOMOMYS Wied.

Thomomys Wied, Nova Acta Acad. Ces. Leop.-Carol., XIX, pt. 1, 378, 1839. Type,
Thomomys rufescens Wied. .

GENERIC CHARACTERS.

Upper and lower molars with both anterior and posterior enamel
plate always present. Upper incisors with plain, slightly rounded ante-
rior surface, a minute groove near inner edge of each. Insome species,
notably T. bulbivorus, T.lewcodon, and T. umbrinus, the groove is very
obscure and can be detected only with a lens; im others, especially the
monticola group, it is relatively deep, well defined, and readily seen
with thenaked eye. Inspecies where the nner groove is obscure there
is often a trace of a median line of depression along the front of the
incisor, but it is a mere trace, difficult to detect and not sufficiently
constant to serve as a diagnostic character. Fore feet relatively much
slenderer and claws lighter than in other genera of the family, but
claws long and well adapted to burrowing.

GROUP RELATIONSHIPS.

In the present paper the species are arranged so far as possible in
accordance with their relationships. There are no superspecific divi-
sions of sufficient value to be accorded subgeneric rank, but related
species have characters in common which distmguish them as a group
from other groups.

The lower outline of the rostrum is the strongest division character,
and this, though slight, seems clearly to separate two great groups.
In one group the rostrum is deep and heavy in side view, and its
lower outline slopes rather evenly from the anterior base of the upper
molars to near the base of the incisors. This group includes 7’. bulb-
worus, T. townsendi, T. botte, T. alpinus, T. perpallidus, T. fulvus,
T. umbrinus, and related species. In the other group the rostrum is
relatively slender in side view and its lower outline is abruptly arched
in front of the upper molars. This group includes T. talpoides, T.
fossor, T. douglas, T. monticola, T. fuscus, and related species. In
the text this division falls on page 96, the slender-rostrum group
beginning with T. talpoides and continuing to the end. The species of
these two groups seem not to intergrade.

HEAVY ROSTRUM GROUPS.

In the heavy rostrum groups are seven divisions, not all of equal
distinctness.

Thomomys bulbivorus stands by itself because of its concave ptery-
goids, small claws, and peculiar fur; but it is not very distant from
the bottz group with which it agrees in arrangement of mamme in 4
pairs.

1915.] | GENUS THOMOMYS. 33

The townsendi group also stands close to the botte group, as shown
by the number and arrangement of mamme; projecting, obscurely
grooved incisors; and short, wide skull. °

The bottx group is large, variable, and not sharply separated from
the perpallidus and fulvus groups; all of them agree in number and
arrangement of mamme, short ears, and many other characters. All
the forms except T. magdalenx and T..b.russeolus are dark or light ochra-
ceous, and it is doubtful whether these two pale forms may not really
belong to the perpallidus group, with which they agree in distinctly
erooved incisors.

The alpinus group includes long, narrow-skulled, dark-colored,
rather large-eared forms from high up in separate mountain ranges.
The mamme are as in the bottz group. The incisors are abruptly
decurved and distinctly grooved.

The perpallidus group includes mainly pale desert forms with very
small ears, but has no very distinctive group characters. The
incisors are abruptly decurved and distinctly grooved.

The fulvus group includes several mountain forms of rather bright
tawny shades, and paler tawny valley forms in the surrounding
country. Some strongly resemble forms of the wmbrinus group, but
all have the mamme in 4 pairs (2 of inguinal and 2 of pectoral). The
incisors are slightly projecting in some species. All are distinctly
orooved.

The umbrinus group is characterized by 3 pairs of mamme (2 of
inguinal and 1 of pectoral); by relatively short, wide skulls; by pro-
jecting, obscurely grooved incisors; and, in most species, by indented
or straight anterior base of zygomata.

SLENDER ROSTRUM GROUPS.

In the slender rostrum groups are five subdivisions, none very
strongly marked, but all with distinctly grooved incisors.

The talpoides group is best characterized by 6 pairs of mamme
(2 of inguinal, 2 of abdominal, and 2 of pectoral), by medium-sized
ears, and by general gray coloration.

The fossor group is distinguished by 5 pairs of mamme (2 pairs of
inguinal and 3 of pectoral) and by generally brownish coloration.
_ In the three mountain forms the ears are rather large, but in the two
_ valley forms (7. quadratus and T. fisheri) the ears are very small.

The douglasi group and the two following are distinguished by 4
pairs of mamme (2 of inguinal and 2 of pectoral).

The monticola group is very similar to the douglasi group, but has
shghtly larger and more pointed ears and generally more slender
skulls.

The fuscus group is distinguished by very small, pe ears and
light brown color.

98121°—15——_3

84 NORTH AMERICAN FAUNA. [NO. 39.-

List of Species and Subspecies, with Type Localities. .

Thomomys bulbivorus group (pp. 40-42):

Thomomys bulbivorus (Richardson)... . . Portland, Oreg.
Thomomys townsendi group (pp. 42-45):

Thomomys townsend: townsendi (Bach-

MAID) oto ene ae eae ems Canes oe Nampa, Idaho (?).
townsendi nevadensis Merriam....... Austin, Nev.

Thomomys bottxe group (pp. 45-63):

Thomomys bottx# botte (Eydoux & Ger-

Wials))=.2 2 bi atpeta ye ee hee ee ee ‘‘Coast of California.”’
bottzlaticeps Baird. 2a) ese Humboldt Bay, Cal.
botte leucodon Merriam........---- Grants Pass, Oreg.
botize navus Memranmt.- 509.20 ee Red Bluff. Cal.
botte mewa Merriam.....-.....---- Raymond, Cal.
boliz minor Bailey 24. - wae eee Fort Bragg, Cal.
boitz diabola Grinnell... --:-2.:2--- Diablo Range, Cal.
bottx angularis Merriam........-.--- Los Banos, Cal.
bottxe pascalis Merriam..........--- Fresno, Cal.
bottx pallescens Rhoads. .....-....-- Grapeland, Cal.
bottx infrapallidus Grinnell......... Carrizo Plain, Cal.
botte nigricans Rhoads........-.---- Witch Creek, Cal.
botts puerte Grinnellae 255-24 La Puerta, Cal.
botix-antia Adlens- poses see nore Santa Anita, Lower California.
botte aliicolussAVlen= 2) : Sierra Laguna, Lower California.
bottse russeolus Nelson & Goldman.. San Angel, Lower California.
magdalenxe Nelson & Goldman... .. Magdalena Island, Lower California. .
aliwallis Rhoadse=26- eee eee San Bernardino Mountains, Cal.
Thomomys alpinus group (pp. 63-68):
Thomomys alpinus alpinus Merriam..... Mount Whitney, Cal.
alpinus awahnee Merriam......-...- Yosemite Valley, Cal.
MEGUECLUS DAllOVe a - eae ee . San Gabriel Mountains, Cal.
jacinteus Grinnell & Swarth........ Round Valley, San Jacinto items.
Cal.
WATT CENSUS AU OMe sane ae eee ee etree San Pedro Martir Mountains, Lower
California.

Thomomys perpallidus group (pp. 68-80):
Thomomys perpallidus perpallidus Mer-

THAT 24 6 BS 5 PAS A eo es Na Palm Springs, Cal.
perpallidus albatus Grinnell....--.- Near Pilot Knob, Cal.
perpallidus chrysonotus Grinnell.... Ehrenburg, Ariz.
perpallidus perpes Merriam.....---- Lone Pine, Cal.
perpallidus canus Bailey....------- Deep Hole, Nev.
perpallidus aureus Allen. ...------- Bluff, Utah.
perpallidus apache Bailey. .-..-.----.- Lake La Jara, N. Mex.
cqbezone#, Mermam..25-24-t osen ase Cabezon, Cal.
OperariusMieniam. = eee ere Keeler, Cal.
latirosinis;) Mera: 225 =) eae ene Tanners Crossing of Little Colorado

: River, Ariz.
cerdinus Alien: sit ai. hie: Melee Phoenix, Ariz.

sinalow Merriam:253: 522 aaeek == Altata, Sinaloa, Mexico.

1915.] GENUS THOMOMYS. 35

Thomomys fulvus group (pp. 80-89):
Thomomys fulvus fuluus (Woodhouse)... San Francisco Mountains, Ariz.

fulvus pervagus Merriam....---.... Espanola, N. Mex.
fulvus desertorum Merriam........-- Mud Spring, Detrital Valley, Ariz.
fulvus intermedius Mearns.....--.-- Huachuca Mountains, Ariz.
Weious texensis Bailey 25.22.22 5-.-- Davis Mountains, Tex.
muiusetoliccus Allen... . 222.22... Colonia Juarez, Chihuahua, Mexico.
MICOGNS IAW 0 a2 ee Nee 8 Tee Animas Valley, N. Mex.
GarleuveMierrmiany. ~. 2.025. 222222. -22 Sierra Blanca, Tex.
facimoutiian Batley 2.0.25. 2: El Paso, Tex.

Thomomys umbrinus group (pp. 89-96): |

Thomomys umbrinus winbrinus (Rich-

FICISOIN) oyccc gs a ae ae ae eee Boca del Monte, Vera Cruz, Mexico (?).
umbrinus orizabe Merriam.........- Mount Orizaba, Puebla, Mexico.
umbrinus peregrinus Merriam....-- Salazar, Mexico, Mexico.
melsonaMleriam:. ..2......- 2.222: Parral, Chihuahua, Mexico.
SHEGONEMODIS.. 3202 a8. 26 2s bet Santa Teresa, Tepic, Mexico.
goldmanit Merriam......-......--- Mapimi, Durango, Mexico.
perditus Merriam........-.......-- Lampazos, Nuevo Leon, Mexico.
GrovariisvNilen nisl Soo s ee 2 Tatemales, Sinaloa, Mexico.

Thomomys talpoides group (pp. 96-110):
Thomomys talpoides talpoides (Rich-

FUISCI ic. 3-5) eg ee ee Fort Carlton, Saskatchewan, Canada.
talpoides rufescens Wied.......-.-- Fort Clark, N. Dak.
talpoides. clusius Coues: 2. 2---.-- Bridgers Pass, Wyo.
talpoides bullatus Bailey......---- Powderville, Mont.
talpoides nebulosus Bailey....-...- Sand Creek (Black Hills), Wyo.
nalpondes cary? Bailey... .....--- - Big Horn Mountains, Wyo.
taupoides pryort Bailey :.-..2. 2. : - Pryor Mountains, Mont.
talpordes agrestis Merriam... .----- Medano Ranch, San Luis Valley, Colo.
coummoianus Bailey... ........---. Touchet, Wash.

OCS NG sain Fort Bridger, Wyo.
idanoensis Merriam. ...........-.- Birch Creek, Idaho.
ugiiecis Merriam... 2... .--- Montpelier Creek, Idaho.
Thomomys fossor group (pp. 111-116):
ihomomys jossor Allen.......22..2.-2: Florida, Colo.
buidecn Mermam:-....5.........- Fort Bridger, Wyo.
| MugnieNentlaml. sh. ee. Uinta Mountains, Utah.
| quadratus quadratus Merriam. ...-- The Dalles, Oreg.
| quadratus fishert Merriam. .......- Beckwith, Cal.
|

Thomomys douglasi group (pp. 116-121):
Thomomys douglasi eougias (Rich-

2 IPG SION) Sy Sas a ea a ae Fort Vancouver, Wash.
) ian, oregonus Mierriam= 1.42 Oregon City, Oreg.
douglasi yelmensis Merriam. ...... Tenino, Wash.
douglast melanops Merriam. ...... Olympic Mountains, Wash.
y mosis Memiam.:. 2.0... 2....--- White Salmon, Wash.
Mmcr MeTmtam 6k Sk Seaton, Oreg.

Thomomys monticola group (pp. 121-126):
Thomomys monticola monticola Allen.. Mount Tailac, Cal.

monticola mazama Merriam........ Anna Creek, near Crater Lake, Oreg.
monticola pinetorum Merriam. . . - - Sisson, Cal.
monticola nasicus Merriam. ...-. - - Farewell Bend, Deschutes River, Oreg.

monticola hellert Elliot........---. Goldbeach, Oreg.

386 NORTH AMERICAN FAUNA. [No. 39.

Thomomys fuscus group (pp. 126-131):

Thomomys fuscus fuscus Merriam... -- - Head of Big Lost River, Idaho.
fuscus saturatus Bailey-...------- Silver (near Saltese), Mont.
juscus loringiisaileyeaiee «see. South Edmonton, Alberta, Canada.
fuscus myops Merriam. .....---.-- Conconully, Wash.
hesperus Merriam ues. hee a. ao Tillamook, Oreg.

Key to Groups of Related Species.

a1, Rostrum deep and evenly sloping in front of upper molars.
b1. Pterygoids concave on inner surface and convex on outer; mamme in 4 pairs,

bulbivorus group (p. 40).
b?. Pterygoids flat and straight.

cl, Mamme in 3 pairs (inguinal, 2-2; pectoral, 1-1)...... umbrinus group (p. 89).
c?, Mamme in 4 pairs (inguinal, 2-2; pectoral, 2-2).
d'. Skull short and wide; color mainly dark or light ochraceous,

botteze group (p. 45).
d?. Skull not conspicuously short and wide.

e!, Skull long and marrow color dark222325--545442eeee alpinus group (p. 63).
e?. Skull not conspicuously long and narrow; color mainly pale.
jf. Color pale buffy yellowish, or gray and black.
g'. Color buffy or yellowish (except apache)... perpallidus group (p. 68).
g?. Colorgray:and black=: 2 433.22 ee eee townsendi group (p. 42).
j?. Color tawny (light or dark tawny): 525: 5222-45" fulvus group (p. 80).
a2. Rostrum slender, abruptly arched in front of upper molars.
b1. Mamme in 6 pairs or more (inguinal, 2-2; abdominal, 2-2; pectoral, 2-2),
} talpoides group (p. 96).
b?, Mamme in 4 or 5 pairs.

cl. Mammee in 5 pairs (inguinal, 2-2; pectoral, 3-3)....-...- fossor group (p. 111).
c?, Mammee in 4 pairs (inguinal, 2—2; pectoral, 2-2).
d!, Ears rather large and rounded at tips...........-.- douglasi group (p. 116).
d?, Fars large or small and pointed at tips.
e!. Ears relatively large and pointed...........--- monticola group (p. 121).
e*, Ears relatively small and pointed..............-..- fuscus group (p. 126).

-—Key to Species and Subspecies of Thomomys.
[Measurements in millimeters.]

a}, Size large; hind foot of ¢ 38 or more.
b1. Color uniformly dark sooty brown above and below; hind foot of ¢ 40 or more,
bulbivorus (p. 40).
6?. Color gray or black (dichromatic); hind foot of ¢ 38, of Q 35.
c!. Color in gray phase dark buily gray (23-52 5. oe eae eee townsendi (p. 42).
c?. Color in gray phase light buffy gray .......-..-...- 2 eae nevadensis (p. 44).
a?, Size medium or small.
b1. Size very small, hind foot of ¢ averaging 26 or less; color buffy gray or brown.
ce}. Color buffy gray.

d'. ‘Hind foot of o averaging 262). 2. . 222) 22ers ocius (p. 107),

a2. Hind foot, of 6 averaging.23-. 32-4422 54 5 ee idahoensis (p. 108).
c?. Color light brown.

d', Hind foot of jf averaging 25-22-2222 -2- see myops (p. 130).

d?, Hind foot of g averaging 22 (smallest of the genus).-.--- pygmeeus (p. 109).

1915.1 GENUS THOMOMYS. 837

6?. Size medium, hind foot of ¢ averaging over 26 and under 36.
e!. Color brown, mainly cinnamon-hazel, chestnut,.or russet. —
d'. Color dark and rich browns.
el. No conspicuous black on nose and face; hind foot of ¢ about 30,

limosus (p. 120).
e?. Much blackish on nose and face.

Ge prze smaller. hind foot of ¢ about27uai.2..l<S ee) - hesperus (p. 131).
f?. Size larger, hind foot of ¢ about 29.
g:. Skull with very small oval interparietal........-- melanops (p. 119).
g?. Skull with large triangular interparietal.............. helleri (p. 126).

d?. Color light, or bright or dull browns.
e!. Ears small but pointed and well formed.
ae aoe light brown; hind foot of ¢ about 27.
Eesuohtiy paler: molars dishter.. 05:2 ist sees! ...-.fuscus (p. 126).
2y slightly darker; molars heavier. ii2..-2.:2:..- --loringi (p. 129).
if: "Color duller and devin: shades of browne size ees
g. Skull very long and narrow; hind foot of ¢ about 29.saturatus (p. 128).
g?. Skull relatively short and wide.
hi. Incisors projecting beyond tip of nasals; hind foot of ¢ 29,

id pryori (p. 104).
h?. Incisors not projecting beyond tip of nasals.

i, Nasals truncate at posterior tip; hind foot of ¢ 27
quadratus (p. 114).
7. Nasals emarginate at posterior tip; hind foot of ¢ 28,

: caryi (p. 104).
e?. Ears relatively long and pointed.

f?. Color dull brown.

g:. Size larger, hind foot of ¢ averagirz 33_.......-.-.- bridgeri (p. 112).
g?. Size smaller, hind foot of ¢ under 33.
i}. Bulle large; posterior tip of nasals rounded........ fossor (p. 111).

k?. Bullze smaller; posterior tip of nasals emarginate..-.-uinta (p. 113).
f?. Color bright, warm browns.
g'. Size larger, hind foot of male averaging about 30.
hM, Skull relatively short and wide; interparietal triangular,
oregonus (p. 117).
h?, Skull long and narrow; interparietal small and oval.

. Zygomatic arches widest anteriorly...-..-- ...-douglasi (p. 116).
' @. Zygomatic arches widest posteriorly. ......-- yelmensis (p. 118).
g?. Size smaller, hind foot of ¢ about 28 or less.
h+. Color russet; bulle full and rounded..........--- mazama (p. 123).
h?. Color hazel; bull relatively small and angular.
. Color dull hazel; braincase very narrow....--- monticola (p. 121).

72. Color more yellowish hazel; braincase wider.
7. Nasals longer and conspicuously widened anteriorly,
nasicus (p. 125).
7. Nasals shorter and merely cuneate ...--..- pinetorum (p. 124).
c. Color not distinctly brown, but of various shades of ochraceous, tawny, buff,
gray, umber, or black.
d. eect strongly ochraceous.
. Incisors projecting conspicuously beyond tip of nasals.
uy 1. Color dark ochraceous, clouded with dusky in winter.

g'. Size large, hind foot of 3 averaging 33......-.-.------ bottze (p. 45).
g’?. Size smaller, hind foot of ¢ not over 30.
ee olor darker than. iV ea. 5 os fais eee sae minor (p. 51).
fColor behter than Ut. ote - tanta et 3 a diaboli (p. 51).

38 NORTH AMERICAN FAUNA. fa AL NOS 39.

f?. Color light, bright, or rusty ochraceous.
g'. Color light or bright ochraceous.
h}. Color bright ochraceous; skull heavy and angular,
angularis (p. 53).
h?. Color pale ochraceous; skull not heavy and angular,

; infrapallidus (p. 55).
g’. Color rusty ochraceous.

h!. Incisors heavy and not greatly projecting. ......-. laticeps (p. 46).
h?. Incisors slender and greatly projecting.
a. Hind footer g30' to s3e oe. es ee leucodon (p. 47).
7. Hind foot of ¢ 27 to:30!32 3 20.) eee navus (p. 49).

e’. Incisors not projecting conspicuously beyond tip of nasals.
jf}. Color light ochraceous.
g'. Size large, hind foot of 3 averaging about 32.

M, Bulle large; skull short and wide...-.-.......-... pascalis (p. 54).
h?. Bulle small; skull long and narrow..........-- pallescens (p. 55).
g?. Size smaller, hind foot of ¢ averaging 30 or less.
h*. Underparts butfy or whitish 92.2 3 eee cabezone (p. 76).
}?, Underparts light ochraceous: 2222222) seas puertz (p. 58).
jf?. Color dark ochraceous, or ochraceous heavily clouded with black.
g'. Size large, hind foot averaging 34...................altivallis (p. 62).

g’. Size medium or small.
h. Size small, hind foot of ¢ about 27.
=a) Skull narrow. :. 22.0 22 ee ee awahnee (p. 64).
?. Skull wide. 22.22) 4..2.. ae mewa (p. 50).
h?. Size medium, hind foot of 3 averaging 30-32.
“, Dorsal outline of skull arched, rostrum decurved.
7. Interpterygoid fossa wide, not sharply V-shaped,

apache (p. 75).
7. Interpterygoid fossa narrow, sharply V-shaped.

ki, Throat usually white, interparietal large and quadrate,
alpinus (p. 63).
k?, Throat rarely white, interparietal small and often triangular.

it. “Premaxille wide... 5.22202 ee alticolus (p. 60).
??, Premaxille narrow.
m!. Skull very long and narrow. ....-- martirensis (p. 67).
m*. Skull not very long and narrow. ..... nigricans (p. 56).

#?, Dorsal outline of skull straight, rostrum not decurved.

7. Zygomatic arches widest posterior to middle; nasals emarginate,

neglectus (p. 65).

j.2 Gygomatic arches widest anterior to middle; nasals usually

trumcate.. 2.22 se. chest 2c) eee eee eee Le US (0-0):

d.? Color not strongly ochraceous. :
e.1 Color tawny, varying from dark to bright and pale tawny.
jf. Size large, hind foot of 3 averaging 31 or more.
g'. Color bright tawny.

ht, Pail, feet, and belly well haired -<\!.723-= 2 =eeee pervagus (p. 82).
h?. Tail, feet, and belly very thinly haired.
7'. Skull short and wide. -...-..22-2-2--- == sinaloz (p. 80).
7. Skull long and relatively narrow--2--- 2-1 oes anitze (p. 59).
g’. Color dull tawny, bordering on buffy and fawn.
hi. Hind foot of g averaging 34 or more: _- 2.222.522: cervinus (p. 79).
h?, Hind foot of 3 averaging about 31 or less.
e. Color dull tawny. 33/57 s.c2 eee ee ee toltecus (p. 86).

7: Color yellowish tawily esc see eee es pa baileyi (p. 87).

1915. ] GENUS THOMOMYS.

f?. Size smaller, hind foot averaging 30 or less.
g'. Color dark tawny, clouded with black in winter pelage.

Ue THITICRaOtOl Ge ADOU L022 5)s251 5 <--biesiee Soh s fe 2 - fulvus (p.

h?. Hind foot of ¢ about 27.
u. Color dark, bordering on umber ...........-. intermedius (p.
?. Color brighter with less black on back.......... texensis (p.

g?. Color bright or pale tawny.
h'. Color very bright tawny.
v'. Incisors not conspicuously protruding beyond nasals,

desertorum (p.

7. Incisors conspicuously protruding beyond nasals.

Giese belly palortawiyee 222 aie i ei eed Oe nelsoni (p.

j?. Belly white or yellow.
ch Belly white 322-22 .2 4: Bh ey seg Le goldmani (p.
hee Belly -yellows<sa-5 -4- sae 1s de: BEE ae mearnsi (p.

h?. Color pale tawny.
u. Color brighter, more buffy tawny............ lachuguilla (p.
i. Color duller, more grayish tawny......-......-- perditus (p.
e?. Color not tawny.
j*. Color buff or orange, varying from light buff to rich orange-buff.
g'. Color bright orange-buff or buffy yellow.

h’. Size large, hind foot of type (¢ ad.) 36........ magdalene (p.

h?. Size smaller, hind foot not over 33.
u'. Zygomatic arches abruptly constricted posteriorly,

operarius (p. ’

?. Zygomatic arches not abruptly constricted posteriorly.

fe SrOStrull Mery Wroad 2. Se Lele latirostris (p.
Byes rostrum less broad. .-2 22-222. so2--41-.-- See aureus (p.
g°. Color buff or grayish buff.
h'. Color grayish buff.
fomoize Jaree. hind. foot of g. about 33....---..--.:-- canus (p.
#. Size smaller, hind foot of ¢ about 30.............- perpes (p.
h?. Color clear yellowish buff.
u. Size large, hind foot of ¢ averaging 34.......--. albatus (p. '

?. Size smaller, hind foot of ¢ about 30.

7. Nasals conspicuously truncate posteriorly. ..-russeolus (p.

7, Nasals usually slightly emarginate posteriorly.

cola marrow.and slender. =. 2 202 - perpallidus (p.
(eS Skalltshort and wide: +4222 7.22222: chrysonotus (p.

f?. Color not buff or orange.
g'. Color gray, varying from light to dark brownish gray.
h’. Color dark brownish gray; claws stout; ears larger.
H, Audital bulle long but not full and rounded.
7), Skull not heavily ridged in old ¢, hind foot about 30,

talpoides (p.

72, Skull heavily ridged in old ¢&, hind foot about 31,

rufescens (p.

7, Audital bulle full and rounded.

39

96).

98).

7 Color telter, pullee largers---ss2s---ces-- = - bullatus (p. 102).
42, Color darker, bulle smaller._.............. nebulosus (p. 103).
h2. Color light brownish gray or pale gray; claws slender; ears smaller.

2. Audital bulle large and globose.

em@oloryery pale:perayecness. sa s2 ot 3 ee = agrestis (p. 105).
fa COLT ROL SO Pale eTay= so. .+ see so eee clusius (p. 100).

40 NORTH AMERICAN FAUNA. [NO. 39. °

#, Audital bulle relatively narrower.
j'. Size larger, hind foot of ¢ about 28... .columbianus (p. 106).
7, Size smaller, hind foot of ¢ about 26 ..._..... fisheri (p. 115).
g’?. Color not gray, but dark or light umber or black. '
h}, Incisors abruptly decurved, inner groove conspicuous; color always
black.22f) 2.22) oo) Ae eee 3 eae niger (p. 121).
h?, Incisors conspicuously projecting beyond nasals, inner groove very
obscure; color not always black. :
v. Color generally black (dark umber in a few individuals),
orizabe (p. 90).
#, Color dark or light umber (black in a few individuals).
7. Color dark umber.
ki. Feet, tail, and belly almost naked... ....atrovarius (p. 95).
k?. Feet, tail, and belly with normal covering of hair.
1+. Skull very short and wide; color very dark,
umbrinus (p. 89).
22. Skull less short and wide; color bright. -peregrinus (p. 91).
9g, Colorlight umber"s 2 2 eee sheldoni (p. 93).

Description and Distribution of Species and Subspecies.
Thomomys bulbivorus Group.
THOMOMYS BULBIVORUS (RicHarpson).

Camas Pocket GOPHER (Camas Rat of Richardson).
(Pl. II, fig. 1; Pl. II, fig. 1.)

Diplostoma bulbivorum Richardson, Fauna Boreali-Americana, I, 206, 1829.

Geomys bulbivorus Richardson, Ann. Rept. Brit. Assn. for 1836, VI, 150, 1837.

Ascomys bulbivorus Wagner, Suppl. Schreber, III, 388, 1843.

Pseudostoma bulbivorum Audubon & Bachman, Quad. N. Am., IIT, 337, 1854.

Geomys (Thomomys) bulbivorus Giebel, Siug. 530, 1855.

Thomomys bulbivorus Brandt, Beit. Kennt. Saug. Russl., 188, 1855.—Allen, Bul. Am.
Mus. Nat. Hist., V, 56, April 28, 1893.—Miller, Proc. Biol. Soc. Washington,
VIII, 113, Aug. 6, 1893.

Thomomys [sub-genus Megascapheus| bulbivorus Elliot, Field Columbian Mus., zool. ser.
IIT, 190, May, 1903.

Type.—Collected on ‘‘Banks of the Columbia River, Oregon,”
probably Portland, the only place near the Columbia River where it
has since been taken. The type is said to have been in the Hudson
Bay Museum, but Oldfield Thomas writes (Mar. 15, 1915) that it
is not now in the British Museum collection.

Mstribution.—Willamette Valley, Oreg., from Portland and Forest
Grove south to Eugene; west to Grand Ronde (fig. 5).

Characters.—Largest known species of the genus; hind foot of
adult male 40-43 mm.; front claws small and weak; external ear
merely a thickened rim; tail almost naked; winter coat long and
furry; summer coat short and harsh; color dark brown or sooty, nearly
concolor above and below; mamme in 4 pairs, inguinal 2-2, pectoral
2-2, with 4 separate mammary glands.

$$ ae

1915.} THOMOMYS BULBIVORUS GROUP. 4]

Color— Winter pelage: Dark sooty brown, the plumbeous basal
color showing through on belly; ears and nose blackish; chin and
throat and usually small anal patch white; feet usually streaked or
mottled with white. Summer pelage: Washed with rusty brown
above and below. Young: Like summer adults but very thinly
haired, the skin showing through on belly. |
_ Skull—Short and wide with zygomatic arches usually widest
posteriorly; rostrum, slender; nasals short, falling considerably short
of premaxille at both ends; pterygoids convexly inflated and
divided by narrow fossa; bulle rather small and narrow but auditory
meatus unusually open or dilated. Dentition rather light; incisors
slender, greatly protruding, and usually white tipped; groove on inner
edge of upper incisors very obscure.

Measurements.—Average of 5 adult males from Salem and Beaver-
ton, Oreg.: Total length, 300; tail vertebre, 90; hind foot, 42. Largest
male: 302, 99, 43. Average of 4 adult females: 271, 81, 39. Skull
(¢ ad.):1 Basal length, 52; greatest length over incisors, 57; nasals,
19; zygomatic breadth, 36.5; mastoid breadth, 30.5; alveolar length
of upper molar series, 10.

Remarks.—This is the most aberrant form of the genus, marking
either an ancestral type or more probably a highly specialized de-
velopment of adaptive characters. Widely separated geographically
from the bottz group, with which it shows the closest affinities, it is
entirely isolated in the humid, rich-soiled valley of the Willamette.

The object of the projecting incisors becomes evident when these
gophers are trapped. During the dry season the soil of the Willam-
ette Valley becomes baked and so hard that digging open gopher
holes with a small shovel, knife, or sharp stick is difficult, and but
for a granular cleavage would be almost impossible. The adobe
earth breaks out somewhat as the segments of a pomegranate, but
in coarser granules. Evidently the little irregular clay balls mak-
ing up the gopher hills are pried out with the projecting incisors,
which by long use have become perfect miners’ picks. The upper
incisors project actually beyond the tip of the gopher’s nose and at a
convenient angle for prying out the earth balls. The lower incisors
project equally far but from farther back. The fur-lined lips close
behind the incisors, so that no earth can get into the mouth. The
short bristly fur surrounding the base of the incisors protects the
lips, and leaves the ‘‘pick and shovel’’ teeth free. That the teeth
are regularly used for digging is evidenced by the much-worn tips and
edges. The white tips are shown by the lens to be due to the wearing
away of the yellow enamel surface. The slight inner groove is usually
worn entirely away before it reaches the tip of the incisor. It seems

1No. 57322, U. S. Nat. Mus., from Salem, Oreg.

49, | NORTH AMERICAN FAUNA. [No. 29.

highly probable that the unusually weak front claws of this gopher
result from their disuse in heavier kinds of digging.

The subgeneric characters ascribed to this species consist of little
more than the convex wings of the pterygoids. While this character
is unique in the genus it involves only the form of a weak and variable
bone and seems too superficial for more than specific value. As there
are no other divisions of subgeneric rank in the genus this one seems
of doubtful value.

The large mounds of these gophers have been observed in Oregon
at Albany, Cottage Grove, Harris, Lyons, and almost continuously
along the roads from Salem to a little south and 12 miles east of
Kugene. |

Specimens examined.—Total number, 62, as follows:

Oregon: Beaverton, 12; Corvallis, 1; Eugene, 8; Gaston, 2; Grand Ronde, 3;
McCoy, 6; Mulino, 6; Portland, 5; Salem, 20; Sheridan, 2.

Thomomys townsendi Group.

THOMOMYS TOWNSENDI TOWNSENDI (Bacuman).
TOWNSEND Pocket GOPHER.
(PL. I, fig. 2; Pl. III, fig. 2.)

Geomys townsend Bachman (from Richardson’s MS.), Journ. Acad. Nat. Sci. Phila-
delphia, VIII, pt. 1, 105, 1839.
Thomomys nevadensis atrogriseus Bailey, Proc. Biol. Soc. Washington, X XVII, 118,

July 10, 1914. Type locality, Nampa, Idaho.

Type.—T ype locality erroneously given as ‘‘Columbia River,” but
probably southern Idaho and very probably near Nampa, where —
Townsend’s party camped to trade with Indians, August 22, 1834.
Type specimen in Acad. Nat. Sci. Philadelphia.

Distribution.—Valley of Snake River in southern Idaho, from
American Falls to Weiser (fig. 5).

Characters.—Size very large; ears small but pointed; claws medium;
mamme 4 pairs, inguinal, 2-2, pectoral 2-2; color dichromatic, a—
dark gray phase and a black phase; skull relatively wide and angular.

Color.—In gray phase: Upperparts dark buffy gray or sooty gray;
nose and face blackish; ear patch black; feet and tail soiled gray;
underparts washed with rich buff; cheek pouches lined with black
and white; chin white. In black phase: Dull, slaty black all over
except white patch on chin and toes and usually on lower part of
feet. There seems to be little seasonal variation except for a general
darkening with wear as the buff tips disappear from the hairs. Two
specimens from Caldwell, Idaho, collected May 30 and June 3, are
clearer, richer buff than any others and much darker and richer
even than any specimens of the subspecies nevadensis. This may be a
normal but rare pelage.

1915.] THOMOMYS TOWNSENDI GROUP. 43

Skull_—tLarge and wide with spreading zygomatic arches, narrow
nasals and rostrum, and short, wide braincase; sagittal and occipital
crests well developed in old males; audital bulle large but depressed to
level of basioccipital, rather angular; basioccipital thick and narrow;
pterygoids high but short and thickened at margins. Dentition
heavy; incisors long and slightly projecting; inner groove obscure.

Measurements.—-Aduit male: Total length, 305; tail vertebra, 100;
hind foot, 38. Adult female: 276, 75, 35. These are about average
measurements. Skull (3 ad.): ibaa length, 49; nasals, 18; zygo-
matic breadth, 35; mastoid breath, 29; inieoronbital Broadus 8;
alveolar length of upper molar series, 10.

Remarks.—The type of Bachman’s Geomys townsendi which evi-
dently was labeled, not by the collector but after it came into -the
possession of the Academy of Natural Sciences of Philadelphia, ‘‘ Rocky
Mts., J. K. Townsend,” is fortunately still extant. Bachman pub-
lished Richardson’s manuscript description of the species in 1839,
and stated that it came from the Columbia River. It was a mounted
specimen with the skull inside, and its identity has caused much
speculation. Through the kindness of Dr. Witmer Stone, of the
Academy of Natural Sciences, the type is now before me and the

skull has been removed and cleaned. The type is an immature female

and much faded, but the skull serves to identify the species at once
as one which I recently described under the name atrogriseus from
southern Idaho. I take pleasure in yielding my recent name in favor
of one which for 76 years has been misapplied. In 1834, J. K.
Townsend made his journey to the coast of Oregon through southern
Idaho, and very probably secured the type of the species named for
him near the present town of Nampa, where his party camped
August 22 to trade with the Indians.!. The type locality of Thomomys
townsendi may therefore be fixed as the vicinity of Nampa, where
these large gophers still abound and from which place we have a fine
series of specimens.

My atrogriseus was described as a subspecies of nevadensis, but now
nevadensis becomes a subspecies of townsend: since the latter is the
earlier name. The nearest relationships are with the botte and
perpallidus groups, but there is uoivelyyy no direct connection with
either at the present time.

Speciumens examvned.—Total number, 63, as follows:

Idaho: American, Falls, 10; Caldwell, 2; Nampa, 19; Payette, 14; Weiser, 4.
Oregon: Ontario, 2; Owyhee, 6; Vale, 6.

1Townsend, J. K., Narrative of a Journey Across the Rocky Mountains to the Columbia River, 136,
1839.

44 NORTH AMERICAN FAUNA. [No. 39.

THOMOMYS TOWNSENDI NEVADENSIS Merriam.
NEVADA PocKET GOPHER.
(Pl. III, fig. 3.)

Thomomys nevadensis Merriam, Proc. Biol. Soc. Washington, XI, 213, 1897.

Type.—Collected at Austin, Nevada (exact locality, bottom of
valley about 5 miles west of the town in moist soil near Reese
River) by Vernon Bailey, November 11, 1890. Type specimen in
U.S. Nat. Mus., Biological Survey collection.

Distribution.—Valleys of central and northern Nevada and south-
eastern Oregon, from Austin and Lovelocks, Nev., north to Alvord
Lake, Oreg. (fig. 5).

Characters.—Size large; hind foot 35-40 mm.; skull heavily ridged
and angular with projecting incisors; colors dichromatic, buffy gray,
or black; in the gray phase more buffy and in the black phase more
plumbeous than in townsendi; mamme in 4 pairs, inguinal 2-2,
pectoral 2-2.

Color.—In gray phase: Light buffy gray, slightly paler on belly,
feet, and tail; nose and cheeks plumbeous; throat white. In black
phase: Bluish black or plumbeous all over except white throat
and usually white feet. There is often a small white spot on top
of head and the white throat patch extends to breast on some speci-
mens. Intermediate specimens are bluish gray or mixed gray and
plumbeous. “sgh

Skull.—Heavy and angular with long nasals and rostrum; lateral
ridges joined in sagittal crest in old males; zygomatic arches wide
spreading with sight median constriction; bulle large but depressed
to plane of basioccipital and with sharp anterior points; basi-
occipital narrowed between bulle; pterygoids high, thin, and arched.
Dentition heavy; incisors slightly projecting; inner groove obscure.

Measurements.—Type (& ad.): Total length, 275; tail vertebre,
90; hind foot, 38. Topotype (@ ad.): 255, 82, 35. Skull (of type):
Basal length, 44; nasals, 15.5; zygomatic breadth, 31; mastoid
breadth, 26; alveolar length of upper molar series, 9.

Remarks.—None of the type series are in the black phase, but
in a series of 15 from Battle Mountain 7 are black, 5 are gray, and
3 are intermediate. Other melanistic specimens come from Halleck,
Argenta, Winnemucca, and McDermitt, Nev., and Lake Alvord, Oreg.
Like townsendi, this gopher inhabits moist fertile valley bottoms and
not the surrounding dry sagebrush land. Its range therefore is along
stream valleys, and while its close relationship with townsendi indi-
cates only subspecific rank, its range is apparently at present not
continuous with that form. On the west it is cut off from its next
nearest relative—griseus—by the Sink of the Humboldt and its desert
margins.

1915.] THOMOMYS BOTTZ GROUP. 45

Specimens examined.—Total number, 93, as follows:

Nevada: Argenta, 1; Austin, 9; Battle Mountain, 15; Big Creek, 3; Halleck, 5;
Independence Valley, 1; Lovelocks, 10; McDermitt, 2; Paradise, 8; Quinn
River Crossing, 14; Winnemucca, 7.

Oregon: Alvord, 1; Alvord Lake, 10; Tumtum Lake, 7.

Thomomys potte Group.
THOMOMYS BOTTA BOTTH (Eypoux & GERVaIs).
CALIFORNIA PocKET GOPHER.
(Pl. II, fig. 3; Pl. III, fig. 4; text fig. 2.)

Oryctomys (Saccophorus) botte Eydoux & Gervais, Mag. de Zool., VI, 23-24, pl. 21,

fic. 4, 1836.

Thomomys botte Baird, Proc. Acad. Nat. Sci. Philadelphia, VII, 335, 1855.
Thomomys bulbivorus Baird, Mamm. N. Am., 389, 1857. Not Diplostoma bulbivorum

Richardson.

Type.—Collected on ‘‘coast of California,’ probably at Monterey
or San Francisco. Type in “Collection de la Faculté des Sciences,”
Paris.

Distribution. Coast region of California, from Sonoma County
(Freestone) south to San Diego (fig. 6).

Characters.—Size large, males much larger than females; average of
hind foot in males, 33 mm., in females, 28 mm; ears short, a mere
thickened rim; color dull, dark yellowish brown, nearly unicolor;
skull well arched, heavily ridged, and angular, with slightly project-
ing incisors; grooves on inner edge of incisors very small; mamme in
4 pairs, inguinal 2-2, pectoral 2-2.

Color— Winter pelage: Upperparts dark ochraceous, heavily clouded
with black-tipped hairs; underparts lightly washed with dull ochra-
ceous over the plumbeous underfur, often almost as dark as upper-
parts; ear patch, nose, and cheeks blackish; lips and lining of pockets
white; tail usually dusky or brownish gray; feet usually white. Sum-
mer pelage: Very similar, but slightly brighter, clearer ochraceous.

Skull.—Adult male: Heavy and angular with long, decurved ros-
trum; lateral ridges occasionally meeting in very old age in sagittal
crest; nasals long, cuneate or slightly spatulate, narrow, and gener-
ally emarginate posteriorly; auditory meatus large and slightly
inflated; bulla rather depressed and angular; upper incisors project-
ing slightly beyond -tip of nasals. Adult female: Much smaller,
slenderer, and less ridged than in males, but with the same arched
outline, slender rostrum, and projecting incisors.

Measurements.—Average of 5 adult males from near Monterey: Total
length, 267; tail vertebrae, 81; hind foot, 33. Average of 5 females:
219, 69, 28. Skull (é ad.):1 Basal length, 45; nasals, 16.5; zygo-

¢

1No. 107856, U. S. Nat. Mus., from near Monterey, Cal.

46 NORTH AMERICAN FAUNA. [No. 39.

matic breadth, 29; mastoid breadth, 24; alveolar length of upper
molar series, 9. Skull of adult female (from same locality): 36, 13,
24, 19, 8.7.

Remarks.—In addition to the unusual discrepancy in size of males
and females of botte there is a great range of local variation in
specimens from widely separated localities. To the south this sub-
species passes into pallescens; to the north it intergrades with minor
and apparently with leucodon; to the east it grades into the larger
bright-colored angularis -of the San Joaquin Valley; and along the
eastern side of the valley it runs into the paler pascalis. But in
the coast region it is fairly typical from Bodega on the north to
San Diego on the south. :

Specimens examined.—Total number, 713, as follows:

California (western): Alameda, 1; Alhambra, 4; Arroyo Seco (Monterey County),
7; Banta, 7; Bear Valley (head of Carmel River), 2; Berkeley, 18; Big Pine
Mountains, 2; Bodega, 11; Carmel River, 1; Carmel Valley, 5; Carpenteria,
2; Colma, 1; Cone Peak, 1; Cuyama Valley (Schoolhouse Canyon), 6; Drum
Barracks (Los Angeles County), 2; Freestone, 12; Gaviota Pass, 3; Glendora;
6; Half Moon Bay, 1; Haywards, 14; Hernandez (San Benito County), 1,
Hueneme, 5; Inverness, 28; Jamesburg, 13; Jolon, 10; Laguna Ranch, 6; La-
gunitas, 1; La Honda, 5; Las Virgines Creek, 6; Little Pine Canyon, 1; Los
Angeles, 14; Los Gatos, 1; Mansfield, 5; Matilija (Ventura County), 9; Mil-
pitas Ranch (south base Santa Lucia Peak), 1; Mono Flats, 9; Monrovia, 1;
Monterey (Pacific Grove and Point Pinos), 34; Morro, 2; Mount Tamalpais,
1; Mountain View (Santa Clara County), 1; Nicasio, 84; Oakland, 1; Olema,
1; Pacific Grove, 1; Palo Alto, 44; Palos Colorados Canyon, 2; Pasadena and
vicinity, 22; Pescadero, 7; Pine Creek (Bailey’s Ranch), 2; Pine Valley
(Monterey County), 7; Petaluma, 6; Pleyto, 6; Point Reyes, 18; Point Sur,
6; Portolo (San Mateo County), 8; Posts, 11; Pozo, 2; Priest Valley, 9; Red-
wood City, 27; Salinas Valley, 4; San Diego, 8; San Francisco (Golden Gate),
9; San Jose, 2; San Lorenzo, 1; San Mateo, 2; San Simeon, 2; Santa Ana Can-
yon, 4; Santa Barbara, 1; Santa Clara County, 6; Santa Cruz, 6; Santa Cruz
Mountains, 1; Santa Inez, 2; Santa Margarita (San Luis Obispo County), 1;
Santa Monica, 3; Santa Paula, 1; Soquel (Santa Cruz County), 2; Stanford,
48; Stanford University, 7; Stony Creek (Monterey County), 1; Sur River, 1;
Tassajara Spring (Contra Costa County), 2; Tomales, 18; Tracy and vicinity,
25; Twin Oaks, 13; Union Island, 1; Ventura, 5; Walnut Creek, 16.

THOMOMYS BOTTA LATICEPS Barrp.

HumpBoupt Bay Pocket GOPHER.
(Pl. II, fig. 6.)

Thomomys laticeps Baird, Proc. Acad. Nat. Sci. Philadelphia, VII, 335, Apr., 1855.
Type.—Collected at Humboldt Bay, California, by Lieut. W. P.
Trowbridge, February 21, 1855. Type specimen in U.S. Nat. Mus.
Distribution.—Coast region of northwestern California, from Smith
River south to Eel River (fig. 6).
Characters.—Size and general appearance of bottz, but color warmer
and brighter and less clouded with black-tipped hairs; skull averaging
slightly wider, and nasals especially wider; mamme in 4 pairs.

1915.] THOMOMYS BOTT GROUP. . 47

Color—Upperparts uniform clear rusty ochraceous or almost
snuff brown, brighter than in bott# and with less black; underparts
light buffy ochraceous, in strong contrast with upperparts; feet and
distal half of tail and usually lips and chin whitish. Summer and
winter pelage identical in all specimens examined.

Skull—Similar to that of botte, but slightly wider in proportion
to length; middle of malar arch more indented; auditory meatus less
inflated; lateral ridges not meeting in old males; nasals wide and not
sharply emarginate posteriorly. Dentition about the same as in botte
or slightly heavier.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
264; tail vertebre, 88; hind foot, 33. Average of 5 topotypes —
(2 ad.): 226, 77,30. Skull (of topotype, ¢ ad.):1 Basal length, 43;
nasals, 16; zygomatic breadth, 30; mastoid breadth, 23; alveolar
length of upper molar series, 9.

Remarks.—The subspecies laticeps is closely related to bottx, but
seems not to intergrade directly with it, as a smaller form of bottz
comes between. It can stand, however, as a well-marked subspecies.
Specimens from the region of Humboldt Bay only are typical, while
those from Crescent City are slightly paler. Two from Rio Dell
may be grading toward leucodon and a small female from Alton
Junction is doubtfully referred to laticeps. Specimens from Cuddeback
and Hoopa are clearly intermediate between laticeps and leucodon,
but are referred to the latter.

Specimens examined.—Total number, 90, as follows:

California (coast region of Humboldt and Del Norte Counties): Alton Junction,

1; Arcata (on Humboldt eps 49; Crescent City, 10; Eureka, 1; Requa, 6;
Rio Dell, 2; Smith River, 2: Table Bluff (on Eimmbold? Bay), 16; Tsenidadd ay

THOMOMYS BOTTA LEUCODON MeErRrR1aAM.

WuitE-TooTHED PocKET GOPHER.
(PL. II, fig. 11; Pl. II, fig. 5.)
Thomomys leucodon Merriam, Proc. Biol. Soc. Washington, XT, 215, July 15, 1897.

Type.—Collected at Grants Pass, Rogue River Valley, Oregon, by
Clark P. Streator, December 17, 1891. Type specimen in U. S. Nat.
Mus., Biological Survey collection.

Distribution.—Portions of northern California and southwestern
Oregon, from Grants Pass, Oreg., south to Fairfield and Placerville,
Cal. (fig. 6).

Characters—Smaller, lighter, and brighter colored than bottz; in
winter pelage, very similar in color and markings to laticeps, much
brighter in summer pelage; skull short and wide, with slender, pro-
jecting incisors, which are generally white or white tipped and with
obscure inner grooves; mamme in 4 pairs.

INo. 58452, U. S. Nat. Mus.

48 NORTH AMERICAN FAUNA. [No. 39.

Color.— Winter pelage: Upperparts uniform dark rusty ochraceous
or snuff brown; nose dusky; underparts light buffy ochraceous; feet,
chin, and often cheeks and spots on belly white. Summer pelage:
Upperparts dark cinnamon; underparts clear bright cinnamon.

Skull.—Low and wide with short, wide braincase, wide-spreading
zygomatic arches, widest posteriorly, and wide, quadrate, or trian-
gular interparietal; nasals short, deeply emarginate; bulle small;
pterygoids narrowly V-shaped. Dentition very light; incisors slender,
mainly white or white tipped and projecting far beyond nasals and
premaxille.

Measurements.—Type (young 3): Total length, 221; tail verte-
bree, 68; hind foot, 29. Topotype (old ¢): 246, 72, 32. Average
of 4 topotypes (females): 188, 60, 28. Skull (of type): Basal length,
36; nasals, 13; zygomatic breadth, 26; mastoid breadth, 19; inter-
orbital breadth, 6.5; alveolar length of upper molar series, 7. Skull
of topotype (old 3): 41, 15.8, 29, 23.5, 6, 8.

Remarks.—Typical leucodon differs widely from typical bottz, but
as the gap is largely bridged by the somewhat divergent characters
of navus, and the chain of intergradation between the two extremes
is complete, there is ample ground for grouping not only leucodon
but navus and mewa as subspecies under botte. Only specimens
from the Rogue River Valley can be considered typical of leucodon,
but in spite of much variation specimens occurring over a wide range
of rough mountain, valley, and foothill country south to Fairfield,
Cal., can better be referred to leucodon than to any other form.
Skins of Fairfield specimens are indistinguishable from topotypes of
leucodon, but the skulls are close to navus. The principal variation —
over this range is in a reversion toward the cranial characters of
bottze to the southward and a brightening of color in the less humid
interior. These brighter-colored specimens are externally very simi-
lar to typical fulvus of the San Francisco Mountain plateau region of
Arizona, but comparison of skulls shows that the resemblance is
only superficial. Specimens from Hoopa Valley and Cuddeback are
clearly intermediate between lewcodon and laticeps but are listed
under leucodon. 'Those from Cuddeback also strongly suggest butte
but are rather small and in this approach mznor.

Specimens examined.—Total number, 313, as follows: |

California: Baird, 3; Battle Creek, 2; Big Valley Mountains, 12; Bartlett Moun-
tain, 5; Berger Creek (Mendocino County), 1; Blue Canyon, 1; Briceland, 6;
Burney, 5; Cahto, 1; Calistoga, 2; Cassel, 1; Cuddeback, 9; Dana, 4; Downie-
ville, 1; Dutch Flat, 2; Edgewood (Shasta Valley), 2; Eel River (head of,
near South Yolla Bolly Mountain), 1; Fairfield, 6; Fair Oaks, 13; Fall River
Mills, 2; Fort Jones, 1; Fyffe, 5; Gazelle, 6; Genesee, 3; Glen Ellen, 2; Gold
Run, 6; Goosenest Mountain (Shasta Valley), 1; Greenville,1; Hayden Hill, 1; -
Helena, 26; Hoopa Valley, 7; Hornbrook, 10; Indian Valley (Lassen County),
2; Lakeport, 1; Laytonville, 1; Lower Lake, 12; Mad River, near Kunz (Trinity
County), 6; Mountain House (Butte County), 8; Mount George, 1; Mount

1915.] THOMOMYS BOTTE GROUP. 49

St. Helena, 3; Mount Sanhedrin (3 miles west of summit), 8; Mount Veeder, 2;
Novato, 6; Old Fort Crook, 20; Petaluma, 3; Picard, 9; Placerville, 2; Post
Creek, 2; Prattville, 1; Quincy, 6; Rumsey, 6; Slippery Ford, 4; Snow
Mountain (Colusa County), 3; Susanville, 3; Tower House, 5; Ukiah, 7; Vaca-
ville (8 miles west), 7.

Oregon: Ashland, 2; Ferren, 3; Grants Pass, 27; Grizzly Peak (west slope), 5.

THOMOMYS BOTTA NAVUS Merriam.

RED PockET GOPHER.
(Pl. IV, fig. 3.)
Thomomys leucodon navus Merriam, Proc. Biol. Soc. Washington, XIV, 112, July 19,
1901.

Type.—Collected.at Red Bluff, California, by Clark P. Streator,
- December 26, 1893. Type specimen in U.S. Nat. Mus., Biological
Survey Peale stion.

Distribution. Sacramento Valley, Cal., froin Battle Creek, Tehama -
County, south to Tracy Lake, San J anes Valley (fig. 6).

Characters.—Size much smaller than bottx; color lighter; skull
relatively shorter and wider, with more projecting incisors, interme-
diate to some extent between. botie and leucodon; mamme in 4 pairs.

Color.— Winter pelage: Upperparts light rusty ochraceous, darker
on back, face, and nose; ears blackish; underparts pale buffy or
ochraceous; feet whitish. Summer pelage (as shown in one July
specimen from Marysville Buttes): Slightly brighter and more fulvous
above and below than in winter.

Skull.Short and wide with zygomatic arches widest posteriorly;
nasals short, cuneate, emarginate posteriorly; bulle not large but
smoothly rounded. From the skull of leucodon it differs in smaller
size, narrower interorbital constriction, orange coloration and rela-
tively heavier incisors. Dentition light; incisors slender, protruding,
and bright orange except the white tips.

Measurements.—Average of 4 topotypes (¢ ad.): Total length,
203; tail vertebre, 67; hind foot, 27.5. Average of 4 topotypes
(@ ad.): 189, 61, 26.5. Skull (of type, ¢ ad.): Basal length, 35;
nasals, 13; zygomatic breadth posteriorly, 25; mastoid breadth, 20;
alveolar length of upper molar series, 7.

Remarks.—By treating leucodon as a subspecies of bottx, 1t becomes
necessary to place navus under bottz, with which it connects at one
angle as it does with leucodon at another. While typical Sacramento
Valley specimens of navus are strongly marked, others would go
with one form as well as with another. Fairfield specimens may with
equal propriety be placed with bottz, navus, or leucodon. Again, on
the east side of the Sacramento Valley, the gradation northward is
toward leucodon, and southward toward mewa. Wheatland and Jack-
son specimens, though having typical navus skulls, are dark and rich
in color, approaching leucodon.

98121°—]5——__4

50 NORTH AMERICAN FAUNA, ENO. 39.

Specimens ecamined.—Total number, 178, as follows:

California (Sacramento Valley): Carbondale, 23; Chico, 19; Colusa, 3; Davis, 1;
Grafton (Knights Landing), 7; Jackson, 4; Leesville (5 miles west), 2; Marys-
ville Buttes, 38; Mokelumne Hill, 2; Nelson, 1; Oroville (6 miles east), 2;
Payne, 1; Red Bluff, 42; Sacramento, 1; St. John, 1; Sites, 8; Tehama, 6;
Tracy Lake (San Joaquin Valley), 9; Wheatland, 4; Willows, 1; Winslow, 8.

THOMOMYS BOTTA MEWA Merriam.
DiacEeR Pine POCKET GOPHER.
(Pl. V, fig. 10.)
Thomomys mewa Merriam, Proc. Biol. Soc. Washington, X XI, 146, June 9, 1908.

Type.—Collected at Raymond, Madero County, Callfornia, by
N. Hollister, June 28, 1904. Type specimen in U. 8S. Nat. Mus.,
Biological Survey collection. ;

Distribution.—Foothill country on east side of San Joaquin Valley,
Cal.,,from Kernville north to Chinese (fig. 6).

Characters.—Smaller than bottx, blacker in winter pelage, more
intensely ochraceous-tawny in summer; skull relatively shorter and
wider with more rounded bulle; mamme in 4 pairs.

Color.— Winter pelage: Whole upperparts dull ochraceous-tawny,
heavily clouded with glossy black; underparts bright ochraceous; feet
whitish; tip of tail whitish or gray. Summer pelage: Upperparts
rich ochraceous-tawny, darkening to blackish on forehead and around
nose and ears; underparts lighter and brighter; nose rufous; feet and
tip of tail whitish.

Skull.—Smaller than in botte and relatively shorter and wider, with
fuller, shorter, more rounded bullae. Size of navus, but with less —
protruding incisors and with zygomatic arches not so conspicuously
widened posteriorly; incisors with well-defined inner groove.

Measurements.—Type (¢ ad.): Total length, 225; tail vertebre,
60; hind foot, 27. Topotype (¢ ad.): 196, 53,25. Skull (of type):
Basal length, 37.4; nasals, 13; zygomatic breadth, 24.7; mastoid
breadth, 20; alveolar length of upper molar series, 7.

Remarks.—This little, richly colored gopher occupies the Upper
Sonoran foothill country from Kernville north to Chinese but farther
north grades into navus, as shown by Wheatland and Jackson speci-
mens. Others from Merced approach pascalis in color but have more
nearly the cranial characters of mewa. Four females from Kernville
are peculiar in abruptly decurved incisors and very rich color. A
series of 13 December specimens from Salt Springs are in the fresh
black winter pelage with an iridescent purple gloss.

Specuomens ecamined.—Total number, 117, as follows:

California (east side of San Joaquin Valley): Bower Cave (near Mariposa), 3;
Camp Badger (Fresno County), 1; Chinese, 2; Coarse Gold, 6; Coulterville,

2; Fresno Flat, 5; Kernville, 4; Merced, 6; Merced Falls, 8; Porterville (8
miles east), 3; Raymond, 58; Salt Springs, 13; Three Rivers, 3; Wawona, 3.

1915.] | THOMOMYS BOTTA GROUP. D1

THOMOMYS BOTTA MINOR Barey.
MENDOCINO PocKET GOPHER.
(PL. IV, fig. 2.)
Thomomys bottz minor Bailey, Proc. Biol. Soc., Washington, XXVII, 116, July 10,
1914.

Type.—Collected at Fort Bragg, Mendocino County, California, by
James H. Gaut, November 30, 1905. Type specimen in U. S. Nat.
Mus., Biological Survey collection.

Distribution.—Coast region of California, from Cape Mendocino
south to Cazadero (fig. 6).

Oharacters.—Size smaller than botte#; color hohe darker; skull
narrower and slenderer, with especially slender rostrum, and deeply
emarginate posterior tip of nasals; mamme in 4 pairs.

Color.—In November pelage, upperparts dark, rich ochraceous or
cinnamon-brown, heavily obscured with black, and in most specimens
becoming almost black on nose, face, and around ears; feet white;
tail grayish; underparts ochraceous-buff; lips and lining of cheek
pouches usually white. Summer pelage but little lighter or brighter.

Skull.—Very slender and rather narrow, with very narrow and
deeply emarginate nasals, slender zygomata, rather large and quad-
rate interparietal, very small and short audital bulle. Dentition:
Incisors slender, ight colored, and projecting about as in botte.

Measurements.—Type (¢ ad.): Total length, 226; tail vertebre,

73; hind foot, 29. Average of 3 adult males: 219, 73,29. Topotype
(2 young ad.): 187, 58, 24. Skull (of type): Basal length, 35;
nasals, 13.4; zygomatic breadth, 23; mastoid breadth, 19; interor-
bital breadth, 4.7; alveolar length of upper molar series, 7.5.
- Remarks.—This little, dark, narrow-skulled gopher of the rugged
coast strip south of Humboldt Bay can not be included with bottz or
leucodon, and it is still more distinct from the big, wide-skulled laticeps.
It closely resembles nigricans, but can be distinguished at a glance
by its slender, protruding incisors.

Specimens examined.—Total number, 56, as follows:

California: Cape Mendocino, 2; Cazadero (7 miles west), 3; Ferndale, 11; Fort
Bragg, 4; Gualala, 9; Gurneyville (1 mile west), 1; Mendocino City, 12;
Petrolia, 4; Punta Arena, 7;* Rockport, 2; Westport, 1.

THOMOMYS BOTTA DIABOLI Grinne tt.
D1aBLo PocKkEeT GOPHER. |
Thomomys diaboli Grinnell, Univ. of Cal., Publ. Zool. XII, 313, Nov. 21, 1914.
Type.—Collected in Diablo Range, Merced County, California, at
Sweeney’s ranch, 22 miles south of Los Banos, by C. H. Richardson

and H. A. Carr, April 2, 1911.. Type specimen in Mus. Vert. Zool.,
Univ. of California.

59 NORTH AMERICAN FAUNA. [No. 39.

Distrrbution.—Inner ridge of the Coast Ranges along west side of
the San Joaquin Valley, Cal. (fig. 6).

Characters.—Size small, about as in navus; color bright yellowish
brown, much as in angularis, brighter than summer pelage of bottx;
skull a miniature of the bottz skull, with short, wide braincase and
projecting incisors. |

Color.—Summer pelage (March and October specimens): Upper-
parts bright sayal brown, with but slight darkening of black-tipped
hairs; ears and ear patches black; nose dusky; underparts clear,
bright ochraceous-tawny; lips and lining of cheek pouches white;
tail buffy; feet whitish. Winter pelage (as shown in-a few not fully
typical October specimens from Pacheco Pass and Pacheco Peak):
Much darker than in summer, but apparently not so black as in
botte.

Skull.—Similar to that of botte but smaller and slenderer, with
especially slender rostrum and zygomatic arches. Dentition: Teeth
very light, incisors slender and projecting well beyond tip of nasals,
pale yellow or whitish.

Measurements—Type (@ young ad.): Total length, 180; tail
vertebree, 60; hind foot, 25. Female topotype: 193, 65, 26.1 Aver-
age adult male from Pacheco: 218, 65, 30; female: 201, 57, 28. Skull
(of type, @ ad.): Basal length, 31; nasals, 10.3; zygomatic breadth,
22; mastoid breadth, 17.9; interorbital breadth, 7; alveolar length
of upper molar series, 7.2. Skull of topotype (¢ ad.): 34, 12, 24,
19.3, 6, 7.2. Skull of old male from Pacheco Pass:? 37, 12, 25, 20.7,
5.5, 7.0. skull of old female: 32, 10; 22) 10 Morar

Remarks.—This depauperate form of bottz seems to be a product —
of the hot and arid Upper Sonoran inner ridge of the Coast Ranges
along the west side of the San Joaquin Valley, in what Dr. Grinnell
calls the “hillside juniper association.” It needs no comparison
with the robust and massive-skulled angularis of the valley along its
eastern border, but could easily be confused with intermediates
between lewcodon and navus from similar arid ridges north of San
Francisco Bay. The skull, however, is more nearly of the true botte
type.

Specumens exammned.—Total number, 19, as follows:

California: McKittrick, divide west of (Kern County, 3,000 feet altitude), 1;
Pacheco Pass (Santa Clara County), 6; Pacheco Peak (Santa Clara County),
6; Sweeney’s Ranch (Diablo Range, Merced County), 6.

—

1 Foot measured dry. 2 No. 150799, Biological Survey collection.

1915.] THOMOMYS BOTTA GROUP. 58

THOMOMYS BOTTA ANGULARIS Merreram.

Los Banos PockET GOPHER
(PI. III, fig. 7.) |
Thomomys angularis Merriam, Proc. Biol. Soc. Washington, XI, 214, July 15, 1897.

Type.—Collected at Los Banos, Merced County, California, by J.
Ellis McLellan, January 1, 1894. Type specimen in U.S. Nat. Mus..
Biological Survey collection:

. Distribution.—West side of San Joaquin Valley, Cal., from sie
south to Santiago Spring; also Santa Clara, San Treen, and Salinas
Valleys (fig. 6).

Characters.—Size large, hind foot in males averaging 33; ears
short and thick; color lighter and brighter than in bottx,; skull short,
wide, angular, and heavily ridged; mamme in 4 pairs. inguinal 2-2,
pectoral 2-2.

Color.—Summer pelage: Rich ochraceous-buff, shghtly darkened
above with black-tipped hairs; ear patch pleck nose and cheeks
dusky or brownish; lining of pockets, lips, and sometimes chin,
white; feet white; tip of tail nearly or quite naked. Winter pelage:
Considerably darkened above by excess of black-tipped hairs;
slightly paler below—not quite so black as bottex.

Skull_—Males: Relatively shorter, wider, more angular, and
more heavily ridged than in bottx, with high sagittal crest in adults;
nasals short and widely spatulate; incisors projecting about as in
bott2. Kemales: Much smaller, not ridged or angular; short and
- wide with arched outline.

Measurements.—Average of 6 topotypes (¢ ad.): Total length,
258; tail vertebre, 80; hind foot, 33. Average of 4 topotypes
(@ ad.): 209, 66, 28.5. Skull (of type, small ¢ ad.): Basal length,
43; nasals, 15.7; zygomatic breadth, 28; mastoid breadth, 24; alveo-
lar length of upper molar series, 8.5. Skull (of topotype, larger
g ad.):! 45, 15, 31, 25, 9

Remarks.—Specimens from the type locality of angularis show
strongly marked characters, and considered alone these would indi-
cate that this is a distinct species not closely related to bottx; others,
however, from the Santa Clara, San Juan, and Salinas Valleys, while
nearest to angularis, show a tendency toward botte in both external
and cranial characters, and suggest the probability of intergradation
along the sides of these valleys with the smaller, darker, slenderer-
skulled bottz.

Specimens examined.—Total number, 101, as follows:

California: Bitterwater, 17; Coalinga, 2; Del Monte (7 miles southeast, in
Salinas Valley), 5; Kings City, 1; Los Banos, 54; Palo Alto, 2; Paraiso
Spring, 1; Paso Robles, 4; Salinas, 2; San Benito, 3; San Miguelito, 1;
Santiago Spring, 2; Tracy (8 miles south), 7.

No. 93787, U. S. Nat. Mus.

54 NORTH AMERICAN FAUNA. [No. 39.

THOMOMYS BOTTA PASCALIS Merriam.

FRESNO PockET GOPHER.
(Pl. IV, fig. 1.)
Thomomys angularis pascalis Merriam, Proc. Biol. Soc. Washington, XIV, 11, July
19, 1901,

Type.—Collected at Fresno, San Joaquin Valley, California, by
Clark P. Streator, March 4, 1892. Type specimen in U.S. Nat. Mus..
Biological Survey spllacidon.

Distribution.—East side of San Joaquin Valley, Cal., from Stockton
south to San Emigdio Canyon and Cuyama Valley Ge 6).

Characters.—Size approximately as in botte; hind foot averaging
32 mm. in adult males and 28 mm. in females; ear short; color dull
ochraceous; skull with broad base and swollen bulle; mamme in 4
pairs; inguinal 2-2, pectoral 2-2.

Color.— Winter pelage: Upperparts dull ochraceous, slightly dark-
ened on nose and face; ear patch inconspicuous; underparts paler
ochraceous; feet white; irregular white spots in many specimens on
legs, throat, and belly. Summer pelage: Brighter, more fulvous
above and below than in winter pelage. The summer pelage is
shghtly paler than in angularis and the winter pelage instead of being
darker than the summer, as in angularis and botte, is still paler.

Skull_—Shorter and wider, with larger bulle and more swollen
auditory meatus than in bottz; smaller than in angularis, less ridged,
with much larger, more rounded bulle and with narrower angular
process of lower jaw, and rarely with sagittal crest, even in old males.

Measurements.—Average of 3 topotypes (¢ ad.): Total length,
219; tail vertebra, 73; hind foot, 32. Average of 4 females from
type locality: 195, 63, 28. Skull (of type, ¢ ad.): Basal length, 41;
nasals, 16; zygomatic breadth, 28; mastoid breadth, 23; alveolar
length of upper molar series, 8.6.

Remarks.—Specimens from Tehachapi, Old Fort Tejon, and San
Emigdio Canyon and Mount Pinos, at the south end of the San
Joaquin Valley, vary from typical pascalis in smaller bulle and
larger auditory meatus, showing probable intergradation with botte of
the adjoining chaparral-covered ranges. Specimens from Cuyama
Valley about equally approach pascalis, bottz, and pallescens. On the
north, three specimens from Stockton might almost as well be
referred to angularis.

Specimens examined.—Total number, 169, as follows:

California (San Joaquin Valley): Alila, 5; Bakersfield, 2; Bakersfield (8 miles
northeast), 2; Bodfish, 11; Bodfish (12 miles below), 2; Buena Vista Lake, 6;
Buena Vista Slough, 1; Buttonwillow, 12; Cuddy Canyon (near Frazer Moun-
tain), 1; Cuyama Valley, 4; Delano, 1; Earlmart, 2; Fresno, 16; Lane
Bridge, 29; Lemoore, 7; Modesto, 5; Mount Pinos (5,500-8,500 feet altitude),
6; Oakdale, 4; Old Fort Tejon, 10; San Emigdio Canyon, 7; Stockton, 3;
Tehachapi, 10; Tejon Pass (8 miles west), 3; Tipton, 9; Tulare, 1; Walker
Basin, 10.

1915.] THOMOMYS BOTT GROUP. 55

THOMOMYS BOTTA PALLESCENS Ruoaps.

GRAPELAND Pocker GorHer.
(Pl. III, fig. 8.) .
Thomomys boitz pallescens Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1895, 36, Feb.
21, 1895. ;

Type.—Collected at Grapeland, San Bernardino Valley, California,
by R. B. Herron, March 22, 1894. Type specimen in Acad. Nat. Sci.
Philadelphia.

Distribution.—San Bernardino Valley, Cal., north to San Fernando
(fig. 6):

Characters.—Size of botte, but paler, and with buffy or whitish belly
and less ridged skull; mamme in four pairs.

Color.—Summer pelage: Upperparts dull ochraceous, darker along
line of back and around nose; ear patch blackish; underparts pale
buffy or soiled whitish. Winter pelage: Upperparts heavily clouded
with blackish; underparts clear pale buff in sharp contrast.

Skull—Long and narrow, with narrower braincase and less spread-
ing zygomatic arches than in bottx; lateral ridges parallel, never meet-
ing in old males; incisors decurved more nearly at right angles to
axis of skull.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
268; tail vertebre, 84; hind foot, 32.3. Average of 5 females from
type locality: 207, 64, 28.- Skull (of topotype, ¢ ad.):' Basal length,
43; nasals, 16; zygomatic breadth, 27; mastoid breadth, 22; alveolar
length of upper molar series, 9.5.

Remarks.—The subspecies pallescens is a large pale form of the
botte group, differmg from both pascalis and altwallis in cranial and
external characters, but combining some of the characteristics of
both with those of bottz. The only typical specimens examined are
from the great dry San Bernardino Valley. Those from San Fernando
are darker colored and almost as near to botte#, while Santa Paula,
Mono Flats, and Little Pine Valley specimens might with almost
equal propriety be referred to pallescens or botte.

Specimens examined.—Total number, 68, as follows (fig. 6):

California: Beaumont, 1; Cajon Pass (west slope), 1; Cajon Wash (near San Ber-
nardino), 10; Cold Water Canyon, 1; El Casco, 4; Glendora, 4; Grapeland, 18;
Jumpa Mountains, 2; Lytle Creek, 3; Réche Canyon (4 miles southeast of
San Bernardino), 3; San Bernardino, 17; San Fernando, 3; West Riverside, 2.

THOMOMYS BOTTA INFRAPALLIDUS Grinne t.
Carrizo Pian PocKET GOPHER.
Thomomys infrapallidus Grinnell, Univ. of Cal. Publ. Zool.; XII, 314, Nov. 21, 1914.
Type.—Collected on Carrizo Plain (Pimento Ranch, 7 miles south

of Simmler) San Luis Obispo County, California, by H. S. Swarth,
May 25,1911. Typespecimen in Mus. Vert. Zool., Univ. of California.

1No. 94013, U. S. Nat. Mus.

56 NORTH AMERICAN FAUNA. | [No. 39.

IMstribution.—Carrizo Plain, Cal. (fig. 6).

Characters.—Size large, set: as in bottx; color pale, belly buff or
whitish; skull similar to that of bottse, but with smaller molars.

ie — Saco pelage (May specimens): Upperparts dull ochra-
ceous-buff, paler than in angularis or pascalis; nose brownish; ears
blackish; underparts pale buffy or whitish (one specimen with almost
pure white belly); feet and tail white.

Skull.—Very similar to that of botte but with conspicuously
smaller molar crowns and smaller bulle. The narrow skull and pro-
jecting incisors readily separate this race from angularis or pascalis,
while the projecting incisors and shorter braincase distinguish it from
pallescens.

Measurements.—Type (& ad.): Total length, 248; tail vertebrae,
76; hind foot, 34. Female topotype: 205, 70, 29. Skull (of type):
Basal length, 40; nasals, 15; zygomatic breadth, 29; mastoid breadth,
23; interorbital breadth, 6.7; alveolar length of upper molar series, 8.

Remarks.—This is evidently a rather local form of the botiz group
occupying the hot, arid, alkaline desert valley known as Carrizo
Plain. It is paler and more strongly differentiated than is the
Cuyama Valley form, in which some of the same characters are less
emphasized. These specimens it seems better to identify with pas-
calis. Dr. Grinnell says of the Carrizo Plain, ‘ Faunally and zonally
it may be considered very arid, high Lower Sonoran. A sparse
prairie. vegetation grows on ground that is more or less strongly
alkaline over most of the area.’’!

Specimens excamrned.—Total number, 9, as follows:

California (Carrizo Plain): Painted Rock (5 miles north),3; Simmler (7 miles
south), 6.

THOMOMYS BOTTA NIGRICANS Ruoaps.

STEPHENS PocKET GOPHER.
(Pl. V, fig. 16.)

Thomomys fulvus nigricans Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1895, 36,
Apr i2Z 1895.

Thomomys aphrastus Elliot, Field Columb. Mus., zool. ser., III, 219, June, 1903.
Type from Santo Tomas, Lower California, 18 miles south of Ensenada at 50-100
feet altitude. Type collected by E. Heller; type specimen in Field Mus.
Nat. Hist.

Type.—Collected at Witch Creek, 2,753 feet altitude (7 miles west of
Julian), San Diego County, California, by F. Stephens, December 22,
1893. Type specimen in Acad. Nat. Sci. Philadelphia.

Distribution. —Southwestern California and northern Lower Cali-
fornia, from the San Jacinto Mountains, Cal., south to Ubar, Lower
California (fig. 7).

1 Univ. of Cal., Publ. Zool., XII, 314, 1914.

1915.] THOMOMYS BOTTH GROUP. 57

Characters.—Size smaller than bottz, with the same small ears and
very similar colors in both winter and summer pelage; skull slenderer,
with shorter bulle and less projecting incisors; mammee in 4 pairs,
meuinal 2—2, pectoral 2-2.

Color.— Winter pelage: Upperparts dark ochraceous-tawny, dark-
ened above by black-tipped hairs; nose, cheeks, and ear patches
blackish; underparts lightly washed with rich ochraceous; tip of -tail,
hind feet, and lining of cheek pouches usually whitish. Swmmer
pelage: Lighter and suffused with tawny above and below. Young,
paler, more buffy above and whitish below.

Skull.—Relatively narrower, slenderer, and less arched than that
of bottz; bulle shorter, more rounded; interparietal more quadrate;
external auditory meatus with less expanded rim. Dentition lighter;
incisors less projecting and more distinctly grooved. It differs from
fulvus, with which it was originally compared, in less spreading
zygomatic arches, slenderer rostrum, and relatively narrower brain-
case.

Measurements.—Topotype (¢ ad.):' Total length, 231; tail, 74;
hind foot, 30. Average of 5 typical males from San Jacinto Mountains:
232, 77, 30.2. Average of 5 females: 199, 65, 27.4. Skull (of topo-
type, ¢ ad.):? Basal length, 34.5; nasals, 13.5; zygomatic breadth,
23; mastoid breadth, 18; interorbital breadth, 6.5; alveolar length of
upper molar series, 7.5.

Remarks.—From botix, nigricans differs in lighter, slenderer skull;
more abruptly decurved and more distinctly grooved incisors; lighter
dentition; and numerous details of cranial and external characters;
but scarcely if at all in color. It is much smaller and darker than
pallescens with which it intergrades at Elsinore and Wildomar,
while Twin Oak specimens are clearly intermediate between nigricans
and botte.

In a series of 18 specimens from Santo Tomas, Lower California, I
find no characters by which to recognize Thomomys aphrastus as even
subspecifically different from nigricans. <A series of 32 specimens of
alticolus from the Victoria Mountains, farther south on the peninsula,
are scarcely distinguishable from nigricans in external characters, but
in cranial characters tend toward anite.

Specimens examined.—Total number, 401, as follows:

California (southern): Ballena, 1; Campo (San Diego County, at 2,500 feet alti-
tude), 8; Cuyamaca Mountains, 16; Dulzura, 14; Escondido, 3; Foster, 5;
Fullers Mill, 1; Grapevine Spring, 1; Hemet Valley, 5; Jacumba, 11; Jamul
Creek, 3; Julian, 15; Kenworthy, 2; La Puerta, 24; Mvuuntain Spring, 1;

3
Pine Mountains, 1; Poway, 1; Rose Canyon, 1; San Jacinto Mountains (south

end at 5,000 to 8,000 feet), 21; Santa Rosa Peak, 1; Santa Ysabel, 15; Schains
Ranch, 4; Strawberry Valley (in central part of San Jacinto range at 6,000
feet), 24; Thomas Mountain, 1; Tia Juana, 2; Tia Juana River (mouth), 1
Warners Pass, 7; Witch Creek (San Diego County at 2,750 feet), 53.

1No.. 148685, U.S. Nat. Mus. 2No. 148688, U.S. Nat. Mus.

58 NORTH AMERICAN FAUNA. [No. 39.

Lower California: Aguaji de las Fresas, 2; Ensenada, 6; Ensenada (canyon 20
miles east), 1; Fresnal, 1; Hanson Laguna, 11; La Grulla (at 7,000 feet alti-
tude), 13; La Huerta (Hanson Laguna Mountains), 1; Mattomi, 1; Milquatay
Valley, 2; Nachoguero Valley, 4; Parral, 5; Pinon (west slope San Pedro
Martir Mountains), 3; Rancho la Progresa, 5; Rancho Santo Tomas (12 miles
south of La Grulla), 3; Rancho Viejo (15 miles south of Alamos), 5; Rosarito,
2; San Antonio, 3; San Francisquito, 2; San Pedro Martir Mountains, 11;
San Quintin, 14; San Telmo, 18; San Ysidro Ranch, 4; Santa Eulalia, 7;
Santa Rosa, 1; Santo Tomas, 18; Socorro, 2; Tecate Valley, 2; Trinidad ~
Valley (northwest base San Pedro Martir Mountains), 2; Vallecitos, 6; Yubay,
4 (not typical).

THOMOMYS BOTTA PUERT GrinneELL.
LA Puerta Pocket GOPHER.

Thomomys nigricans puertxe Grinnell, Univ. of Cal., Publ. Zool., XII, 315, Nov. 21, 1914.

Type.—Collected at La Puerta (Mason’s Ranch, 5 miles west of Val-
lecitos, at the lower end of La Puerta Valley), San Diego County, Cali-
fornia, by F. Stephens, May 31, 1909. Type specimen in Mus. Vert.
Zool., Univ. of California.

Distribution.—La Puerta and San Felipe Valleys, Cal. (fig. 6).

Characters.—In size, proportions, and cranial characters, very
similar to nigricans, but much paler in both summer and winter
pelage; considerably brighter than cabezone.

Color.—Summer pelage: Upperparts pale ochraceous-tawny, be-
coming darker along median line of back; ears and postauricular spot
black; nose and face brownish or dusky; underparts pale or bright
cinnamon; feet and tail buffy or soiled whitish. Winter pelage:
Duller and more grayish, fading to pinkish- or vinaceous-salmon in
bright-colored individuals.

Skull.—N ot readily distinguishable from that of nigricans from the
surrounding chaparral slopes.

Measurements.—Type (3 young ad.): Totallength, 203; tail verte-
bre, 80; hind foot, 28. Skull (of type): Basal length, 32; nasals, 12;
zygomatic breadth, 21.2; mastoid breadth, 18; interorbital breadth,
6; alveolar length of upper molar series, 7.

Remarks.—The series of 43 specimens in the Museum of Vertebrate
Zoology labeled La Puerta were collected by F. Stephens, and Grinnell
says, ‘‘According to his field notebook, the majority were caught in
cultivated land on Mason’s Ranch, which is located at-the lower end
of La Puerta Valley. Others, however, were taken on surrounding hill-
sides.”’! The 43 specimens divide readily into 21 of the present pale
form and 22 of the typical dark-colored mgricans. These I assume
represent a pale desert form from the valley bottom where creosote
bush, mesquite, and cactus indicate extreme arid Lower Sonoran
conditions; and typical nigricans from the Upper Sonoran chaparral-

1 Univ. of Cal. Pub. Zool., XII, 315, Nov. 21, 1914.

1915.] THOMOMYS BOTT GROUP. 59

covered slopes along the sides of the valley. In a valley so narrow
the two forms undoubtedly intermingle to some extent along the
boundaries of their respective habitats, but most of the specimens
show no intermediate characters.
Specimens examined.—Total number, 23, as follows:
California: La Puerta, 21; San Felipe Valley, 2.

‘THOMOMYS BOTT ANITA Aten.

Cape SAN Lucas Pocket GopHeER.
(PL. V, fig. 11.)
Thomomys fulvus anitz Allen, Bul. Am. Mus. Nat. Hist., X, 146-147, 1898.

Type.—Collected at Santa Anita, Lower California, by Dane
Coolidge, May 28, 1896. Type specimen in British Museum.

Distribution.—Southern part of Lower California from Santana
south to Cape San Lucas (fig. 7).

Characters.—Slightly smaller than botte; larger than nigricans;
thin haired, bright tawny all over at all seasons, but little darker above
than below; skull heavy, with abruptly decurved incisors; mamme in
4 pairs. :

Color.—For most of the year bright tawny, nearly concolor; under-
parts slightly lighter because thinly haired; feet and tail half naked.
In new autumn pelage, apparently from September to December,
slightly duller, more grayish. Young, slightly paler, with whitish
belly. puede

— Skull.—Relatively heavier than in botts or nigricans, with short,
heavy rostrum, abruptly decurved incisors, short thick pterygoids,
and full rounded bulle.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,

- 243; tail vertebre, 78; hind foot, 32.8. Average of 5 topotypes

(? ad.): 208, 67, 30. Skull (of topotype, ¢ ad.):' Basal length,
40; nasals, 14; zygomatic breadth, 25.5; mastoid breadth, 21; inter-

- orbital breadth, 6.5; alveolar length of upper molar series, 8.8.

Remarks.—EKvidently intergradation is complete through local
forms up the peninsula through nigricans to bottz. The subspecies
anite strikingly resembles fulvus, however, and but for its direct
connection with bottx [should not change Dr. Allen’s original arrange-
ment, making it a subspecies of fulvus. The bottxe and fulvus groups
are widely separated geographically. Externally anit is not easily
distinguishable from sinalox, just across the gulf, but the skulls are
different.

Specimens examrned.—Total number, 99, as follows:

Lower California: Cape San Liicas, 11; La Paz, 12; Las Palmas, 1; Matancita,

6; Miraflores, 5; Rosarito, 20; San Jorge, 5; San Jose del Cabo, 10; Santana, 8;
Santa Anita, 16; Tres Pachitas, 1; Triunfo, 4.

1 No. 146831, U. S. Nat. Mus.

60 NORTH AMERICAN FAUNA. INO. 39.

THOMOMYS BOTTA ALTICOLUS Auten.
SreRRA Lacuna Pocket GOPHER.
(BIS L VE fies s5)
~ Thomomys fulvus alticolus Allen, Bul. Am. Mus. Nat. Hist., XII, 13, Apr. 4, 1899.

Type.—Collected at Sierra Laguna, Lower California, Mexico
(altitude 7,000 feet), by Dane Coolidge, July 10, 1896. Type specimen
in British Museum.

Distribution.—Victoria Mountains, southern Lower California (fig. 7).

Characters.—Size and proportions of anite or slightly smaller;
well furred and dark colored; skull shghtly slenderer than in anite,
but very similar; mamme in 4 pairs. |

Color.— Winter pelage: Upperparts dark ochraceous or snuff brown,
heavily clouded with dusky, usually darkest along median line of
back; nose, ear patch, and cheeks blackish; underparts washed with
bright ochraceous-tawny; lining of cheek pouches, feet, and tip of
tail usually whitish. Swmmer pelage: Clear ochraceous, scarcely
darker above than below, or varying to almost black in specimens
with a strong tendency toward dichromatism.

Skull.—Slightly slenderer than that of anitex, less slender than in
typical nigricans, and with larger bulle, wider occiput, and wide
premaxille.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
225; tail vertebree, 72; hind foot, 30. Average of 5 topotypes ( ? ad.):
203, 67,28. Skull (of topotype, ¢ ad.): Basai length, 38; nasals, 13.5;
zygomatic breadth, 24; mastoid breadth, 20; interorbital breadth,
6.5; alveolar length of upper molar series, 8.5.

Remarks.—This is a dark, mountain form of anitz from the higher
zone of nut pines and oaks. The two forms are so closely related
that the skulls are practically indistinguishable, but the skins greatly
differ in color. Compared with nigricans, the skins are scarcely dis-
tinguishable, but the skulls are strikingly different.

Specumens examined.—Total number, 52, as follows:

Lower California (southern): La Laguna, 20; Victoria Mountains, 32.

THOMOMYS BOTTA RUSSEOLUS Netson & GOLDMAN.
San ANGEL Pocket GOPHER.
(Pl. IV, fig. 9.)

Thomomys bottx russeolus Nelson & Goldman, Proc. Biol. Soc. Washington, X XII, 25,
Mar. 10, 1909.

Type.—Collected at San Angel, Lower California, at 100 feet
above sea level, by E. W. Nelson and E. A. Goldman, October 15, 1905.
Type specimen in U. S. Nat. Mus., Biological Survey collection.

1 No. 146836, U.S. Nat. Mus.

1915.1] THOMOMYS BOTTZ GROUP. 61

Distribution.—Known only from type locality (fig. 7).

Characters —Smaller than anit# or nigricans, with tiny ears, slender
skull, thin hair, and pale buffy coloration.

Color.—About as in perpallidus, or shghtly brighter. October
specimens: Upperparts varying from pale to bright buffy; nose
grayish; small ear patch dusky; underparts pale buffy, becoming
whitish on throat; feet and tail thinly clothed with whitish hairs.

Skull.—Light and slender, with narrow braincase, short narrow
posteriorly truncate nasals, wide premaxille, slender pterygoids,
full rounded bull, and narrowly constricted shaft of basioccipital.
Dentition: Incisors abruptly decurved at right angles to axis of skull.

Measurements.—Type (3 young ad.): Total length, 208; tail
vertebre, 73; hind foot (dry), 30. Topotype (¢ young ad.): 225,
82, 27 (dry). Topotype (2 im.): 88, 60, 27 (dry). Skull (of type):
Basal length, 35.5; nasals, 12.7; zygomatic breadth, approximately
24; mastoid breadth, 19; interorbital breadth, 6; alveolar length of
upper molar series, 7.5.

Remarks.—While in color practically identical with magdalene and
very similar to perpallidus, this desert form differs in cranial charac-
ters from magdalene and anitz in light slender skull, slender ptery-
goids, large rounded bulle, and narrow basioccipital; and from
perpallidus in narrow braincase, wider premaxille, and uniformly
truncate nasals. The characters strongly suggest a little slender-
skulled form of perpallidus. The type specimens are from an oasis
in a sandy desert where they were apparently isolated, but speci-
mens from Rosarito on the north and San Jorge on the south appa-
rently intergrade with anite.

One specimen from San Jorge and two from Rosarito might be’
thrown with russeolus (though the majority from the same places
are nearer anitx), and on this account this form was made a sub-
species of bottz rather than of perpallidus.

Specimens examined.—Four, from type locality.

THOMOMYS MAGDALENE Netson & GoLpMAN.
MaGDALENA PocKET GOPHER.
(PL. V, fig. 12.)
Thomomys magdalene Nelson & Goldman, Proc. Biol. Soc. Washington, XXII, 24,
Mar. 10, 1909. :

Type.—Collected at Magdalena Island, near west coast of Lower
California, by E. W. Nelson and E. A. Goldman, December 3, 1905.
Type specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Magdalena Island, Lower California (fig. 7).

Characters.—Larger than anitz, with heavy, angular skull; color
paler; hair thin.

62 NORTH AMERICAN FAUNA. [No. 39.

Color.—Type (collected December 3): Upperparts bright buffy
orange; underparts, feet, and tail pale buffy or whitish; a scarcely
naneasit he dusky ear spot.

Skull.—Old male: Very heavy and angular, with narrow brain-
case, short heavy rostrum, abruptly decurved incisors, angular and an-
teriorly spreading ome arches, deep lateral pits of palate, promi-
nent and revolutely margined pterygoids.

Measurements.—Type: Total length, 255; tail vertebre, 87; hind
foot, 36. Skull: Basal length, 41.5; nasals, 15; zygomatic breadth,
29; mastoid breadth, 23; interorbital breadth, 6; alveolar length of
upper molar series, 9.

Remarks.—The type and only specimen of magdalene is a fine old
male from this barren, sandy island off the coast of Lower California.
While having strongly marked affinities with anitx of the mainland,
the single specimen is so different as to indicate a well-marked insular
species. ‘The general characters are those of anite greatly accentu-
ated. It evidently belongs in the botte group and is not more different
from that species than is angularis, just over the ridge from the type
region of botte.

Specumen examined.—One, the type.

THOMOMYS ALTIVALLIS Raoaps.

SAN BERNARDINO MounTAIN PocKET GOPHER.
(Pl. III, fig. 9.)

Thomomys altivallis Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1895, 34, Feb. 21, 1895.

Type.—Collected on San Bernardino Mountains, California, (altitude
5,000 feet), by R. B. Herron, August 10, 1894. Type specimen in
Acad. Nat. Sci. Philadelphia.

Distribution.—San Bernardino Mountains, Cal. (fig. 6).

Characters.—Size large, hind foot in males averaging 34, in females
30; ears short; color dull and dark; skull long, well ridged, with
heavy dentition and sloping posterior base of zygoma; mammee in 4
pairs, inguinal 2-2, pectoral 2-2 (one individual with 2 pairs of
abdominal also).

Color.—Paler than in botte, slightly duller and darker than in pal-
lescens; upperparts dull ochraceous with trace of darker dorsal stripe;
ear patch, nose, and cheeks blackish; underparts, feet, and most of
tail buffy or soiled whitish. Pelage apparently brighter ochraceous in
summer than in winter, but only traces of much-worn winter pelage
are shown in the specimens examined.

Skull—tLonger and less spreading than in botte, less sharply ridged,
with less inflated auditory meatus, and heavier rostrum; larger than in
pallescens, with heavier rostrum and less sharply constricted posterior
base of zygomatic arch.

1915. ] THOMOMYS ALPINUS GROUP. 63

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
269; tail vertebre, 78; hind foot, 34. Average of 5 topotypes (9
ad.): 224, 64, 30. Skull (of topotype, old ¢):*' Basal length, 46;
nasals, 17; zygomatic breadth, 30; mastoid breadth, 24; alveolar
length of upper molar series, 10.

Remarks.—Thomomys altivallis apparently has its nearest relative in
pallescens, which stands intermediate in characters and geographic
position between it and botte. It is probably arobust, mountain form
of the botte group occupying the higher levels of the San Bernardino
Mountains. The exact type locality is unknown other than as 5,000
feet altitude in the San Bernardino Mountains. Herron, who col-
lected it, is no longer living, and S. N. Rhoads, who described the
species, has no further information as to its location. The type speci-
men, however, is of the large, high-mountain form, agreeing with the
large series from Bluff Lake and vicinity.

Specomens ecamined.—Total number, 155, as follows:

California (at various altitudes from 2,700 to 9,200 feet in the San Bernardino
Mountains): Bear Lake, Bear Valley, Bluff Lake, Doble, Dry Lake, Fish
Creek, Santa Ana River, Seven Oaks, Sugarloaf, Waterman Canyon, and
Fawnskin Park.

Thomomys alpinus Group.
THOMOMYS ALPINUS ALPINUS Merriam.

Mount WuitNEyY Pocket GOPHER.
(P1. V, fig. 13.)

Thomomys alpinus Merriam, Proc. Biol. Soc. Washington, XI, 216, 1897.

Type.—Collected on Big Cottonwood Meadows (8 miles southeast
of Mount Whitney), at 10,000 feet altitude in the Sierra Nevada,
California, by B. H. Dutcher, August 6, 1891. Type specimen in
U.S. Nat. Mus., Biological Survey collection. |

Distribution.—Southern part of the Sierra Nevada, Cal., at alti-
tudes between 6,000 and 11,000 feet, from Mount Whitney south to
Siretta Meadows (fig. 6).

Characters.—Size large; hind foot about 30 mm. in both males and
females; sexes very similar; ears large and conspicuous; colors dull
and dark; skull not ridged or angular; mamme in 4 pairs, inguinal
2-2, pectoral 2-2.

Color—Summer pelage: Dull dark ochraceous (much as in bottz),
becoming blackish along middle of back, paler on belly; nose blackish;
throat white; feet and distal two-thirds of tail whitish. One immature
specimen is paler, more ochraceous above and buffy below. Winter
pelage (long and faded on rump and sides in July specimens): Dull
dark clay color.

1 No. 89968, U. S. Nat. Mus.

64 NORTH AMERICAN FAUNA. [No. 39.

Skull——Compared with that of botte the skull is light and slender,
without conspicuous ridges or angles in old age; the incisors are
decurved; interorbital region wide and flat; interparietal quadrate;
auditory meatus smaller; zygomatic arches less spreading; and nasals
conspicuously widened anteriorly. From that of fulvus the skull
differs less in size and proportions, but has relatively heavier dentition ;
smaller bull; narrower basioccipital; larger, more quadrate inter-
parietal; and narrower posterior part of nasals.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
222; tail vertebre, 61; hind foot, 30.3. Average of 5 topotypes
(2 ad.): 222; 63, 30. Skull’ (of. type, :¢.ad.): Basal@leneth, 36;
nasals, 14; zygomatic breadth, 25; mastoid breadth, 19; interorbital
breadth, 6.6; alveolar length of upper molar series, 8.

Remarks.—Typical alpinus seems not readily to fall into any of
the larger groups of species. From fulvus it differs more than from
botte, and from the little slender-skulled monticola still more widely,
except in the large ears. From the perpallidus group it differs in
long, narrow skull, large molars, and in so many other details as to in-
dicate no close connection. From nigricans it differs less in -either
color or cranial characters than from bott2 but shows larger molars,
more spreading nasals, greatest width of zygomata posterior instead
of anterior to middle, and other distinctions. For the present it
may well be retained as the center of a group to which awahnee and
neglectus belong, and possibly also jacunteus and martirensis.

Specimens examined.— Total number, 158, as follows:

California (southern High Sierra): Bie Cottonwood Meadows (10,000 feet alti-
tude), 30; Cottonwood Creek (9,500 feet), 2; Cottonwood Lakes (11,000 feet),
25; Jackass Meadows (7,000 feet), 12; Jordan Hot Springs (6,700 feet), 8;
Menache Meadows (8,000 feet), 17; Olanche Peak (9,000-9,750 feet), 6;

Ramshaw Meadows, 3; Red Rock Meadows (9,000 feet), 2; Siretta Meadows
(9,000 feet), 1; Trout Creek (6,000 feet), 19; Whitney Meadows (9,000 feet), 33.

THOMOMYS ALPINUS AWAHNEE Merriam.

YOSEMITE PocKET GOPHER.
(Pl. V, fig. 9.)
Thomomys alpinus awahnee Merriam, Proc. Biol. Soc. Washington, X XI, 146, June 9,
1908.

Type.—Collected in Yosemite Valley, Mariposa County, California,
(at 4,000 feet altitude in bottom of valley, near the Sentinel Hotel),
by N. Hollister, June 14, 1904. Type specimen in U. S. Nat. Mus.,
Biological Survey polleeeiant

Distribution.—Western slopes of the Sierra Newman Cal., from
Sequoia, Tuolumne County, south to Tehachapi Peak ae 6).

Characters.—Size small; colors dull and dark; skull slender and nar-
row, with short rostrum and abruptly decumneds incisors; Mammee in
4 pairs.

1915.] THOMOMYS ALPINUS GROUP. 65

Color.—Summer pelage: Upperparts dull dark ochraceous; under-
parts varying from pale buffy to ochraceous; belly irregularly marked
and spotted with white in all Yosemite specimens; feet and distal
half of tail whitish. Winter pelage: Darker, more dusky ochraceous.

Skull—But little arched; not ridged or angular; rostrum short
and incisors abruptly decurved; interorbital region wide and flat;
interparietal usually short and wide; bullz small and well rounded.

Measurements —Topotype (¢ ad.): Total length, 220; tail verte-
bre, 75; hind foot, 27. Type (@ ad.): 194, 54, 26. Skull (of type):
Basal length, 32; nasals, 12; zygomatic breadth, 21; mastoid breadth,
17; alveolar length of upper molar series, 7. Skull of male from
Merced River: 34, 12.4, 22, 18.5, 7.5.

Remarks.—This form is not represented by a good series of adult
male specimens from any one locality, but specimens from scattered
localities throughout the Transition Zone, from Crockers (north of
the Yosemite Valley) south to the vicinity of Mineral King, show the
same general characters. Those from near Mineral King are some-
what darker than specimens from the Yosemite Valley and suggest
an approach to alpinus, while those in the Yosemite are paler and
have uregular white markings. Others from Kern River Lakes
and Piute Mountain and one from Tehachapi Peak are also referred
to it, but they show considerable variation.

Specumens examined.—Total number, 116, as follows:

California (southern and western slopes of the Sierra Nevada): Alta Mead-
ows (Giant National Forest), 2; Connell Meadow (Tulare County, 7,500
feet altitude), 11; Halstead Meadows (in Sequoia National Park, 7,000 feet), 5;
Horse Corral Meadows (in Sequoia National Park, 8,000 feet), 2; Huckleberry
(near Giant National Forest), i; Kaweah River (east fork, 5,600 and 8,900 feet),
2; Kern River Lakes, 2; Mineral King, 8; Piute Mountain (5,000 feet?), 31;
Sequoia (Crockers), 6; South Fork Merced River (4,000 feet), 1; Taylor
Meadow (Tulare County, 7,000 feet), 25; Tehachapi Peak, 1; Wawona, 2;
Yosemite Valley, 17.

THOMOMYS NEGLECTUS Batrey.

SAN GABRIEL POCKET GOPHER.
(P1. V, fig. 15.)

Thomomys neglectus Bailey, Proc. Biol. Soc. Washington, X XVII, 117, July 10, 1914.

Type.—Collected on San Antonio Peak (Bear Flat Meadows at
6,400 feet altitude) in the San Gabriel Mountains, California, by
James H. Gaut, July 23, 1905. Type specimen in U.S. Nat. Mus.,
Biological Survey collection.

Distribution.—Known only from type locality (fig. 6).

Characters.—Size medium; colors grizzled with a dark mixture of
ochraceous and black; skull long and narrow, with almost straight
dorsal outline; mammez: in 4 pairs, inguinal 2-2, pectoral 2-2.

98121°—15——5

66 NORTH AMERICAN FAUNA. [No. 39.

Color.—July pelage, in which 3 molts are shown: Upperparts dark
eray produced by a heavy mixture of black-tipped hairs over dull
ochraceous; nose, face, and ear patch blackish; tail gray; feet whitish;
underparts dull buffy.

Skull.—Adult males: Long and narrow, with narrow braincase;
nasals and rostrum long and straight; zygomatic arches slender;
bullz small and especially short and rounded; interparietal small and
triangular.

Measurements.—Type (3 ad.): Total length, 229; tail vertebre,
77; hind foot, 32 (measured dry). Topotype (¢ ad.): 206, 64, 31.
Female topotype: 211, 65, 31. Skull (of type): Basal length, 37;
nasals, 14.5; zygomatic breadth, 24; mastoid breadth, 19.5; inter-
orbital breadth, 6; alveolar length of upper molar series, 7.5.

Remarks.—Externally this species differs very little from altivallis
of the San Bernardino Mountains, but the skulls show such striking
differences that the two forms can not be considered closely related.
Neither does the present form show any characters of pallescens on
the south or of perpes on the north slopes of its range, but appears to
approach alpinus and awahnee of the southern Sierra Nevada. Its
actual range and real relationship present an -interesting problem
still to be worked out.

Specumens examined.—Three, from type locality.

THOMOMYS JACINTEUS GRrRInNNELL & SWARTH.

San Jacinto PockET GOPHER.
(Pl. V, fig. 14.)
Thomomys jacinteus Grinnell & Swarth, Proc. Cal. Acad. Sci., 4th ser., IV, 153-160,
Dec. 30, 1914.

Type.—Collected on San Jacinto Mountains (Round Valley, at
9,000 feet altitude), California, by H. S. Swarth, September 15, 1914.
Type specimen in Mus. Vert. Zool., Univ. of California.

Distribution.—Upper slopes of San Jacinto Mountains, Cal. (fig. 6):

Characters.—Size medium, slightly larger than nigricans, much
smaller than altivallis, and about the size of martirensis and neglectus;
colors dark; skull long and narrow with relatively straight dorsal
outhne; mamme in 4 pairs.

Color.—Summer pelage: Upperparts dark rich ochraceous, darkened
with black-tipped hairs, darkest along median line of back; nose dusky;
ears and ear patches black; underparts brighter ochraceous without
black-tipped hairs; lining of cheek pouches and usually lips white;
base of tail brownish; tip of tail and feet whitish. Winter pelage:
More clouded by black above.

Skull.—Long, low, and narrow, with especially long narrow brain-
case; zygomatic arches widest at anterior angle and narrowed pos-
teriorly with the narrowing of the braincase; nasals truncate poste-
riorly, generally long and narrowly cuneate, but in the type rather

1915.] THOMOMYS ALPINUS GROUP. 67

wide and spatulate; interparietal rather small, quadrate, narrowing
with age; pterygoids long and low and nearly parallel; interpterygoid
fossa not narrowly V-shaped. Dentition relatively heavy; incisors
curved abruptly downward and not extending much beyond tip of
nasals and premaxille.

Measurements.—Type (¢ ad.): Total length, 240; tail vertebre,
82; hind foot, 32. Topotype (¢ ad.): 234, 80, 31.5. Topotype
(2 ad.): 232, 76,29. Skull (of type): Basal length, 40; nasals, 14.5;
zygomatic breadth, 24.7; mastoid breadth, 20; interorbital breadth,
6; alveolar length of upper molar series, 8.8.

Remarks.—At first I was reluctant to separate this high-mountain
form from nigricans by skull characters alone, as the skins are indis-
tinguishable, but the series of specimens recently collected in Round
Valley in the San Jacinto Mountains (at 9,000 feet altitude) by H.S.
Swarth show that the skull characters are well marked and constant.
The series of 8 males and 5 females from the type locality, now before
me, have the long, low, narrow, straight skulls of cabezone with the
same peculiar pterygoids. They differ from cabezone in larger size,
heavier dentition, and much darker color, but show a decided rela-
tionship with that species. The resemblance to nigricans may be
entirely superficial just as it is with fulvus, the three forms being very
similar in coloration while evidently quite distinct. A close rela-
tionship with neglectus is indicated by the general type of skull, but
that form is much more extreme in its characters. From martirensis
the present form differs in a rather straight instead of well-arched
outline of skull. The two other high-mountain species with which
comparison is suggested are altivallis and alpynus from similar alti-
tudes in the San Bernardino Mountains and the Sierra Nevada. Of
these, altwallis is at once eliminated on account of its larger size and
its broad, well-arched, and massive skull, protruding incisors, and
narrowly contracted pterygoids. Thomomys alpinus, while so similar
in general size and appearance as strongly to suggest relationship,
has a very different type of skull; shorter, wider, and more arched
zygomata (widest posteriorly); and too many detailed differences to
admit of close relationship with the present form.

Specumens examined.—Total number, 25, as follows:

California (higher levels of San Jacinto Mountains): Round Valley (9,000 feet alti-
tude) 8; San Jacinto Peak (10,200 feet), 1; Tahquitz Valley (8,000 feet), 13;
Tamarack Valley (9,400 feet), 2; uncertain locality along the trail (8,500 feet), 1.

THOMOMYS MARTIRENSIS Aten.
San PepRo Martir Pocket GOPHER.
Thomomys fulvus martirensis Allen, Bul. Am. Mus. Nat. Hist., X, 147, 1898.

Type.—Collected on San Pedro Martir Mountains (at 8,200 feet
altitude), Lower California, by A. W. Anthony and E. C. Thurber, in
May, 1893. Type specimen in Am. Mus. Nat. Hist.

68 NORTH AMERICAN FAUNA. [ NO. 39.

Distribution.—San Pedro Martir Mountains, Lower California (fig. 7).

Characters.—Size considerably larger than mgricans; skull longer;
colors grayer in both winter and summer pelage.

Color.— Winter pelage: Very long and dark gray, not so black as in
ngricans; underparts buffy gray; tail and feet grayish; lining of
cheek pouches white; no white on chin and throat. Summer pelage:
Dull fulvous, not so intense as in nigricans or fulvus.

Skull.—Adult males: Very long and narrow, with especially nar-
row zygomatic arches and long, narrow braincase, in these characters
apparently exceeding neglectus and jacinteus; dorsal outline arched,
and rostrum depressed about as in nigricans.

Measurements.—Type (¢ ad.): Total length, 248; tail vertebra,
67; hind foot (measured dry), 31. The tail measurement of type is
short; other specimens have longer tails, 8 females averaging 70.
Skull: Basal length, 41.6; nasals, 15; zygomatic breadth, 25; mas-
toid breadth, 21; interorbital breadth, 6.5; alveolar length of upper
molar series, 8.

Remarks.—This is evidently a very local form occupying some high
valley or the crest of the range, since other series from not far distant
in the same range and only a little lower down, as at Vallecitos at
8,000 feet altitude, La Grulla at 7,000 feet, and Santo Tomas at 6,000
feet, are typical of mgricans. In December, 1914, A. W. Anthony
wrote me that the type series of specimens was collected at La Grulla
Meadows at 8,200 feet altitude in the San Pedro Martir Mountains, but
the series of specimens collected by Nelson and Goldman at La
Grulla (7,000 feet) are nigricans.

As the mountains rise to 10,200 feet, this may be a mountain-top
form of considerable range. The true relationship of these high-
mountain forms from alpinus to martirensis is not very clear and will
not be until definite field study shows whether they merge into lower
forms or are distinct and isolated.

Specumens examaned.—Fourteen, from type locality.

Thomomys perpallidus Group.
THOMOMYS PERPALLIDUS PERPALLIDUS Merriam.

PaLM SPRINGS PocKET GOPHER.
(P1. V, fig. 2.)
Thomomys talpoides perpallidus Merriam, Science, VIII, 588, Dec. 24, 1886.
Type.—Collected at Palm Springs (Agua Caliente) Riverside County,
California (at sea level on the edge of the Colorado Desert, 6 miles
south of Palm Springs Station), by F. Stephens, March or April,
1886.1 Type specimens in U.S. Nat. Mus., Merriam collection.

1 Tn the original description no type was designated by number, but Dr. Merriam refers to several skins
collected by F. Stephens in March and April on the Colorado Desert. In the Merriam collection are 4
skins and skulls labeled in accord with this statement, Nos. 22389, 22129, 33289, and 22219. Of these
No. 2218 is the only fully adult male and should have been made the type.

1915.4 THOMOMYS PERPALLIDUS GROUP. 69

Distribution.—Colorado Desert, southern California, from White-
water south to Salton Sea (fig. 6).

Characters.—Size medium, hind foot 31 mm: in adult male topo-
type; tail long; ears small; color pale buffy gray or whitish; mamme
in 4 pairs, inguinal 2-2, pectoral 2-2.

Color—Summer pelage: Upperparts buffy or cream color; ear
patches dusky; nose and cheeks grayish brown; underparts, feet, and
tail whitish, thinly haired. Winter pelage: Paler than in summer,
upperparts creamy or whitish. Young, buffy gray.

Skull.—Rather light and slender, without conspicuous ridges or
angles; zygomatic arches slender and parallel; nasals rather short,
cuneate, truncate, or slightly emarginate at posterior tips; inter-
parietal small and irregularly triangular, oval, or quadrate; bulle
short but full and rounded. Dentition rather light; incisors abruptly
decurved.

Measurements.—Average of 5 topotypes( ¢ ad.): Total length, 241;
tail vertebre, 84; hind foot, 31.5. Average of 4 topotypes ( ? ad.): 215,
78, 30. Skull (of topotype, ¢ ad.): Basal length, 37; nasals, 15;
zygomatic breadth, 26; mastoid breadth, 21; interorbital breadth,
6.4; alveolar length of upper molar series, 7.7.

Remarks.—This very pale, desert species occupies the scattered
moist oases in the hottest and driest of our deserts. Apparently its
range is not continuous, as no gopher hills are to be found over wide
areas of bare hot desert. The numerous colonies are to some extent
isolated, and differences too slight for even subspecific recognition
mark the specimens from almost every locality, but the main char-
-acters hold true over a wide area. Specimens from Carrizo Creek
srade toward the more robust form albatus of the Colorado River
bottoms on the California side, west of Yuma. A small series from
Salt Creek, north of the Salton Sea, seem to go better with perpallidus
than with any of the other three forms to which they might almost
as well be referred. Direct intergradation is shown with perpes and
through it with griseus and aureus.

The aureus group as heretofore considered now becomes the
perpalliidus group and includes perpallidus, albatus, chrysonotus,
perpes, canus, aureus, apache, cabezonx, operarius, and probably also
latirostris, cervinus, and sinalox.

Specimens examined.—Total number, 68, as follows:

California: Agua Dulce, 1; Baregas Spring (8 miles east), 5; Colorado Desert

(no specific locality), 5; Fish Spring, 2; Mission Creek, 1; Palm Springs, 35;
Salt Creek (Riverside County), 8; Salton Sea (west side), 1; Whitewater, 10.

70 NORTH AMERICAN FAUNA. [No. 39.

THOMOMYS PERPALLIDUS ALBATUS GRrRINNELL.

Wuitrt Pocket GOPHER.
(P1..V, fig. 1.)

Thomomys albatus Grinnell, Univ. of Cal., Publ. Zool., X, 172, 1912.

Type.—Collected on west side of Colorado River at old Hanlon
Ranch near Pilot Knob, Imperial County, California, by J. Dixon, May
7,1910. Type specimen in Mus. Vert. Zool., Univ. of California.

Distribution.—Southeastern California and northeastern Lower
California fiom Carrizo Creek south to Gardner’s Lagoon, Salton
River (fig. 6).

Characters —Size considerably larger than perpallidus; color
lighter; ears small; tail long; feet stout; mamme in 4 pairs.

Color.—Summer pelage (May and June): Upperparts pale buff or
cream color; nose and ears grayish brown; underparts white or
creamy with little or no plumbeous base to fur; feet and tail almost
naked but with scattered short white hairs. Winter pelage: Prac-
tically as in summer, but with more evident plumbeous base to fur on
belly. Young: Less buffy, more grayish white. :

Skull—tIn old males rather heavy, ridged, and angular, widest at
anterior angle of zygomata; rostrum short and heavy; incisors not
so abruptly decurved as in perpallidus; bulle full and rounded;
occiput rather sloping, not. so abruptly truncate as in aureus and
per pes.

Measurements.—Type(¢& ad.): Total length, 272; tail vertebre, 100;
hind foot, 35. Topotype (¢ ad.): 264, 91, 34. Topotype (¢ ad.):
229, 79,34. Skull (of type): Basal length, 41.5; nasals, 15; zygomatic :
breadth, 28; mastoid breadth, 22; interorbital breadth, 6.8, alveolar
length of upper molar series, 8.5. Skull of old female: 38, 13.5, 25,
DiiGeoysnen:

Remarks.—This is a robust gopher very close to perpallidus in
color, but in size and skull characters close to aureus. It occupies
the moist, white, river sand of the flats where there is an abundant
food supply. Specimens from Carrizo Creek are not typical but
nearer albatus than perpallidus. Others from Baregas Spring and
west of the Salton Sea are nearer perpallidus.

Specimens examined.—Total number, 72, as follows:

California: Carrizo Creek, 18; Colorado River bottoms opposite Yuma, 37; Salt

Creek (Imperial County), 11; Salton Sea (Imperial County), 6.
Lower California: Salton River (Gardners Lagoon), 1.

1915.] THOMOMYS PERPALLIDUS GROUP. 71

THOMOMYS PERPALLIDUS CHRYSONOTUS GEINNELL.
YELLOW-BACKED PocKET GOPHER.

Thomomys chrysonotus Grinnell, Univ. of Cal:, Publ. Zool., X, 174, 1912.

Type.—Collected at Ehrenburg, Arizona, (on dry mesa back from
river bottoms), by F. Stephens, March 27, 1910. Type specimen in
Mus. Vert. Zool., Univ. of California.

Distribution.—Southwestern Arizona and northwestern Sonora,
from Ehrenberg south to near mouth of Colorado River, and east to
Quitobaquiio (figs. 7 and 8).

Characters.—Size smaller than perpallidus; ears minute; skull
shorter, and bulle more globose; color about the same; mamme in
4 pairs.

Color.—Summer pelage: Upperparts bright to pale buff; ears brown
or dusky at tips; postauricular patch generally inconspicuous;
nose brownish; underparts and well up on sides whitish with pale
plumbeous base of hair; feet and tail thinly clothed with short white
hairs. Winter pelage: More grayish, fading to very pale buff in early
spring.

Skull._-Short and wide; rostrum relatively shorter and wider than
in perpallidus; bulle fuller and more rounded; upper incisors more
abruptly decurved; premaxille in type ending approximately even
with posterior tip of nasals, but in the majority of specimens extend-
ing well back of nasals; interparietal small and nearly triangular;
pterygoids short, thick, low, and wide apart..-

Measurements.—Type (¢ ad.): Total length, 217; tail vertebra,
73; bind foot, 30. Adult female from Yuma, Ariz.: 203, 65, 27.
Skull (of type): Basal length, 33.5; nasals, 13.5; zygomatic breadth,
23; mastoid breadth, 19; interorbital breadth, 7; alveolar length of
upper molar series, 8.

Remarks.—This seems to be a dry-mesa form more nearly agreeing
with typical perpallidus than with the more robust albatus of the
moist bottoms just across the Colorado River. Originally I identified
the type for Dr. Grinnell as perpallidus, but since that time he has
collected a fine series of topotypes of perpallidus at Palm Springs,
which show better diagnostic characters than do the series in the
National Museum collection, and fully warrant the separation of
the form ranging in the dry hot desert east of the Colorado River
in southwestern Arizona and northwestern Sonora.

Specimens examined.—Total number, 25, as follows:

Arizona: Ehrenburg, 1; Monument No. 204, east side of Colorado River, 1;
Quitobaquito, 2; Tacna, 1; Tule Wells (in Tule Mountains near Sonora line),
5; Yuma, 12.

Sonora: Cienega Well, 2; Mesa on east side of Colorado River, 20 miles below the
Arizona line, 1.

792 NORTH AMERICAN FAUNA. [NO. 39.

THOMOMYS PERPALLIDUS PERPES Merriam.

Lone Pine Pocket GOPHER.
(Pl. V, fig. 3.)

Thomomys aureus perpes Merriam, Proc. Biol. Soc. Washington, XIV, 111, July 19,
1901:

Thomomys scapterus Elliot, Field Columb. Mus., zool. ser. III, 248, 1903. Type
collected by E. Heller, in Hanopee Canyon, Panamint Mountains, California.
Type specimen in Field Mus. Nat. Hist.

Type.—Collected at Lone Pine, Owens Vailey, Inyo County, Cali-
fornia, by E. W. Nelson, December 23, 1890. Type specimen in U.S.
Nat. Mus., Biological Survey collection. .

Distribution.—Upper Sonoran desert valleys and mountain slopes
of eastern California, from near head of Owens Valley south to Hes-
peria and Morongo Valley, into the valley of Kern River, and east to
the Providence Mountains, Cal., and the Grapevine Mountains, Nev.
(fig. 6).

Characters.—About the size of perpallidus or slightly smaller, with
shorter tail, larger ears, slenderer skull, and brighter, stronger color;
smaller and grayer than aureus; mamme in 4 pairs. d

Color.—Summer pelage: Upperparts bright buffy ochraceous; nose
grayish brown; ear patch dusky; underparts buffy or creamy white;
throat often pure white; feet and tail thinly covered with short
silvery hairs. Winter pelage: Upperparts duller, darker, and more
grayish buff. Young, very similar to adults.

Skull.—Slender and light with rarely a trace of lateral ridges; well
arched; narrower and slenderer than in perpallidus, with especially
slenderer rostrum; palate slightly more arched than in perpallidus,
much less arched than in aureus; interparietal generally quadrate;
bulle full and rounded; nasals slightly emarginate or occasionally
truncate. Dentition rather ight; upper incisors decurved at right
angles to axis of skull.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
215; tail vertebre, 66; hind foot, 28.7. Average of 5 topotypes
(¢@ ad.): 210, 67, 28.5. Skull (of type): Basal length, 33; nasals,
12.3; zygomatic breadth, 22; mastoid breadth, 18.6; interorbital
breadth, 6.5; alveolar length of upper molar series, 7.5.

Remarks.—This small gray form of the perpallidus group occupies
Upper Sonoran desert valleys and slopes of eastern California. In
parts of the Mohave Desert where it extends down ‘nto Lower
Sonoran valleys it becomes slightly larger and paler than normal and
might almost as well be placed with aureus; while in the foothills of
the Sierra Nevada, San Gabriel, and San Bernardino Mountains it
becomes slightly darker than the typical form and may grade into
awahnee. It does not occur in Death Valley or Panamint Valley,

1915.1 THOMOMYS PERPALLIDUS GROUP. le

and gives place to operarius in the bottom of Owens Valley, and to
aureus in Ash Meadows and the Amargosa Valley. Its connection
with perpallidus on the south has been fully established by recent
collections, so that both perpes and aureus must stand as subspecies
of this earliest-described form of the desert group. Thomomys scap-
terus of Elliot proves to be typical of perpes.

Specimens examined.—Total number, 344, as follows:

California: Ash Creek (west of Owens Lake), 1; Barstow, 16; Benton Station, 4;
Bishop, 1; Cactus Flat, 1; Carroll Creek, 1; Copper City, 1; Coso, 19; Cushen-
bury Spring, 2; Daggett, 2; Fairmont, 1; Granite Spring, 1; Grapevine Ranch,
3; Haway Meadows (south of Owens Lake), 2; Hesperia, 1; Independence, 13;
Independence Creek, 2; Inyo Mountains, 1; Isabelle, 6; Kern River (South
Fork, near Onyx), 17; Lone Pine, 64; Lone Willow Spring, 7; Ludlow, 2;
Maturango Spring, 1; Mohave River, 4; Morongo Pass, 1; Mount Waterman
(north slope in San Gabriel Mountains), 1; New York Mountain (in Provi-
dence Range), 1; Orogrande, 16; Panamint Mountains (Johnson Canyon,
Perognathus Flat, Coal Kilns, Hanopee Canyon, head of Willow Creek, and
Cottonwood Creek), 23; Resting Spring, 24; Tuttle Creek, 5; Twelve Mile
Spring, 1; Victorville, 26; Walker Pass, 28; Warrens Ranch (in Morongo
Valley), 1; Warrens Well (east of Morongo Valley), 2; Weldon, 37; White
Mountains (pass between Deep Spring and Owens Valley), 1.

Nevada: Grapevine Mountains, 3; Thorps Mill, 1.

THOMOMYS PERPALLIDUS CANUS Batzey.

Gray Pocket GOPHER.
(Pl. V, fig. 6.)
Thomomys canus Bailey, Proc. Biol. Soc. Washington, X XIII, 79, May 4, 1910.

Type.—Collected at Deep Hole, at north end of Smoke Creek
Desert, Nevada, by Clark P. Streator, May 14, 1896. Type specimen
m U.S. Nat. Mus., Biological Survey collection.

Distribution.—Valleys of western and central Nevada, from Flow-
ing Springs, western Humboldt County, south to Cloverdale and
Monitor Valley, west to Honey Lake, Cal. (fig. 5).

Characters —Considerably larger than perpallidus, with larger ears
and shorter tail; size of aureus or a little larger; hind foot 30-33
mm.; colors buffy gray; skull wide, palate flat, not arched between
molar series; Mamme in 4 pairs, inguinal 2—2, pectoral 2-2.

Color.—Upperparts pale buffy gray with dusky ear patch and
brownish nose; underparts, feet, and tail whitish. The remnant of a
more yellowish pelage on the rump of a topotype collected May 16
would indicate a darker winter pelage, while an old female collected
at Amadee July 24 is coming into a still more ashen gray pelage.

Skull.Much heavier and wider than that of perpallidus; like that
of aureus but lower and wider with flat instead of arched palate;
interparietal larger and more quadrate; bulle slightly larger; anterior
points of frontals less acute; lateral pits of palate deeper.

1A. NORTH AMERICAN FAUNA, [NO. 39.

Measurements.—Type (¢ ad.): Total length, 242; tail vertebre,
64; hind foot, 33. Average of 3 topotypes (2 ad.): 216, 66, 30.3.
Skull (of type): Basal length, 41; nasals, 15; zygomatic breadth, 28;
mastoid breadth, 23; alveolar length of upper molar series, 9.

Remarks. closely resembles the gray phase of its
near neighbor nevadensis, but shows no relationship with it in cranial
characters, while it does agree closely with the perpallidus group. It
inhabits moist fertile soil of alkaline valleys, with many interruptions
in range and some variation in characters. Specimens from Clover-
dale, Peavine, and Monitor Valley, Nev., gous almost as well be
waleeantadl to aureus.

Specimens examined.—Total number, 43, as follows:

California: Amedee (Lassen County), 1; Fort Sage, 1.

Nevada: Carson River (Ragtown), 2; Carson Sink, 2; Cloverdale, 7; Deep Hole,
5; Fallon, 3; Flowing Spring, 1; Granite Creek, 2; Monitor Valley, 1; Pea-
vine, 4; Pyramid Lake (south end), 9; Smoke Creek. 4; Wadsworth, 1.

THOMOMYS PERPALLIDUS AUREUS Atten.

YELLOW PocKET GOPHER.
(BI, fig--4seRIWeotio- oe)
Thomomys aureus Allen, Bul. Am. Mus. Nat. Hist., V. 49, Apr. 28, 1893.

Type.—Collected at Bluff, San Juan County, Utah, by Charles P.
Rowley, May 12, 1892. Type specimen in Am. Mus. Nat. Hist.

Distribution.—Desert region of southern Nevada, southern Utah,
western Colorado, central and northwestern New Mexico, and north-
ern and western Arizona (fig. 8).

Characters.—Size rather large; hind foot 29-32 mm.; ear relatively
small; color golden buff; skull heavy, long, high, and narrow, with
palate greatly arched between the molar series; mamme in 4 pairs,
inguinal 2-2, pectoral 2-2.

Color.— Winter pelage: Upperparts beautiful orange-buff, varying
to paler and darker shades and sometimes with a wash of dusky along
the back; ear patch and nose blackish in the darker and slightly
dusky in the lghter individuals; underparts, feet, and tail creamy
white. Summer pelage: Shghtly darker. Young, more grayish.

Skull.—Long and narrow, with light lateral ridges parallel or some-
times nearly meeting in a sagittal crest in extreme old age; outline of
palate strongly arched; bulle full and rounded; interpterygoid fossa
normally U-shaped with a central point or spicule extending from
median ridge of palate; nasals cuneate and usually truncate pos-
teriorly. Dentition heavy; upper incisors abruptly decurved at right
angles to skull. |

Measurements.—Average of 5 old male topotypes: Total length,
240; tail vertebre, 73; hind foot, 31. Average of 5 old females:
294, 78, 30. Skull (of topotype, ¢ ad.):' Basal length, 42; nasals,

1No. 57162, U.S. Nat. Mus.

1915.] THOMOMYS PERPALLIDUS GROUP. 15

17; zygomatic breadth, 27; mastoid breadth, 22; interorbital breadth,
ie Bitola length of upper molar series, 9.

Remarks. iyi, subspecies aureus is an Upper and Lower Sonoran
desert form occupying the bottoms of open sandy valleys, many of
which are separated by mountainous or rough country in which the
subspecies does not occur. In almost every valley shght variations
of characters can be detected, but in none do they become so marked
as to require recognition by name. In eastern California this form
grades into the smaller perpes, which is mainly Upper Sonoran in
range, while aureus occupies the Lower Sonoran valleys. On the
south it evidently grades into albatus along the Colorado River bot-
toms, and thence into perpallidus of the Colorado Desert. Two speci-
mens from Wickenburg, Ariz., are referred with some hesitation to
aureus. Series of specimens from Manti and the Pine Valley Moun-
tains, Utah, are by no means typical but may be called aureus rather
than perpes. In New Mexico the subspecies reappears in the Rio
Grande Valley from Bernalillo to San Marcial, and while growing
darker to the southward is perfectly typical of aureus at Albuquerque.

Specumens ecamined.—Total number, 306, as follows:

Arizona: Chin Lee, 2; Jacobs Pool, 4; Keams Canyon, 6; Parker, 16; Wicken-
burg, 2

California: Amargosa River (near Nevada line), 3

Colorado: Ashbaugh Ranch, 1; Coventry, 8; Grand Junction, 1; Los Pinos, 1;
Mesa Verde, 1.

Nevada: Ash Meadows, 51; Charleston Mountains (east base), 4; Colorado River
(head of Black Canyon), 1; Oasis Valley, 3; Pahranagat Valley, 1; Pahrump
Valley, 23; St. Thomas, 5; Vegas Valley, 10.

New Mexico: Acoma, 2; Albuquerque, 4; Bear Spring Mountains, 6; Belen, 6;
Bernalillo, 4; Chusca Mountains (west slope), 1; El Vado (Chama River
Valley), 1; Fruitland, 4; Gallina (Chama River Valley), 1; Gallup, 2;
Juan Tofoya, 2; Laguna, 4; Riley, 4; San Augustine Plain (12 miles north-
west of Monica Spring), 6; San Marcial, 5; Shiprock, 2; Socorro, 13; Win-
gate, 1.

Utah: Bluff (San Juan County), 48; Hanksville, 6; Hebron, 1; Henry Moun-
tains, 3; Kanab, 4; Manti, 13; Mountain Meadows, 2; Pine Valley, 4; Pine
Valley Mountains, 7; St. George, 5; Santa Clara, 2

THOMOMYS PERPALLIDUS APACHE Batey.

JICARILLA POCKET GOPHER.
CBIR fst 65)

Thomomys apache Bailey, Proc. Biol. Soc. Washington, XXIII, 79, May 4, 1910.

Type.—Collected at Lake La Jara (7,500 feet altitude) on the
Jicarilla Apache Indian Reservation, N. Mex., by James H. Gaut,
September 19, 1904. Type specimen in U.S. Nat. Mus., Biological
Survey collection.

Distribution.—Transition Zone in northeastern Arizona, north-
western New Mexico, and southwestern Colorado (fig. 7).

Characters.—Size large, hind foot 33-34 mm.; color dark; hind
feet and tip of tail conspicuously white; mamme in 4 pairs.

76 NORTH AMERICAN FAUNA. — LNO. 39.

Color.—Upper and under parts nearly uniform dull sooty gray
shehtly washed with dull ochraceous; back with an ill-defined stripe
of blackish from tip of nose to base of tail; basal half to three-quar-
ters of tail brownish or blackish, the rest abruptly white; hind feet
white; lips usually and chin rarely white. Young: One half-grown
individual is hghter, more buffy, with white belly. st

Skull.—Sumilar in form and general characters to that of aureus;
bullz full and rounded; pterygoids U-shaped, with spicule point of
palatal ridge; nasals normally shghtly emarginate with’ doubly
rounded posterior tips, instead of normally truncate as in-aureus,
or widely emarginate as in fulvus. Dentition: Upper incisors white-
tipped and decurved at nearly right angles to axis of skull.

Measurements.—Type (¢& ad.): Total length, 229; tail vertebree,
74; hind foot, 34. Topotype (¢ ad.): 229, 74, 33. Skull (of type):
Basal length, 41; nasals, 14; zygomatic breadth, 28; mastoid
breadth, 23; alveolar length of upper molar series, 8.5.

Remarks.—This is a large, dark form of the perpallidus group with
conspicuous markings, but with cranial characters close to those of
aureus. It seems to be a dark local Transition Zone form of the
yellow aureus of the surrounding valleys.

Specomens ecamined.—Total number, 36, as follows:

Arizona: Canyon de Chelly (7 miles above mouth), 1; Fort Defiance (sawmill
12 miles northwest), 2.

Colorado: Arboles, 1; Bayfield, 1.

New Mexico: Boulder Lake, 4;1 Chama River (near Gallina), 1 (not typical);
Chusca Mountains (around lakes at top of range, 8,000—9,000 feet altitude), 13;
Lake La Jara, 2;! Stinking Lakes, 11.!

THOMOMYS CABEZONZ MeErrRIaAm.

CABEZON POCKET GOPHER.
(PL. V, fig. 8.)
Thomomys cabezone Merriam, Proc. Biol. Soc. Washington, XIV, 110, July 19, 1901.

Type.—Collected at Cabezon, San Gorgonio Pass, California, by
Clark P. Streator, June 3, 1893. Type specimen in U.S. Nat. Mus.,
Biological Survey collection.

DPistribution.—San Gorgonio Pass, southern California, south to
Cabezon (fig. 6).

Characters.—Size medium, hind foot 30 mm.; ears rather large;
color dull ochraceous; skull long and narrow.

Color.—Summer pelage: Upperparts dull ochraceous, varying from
buffy ochraceous to dull brownish; ear patch black; nose and lips
dusky or plumbeous; underparts creamy white to buffy or salmon.
In 8 of 10 topotypes collected in June, a paler, more buffy gray
pelage (probably winter) is disappearing on the rump.

1On the Jicarilla Apache Indian Reservation in northern New Mexico.

1915.] THOMOMYS PERPALLIDUS GROUP. 14

Skull—tLong, narrow, and slender, with nearly parallel lateral
ridges;. zygomata parallel or slightly wider anteriorly; nasals long
and narrow; interparietal rectangular or oval, wider than long in
immature skulls; pterygoids long, low, and thin, with rounded
symphysis.

Measurements.—Average of 7 male topotypes: Total length, 221;
tail vertebre, 79; hind foot, 30. Average of 3 female topotypes:
208, 71, 28.7. Skull (of type): Basal length, 35; nasals, 14.5; zygo-
matic breadth, 23; mastoid breadth, 18; alveolar length of upper
molar series, 7.

Remarks.— Thomomys cabezone seems to be a local form not closely
connected with surrounding species. From perpallidus, its nearest
neighbor on the east, and pallescens on the west, it is sharply differen-
tiated. In genera: appearance it most nearly resembles perpes, from
which it differs strikingly in large ears and in many cranial characters.
Two specimens from Whitewater show an overlapping of range
with perpallidus, but Gaut reports cabezone in irrigated fields, and
perpallidus in the sagebrush. A single specimen of nigrescens from
2,500 feet altitude near Banning, and 7 specimens of cabezone, show
a meeting or overlapping of ranges along the edge of the valley.
The narrow braincase suggests relationship with j7acinteus, and with
neglectus of which it may be a pale valley form.

Specimens examined.—Total number, 55, as follows:

California (southern): Banning, 12; Cabezon, 27; Schains Ranch (4,500 feet
altitude), 4; Snow Creek (near Whitewater), 2; Whitewater, 10.

THOMOMYS OPERARIUS Merriam.
OwENS LAKE PocKkET GOPHER.
(Pl. V, fig. 4.)

Thomomys operarius Merriam, Proc. Biol. Soc. Washington, XI, 215, July 15, 1897.

Type.—Collected at Keeler, California, on east side of Owens Lake,
at 3,600 feet altitude, by E. W. Nelson, November 29, 1890. Type
specimen in U. 8. Nat. Mus., Biological Survey collection.

Distribution.—Known only from type locality (fig. 6).

Characters.—Size medium, hind foot 30 mm.; color rich buff;
skull short and wide; mamme in 4 pairs.

Color.—Upperparts rich buff or pale ochraceous; ear patch gray
or plumbeous; underparts, feet, and tail creamy white. Young,
more grayish.

Skull.—Short and wide with short, heavy rostrum; zygomata
abruptly spreading; upper incisors sharply decurved or slightly
incurved; maxillaries short and heavy.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
223; tail vertebre, 69; hind foot, 30.6. Average of 5 females: 217,
66, 29.6. Skull (of type): Basal length, 38; nasals, 13.5; zygomatic

78 NORTH AMERICAN FAUNA. [No. 39.

breadth, 26.7; mastoid breadth, 21.5; interorbital breadth, 7; alveolar
length of upper molar series, 8.5. |

Remarks.—Thomomys operarvus seems to be a very local form,
quite distinct from perpes, by which its range is closely surrounded,
but more nearly resembling albatus in short rostrum and heavy
skull, and aureus in color. While clearly of the perpallidus group,
it seems not to have direct connection with any of the forms and
may well stand as described, a full species. It is confined to the
sandy alkaline soil along the east side of Owens Lake.

Specvmens examined.—Seventy-nine, from type locality.

THOMOMYS LATIROSTRIS Merriam.

PaInTED DESERT PocKET GOPHER.
(PL. V, fig. 7.)
Thomomys latirostris Merriam, Proc. Biol. Soc. Washington, XIV, 107, July 19, 1901.

Type.—Collected at Little Colorado River, Painted Desert, Arizona
(at Tanners Crossing, northeast of San Francisco Mountain), by
C. Hart Merriam and Vernon Bailey, September 22, 1899. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Painted Desert, Ariz. (fig. 8).

Characters.—Size medium, hind foot 33 mm.; skull flattened;
rostrum short and broad; ears very small; color orange-buff, much
as In aureus.

Color.—Upperparts bright buffy yellow or pale orange-buff, without
trace of black-tipped hairs; ears dusky; nose grayish brown; under-
parts, feet, and tail white, with a tinge of sulphur yellow stain on
belly of type. Young, slightly less yellow, more buffy.

Skull.—N arrow and flat, with wide, flat rostrum; nasals narrow in
the middle, spreading anteriorly; premaxille unusually wide at
base of rostrum; incisors abruptly decurved at right angles to axis
of skull; interparietal broadly pentagonal; bullze small, about as
in fulvus; basioccipital wide, flat, and keeled.

Measurements.—Type (¢ ad.): Total length, 232; tail vertebre,
79; hind foot, 33. Skull (of type): Basal length, 39; nasals, 14;.
zygomatic breadth, 26; mastoid breadth, 21; alveolar length of upper
molar series, 8.

Remarks.—While unique in characters, latirostris can best be placed
in the group with aureus (perpallidus), with which it agrees in
abruptly decurved incisors and U-shaped pterygoids. It is evidently
an isolated form of restricted range. The type specimen was caught
in the sand dunes in the Painted Desert along the Little Colorado
River at Tanners Crossing, between Black Tank and Moencopie.
It is a very old male in thin summer pelage, a worn and faded por-
tion of which is disappearing over the posterior half of the body,

1915.1 THOMOMYS PERPALLIDUS GROUP. 719

while a new, bright coat is replacing it over the anterior half. An
immature female taken at Tuba, August 16, 1909, by Clarence Birds-
eye, is the only other known specimen. The distinguishing charac-
ters of the species are maintained in this young specimen, which
disproves the idea that the type was abnormal. Twoskulls of imma-
ture females without skins, from Winslow, do not possess char-
acters sufficiently well marked for satisfactory determination, but
are provisionally referred to latirostris. Apparently this is a Painted
Desert form somewhat related to aureus.
Specumens examined.—Total number, 4, as follows:
Arizona: Painted Desert. 1; Tuba, 1; Winslow, 2.

THOMOMYS GERVINUS Axten.
PHa@NIxX PocKET GOPHER; FAWN-COLORED POCKET GOPHER.
(PL. VI, fig. 2.)
Thomomys cervinus Allen, Bul. Am. Mus. Nat. Hist., VII, 203, June 29, 1895.

Type.—Collected at Phoenix, Arizona, by J. Diefenbach, October
20, 1894. Type specimen in Am. Mus. Nat. Hist.

Distribution.—Upper Gila Valley, Ariz. (figs. 7 and 8).

Characters.—Size large, hind foot 33-36 mm.; color pale fawn;
skull heavy, ridged, and angular; mamme in 4 pairs, inguinal 2-2,
pectoral 2-2. :

Color.—Summer pelage: Upperparts pale fawn, with conspicu-
ously blackish ear patch; nose and cheeks brown or blackish; under-
parts, feet, and tail a still paler shade of fawn; lining of cheek pouches
white. Young, with whitish bellies.

Skull.—Long and narrow, as in aureus; palate only slightly arched;
bulle large and truncate instead of smoothly rounded anteriorly;
pterygoids variable, narrowly constricted or wide apart, with thick-
ened margins, and often inclosing a postpalatal spicule.

Measurements.—Average of 3 topotypes (¢ ad.): Total length,
253; tail vertebrae, 84; hind foot, 34.6. Average of 5 topotypes ( 9
ad.): 245, 77, 34. Skull (of topotype, ¢ ad.):! Basal length, 42;
nasals, 15.5; zygomatic breadth, 28; mastoid breadth, 20.5; inter-
orbital breadth, 6.5; alveolar length of upper molar series, 8.

Remarks.—This is evidently a large form of the perpallidus group,
occupying the Gila Valley. From albatus of the Colorado River
valley it differs in darker color and longer, slenderer skull and ros-
trum. It also suggests relationship with the fulvus group through
toltecus of the valleys farther east.

Specumens examined.—Total number, 19, as follows:

Arizona: Phoenix, 16; Sacaton, 1; Tempe, 2.

1No. 23707, U. S. Nat. Mus.

80 NORTH AMERICAN FAUNA. [No. 39.

THOMOMYS SINALOZ Merriam.

SINALOA POCKET GOPHER.
(Pl. IV, fig. 7.)

Thomomys sinaloe Merriam, Proc. Biol. Soc. Washington, XTV, 108, July 19, 1901.

Type.—Collected at Altata, Sinaloa, Mexico (altitude 10 feet above
sea level), by E. A. Goldman, March 28, 1899. Type specimen in
U.S. Nat. Mus., Biological Survey collection.

Distribution. Ae A Sonora and Sinaloa, from Hermosillo south
to Albata (fig. 7).

Characters.—Size rather large, hind foot 30-32 mm.; tail long;
pelage thin; color bright tawny; skull heavy, wide, and angular;
mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color.—Upperparts bright tawny or dull chestnut with a trace of
dusky along back; underparts plain hght tawny; nose grayish brown;
feet and tail half-naked, half-covered with buffy hairs. Young,
paler, more buffy. Summer and winter colors not very different.

Skull.Heavy, wide, and angular; rostrum wide; nasals short and
normally truncate posteriorly; zygomata abruptly spreading; bulle
full and rounded; basioccipital narrow between bulle. Dentition
heavy; incisors sbarotLe decurved.

Measurements.—Type (¢ ad.): Total length, 233; tail mitted
73; hind foot, 31. Average of 3 topotypes (¢ ad. ): DAS. FO, 18 eo:
Average of 3 adult females: 214, 73, 31.3. Skull (of type): Basal
length, 39.5; nasals, 14.5; zygomatic breadth, 28.3; mastoid breadth,
22; interorbital breadth, 6.2; alveolar length of upper molar series, 8.2.

Remarks.—Despite fulvous coloration the closest affinities of this
form appear to be with the perpallidus group. From its nearest
relative, cervinus, it differs in thin pelage, darker coloration, and short
wide skull. Two small females from Magdalena, Sonora, are not
typical and can only provisionally be referred to this species.

Specimens ecamined.—Total number, 22; as follows:

Sinaloa: Altata, 6.
Sonora: Camogz, 11; Hermosillo, 2; Magdalena, 2; Ortiz, 1

Thomomys fulvus Group.
THOMOMYS FULVUS FULVUS (WoopHoUSE).
Futvous Pocket GOPHER.
(Pl. VI, fig. 1.)
Geomys fuluus Woodhouse, Proc. Acad. Nat. Sci., Philadelphia, VI, 201, 1852.
Thomomys fuluus Baird, Mamm. N. Am., 402, 1857.
Type.—Collected on San Francisco Mountain, Arizona, by Dr. 5S. W.
Woodhouse, October, 1851. Type specimen in U.S. Nat. Mus.
Distribution.—Transition Zone in northern and central Arizona
from the Trumbull Mountains to the White Mountains; east in New

1915.] THOMOMYS FULVUS GROUP. 81

Mexico to Sierra Grande, and the White and Guadalupe Mountains;
north to Fisher Peak, southeastern Colorado (fig. 7).

Characters. color dark
tawny or dull chestnut; ears and claws peau mamme in 4 pairs,
inguinal 2—2, pectoral 2-9.

Color—Summer pelage: Upperparts clear dark tawny or light
chestnut, often dusky or blackish along back; ear patch, nose, and
cheeks dusky or blackish; underparts lighter; chin, lining of eheale
pouches, and feet usually white or whitish; tail tawny or dusky
above, grayish below. Winter pelage: Duller darker tawny, more
uniformly dusky above. Young very similar to adults.

Skull.—Heavier and relatively wider than in the fuscus or fossor
groups; lighter and slenderer than in perpallidus or aureus. Lateral
ridges slightly developed and approximately parallel at all ages;
nasals broadly emarginate posteriorly; bulle medium; basioccipital
broadly triangular; pterygoid fossa normally V-shaped. Dentition:
Incisors not projecting beyond tip of nasals; distinctly grooved.

Measurements.—Old male topotype: Total length, 231; tail verte-
bre, 74; hind foot, 30. Average of 5 male topotypes: 219, 70,
30. Average of 5 female topotypes: 209, 66, 29.2. Skull (of topo-
type, ¢ ad.):* Basal length, 36; nasals, 15; zygomatic breadth, 24;
mastoid breadth, 19; interorbital breadth, 6; alveolar length of upper
molar series, 8.

Remarks.—The typical subspecies of Thomomys fulvus seems to
occur only in the Transition Zone, above which it rarely extends and
below which it extends mainly in modified forms, several of which
have been separated. The name fulvus stands not only for a species
with its six subspecies, but for the center of a widely distributed
group, including mearnsi, baileyi, and lachuquilla. Its relationship
with the botte group of the west coast is genetically close but
geographically the two are widely separated. If connected with the
perpallidus group it is through toltecus and cervinus, but this con-
nection seems doubtful and by no means close.

Specumens examined.—Total number, 305, as follows:

Arizona: Aubrey (10 miles south of Pine Spring, 6,000 feet altitude), 2; Beaver
Creek (near Fort Verde), 1; Bill Williams Mountain, 2; Blue (Greenlee
County, 6,000 feet), 2; Bradshaw City, 17; Flagstaff, 2; Fort Verde, 5; Fort.
Whipple, 2; Fossil Creek, 3; Grand Canyon (Canyon Spring), 1; Little Spring
(18 miles northwest of Flagstaff), 1; Pine Spring (5 miles north, 7,000 feet),
1; Prescott, 7; Prieto Plateau (south end Blue Range, Greenlee County,
7,000-9,000 feet), 6, San Francisco Mountain, 20; Springerville (7,000 feet),
29; Trumbull Mountains, 3; White Mountains, 3.

~ Colorado: Fisher Peak, 1.

@ 1No. 24506, U. S. Nat. Mus.
98121°—15——-6 >

89 NORTH AMERICAN FAUNA. . [No. 39.

New Mexico: Burro Mountains, 2; Capitan Mountains, 36; Chloride (10 miles
east), 1; Cliff, 1; Cloudcroft, 7; Cloudcroft (10 miles north), 1; Coppermines,
1; Copperton, 6; Corona, 6; Carasal (Bernalillo County), 3; Datil Mountains,
7; Fairview, 1; Folsom, 2; Fort Wingate, 2; Gallina Mountains, 1; Gila, 5;
Glenwood, 1; Guadalupe Mountains, 6; Halls Peak, 7; Head of Mimbres, 2;
Hoskins Ranch (Colfax County), 2; Jicarilla Mountains, 5; Kingston (about
4 miles west, 9,500 feet altitude), 2; Long Canyon (3 miles north of Catskill),
1; Luna, 3; Luna (6 miles southwest, 7,000 feet), 5; Magdalena Mountains
(Copper Canyon, 8,200 feet), 1; Magdalena Mountains (Water Canyon, 6,500
feet), 1; Manzano Mountains, 12; Mimbres River, 1; Mogollon Mountains, 7;
Mora (10 miles south), 4; Mount Capitan (east base), il Pecos, 2; Pleasanton, 3;
Quemado (10 miles eutuhireahy: 2; Raton Range (teas Folder); 3; Rio Ala-
mosa (15 miles north of Ojo alent 6,900 feet), 1; Ruidosa, 7; San Andres
Mountains, 6; San Mateo Mountains (Monica conem 8,000 eae 2; San
Pedro, 2; Sierra Grande, 4; Silver City, 12; Fort Stanton, 3; Fort Union, 1;
Zuni Mountains, 5; Zuni River, 1.

THOMOMYS FULVUS PERVAGUS Merriam.
EspaNoLta PockreT GOPHER,
(Pl. IV, fig. 5.)

Thomomys aureus pervagus Merriam, Proc. Biol. Soc. Washington, XIV, 110, 1901.

Type.—Collected at Espanola, New Mexico, by J. Alden Loring,
January 4, 1894. Type specimen in U. S. Nat. Mus., Biological
Survey collection.

Distribution.—Upper Rio Grande and San Luis Valleys in northern
New Mexico and southern Colorado (fig. 8).

Characters.—Size large, hind foot 31-33 mm.; color lighter than in
fulvus; skull longer and heavier; mamme in 4 ae

Color.—Summer pelage: Dppenene plain bright oe nose,
cheeks, and ear patch blackish; underparts paler tawny; feet whit-
ish; tail dusky above, usually to tip. Winter pelage: Shghtly duller
above and below.

Skull.—As in fulvus, but longer, narrower, and with less-spreading
zygomatic arches, and slightly larger audital bulle; differmg from
aureus in less arched palate, emarginate nasals, and V-shaped
pterygoids.

Measurements.—Type (¢ ad.): Total length, 244; tail vertebrae,
76; hind foot, 31. Average of 4 topotypes (¢ ad.): 239, 73, 32.3.
Average of 5 females: 224, 68, 31.4. Skull (of type): Basal length,
42; nasals, 15.5; zygomatic breadth, 27; mastoid breadth, 21; alve-
olar length of upper molar series, 8.

Remarks.—In characters pervagus stands so nearly between fulvus
and aureus as to suggest that it forms a connecting link between the
two groups, but in a large number of specimens, collected since the
species was described, the affinities are uniformly with fulvus instead
of aureus. It is evidently a paler, more robust, Upper Sonoran val-
ley form of fulvus inhabiting the upper Rio Grande and San Luis
Valleys.

1915.] THOMOMYS FULVUS GROUP. 83

Specimens examined.—Total number, 50, as follows:

Colorado: Antonito, 8; Conejos River (6 miles west of Antonito), 2; Gardiner, 2;
Salida, 9.

New Mexico: Abiquiu, 1; Arroyo Hondo, 4; Chama River Canyon (above
Abiquiu), 1; Espanola, 8; Fort Burgwyn, 5; Questa, 4; Rinconada, 2; Santa
Clara Canyon, 2; Santa Fe, 1; Santa Fe (10 miles north), 1.

THOMOMYS FULVUS DESERTORUM Merriam.
DeESERT PocKET GOPHER.
(Pl. VI, fig. 6.)

Thomomys desertorum Merriam, Proc. Biol. Soc. Washington, XIV, 114, July 19, 1901.

Type.—Collected at Mud Spring (10 miles southwest of Mineral
Park, southern end of .Detrital Valley), Arizona, at about 3,400 feet
altitude in Lower Sonoran Zone, by Vernon Bailey, February 21,
1889. Type specimen in U.S. Nat. Mus., Merriam collection.

Distribution.—Detrital and Big Sandy Valleys, northwestern
Arizona; east in the Grand Canyon to Prospect Valley (figs. 7 and 8).

Characters.—Size small, hind foot 26 mm.; color bright tawny;
skull small, light, and smooth, with abruptly decurved incisors; sexes
very similar; mamme in 4 pairs.

Color.—Summer pelage: Upperparts rich orange-tawny, or orange-
cinnamon, lighter and brighter than in fulvus; ear patch, nose, and
cheeks dusky; underparts creamy white to light cinnamon; feet and
tail soiled whitish or buffy. Wnter pelage: Duller and more yellow-
ish; upperparts bright cinnamon; underparts whitish to buffy ochra-
ceous.

Skull.—Small and light, smoothly rounded, not ridged or angular;
bullz small, but full and rounded, not flattened and angular as in ful-
vus, nor large and prominent, as in aureus; basioccipital, as in fulvus,
wide with sharp keel. Dentition: Incisors bent downward at right
angles to axis of skull. ;

Measurements—Type (3 ad.): Total length, 200; tail vertebre,
68; hind foot, 26. Average of 5 male topotypes: 195, 63,26. Aver-
age of 4 female topotypes: 190,60,25.5 Skull (of type): Basal length,
33; nasals, 12; zygomatic breadth, 22; mastoid breadth, 17.5; alve-
olar length of upper molar series, 6.5.

Remarks.—The color of this little gopher in summer pelage suggests
relationship to fulvus, but its cranial characters are so strongly
marked that it was originally described as a species. It may, how-
ever, probably be considered a depauperate Lower Sonoran desert

1Jn the mammal collection of the Academy of Natural Sciences of Philadelphia is a mounted skin with-
out skull (No. 145) of a typical Thomomys pervagus. It is labeled ‘‘Columbia River,’’ but the label has
been copied from the catalogue where originally no locality was indicated, and the locality of the pre-
ceding specimen has been given to it. This is evidently the specimen referred to (Proc. Acad. Nat.
Sci. Phila., VI, 1852, p. 53, Pl. LXIV) as Geomys rufescens, donated by Col. Geo. A. McCall. McCall’s
specimens apparently were not labeled, but this collection of 70 mammals and bird skins was said to be
from California and Oregon. As pervagus is neither a California nor an Oregon form, the specimen in
question probably came from Santa Fe, N. Mex., where McCall was previously stationed.

84 NORTH AMERICAN FAUNA. - TNO. 39.

form of the fulvus group, with a range corresponding to that of the
creosote bush in the Detrital and Big Sandy Valleys, and extending
well up into the Grand Canyon.

Specumens ecamined.—Total number, 34, as follows:

Arizona: Big Sandy Creek (at 2,000 feet altitude), 7; Dolans Spring, 7; Grand
Canyon (at 4,500 feet in lower end of Prospect Valley on Hualpai Indian
Reservation), 4; Little Meadows (in Sacramento Valley), 2; Mineral Park,
3;1 Mud Spring,:8;1 Willow Spring, 3.!

THOMOMYS FULVUS INTERMEDIUS Mearns.

MountTAin-Torp Pocket GOPHER.
(Pl. VI, fig. 5.)

Thomomys fulvus intermedius Mearns, Proc. U. 8. Nat. Mus., XIX, 1897, 719, July 30,
1897.

Type.—Collected on summit of Huachuca Mountains (9,000 feet
altitude) southern Arizona, by F. X. Holzner, September 6, 1893.
Type specimen in U.S. Nat. Mus.

Iistribution.—Upper slopes of mountains in southeastern Arizona
and extreme southwestern New Mexico (fig. 8).

Characters—Smaller than fulvus, with conspicuous black back
and bright brown sides; skull small and slender; ears and claws
medium; mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color.—Upperparts dark tawny shading into black along the back
from nose to tail, in some specimens forming a broad and clear black
band and in others indistinct and brownish black; underparts pale
tawny; tail brown, with whitish tip; feet soiled whitish. May, August,
and September specimens show practically the same coloration. :

Skull.—Small and slender; zygomata very slender and strictly
parallel; nasals narrow, deeply emarginate; bulle small and flattened;
basioccipital triangular; pterygoids thickened at base and with
narrow fossa; interparietal small and generally triangular. Denti-
tion light.

Measurements—Type (¢ ad.): Total length, 200; tail vertebree,
66; hind foot, 26. Adult male from Fort Huachuca: 200, 59, 27.
Adult female from Fort Huachuca: 198, 62, 25. Skull (of type):
Basal length, 34; nasals, 13; zygomatic breadth, 22; mastoid
breadth, 18; interorbital breadth, 7; alveolar length of upper molar
series, 6.5.

Remarks.—This small, black-backed form of the fulvus group has
a wide and scattered range with considerable variation on the different
mountains. On top of the Chiricahua Mountains it is a little larger
than typical; on the crest of Animas Peak, a little smaller; on the

1In Detrital Valley.

1915. 1 THOMOMYS FULVUS GROUP. 85

Santa Catalina Mountains, slightly larger and darker. Variation in
size corresponds somewhat with the extent of high mountain areas
which the animals inhabit, the smallest specimens bemg from the
very restricted summit of the Animas Mountains. The difference
between those from the tops of the Animas and the Huachuca
Mountains is slight, however, compared with that between specimens
of intermedius from the summit and toltecus from the base of either
range.
Specimens examined.—Total number, 63, as follows:

Arizona: Chiricahua Mountains, 22; Fly Park (at 9,000 feet altitude), 1; Graham
Mountains (8,500 to 9,200 feet), 8; head of Rucker Canyon, 2; Huachuca
Mountains (up to 9,000 feet), 19; Mount Graham, 3; Santa Catalina Moun-
tains (8,000 feet), 6.

New Mexico: Animas Mountains (7,000 feet), 2.

THOMOMYS FULVUS TEXENSIS Battey.

Davis Mountain Pocket GOPHER.
(P1. IV, fig. 4.)
Thomomys fulvus texensis Bailey, Proc. Biol. Soc. Washington, XV, 119, 1902.

Type.—Collected on Davis Mountains, Texas (head of Limpia Creek,
at 5,500 feet altitude, in edge of Transition Zone), by Vernon Bailey,
January 7, 1890. Type specimen in U. S. Nat. Mus., Biological
Survey collection.

Distribution.—Known only from type locality (fig. 7).

Characters.—Similar to fulvus, but smaller; hind foot 24-27 mm.;
skull relatively narrower; color shghtly hghter; mamme in 4 pairs.

Color.—Summer pelage: Upperparts tawny-gray ; underparts brighter
clearer tawny; ear patch, nose, and cheeks blackish; feet and lips
gray; liming of pouches white. Winter pelage: Slightly duller and
darker than in summer.

Skull—tlLight and slender, distinguished from that of fulvus by
high narrow braincase, shallower lateral pits of palate, and shorter
pterygoids.

_Measurements—Type (éad.): Total length, 204; tail vertebre,
63; hind foot, 26. Average of 5 topotypes (¢ad.): 208, 65, 26.6.
Average of 5 females: 187, 61, 25.8. Skull (of type): Basal length,
34.5; nasals, 13; zygomatic breadth, 22; mastoid breadth, 18;
alveolar length of upper molar series, 7.

Remarks.—This is evidently a small local race of fulvus, as shown
by similarity of skull characters and also by color and general char-
acters. It is evidently isolated in the Transition Zone area of the
Davis Mountains, but not widely separated from more nearly typical
fulvus of the Guadalupe Mountains. |

Specumens examined.—Twenty, from type locality.

86 NORTH AMERICAN FAUNA, tNo. 39.

THOMOMYS FULVUS TOLTECUS Aten.
Tottec Pocket GoPHER.
(Pl. VI, fig. 3.)
Thomomys toltecus Allen, Bul. Am. Mus. Nat. Hist., V, 52, Apr. 28, 1893.

Type.—Collected at Colonia Juarez, Mexico (on the Casas Grandes
River at about 4,500 feet altitude in northwestern Chihuahua), by
F’. Robinette, 1890. Labeled ‘Juarez, N. Sonora.’’ Type specimen
in Am. Mus. Nat. Hist.

Distribution.—Lower Sonoran valleys and deserts of southeastern
Arizona, southwestern New Mexico, and adjacent parts of Chihuahua
and Sonora, south to Colonia Juarez, Chihuahua (fig. 7).

Characters.—Size about as in fulvus or slightly larger; colors
paler and grayer; ears and claws medium; mamme in 4 pairs.

Color.—Summer pelage (June specimens from Casas Grandes):
Upperparts dull ochraceous-tawny; nose brownish; ear patch black;
underparts lighter ochraceous-tawny; tail buffy gray; feet whitish.
Winter pelage: Upperparts dark gray with a tawny suffusion over
sides, and usually an indistinct dusky median line along back; nose
and ear patch dusky; underparts light fawn or pinkish buff.

Skull.—Very lke that of fulvus but more angular, less arched,
with wider nasals, larger bulle, and about the same dentition.

Measurements.—Adult male from Casas Grandes: Total length,
221; tail vertebrae, 72; hind foot, 31.5. Adult female from same
locality: 205, 66, 29. Skull (of male): Basal length, 37; nasals, 13;
zygomatic breadth, 25; mastoid breadth, 20.5; interorbital breadth,
7; alveolar length of upper molar series, 7.8. Skull (of type): 41, 14,
26, lil 18: 3 |

Remarks.—This is a light-colored valley form of the fulvus group
occupying the upper edge of Lower Sonoran deserts of southern
New Mexico and Arizona and northern Chihuahua. It is readily
distinguishable from the smaller and grayer lachuguilla of the Rio
Grande Valley, which occurs also at Casas Grandes and seems to
be distinct. In Arizona it may grade into cervinus of the Gila Valley
and in Sonora possibly into sinaloz.

Specimens examined.—Total number, 175, as follows:

Arizona: Calabasas, 5; Chiricahua Mountains, 4; Dos Cabesas, 1; Fairbank,
24; Fort Bowie, 6; Fort Grant, 4; Fort Huachuca, 5; Fort Lowell, 12; Graham
Mountains (Ash Creek, at 6,100 feet altitude), 3; Huachuca Mountains,
9; La Osa, 11; Mammoth, 1; Oracle, 2; Safford, 9; San Bernardino Ranch
(Mexican boundary), 33; San Xavier, 1; Tucson, 2; Wilcox, 2.

Chihuahua: Casas Grandes, 2; Colonia Diaz, 3; Colonia Juarez, 6; Espia, 1.

New Mexico: Adobe Ranch (north base Animas Mountains), 1; Cuchillo, 1;
Deming, 3; Garfield, 2; Hachita, 3; Lake Valley, 2; Las Palomas, 2; Mimbres
River, 1; Monument No. 40, Mexican boundary, (upper corner, 100 miles
west of El Paso), 3; Red Rock, 3.

Sonora: Santa Cruz River, 8.

1 The original label bears the name F. Robinette, but in the original description A. D: Meed was given
as the collector.

1915.4 THOMOMYS FULVUS GROUP. 87
THOMOMYS MEARNSI Barey.
MEARNS Pocket GOPHER.
(Pl. VI, fig. 4.)
Thomomys mearnsi Bailey, Proc. Biol. Soc. Washington, XX VII, 117, July 10, 1914.

Type.—Collected at Gray’s Ranch in Animas Valley, southwest
corner of New Mexico (about 20 miles north of the Mexican line),
by E. A. Goldman, August 10, 1908. Type specimen in U.S. Nat.
Mus., Biological Survey collection.

Distribution.—Known only from type locality (fig. 7).

Characters.—Slightly smaller than fulvus; less dusky in color;
skull short and wide with very slender, projecting incisors; ears
medium; mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color—Summer pelage: Upperparts dull cinnamon or light tawny,
with blackish nose and ear patches; underparts pale cinnamon; tail
buffy gray; feet whitish. Winter pelage: Evidently more grayish as
indicated by some worn patches of old long fur.

Skull—Short and wide, with arched zygomata, short braincase,
short rostrum, and protruding upper incisors; bulle very short and
round; pterygoids V-shaped; nasals short and truncate or slightly
emarginate; anterior base of zygoma convex against frontal and
premaxille. Dentition: Incisors very slender, pale, and protruding
far beyond nasals; distinctly grooved.

Measurements—Type: Total length, 220; tail vertebrae, 67; hind
foot, 31. Topotype (¢ ad.): 201, 65, 29. Skull (of type): Basal
length, 37; nasals, 12.4; zygomatic breadth, 26; mastoid breadth,
21.5; interorbital breadth, 6; alveolar length of upper molar series, 7.

Remarks.—This seems to be a local form in the bottom of the
Animas Valley, somewhat resembling nelsoni but not of the umbrinus
group. The four pairs of mamme place it in the fulvus group, but
it seems to be distinct from the surrounding toltecus. Specimens
were taken around the moist edges of a large marsh on the Gray
Ranch and the moist, alkaline soil may be the environmental factor
that has produced a local form in the midst of wide deserts. It is
quite probable that the form occupies similar situations all along
the Animas and San Simon Valleys.

Specumens examuned.—Your, from type locality.

THOMOMYS BAILEYI Merriam.
SIERRA Bianca POCKET GOPHER.
(PL. VI, fig. 7.)
Thomomys bailey: Merriam, Proc. Biol. Soc. Washington, XIV, 109, July 19, 1901.

Type.—Collected at Sierra Blanca, El Paso County, Texas, by
Vernon Bailey, December 28,1889. The type and topotypes were
caught all around the railway station where the Texas Pacific joins

the Southern Pacific. Type specimen in U.S. Nat. Mus., Biological
Survey collection.

88 NORTH AMERICAN FAUNA. [No. 39. _

Mstribution.—Sierra Blanca, western Texas, north to Tularosa,
N. Mex. (fig. 7).

Characters.—Size medium, hind foot 29-32 mm.; color, dull ochra-
ceous-tawny; skull short and wide with projecting incisors; mammsz
in 4 pairs, inguinal 2-2, pectoral 2-2.

Color.—Fresh winter pelage: Upperparts dull ochraceous-tawny
or buffy fulvous, with dusky ear patch and nose; underparts pale
salmon or creamy white; tail buffy to the tip; feet soiled whitish.
Summer pelage: Not represented.

Skull—sShort and wide, with greatly protruding incisors; basi-
occipital wide between the narrow bulle; premaxille extending but
slightly back of nasals; interparietal quadrate; anterior base of
zygoma convex; coronoid process of mandible curved, and post-
coronoid notch widely circular.

Measurements.—Topotype (3 ad.): Total length, 215; tail ver-
tebree, 64; hind foot, 31. Average of five females: 212, 69,29. Skull
(of type): Basal length, 37; nasals, 17.7; zygomatic breadth, 26.5;
mastoid breadth, 19; alveolar length of upper molar series, 7.5.

Remarks.—In wide skull and projecting incisors baileyi suggests
the umbrinus group, but as no other detailed characters of the group
appear the resemblance is evidently accidental. Nine specimens
from Tularosa, N. Mex., average larger and darker colored than those
of the type series, but show the prominent cranial characters.
There is nothing to indicate that the species has a continuous range
between these localities, but it is not improbable that it follows
along the west base of the Sacramento Mountains in the Upper |
Sonoran or the upper edge of the Lower Sonoran Zone. The Tularosa
specimens suggest a distant connection with the fulvus group.

Specomens examined.—Total number, 15, as follows:

New Mexico: Tularosa, 9.
Texas: Sierra Blanca, 6.

THOMOMYS LACHUGUILLA Batrey.

LACHUGUILLA PocKET GOPHER.
(Pl. VI, fig. 8.)
Thomomys aureus lachuguilla Bailey, Proc. Biol. Soc. Washington, XV, 120, June 2,
02

Type.—Collected at El Paso, Texas (in a dry wash in the mountains
a mile northeast of the town), by Vernon Bailey, September 24, 1901.
Type specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Arid Lower Sonoran mesas in extreme western Texas
and southern New Mexico, south to Casas Grandes, Chihuahua (fig. 7).

Characters.—Size small, hind foot 27-29 mm.; color, dull ochra-
ceous-tawny; skull narrow with abruptly decurved incisors; mamm<ze
in 4 pairs, Inguinal 2-2, pectoral 2-2.

1915.1 THOMOMYS UMBRINUS GROUP. 89

Color.—Upperparts dull ochraceous- -tawny or erayish buff; nose
and ear patch dusky; underparts pale cinnamon to soiled “Erin:
feet whitish.

Skull.—Resembling that of aureus in general form, with full roundéd
bull, narrow basioccipital, and slightly arched pains but differ-
ing in its smaller size, slenderer form, and less-ridged surface; inter-
parietal normally quadrate; slender points of premaxille extending
far back of truncate or slightly emarginate tip of nasals.

Measurements.—Type (3 ad.): Total length, 215; tail vertebre,

65; hind foot, 29. Average of five topotypes (¢ ad.): 202, 61, 27.
Average of three females: 195, 62,26. Skull (of type): Basal length,
35; nasals, 14; zygomatic breadth, 22; mastoid breadth, 19; alveolar
length of upper molar series, 7.
- Remarks.—This form needs no comparison with barleyz on the
east or perditus on the south. It was originally, but probably erro-
neously, considered a small, dull-colored desert form of aureus. It
occupies the arid Lower Sonoran mesas, but apparently does not occur
in moist and fertile bottom lands as do the more robust aureus and
toltecus. Specimens from Presidio County and the Great Bend region
are not typical, but they vary so much with each locality that
to avoid making more subspecies I have referred them to this, the
nearest form. It seems probable that the great variation in the
gophers of this region is due to the scarcity of individuals and the
isolation of colonies.

Specimens examined.—Total number, 33, as follows:

Chihuahua: Casas Grandes, 2

New Mexico: Organ, 2; Oran Mountains, 2.

Texas: Alpine, 1; inert Tine, 2; El Paso, 12; Franklin Mountain (10 miles north
of El Paso, 4; al 15 miles sineith 2), 6; Marathon (15 miles south), 1; Paisano,
ie Buosalte (panty (35 miles south of Marfa), 4

Thomomys umbrinus Group.

THOMOMYS UMBRINUS UMBRINUS (RicHarpson).
SOUTHERN POCKET Goruer.
(Pl. I, fig. 13; Pl. VI, fig. 13; text fig. 2.)
Geomys umbrinus Richardson, Fauna Boreali-Americana, I, 202, 1829.
Thomomys umbrinus Bailey, Proc. Biol. Soc. Washington, XTX, 3, Jan. 29, 1906.

Type.—In British Museum, said to have come from “ Cadadaguios,
a town in southwestern Louisiana,” but no such town has ever been
located, except the Kadhoadacho Indian settlements on the Red
River, in western Louisiana, where Thomomys does not occur. The
type locality is now restricted to Boca del Monte, Vera Cruz. Type
identified with specimens from Vera Cruz.

1 Proc. Biol. Soc. Washington, XIX, 3, Jan. 29, 1906.

90 NORTH AMERICAN FAUNA. fo. 39.”

Distribution.—Known only from Boca del Monte and Xuchil,
Vera Cruz (fig. 9).

Characters.—Size medium, hind foot 27-28 mm.; color dichromatic,
dull umber, brown, or black; skull short and wide with emarginate
anterior base of zygoma and strongly projecting incisors; groove on
inner margin of incisors very obscure; ears small; claws very slender;
mamme in 3 pairs, inguinal 2—2, pectoral 1-1.

Color.—Upperparts in brown phase varying from dull burnt umber,
or Prout’s brown, to nearly black; underparts lightly washed with
pale buff or whitish; chin, feet, and tip of tail white. In black phase,
slaty black all over except white chin, feet, and tip of tail.

Skull—Light and smooth, not ridged or angular, short and wide,
with very short rostrum and greatly projecting incisors; anterior
base of zygoma emarginate or concave in outline where it joins the
frontal; lachrymal long and attached for all or most of its length to
zygoma; premaxille widening posteriorly and terminating approxi-
mately even with nasals; palate flat, not arched between molar
series; lateral pits deep; base of pterygoids constricted. Dentition
light; incisors slender, projecting, pale or whitish, with indistinct or
obsolete inner groove; anterior lower incisor with anterior enamel
plate minute or often entirely wanting.

Mcasurements.—Average of 4 topotypes (¢ ad.): Total length,
193; tail vertebrae, 58; hind foot, 27.2. Average of 3 adult females:
195, 57, 27. Skull (4 ad.):! Basal length, 33.7; nasals, 12.4; zygo-
matic breadth, 24; mastoid breadth, 17.6; alveolar length of upper
molar series, 7.3.

Remarks.—It would be useless to list the various species of Tho-
momys to which authors have applied the name wmbrinus. For over
half a century the name was shifted about to one and another species,
and proved a stumbling block to every naturalist working on the
genus. Fortunately the type is still extant and has been identified
with a sufficiently restricted species to give it a definite location.
The species thus becomes the type of a well-defined group, including
umbrinus, orizabe, peregrinus, nelsoni, sheldoni, goldmam, perdatus,
and atrovarius. 7

Specomens examined.—Total number, 15, as follows:

Vera Cruz: Boca del Monte, 8; Xuchil, 7.
THOMOMYS UMBRINUS ORIZABA Merriam.
OrizABA POCKET GOPHER.
(Pl. VI, fig. 15.)
Thomomys orizabxe Merriam, Proc. Biol. Soc. Washington, VIII, 145, Dec. 29, 1893.

Type.—Collected on Mount Orizaba (9,500 feet altitude), Puebla,
Mexico, by E. W. Nelson and E. A. Goldman, April 25, 1903. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

1No. 66091, U. S. Nat. Mus., from Boca del Monte, Vera Cruz.

1915.] THOMOMYS UMBRINUS GROUP. O]

Distribution.—Known only from type locality (fig. 9).

Characters.—Considerably larger than wmbrinus or peregrinus;
hind foot 27-31 mm.; skull heavier; colors dichromatic, usually
black, rarely bright umber; ears small; claws slender; mammez in
3 pairs.

Color.—In 14 of 16 topotypes: Glossy iridescent black all over
except white on tip of tail, toes, and usually chin and inside of
pockets. Two topotypes in brown phase: Upperparts bright burnt
umber in the adult, darker in the immature specimen; underparts
washed with paler shade of the same or ochraceous-buff; tip of
tail, toes, and inside of pockets white.

Skull.—Relatively more elongated, angular, and ridged than in
umbrinus or peregrinus; base of zygoma deeply concave, or emargi-
nate; lachrymal long and borne mainly on zygoma; premaxille
widening posteriorly and terminating approximately even with
nasals; palate flat between molars, but with deep lateral pits and
constricted base of pterygoids. Dentition heavy; incisors less pro-
jecting than in peregrinus, and plain orange, rarely with white tips;
anterior enamel plate of front lower incisor small.

Measurements.—Average of 4 topotypes (¢ ad.): Total length,
219; tail vertebre, 69; hind foot, 29.3. Average of 5 females: 132, 66,
29. Skull (of type, @ ad.): Basal length, 35; nasals, 13; zygomatic
breadth, 25; mastoid breadth, 19; alveolar length of upper molar
series, 7.7. One topotype (¢ ad.):1 38.5, 14, 25, 19.5, 8.

Specimens examined.—Sixteen, from type locality.

THOMOMYS UMBRINUS PEREGRINUS MErgziam.

Mexican Pocket GOPHER.
(P1. VI, fig. 14.)

Thomomys peregrinus Merriam, Proc. Biol. Soc. Washington, VIII, 146, Dec. 29, 1893.
Type.—Collected at Salazar, Federal District of Mexico, Mexico, at
10,300 feet altitude, by E. W. Nelson and E. A. Goldman, October 24,
1892. Type specimen in U.S. Nat. Mus., Biological Survey collection.
Distribution.—Mountain slopes in the Federal District of Mexico,
east to San Martin, Puebla, and north to El Chico, Hildago (fig. 9).

Characters.—Size medium, slightly larger than umbrinus; hind foot
27-29 mm.; color dark rich umber; skull with projecting incisors
and concave base of zygoma; mamme in 3 pairs.

Color.—Upperparts dark rich burnt umber, becoming blackish
along the back in many specimens; underparts washed with ochra-
ceous-buff; feet, tip of tail, and sometimes chin, whitish. A fully
melanistic specimen from Santa Rosa, Guanajuato, suggests partial
dichromatism.

1No. 53611, U. S. Nat. Mus.

99 NORTH AMERICAN FAUNA. [No. 39.

Skull.—Less markedly short and wide than in umbrinus and with
slightly less projecting incisors; base of zygomaslightly concave; palate
flat between molar series, and unique in having shallow lateral pits
and wide base of pterygoids. Dentition light; incisors bright orange
with white tips; anterior enamel plate on front lower incisors small but
distinct and constant. ,

Measurements.—Average of 3 topotypes (¢ ad.): Total length,
208; tail vertebre, 61; hind foot, 28. Average of 5 females: 189,
59, 27.3. Skull (of type, ? ad.): Basal length, 34; nasals, 12; zygo-
matic breadth, 23; mastoid breadth, 18; alveolar length of upper molar
series, 7. Skull of topotype (4 ad.):1! 34, 12.3, 24, 19, 7.

Specimens excamined.—Total number, 43, as follows:

Hidalgo: El Chico (Sierra de Pachuca), 3; Real del Monte, 3; Tulancingo, 1.

Mexico: Popocatepetl, 1, Salazar, 17; Toluca Valley, 1; Volcan Toluca (north
slope), 15.

Puebla: San Martin, 2.

THOMOMYS NELSONT MERRIAM.

NELSON POCKET GOPHER.
(P1. VI, fig. 11.)
Thomomys nelsoni Merriam, Proc. Biol. Soc. Washington, XIV, 109, July 19, 1901.

Type.—Collected at Parral, Chihuahua, Mexico, at 6,000 feet alti-
tude, by E. W. Nelson and E. A. Goldman, September 18,1898. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Central Chihuahua, from Gallego south to Parral
(fig. 9).

Characters.—Size small, hind foot 28 mm.; color bright tawny;
skull short and wide, with projecting incisors.

Color.— Winter pelage: Upperparts light russet or bright tawny,
darker along back and on face; nose and ear patch blackish; ankle
and base of tail brownish, lower part of feet and terminal part of tail
white; belly pale tawny. Summer pelage: A shade brighter above
and below.

Skull.—Short and wide with abruptly spreading zygomata and
strongly projecting, lightly grooved, white-tipped incisors; nasals
narrow, cuneate, slightly notched, and terminating a little short of
the wide posterior tips of premaxille; basioccipital wide and flat;
frontals wide, crowding slightly into base of zygomata; lachrymal
-mainly adnate to zygoma.

Measurements.—Type (¢ ad.):* Total length, 196; tail vertebre,
60; hind foot, 28. Topotype (¢ ad.): 207, 59, 28.5. Skull (of type):
Basal length, 37; nasals, 13; zygomatic breadth, 24; mastoid breadth,

1No. 50122, U. S. Nat. Mus.
2The type of nelsoni was marked “ ?’’ on the original label and referred to in original description as
female. It is, however, a fine old male with prominent sex marks showing in the skin and with the skull

characters of the male.

1915.] THOMOMYS UMBRINUS GROUP. 93

21; alveolar length of upper molar series, 7. Skull of male topotype:
34, 12, 24, 19.5, 7.

Remarks.—The short, wide skull, projecting incisors, and white slip-
pers and tip of tail reveal relationship with the wmbrinus group,
although in the specimens collected there are no females to show the
arrangement of mammez. However, nelsoni is evidently a northern
form of the group.

Specimens exramined.—Total number, 3, as follows:

Chihuahua: Gallego, 1; Parral, 2.

THOMOMYS SHELDONI.! sp. Nov.
SHELDON POCKET GoPHER.

(Pl. VI, fig. 12.)

Type from Santa Teresa (6,800 feet altitude), Tepic, Mexico.
Adult ¢, No. 90819, U. S. Nat. Mus., Biological Survey collection.
Collected August 10, 1897, by E. W. Nelson and E. A. Goldman; col-
lectors’ number, 11443.

Distribution. Er neition Zone on the Sierra Madre, Mexico, from
northern Chihuahua south to Guanajuato (fig. 9).

Characters.—In size slightly larger than umbrinus or peregrinus,
and more brightly colored than either; ears small; claws slender;
skull relatively long, with long nasals and not very protruding incisors;
mamme in 3 pairs. : |

Color.—Summer pelage: Upperparts light umber, or russet-brown,
darker along median line of back, on nose, and about ears; under-
parts lighter, more tawny; lower half of feet and tip of tail usually
white; rarely any white on chin or throat in typical series; white
Se of cheek pouches scarcely showing. Young duller and. fade

Skull—Relatively longer than in other members of the group,
with longer, narrower nasails and slender rostrum; interorbital region
wide and flat; interparietal triangular; nasals slightly emarginate,
usually fallme short of posterior tips of premaxille; anterior base of
zygoma nearly straight, in some slightly concave and in others
slightly convex; lachrymal adnate to zygoma for over half its length.
Dentition: Incisors moderately projecting, dark orange, with small
but distinct grooves along inner margins.

Measurements—Type (& ad.): Total length, 210; tail vertebra,
64; hind foot, 29. Average of 4 topotypes (¢ ad.): 200, 62, 28.8.
Average of 4 adult females: 191, 61, 28.5. Skull (of type): Basal
length, 37; nasals, 14.5; zygomatic breadth, 25; mastoid breadth,
19; interorbital breadth, 7; alveolar length of upper molar series, 8._

Remarks.—This is a northern form of the wmbrinus group, occupy-_
ing the Transition Zone of the main Sierra Madre of Mexico, and

1Named for Charles Sheldon in recognition of his interest in mammals, large and small, and his many
contributions of specimens and of notes on their habits.

94 NORTH AMERICAN FAUNA. [NO. 39.

probably separated by lower country from the range of peregrinus,
its nearest relative. There is considerable variation in its extension
northward to northern Chihuahua but no indication of its intergra-
dation with intermedius, which is very similar in general appearance
but which with its 4 pairs of mamme and narrow, slender skull,
belongs to the fulvus group.

Specuomens examined.—Total number, 117, as follows:

Chihuahua: Chuichupa, 1; Colonia Garcia (6,400 feet altitude), 13; near Guad-
elupe y Calvo (at 7,000 feet), 6; Pacheco, 11; Sierra Madre (65 miles east of
Batopilas at 7,000 feet), 5.

Durango: Chacala (3,000 feet), 1, immature; Coyotes, 3; El Salto (8,400 feet),
10; La Boca, 1; Rio Sestin, 1.

Guanajuato: Santa Rosa (8,500 feet), 9.

San Luis Potosi: La Tinaja (6,000 feet), 8; San Luis Potosi, 1.

Sinaloa: Sierra de Choix (50 miles northeast of Choix at 5,000 feet), 3.

Sonora: Alamos (1,200 feet), 5.

Tepic: Santa Teresa (6,800 feet), 10.

Zacatecas: Berriozabal (6,600 feet), 10; Plateado, 4; Sierra Madre (8,500 feet),
5; Valparaiso Mountains (8.700 feet), 10.

THOMOMYS GOLDMANI Merriam.
GOLDMAN POCKET GOPHER.
(Pl. VI, fig. 10.)

Thomomys goldmani Merriam, Proc. Biol. Soc. Washington, XIV, 108, July 19, 1901.

Type.—Collected at Mapimi, Durango, Mexico, by E. A. Goldman,
December 15, 1893. Type specimen in U. S. Nat. Mus., Biological
Survey aolllectara.

Distribution.—EKastern Durango, from Mapimi south to aranaa
City (fig. 9).

Characters.—Size small, hind foot 27-30 mm.; color bright tawny
above, white below; ears minute; skull short and wide, with project-
ing incisors; Mamme in 3 pairs, Sheil Di. Perot ial

Color.—Upperparts bright tawny, darkening on head and becoming
dusky around nose and ears; underparts, tip of tail, and lower part
of feet, white.

Skull—sShort and wide with very short nasals and projectmg in-
cisors; zygomatic arches slender, nearly parallel, abruptly con-
stricted at posterior base; basioccipital wide and flat; nasals nearly
truncate, falling slightly short of posterior tips of premaxille; front-
als wide, pushing slightly into base of zygomata; lachrymal lyimg
mainly against base of zygoma, as In wmbrinus.

Measurements.—Type (¢ ad.): Total length, 208; tail vertebre,
68; hind foot, 30. Female topotype: 190, 60, 27. Skull (of type):
Basal length, 33; nasals, 12; zygomatic breadth, 22; mastoid breadth,
18; alveolar length of upper molar series, 7.

Remarks.—The specimens from Mapimi exhibit the extreme differ-
entiation of the form. Those from Durango are darker, more nearly

1915.] THOMOMYS UMBRINUS GROUP. 95

like peregrinus, through which goldmani evidently connects with the
umbrinus group.

Specimens examined.—Total number, 9, as follows:

Durango: Durango, 6; Mapimi, 2; San Gabriel, 1.
THOMOMYS PERDITUS Merriam.
Lirrte Gray PocKET GOPHER.
(Pl. VI, fig. 9.)
Thomonys perditus Merriam, Proc. Biol. Soc. Washington, XIV, 108, July 19, 1901.

Type.—Collected at Lampazos, Nuevo Leon, Mexico, by Clark P.
Streator, January 22, 1891. Type specimen in U. S. Nat. Mus.,
Biological Survey collection.

Mstribution.— Eastern Coahuila and western Nuevo Leon, north
to Rock Springs and Castle Mountains, western Texas (fig. 9).

Characters.—Size small, hind foot 25 mm.; color buffy gray or pale
tawny; skull short and narrow, with shane: projecting incisors;
mamme in 3 pairs, inguinal 2~—2, pectoral 1-1.

Color.—Upperparts dark buffy gray or pale duil tawny, darkest
along back; sides brighter; belly and feet pale buffy or creamy white;
ear patch, nose, and lips dusky; inside of pouches white.

Skull.—sShort, with smooth, rounded braincase and rectangular
zygomatic arches; incisors projecting slightly beyond nasals; nasals
sharply notched posteriorly and terminating on a plane with short
premaxillze; interparietal normally quadrate, wider than long; ante-
rior base of zygoma strongly convex in outline, unlike most species of
the wmbrinus group.

Measurements —Type (é ad.): Total length, 195; tail vertebre,
59; hind foot, 26.5. Average of 2 topotypes (¢ ad.): 192, 58, 26.8.
Average of 2 females: 180, 54, 24.5. Skull (of type): Basal length,
33; nasals, 12; zygomatic breadth, 21; mastoid breadth, 18; alveo-
lar length of upper molar series, 7.

Remarks.—The relationships of this species, as exhibited by the
cranial characters, seem not to be very close with any of the surround-
ing forms, but the number and arrangement of mamme place it in
the umbrinus group.

Specimens examined.—Total number, 24, as follows:

Coahuila: Carneros, 5; Jaral, 1; Sierra Encarnacion, 1; Sierra Guadalupe, 4.
Nuevo Leon: Lampazos, 5; Villadama, 2.

Texas: Castle Mountains, 1; Comstock, 2; Devils River, 1; Rock Springs (35
miles east), 1; Samuels, 1

THOMOMYS ATROVARIUS ALLEN.
BLACK-AND-BROWN POCKET GOPHER.
(Pl. VI, fig. 16.)
Thomomys atrovarius Allen, Bul. Am. Mus. Nat. Hist., X, 148, 1898.

Type.—Collected at Tatemales (near Rosario), Sinaloa, Mexico, by
P. O. Simons, May 15, 1897. Type specimen in the British Museum.

96 NORTH AMERICAN FAUNA. [NO. 39.

Disiribution.—Coastal Plain of southern Sinaloa and Tepic, from
Mazatlan south to Colomo (fig. 9).

Characters.—Size medium, hind foot 30 mm.; pelage thin and
harsh; color dark brown or blackish; skull short and wide, with pro-
jecting incisors; ears small; claws slender; mamme in 3 pairs.

Color.—Dark umber, brown, or brownish black; underparts brown-
ish or plumbeous; ears, feet, and tail nearly naked, but lower part of
feet and tip of tail usually white when there is hair enough to show
color.

Skull.Sheghtly heavier and longer than in umbrinus, but of the
same general characters, with short nasals and less-projecting incisors;
base of zygoma slightly indented and the greater part of lachrymal
adnate to it; posterior tips of premaxille narrowed and extending
slightly back of truncate tip of nasals; pterygoids wide apart.
Dentition slightly heavier than in wmbrinus; incisors dark orange and
very lightly grooved.

Measurements.—Average of 3 females from Rosario: Total length,
196; tail vertebrae, 68; hind foot, 29. Adult male from Mazatlan:
208, 66, 31. Skull (of adult male from Mazatlan): Basal length, 36;
nasals, 13; zygomatic breadth, 25; mastoid breadth, 20; alveolar
length of upper molar series, 8.

Remarks.—Thomomys atrovarwus is a well-marked, thin-haired,
nearly concolor, tropical species in which the skull characters show
close affinities with the wmbrinus group.

Specomens examined.—Total number, 14, as fellows:

Sinaloa: Mazatlan, 2; Plomosas, 2; Rosario, 5; Tatemales, 1.
Tepic: Colomo, 1; Navarete, 2; Pedro Pablo, 1.

Thomomys talpoides Group.
THOMOMYS TALPOIDES TALPOIDES (RicHarpson).
SASKATCHEWAN POCKET GOPHER.

Cricetus talpoides Richardson, Zool. Journ., III, 518, 1828.

Geomys talpoides Richardson, Fauna Boreali-Americana, 204, 1829.

S{accophorus| talpoides Fischer, Synopsis Mammalium, 588 (‘‘388”), 1830. Taken
from Richardson.

Geomys borealis Richardson,! Sixth Ann. Bees Brit. Assn. for PS3b52V—. 150, lob,
1837. Renaming of talpordes.

1 Geomys borealis of Enenerdeen has been generally considered a nomen nudum and the name dated
from Bachman, two years later, based on a different species. Richardson first used the name (loc. cit.,
150) as ““Geomys borealis, Rich. nov. sp.”? On page 156 he states that “‘Geomys borealis inhabits the Plains
of the Saskatchewan,’’ and on page 157 he gives good descriptive characters in comparison with several
other species. He says ‘‘borealis and talpoides have a very fine groove close to the inner margin of each
upper incisor,’’? but evidently failed to detect the much finer grooves on the incisors of bulbivorus and
umbrinus with which he was making the comparison. As Dr. Allen has pointed out (Bul. Am. Mus.
Nat. Hist., V, 61, 1893) Richardson was evidently renaming his talpoides, of which he here (p. 156) very
strangely changes the habitat to Florida. It therefore seems necessary to consider borealis a synonym
of talpoides. Bachman’s borealis (1839), of which the type is extant and perfectly CISTI, is tine same
as clusius of Coues (1875).

Since writing the foregoing I have received a letter (Mar. 15, 1915) from Mr. Oldfield Thomas, Gs the
British Museum, expressing his opinion that borealis of Bachardsdn is not a nomen nudum.

1915.] THOMOMYS TALPOIDES GROUP. 97

Ascomys borealis Wagner, Suppl. Schreber Saug., III, 391, 1843. Taken from
Richardson.
Geomys unisulcatus Gray, List Mamm. Brit. Mus., 149, 1843. Nomen nudum.
Saccophorus borealis Gray, List Mamm. Brit. Mus., 149, 1843. From Richardson.
Ascomys talpoides Wagner, Suppl. Schreber Saug., III, 390, 1843. From Richardson.
Ascomys borealis Wagner, Suppl. Schreber Saug., III, 391, 1843. From Richardson.
Pseudostoma talpoides Audubon & Bachman, Quad. N. Am., III, 43-45, 1854; de-
scribed and figured from type specimen in collection Zool. Soc., London. |
Geomys (Thomomys) talpoides Giebel, Saiug., 530, 1855. Compiled from Richardson.
Thomomys talpoides Baird, Mamm. N. Am., 403, 1857.

Type.'—Type locality fixed at near Fort Carlton (Carlton House) on
the Saskatchewan River, Saskatchewan. ‘Type specimen in British
Museum, a stuffed skin and anterior part of skull, which was removed
from the skin in 1893.

Distribution.—Plains of Saskatchewan and Alberta; south in
Montana to Great Falls and the Big Snowy Mountains (fig. 10).

Characters.—Size medium, between that of rufescens and clusius;
ears prominent and pointed; claws slender; color dull and dark
gray; skull not heavily ridged and with shght temporal ridges often
converging anteriorly or in the middle; nasals emarginate posteriorly;
rostrum slender; mammze normally in 6 pairs, inguinal 2-2,
abdominal 2—2, pectoral 2-2.

OColor.—Summer pelage: Upperparts dull grayish brown with
plumbeous nose and blackish ear patch; underparts buffy or soiled
whitish gray; chin, and often throat and breast, pure white; feet
whitish; tail white tipped or sometimes all white. Winter pelage:
Upperparts dull dark gray, with little brownish; ear patch black;
underparts washed with whitish.

Skull—Rather short and wide, with anteriorly converging tem-
poral ridges in all but very old individuals; nasals emarginate at
posterior tips or rarely truncate; interparietal triangular; audital
bull full and wide but low, with anterior arm of basioccipital narrow
between them. VDentition noticeably lighter than in rufescens; upper
incisors abruptly decurved and distinctly grooved.

Measurements.—Largest male from Borden, Saskatchewan: Total
length, 214; tail vertebra, 60; hind foot, 28. Largest female from
same place: 210,60, 29. Skull (¢ ad. from Borden, Saskatchewan):
Basal length, 34.5; nasals, 14; zygomatic breadth, 23; mastoid
breadth, 19; interorbital breadth, 6; alveolar length of upper molar
series, 7. |

1 The type of talpoides, a stuffed skin with part of Sicall, Hoag pincaredit > mt PeaaneaieS panonaon
dealer, and was recorded from ‘Hudson Bay.’”’ This evidently meant the country of the Hudson’s Bay
Co., or western Canada.. There are no gophers known nearer the shores of Hudson Bay than on the
Saskatchewan, where, as Dr. Allen has pointed out, Richardson refers to them in his original description.
Carlton House or Fort Carlton, where Richardson remained for some time, and apparently the highest
point he reached on the Saskatchewan River, is near the easternmost point at which these gophers are

known to occur along the river. As other species are found farther west on the Saskatchewan, it is neces-
Sary to restrict the type locality of talpoides still further to the vicinity of Carlton House.

98121°—15 7

98 NORTH AMERICAN FAUNA. [ NO. 39.

Remarks.—The name talpoides has long been applied to the gophers
of North Dakota and Manitoba on the supposition that they were
the same as those from the Saskatchewan. A series of specimens
recently collected at Borden, Saskatchewan, the lowest point at
which they could be found along the North Saskatchewan River,
and only about 40 miles above the site of old Fort Carlton, prove to
be separable from the Dakota animal, which therefore takes the
name rufescens. The fact that Richardson did not get specimens or
even see the gopher which he described is probably due to their
absence from the immediate vicinity of Fort Carlton where he records
them in an indefinite way. He may have seen their molelike mounds
on a short trip up the river or may only have heard of them from
trappers and other explorers in that region, possibly from Drum-
mond or Douglas. It, therefore, seems safe to assume that speci-
mens from Borden are typical of talpoides of Richardson.

Loring reports gopher hills 20 miles, and Preble 40 miles, north of
Edmonton.

Specurmens examined.—Total number, 98, as follows:

Alberta: Blindman River, 4; Buffalo Lake, 2; Calgary, 2; Didsbury, 2: Ed-
monton, 22; Irwin Lake, 1; Red Deer, 23; St. Albert, 2; Waghorn, 1.
Montana: Bearpaw Mountains, 1; Big Snowy Mountains, 2; Blackfoot, 12;

Highwood, 3; Zortman, 1.
Saskatchewan: Borden, 16; Indian Head, 4.

THOMOMYS TALPOIDES RUFESCENS Wien.
PRarIRIE PockEeET GOPHER; DAKOTA POCKET GOPHER.
(PING PI, figeos Ril Ville ners texbeissee2io)

Thomomys rufescens Wied, Nova. Acta. Phys. Med. Acad. Ces. Leop.-Carol, XIX,
pt. 1, 378, 1839.

Type.—Collected at ‘‘The Minnetaree Village,’ now Old Fort
Clark, Oliver County, North Dakota (about 6 miles south of Stanton),
on the west side of the Missouri River, by Maximilian, Prince of
Wiedin. 1833. Type specimen in Am. Mus. Nat. Hist.

Distribution.—Greater part of North Dakota, eastern South Dakota,
and southwestern Manitoba (fig. 10).

Oharacters.—Size large; feet large and claws stout; ears prominent
and pointed, not a mere thickened rim as in some western species;
color dull and dark gray; skull heavy, with parallel temporal ridges
and wide basioccipital; rostrum slender; incisors abruptly decurved,
distinctly grooved; mamme in 6 pairs, inguinal 2-2, abdominal 2-2,
pectoral 2-2.

Color.—Thin summer pelage ane and August): Upperparts dull
brownish gray; ear and surrounding fur black; underparts buffy gray
with more or less pure white on chin, throat, aan breast; feet whitish;
tail whitish or gray at base. Early winter pelage (October): Dark
buffy gray above, almost as dark as in summer. full winter pelage

1915. ] THOMOMYS TALPOIDES GROUP. 99

(November to May): Bright buffy gray above, fading to paler buffy
in spring. Young: Upperparts duller, more plumbeous, fading to
buffy when half grown, changing to dark fall pelage in October.
[See section on molt (p. 18); also five molts shown on one animal in
mee, No. 1.]

Skull—Long and straight, with shallow rostrum and strongly
developed and approximately parallel temporal ridges; nasals long
and truncate or rounded at posterior tip; bullz long and rather narrow
(becoming very narrow in specimens from the eastern part of the
range), basioccipital with wide, heavy shaft between bulle; auditory
meatus slender. Dentition: Molars heavy; incisors decurved at right

angle to axis of skull.

— _Measurements.—Adult male from near Fort Clark, N. Dak.: Total
length, 240; tail vertebre, 70; hind foot, 31. Adult female from same
locality: 230, 70, 31. Skull (of adult male from near Fort Clark):
Basal length, 40; nasals, 15.5; zygomatic breadth, 25; mastoid
breadth, 22; interorbital breadth, 7; alveolar length of upper molar
series, 8. :

Remarks.—There is now ample material from the type locality and
its general region to show the characters of rufescens, which proves to
be the large, dark-colored gopher occupying the greater part of
North Dakota, eastern South Dakota, and southwestern Manitoba.
Specimens from. the type region in early winter fur (November or
December) are still needed to complete our knowledge of seasonal
variation in pelage, but this pelage will probably be found to agree,
as do most other characters, with specimens from the eastern part of
the State.

The type, a mounted specimen with skull inside, much faded but
otherwise in good condition, is in the American Museum of Natural
History. It has been removed from the base, and the old label, once
tacked on the base, is now attached to the foot. It bears the legend
“Thomomys rufescens Wied. Machtohpka indigen”’ and also ‘‘Mas
Missouri,” not very legible. It hardly seemed necessary to remove
the skull from so old and fragile a specimen, as there are good topo-
types, and the general size and large claws are good characters. The
hind foot measures 30 mm., dry. It is marked “3” on the museum
label, which also is attached to the foot. :

Specumens examined.—Total number, 159, as follows:

Manitoba: Aweme, 4; Carberry, 11; Selkirk Settlement, 1.

North Dakota: Bismarck, 4; Bottineau, 1; Braddock, 1; Cannonball River, 6;
Devils Lake, 1; Dickinson, 2; Fish Lake (in Turtle Mountains), 8; Fort Clark,
5; Fort Rice, 1; Glen Ullin, 1; Grafton, 3; Hensler, 6; Jamestown, 6; Lari-
more, 7; Mandan, 10; Minnewauken, 3; Minot, 1; Oakdale, 8; Pembina, 25;
Portland, 1; Sherbrooke, 6; Souris River, 2; Steel 1; Streeter, 2; Stump
Lake, 1; Valley City, 9; Wade, 2; Walhalla, 2; Washburn, 3.

South Dakota: Aberdeen, 2; Armour, 3; Fort Randall, 2; Fort Sisseton; 3;
Highmore, 1; Pierre, 3; White Lake, 1.

100 — NORTH AMERICAN FAUNA, [No. 39.

THOMOMYS TALPOIDES CLUSIUS Covzs.

Cours PockEtT GOPHER.
(PI. II, fig. 6; Pl. VII, fig. 5; text fig. 3.)

Geomys borealis Bachman,! Journ. Acad. Nat. Sci. Phila. VIII, pt. 1, 103, 1839.
Type locality said to be the ‘‘Columbia River,’’ probably in Colorado near the
southeast corner of Wyoming. (Not of Richardson.)

Pseudostoma borealis Audubon & Bachman, Quad. N. Am., IIT, 198, pl. 142, 1853.
(Not Geomys borealis of Richardson.)

Thomomys clusius Coues, Proc. Acad. Nat. Sci. Philadelphia, 1875, 138, June 15, 1875.

Type.—Collected at Bridgers Pass, Carbon County, Wyoming, (18
miles southwest of Rawlins), by Dr. W. A. Hammond, July 28, 1857.
Type specimen in U. S. Nat. Mus.

Distribution.—Central and southeastern Wyoming (north to Park-
man, Sheridan County), and eastern Colorado south to Colorado
Springs (fig. 10).

Characters.—Considerably smaller than talpoides, with slenderer
claws; color slightly more rufescent; skull with the same shallow
rostrum,-but differing in having smaller size, large rounded bulle, and
lighter dentition; mamme in 6 or 7 pairs, inguinal 2-2, abdominal
2-2, pectoral 2-2, or sometimes 3-3; embryos 5 to 7.

Color.—Summer pelage: Upperparts light buffy- or hazel-gray,
brightest on crown, paler along sides; cheeks clear gray; ear patch
blackish; underparts whitish or buffy; chin sometimes white; feet
and tip of tail whitish. Autumn pelage (September): Upperparts
duller and darker; crown and back dull hazel darkened with black-
tipped hairs that apparently fade or wear off and leave a grayish or »
buffy brown in winter; black ear patch conspicuous. Young paler,
more buffy gray.

Skull—Light and slender, not angular or heavily ridged; temporal
ridges parallel in fully adult specimens; audital bullze much larger

1 Bachman’s description of Geomys borealis, which he credits to Richardson, was based mainly on a
specimen collected by Townsend, of which he says, “Mr. Townsend’s specimens were prccured on the

Columbia River.’’? The very full description of borealis, including measurements, agrees perfectly with

one (No. 146) of the three specimens brought back by Townsend and now before me through the kindness

of Dr. Witmer Stone, of the Academy of Natural Sciences of Philadelphia. I have had the skulls removed
from these three mounted specimens, and for the first time it is possible to identify them. The old labels
which were formerly tacked on the stands are now attached to these specimens, and Dr. Stone thinks
they are in the handwriting of Le Conte, who published on them in 1852, 18 years after they were collected.

(Le Conte also described a Geomys bursarius as Geomys oregonensis from aspecimen labeled ‘Columbia

River, J. k.Townsend.’’) Itis probable that the specimens came into the collection without labels, and

were supposed to have come from the Columbia River, but none of the three of Thomomys were collected

anywhere near the Columbia. No. 147 is Bachman’s type of townsendi and No. 144 is an adult specimen
of pygmzxus which Bachman considered a young of borealis. Bachman’s type of borealis proves to be
identical with clusius of Coues. Townsend in his Narrative (p. 59) mentions a pocket gopher which he
picked up near Scott’s Bluff, Nebr., at a point near the southeastern corner of what isnow Wyoming. As
this specimen was preserved and is the only specimen mentioned in the narrative, it may well be the type
of Bachman’s borealis. As Richardson two years earlier applied the name borealis to another species (¢al-
poides), Bachman’s name has no standing in this connection. The fact that Richardson had identified

this specimen as his borealis, and on it had drawn up a full description which was used by Bachman, does
not make available the name which was already a synonym of talpoides,

1915.1 THOMOMYS TALPOIDES GROUP. 101

and more rounded than in talpoides; anterior arm of basioccipital
very narrow; nasals slenderer, especially at posterior tips, which are
variously rounded, truncate, or slightly emarginate in the topotype
series; interparietal small and triangular. Dentition conspicuously
lighter than in talpoides.

Measurements.—Topotype (éad.): Total length, 205; tail verte-

bre, 66; hind foot, 28.5. Average of 7 female topotypes: 204, 60, 27.
Skull (topotype, gad.):' Basal length, 33.4; nasals, 13; zygomatic
breadth, 22; mastoid breadth, 19.5; interorbital breadth, 11.5:
alveolar length of upper molar series, 7. :
_ Remarks.—In general appearance clusius is nearer talpoides than
is the larger and paler bullatus, which separates their ranges and evi-
dently grades into one on the north and the other on the south. It
is not improbable, however, that clusius and talpoides meet in a
narrow Transition Zone strip along the eastern base of the Rocky
Mountains above the Upper Sonoran limits of bullaius. At the type
locality and over most of its range clusiws is found in Transition
Zone sage-brush country.

Specumens examined.—Total number, 106, as follows:

Colorado: Avalo, 5; Boulder, 1; Canadian Creek (North Park), 6; Colorado
Springs, 7; Estes Park, 1; Flagler, 7; Gold Hill, 1; Limon, 1; Pawnee Buttes,
1; Siebert (8 miles south), 1.

Wyoming: Bear Creek (3 miles southwest of Laramie Peak, at 7,500 feet alti-
tude), 1; Beaver, 1; Bridgers Pass, 14; Casper (40 miles southwest), 4; Chey-
enne, 7; Dayton, 1; Douglas, 2; Ferris Mountains (7,800 to 9,400 feet), 4;
Fetterman, 1; Fort Russell, 1; Fort Steele, 3; Green Mountains (8,000 feet),
2; Islay, 2; Laramie Mountains (east of Laramie, at 8,500 and 9,000 feet), 3;
Little Bear Creek (20 miles southeast of Chugwater), 1; Miners Delight, 1;
Myersville, 2; Myersville (22 miles southeast), 1; Pass (now Parkman), 2;
Pine Bluffs, 2; Rattlesnake Mountains (north base), 3; Rawhide Butte
(5,400 feet), 1; Red Bank, 1; Rock Creek, 1; Sherman, 2; Shirley Mountains
(7,600 to 8,800 feet), 4; Springhill (12 miles north of Laramie Peak), 2; Sun,
3; Wheatland (15 miles southwest, at 5,200 feet), 1; Woods, 2.

THOMOMYS TALPOIDES BULLATUS Baitey.
SAGE PockET GOPHER.
(Pl. VI, fig. 2.)
Thomomys talpoides bullatus Bailey, Proc. Biol. Soc. Washington, XXVII, 115, July
10, 1914.

Type.—Collected at Powderville, Custer County, Montana, by Dr.
A. K. Fisher, July 21, 1893. Type specimen in U. S. Nat. Mus.,
Biological Survey collection.

Disiribution.—Plains of eastern Montana, northeastern Wyoming,
and western South Dakota; north to Medicine Hat, Alberta (fig. 10).

Characters.—Size of talpoides, but with larger audital bulle, lighter
dentition, and brighter colors; mamme usually in 6 pairs; young, 5 to 7.

1 No. 25534, U. S. Nat. Mus.

102 NORTH AMERICAN FAUNA. [No. 39.

Color..—Summer pelage: Practically the same as in clusius, with
less gray on cheeks; upperparts light buffy- or hazel-gray, brightest
on crown; ear patch blackish; underparts buffy, sometimes with
white on chin; feet and tail whitish. Winter pelage (November
specimen, from Fort Custer): Upperparts much paler, light buffy
eray; underparts creamy white.

Skull.—Size and general form nearly as in talpoides, but less mdeed
and with smaller molars, much larger and more rounded bulle, and
narrower anterior shaft of basioccipital.

Measurements.—Type (¢ ad.): Total length, 238; tail vertebre,
72; hind foot, 30. Male topotype: 242, 76, 30; female topotype:
225, 78, 29. Skull (of type): Basal length, 37.6; nasals, 15.5; zygo-
matic breadth, 24; mastoid breadth, 20.3; interorbital breadth, 6.5;
alveolar length of upper molar series, 8.

Remarks.—This is a pale Upper Sonoran form, not entirely inter-
mediate between talpoides and clusius, both of which are Transition
Zone forms (one of the humid grass-prairies, the other of arid sage-
brush). To some extent bullatus combines the characters of both,
but is not quite intermediate and has a well-defined range. The
beautiful series of specimens from Medicine Hat, Alberta, mainly
immature females, are in the light gray, early winter coat. They are
provisionally placed with this form until better characters can be
observed from skulls of adult males and from summer skins.

Specimens examined.—Total number, 82, as follows:

Alberta: Medicine Hat, 10 (not typical).

Montana: Alzada, 1; Cedar Creek (15 miles north of Terry), 3; Darnalls (on south
bank Missouri River south of Glasgow), 1; Fort Assiniboine, 4; Fort Custer, 6;
Fort Thorne, 1; Johnson Lake, 2; Piney Buttes, 2; Powderville, 4; Red
Lodge, 1; Terry, 1.

North Dakota: Buford, 4; Yellowstone River (mouth), 1

South Dakota: Buffalo Gap, 3; Corral Draw (Pine Ridge Indian Reservation),
4; Crow Buttes (Harding County), 1; Elk Mountain, 10; Fort Meade, 1;
Rapid City, 2; Smithville, 2

Wyoming: Clearmont, 1; Devils Tower, 1; Ishawooa Creek, 4; Moorcroft, 3;
Newcastle, 5; Powder River crossing, 2; Sage Creek (west of Fort Washakie), .
1; Wind River (north of Washakie), 1

THOMOMYS TALPOIDES NEBULOSUS Batey.

Biack Hitts Pocket GOPHER.
(Pl. VIL, fig. 3.)
Thomomys talpoides nebulosus Bailey, Proc. Biol. Soc. Washington, XX VII, 116, J uly
10, 1914.

Type.—Collected in Sand Creek Canyon, Black Hills, Wyoming
(at Jack Boyden’s ranch, 5 miles above mouth of Canyon, at 3,750 feet
altitude), by Vernon Bailey, August 25, 1913. Type specimen in
U.S. Nat. Mus., Biological Survey collection.

1915.3 THOMOMYS TALPOIDES GROUP. 103

Distribution.—Black Hills, S. Dak., and Bear Lodge Mountains,
Wyo. (fig. 10). :

Characters.—Size of talpoides; color more brownish gray; skull
slender, lightly ridged; dentition light; bullse well rounded; not
very different externally from fossor, but mamme in 6 pairs, inguinal
2-2, abdominal 2-2, pectoral 2-2.

Color—Summer pelage (July and August): Upperparts dull gray-
ish brown; nose plumbeous or dusky; ear patch blackish; underparts
buffy, generally with white on chin and sometimes on breast; feet
and tail whitish gray or buffy. Wunter pelage (held over in April
specimens from Redfern, 8. Dak.): Dark buffy gray, darker than in
similar coat of talpoides, not so rufescent as in fossor; underparts
light buff. Young (balf-grown July specimen from Custer, S. Dak.):
Dull and dark, as in talpordes.

Skull.Larger and more heavily ridged than in talpoides but less
heavily ridged than in rufescens; temporal ridges parallel in adults,
and interparictal triangular; audital bulle full and well rounded but
not so large as in bullatus; basioccipital slender. Dentition conspicu-
ously light. Compared with fossor, the skull is larger, relatively
wider, and more robust. i

Measurements.—Type: Total length, 230; tail vertebre, 66; hind
foot, approximately 32 (27 without toenails). Average of 4 female
topotypes: 226, 65, 31.5. Skull (of type): Basal length, 37.7; nasals,
14.3; zygomatic breadth, 24.3; mastoid breadth, 20; interorbital
breadth, 6.5; alveolar length of upper molar series, 7.7.

Remarks.—This is a well-marked form of talpoides, occupying the
higher parts of the Black Hills and Bear Lodge Mountains, and
coming down in wooded canyons to the base of both ranges in the
Canadian and Transition Zones. In the Bear Lodge Mountains it is
less strongly differentiated from talpoides and shows a tendency to-
ward bullatus, but can best be placed with the Black Hills form.

Specumens ecamined.—Total number, 27, as follows:

South Dakota: Beaver Creek, 1; Custer, 4; Redfern, 4; Spring Creek, 1; Ti-
gerville, 2.

Wyoming: Bear Lodge Mountains (at 6,000 and 6,200 feet altitude), 6; Rattle-
snake Creek (head, at 6,000 feet in the Black Hills), 2; Sand Creek Canyon, 5;
Sundance (in canyon at base of Bear Lodge Mountains), 2.

THOMOMYS TALPOIDES CARYI Battry.
BigHORN PocKET GOPHER.
(Pl. VII, fig. 6.)

Thomomys talpoides caryi Bailey, Proc. Biol. Soc. Washington, X XVII, p. 115, July
10, 1914.
Lype.—Collected at head of Trapper Creek, at 9,500 feet altitude,

in the Bighorn Mountains, Wyoming, by Merritt Cary, July 10, 1910.

Type specimen in U.S. Nat. Mus., Biological Survey collection.

104 NORTH AMERICAN FAUNA. [NO. 39.

Distribution.—Canadian Zone on Bighorn Mountains, Wyo. (fig. 10).

Characters.—Size about the same as clusius or slightly smaller; color
darker, more rufescent, without gray cheeks and sides; skull with
smaller bullae and wider interparietal; mamme in 6 pairs.

Color.—Summer pelage (September): Upperparts warm grayish
brown, with plumbeous nose and black ear patch; underparts rich
buffy with white on chin and sometimes on breast; feet soiled whitish;
tail gray or buffy with whitish tip. Wznter pelage (on rump of type,
June 10): Dark buffy gray, underparts creamy.

Skull—General form similar to that of clusius but with shorter,
wider interparietal, slightly smaller bullae, and generally wider and
more emarginate nasals.

Measurements.—Type (2 young ad.): Total length, 196; tail ver-
tebree, 54; hind foot, 26. Topotype (? more fully ad.): 203, 58, 28.
Skull (of er Basal length, 32.5; nasals, 12; zygomatic breadth,
20.5; mastoid breadth, 18; ere nal becadell 6; alveolar length
of upper molar series, 6.2.

Remarks.—This form is poorly represented, but shows characters
that exclude it from any of the neighboring forms. Externally it
most closely resembles the Black Hills gopher, but in cranial charac-
ters stands nearest to clusius, of which it may be considered a dark-
colored mountain race. <A good series of specimens including adult
males from the high central area of the Bighorn Mountaims will
doubtless show more pronounced characters. The details of the
distribution of the form remain to be worked out. The specimens
examined are all females and from the Canadian Zone.

Specumens exanvned.—Total number, 4, as follows:

Wyoming: Bighorn Mountains (9,000 and 9,500 feet altitude, in central part),
2; Bighorn Mountains (8,400 and 9,000 feet, in southern part), 2.

THOMOMYS TALPOIDES PRYORI Batrey.

Pryor Mountain Pocket GOPHER.
(Pl. VII, fig. 7.)

Thomomys pryorit Bailey, Proc. Biol. Soc. Washington, X XVII, 116, July 10, 1914.

Type.—Collected on Pryor Mountains, Montana, at head of Sage
Creek (6,000 feet altitude), by Vernon Bailey, July 16, 1894. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Pryor Mountains, Mont., east to the Bighorn River,
near Fort Custer (fig. 10).

Characters.—Size of clusius, but darker colored and with shorter
nasals and more projecting incisors; mamme in 6 pairs, inguinal 2-2,
abdominal 2-2, pectoral 2-2.

Color. eae pelage: Upperparts dull walnut-brown, about as
in fossor; nose plumbeous; cheeks dark gray; ear patch black;

1915.} THOMOMYS TALPOIDES GROUP. 105

underparts dark buff with no trace of white in topotype series; feet
and tail soiled whitish or buffy. Woanter pelage (as shown in long
fur retained on rump of type and one topotype): More grayish
brown.

Skull_—Low and wide, with very short and posteriorly pointed
nasals; temporal ridges parallel; interparietal triangular; audital
bullz rather small; basioccipital wide between bull; interpterygoid
fossa narrow and sharp. Dentition light; upper incisors strongly
projecting; inner grooves not so distinct as in talpoides.

Measurements.—Type (¢ ad.): Total length, 210; tail vertebra,
60; hind foot, 29. Average of 2 female topotypes: 201, 53, 28.5.
Skull (of type): Basal length, 34; nasals, 12.5; zygomatic breadth,
22; mastoid breadth, 18.5; interorbital breadth, 5.7; alveolar length
of upper molar series, 7.

Remarks.—This seems to be a very local form of the higher slopes
of the Pryor Mountains, and undoubtedly grades into some of the >
other forms below. It seems to be distinct from caryi, the gopher
on the Bighorn Range, from which it is separated by the deep canyon
of the Bighorn River, and also from bullatus, from which it is sepa-
rated at Fort Custer by the river. The shallow, slender rostrum and
arrangement of mamme place it in the talpoides group, but the pecu-
liar nasals and projecting incisors are not found in any of the other
forms.

Specimens examined.—Total number, 7, as follows:

Montana: Bighorn River (west side, near Fort Custer), 2; Pryor Mountains
(cold slopes, edge of Canadian Zone), 5.

THOMOMYS TALPOIDES AGRESTIS Merriam.
San Luts Pocket GOPHER.
(Pl. VI, fig. 4.)
Thomomys talpoides agrestis Merriam, Proc. Biol. Soc. Washington, X XI, 144, June
9, 1908.

Type.—Collected at Medano Ranch (15 miles northeast of Mosca),
San Luis Valley, Colorado, by Merritt Cary, October 29, 1907. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—San Luis Valley, Colo. (fig. 8).

Characters.—Decidedly larger and paler than typical clusius;
skull heavier and more conspicuously ridged; more nearly resembling
bullatus, but with relatively narrower skull, and larger, more pointed
nasals.

Color.—Summer pelage: Upperparts buffy or brownish gray,

richest along median line of back; ear patch blackish; nose gray;

underparts buffy; chin white in two specimens; a white spot on
breast in one; feet and tail whitish or buffy gray. Winter pelage:
Upperparts lighter, more grayish; underparts whitish or creamy.

106 NORTH AMERICAN FAUNA. [No. 39. .

Skull—tLong and narrow, with uniformly arched outline; temporal
ridges prominent and parallel in old age; nasals long and narrow,
and pointed at posterior tip, with a slight expansion back of middle;
interparietal triangular; auditory meatus slender; bullae medium; not
so full and rounded as in bullatus. Dentition medium; incisors bent
at right angles to axis of skull.

Measurements.—Type (2 ad.): Total length, 220; tail vertebre,
57; hind foot, 30. .Topotype (¢ not fully ad.): 205, 50, 30. Skull
(of type): Basal length, 38; nasals, 15; zygomatic breadth, 23.5;
mastoid breadth, 19.7; interorbital breadth, 6.5; alveolar length of
upper molar series, 8. 7

Remarks.—This is a well-marked local form of the talpoides group
occupying the great arid San Luis Valley, in open sage brush of the
Transition Zone. At present it may be isolated from other forms of
the group except fossor, by which it is surrounded in the timbered
mountains and from which it may have been derived. More prob-
ably, however, it is a more recent derivative of clusvus which has
entered the valley through one of the passes.

Specumens examined.—Total number, 18, as follows:

Colorado: Blanco, 1; Creston, 1; Fort Garland, 2; Medano Ranch, 5; Saguache
(12 miles northwest), 1; San Acacia, 8.

THOMOMYS COLUMBIANUS Battey.

CoLtuMBIA POCKET GOPHER.
(PL. VIL, fig. 12.)
Thomomys fuscus columbianus Bailey, Proc. Biol. Soc. Washington, X XVII, 117, |
July 10, 1914.

Type.—Collected at Touchet, Walla Walla County, Washington, by
Clark P. Streator, September 10, 1890. Type specimen in U. S.
Nat. Mus., Biological Survey collection. |

Distribution.—Plains of southeastern Washington and northern
Oregon (fig. 5).

Characters.—Size slightly smaller than talpoides, shghtly larger than
quadratus, colors paler than either, about as in fisheri; skull heavier
than in quadratus; claws moderately stout; mamme in 6 pairs,
inguinal 2—2, abdominal 2-2, pectoral 2-2.

Color.—Summer pelage: Upperparts light wood brown or buffy
gray; sides and belly pale buffy gray; ear and postauricular patch
black; nose slaty; tail gray with white tip; feet whitish. Woanter
pelage slightly grayer. Young very gray.

Skull.—Smaller than that of talpoides, with shorter, wider nasals;
larger and longer than that of guadratus, rather heavy and angular;
lateral ridges well developed in adults, approximately parallel; bull

1915.] THOMOMYS TALPOIDES GROUP. LOZ

large and rounded, widening base of skull and narrowing shaft of
basioccipital; interparietal more or less triangular.

Measurements.—Type (3 ad.): Total length, 209; tail vertebre,
60; hind foot, 28. Topotype (¢@ ad.): 208, 68, 27. Skull (of type):
Basal length, 34; nasals, 13; zygomatic breadth, 22.7; mastoid
breadth, 18.7; alveolar length of upper molar series, 8.

Remarks.—The relationships of this form are doubtful. In many
respects it resembles the talpoides group, especially in arrangement
of mamme and in general skull characters. In small ears and rather
square-built skull it seems close to qguadratus, and there is some ques-
tion whether along the borders of its range it does not interbreed with
both quadratus and fuscus. ‘Two young females from Asotin, Wash.,
are probably typical of columbianus, but are too young to afford reliable —
characters. An old male from Willows Junction, Oreg., is very small,
but is provisionally referred to columbianus.

Specimens examined.—Total number, 37, as follows:

Oregon: Pendleton, 4; Umatilla, 9; Willows Junction, 1.
Washington: Asotin, 2; Baird, 9; Fort Walla Walla, 3; Prescott, 4; Touchet, 4;
Wallula, 1.
THOMOMYS OCIUS Merriam.

GREEN River Pocket GoPHER.
(PI. I, fig. 12; Pl. VIL, fig. 9.)
Thomomys clusius ocius Merriam, Proc. Biol. Soc. Washington, XIV, 114, July 19,
1901.

Type.—Collected near Old Fort Bridger, Wyoming (exact locality,
dry sagebrush mesas at Harvey’s Ranch, on Smiths Fork, 6 miles
southwest of Fort Bridger), by Vernon Bailey, May 24, 1890. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Green River Basin of southwestern Wyoming, north-
western Colorado, and northeastern Utah (fig. 10).

Characters.—Size small; color very pale; skull slender, with very
large and rounded audital buile and sharply incurved upper INCISOTs ;
ears very small but pointed; mamme normally 1 in 7 pairs, snenit el
2—2, abdominal 2-2, pectoral 3-3.

Cater —Summer ae Upperparts light buffy gray, more strongly
buffy or tinged with brownish on crown and back; sides clear gray;
cheeks darker gray; ear patch blackish but small; underparts, feet,
and tail soiled whitish or creamy. Winter pelage: Upperparts lighter
buffy gray; nose and cheeks clearer gray; underparts whitish or
creamy.

Skull——Rather narrow, with slightly arched dorsal outline; inter-
parietal short and wide; temporal ridges slightly converging ante-
riorly; posterior tips of nasals rounded; audital bullz very large and

108 NORTH AMERICAN FAUNA. [No. 39. -

smoothly rounded, with anterior arm of basioccipital slender and
narrow between them. Dentition: Upper incisors sharply incurved
so that the cranium can be readily suspended by them, and lower
incisors also curved very abruptly upward.

Measurements.—Type (a large old male): Total length, 204; tail
vertebrae, 60; hind foot, 26. Topotype (¢ ad.): 188, 51, 24. Aver-
age of 6 topotypes (2 ad.): 194, 58, 24.6. Har: From crown, meas-
ured in flesh, 5. . Skull (of type): Basal length, 33; nasals,12; zygo-
matic breadth, 20; mastoid breadth, 17; interorbital breadth, 5.5;
alveolar length of upper molar series, 7.

Remarks.—This is a little, pale, desert species of the Green River
Basin, mainly in the Upper Sonoran Zone. It occupies the dry, sage-
brush mesas, while bridgert occupies the fertile valleys close by.
Fort Bridger seems to be the extreme upper limit of ocius, and here
its range meets that of the much larger bridgeri and almost or quite
meets that of the smaller, darker-colored pygmzus. On the east it
meets the range of cluswus, but I find no signs of intergradation, and
the skull characters are so strongly marked that it seems best to
treat ocius as a full species. Its nearest relatives are idahoensis of
southern Idaho, and more remotely the little pygmeus, along its
western border in the Transition Zone. ‘These three forms are the
only members of the genus in which the crania can be readily sus-
pended by hooking the upper incisors over a wire or string. The
ears of ocius are very small, measuring uniformly 5 mm. from pos-
terior base to tip in fresh specimens, while in bridgeri from the same
locality they measure 8mm. The females have normally 14 mamme
and contained 7 fetuses, contrasted with 10 mamme and _ 5 fetuses
in bridgeri, collected at the same time and place.

Specimens ecamined.—Total number, 38, as follows:

Colorado: Douglas Spring, 1; Ladore, 2; Lay, 1; Lily, 2; Rangely, 2; Snake
River (20 miles west of Baggs), 2.
Utah: Uncompahgre Indian Reservation, 2.
Wyoming: Bitter Creek, 2; Eden, 1; Fontenelle, 1; Fort Bridger, 9; Green
River (junction of New Fork), 3; Henrys Fork, 1; Maxon (5 miles south-
west), 8; Opal, 1.
THOMOMYS IDAHOENSIS Merriam.
IpaHo PockET GOPHER.
(PL. I, fig. 7; Pl. VII, fig. 10.)
Thomomys idahoensis Merriam, Proc. Biol. Soc. Washington, XIV, 114, July 19, 1901.

Type.—Collected at Birch Creek (10 miles south of Nicholia, at about
6,400 feet altitude), Fremont County, Idaho, by Clark P. Streator,
August 8, 1890. Type specimen in U.S. Nat. Mus., Biological Sur-
vey collection.

Distribution.—Snake River Plains, southeastern Idaho (fig. 10).

1915.] THOMOMYS TALPOIDES GROUP. 109

Characters.—Size very small; color pale yellowish gray; ears small;
skull light and smooth, with very large and globose bulls and incurved
upper incisors; Mamme in 6 pairs, inguinal 2-2, abdominal 2-2, pec-
toral 2-2.

Color.—Summer pelage: Upperparts pale dull olive-buff or buffy
eray; ear with only a trace of dusky patch; nose yellowish; under-
parts pale buff or soiled whitish, occasionally with white chin; fect
and tail very hairy, pale buffy or whitish. Young, more ashy gray.
Winter pelage unknown. ‘The summer pelage is decidedly paler than
the corresponding pelage of ocius, hence the winter pelage is probably
lighter colored than in any other northern species of the genus.

Skull.—Of much the same general form as in ocius, but conspicu-

ously smaller and smoother with smaller, more oval interparictal;
nasals minutely notched at posterior tip and ending approximately
even with premaxille; audital bull very large and globose, sometimes
actually meeting over narrow shaft of basioccipital. While the ex-
ternal ear is very small, the auditory meatus and audital bulle are
unusually developed, the bullze being relatively the largest of any
species of the genus.

Measurements.—Type (é ad.): Total length, 179; tail vertebre,
47; hind foot, 23. Average of 10 male topotypes: 179, 50, 22.5.
Average of 9 female topotypes: 166, 49, 22.4. Har, from base, meas-
ured in flesh, 4.5. Skull (of type): Basal length, 29.5; nasals, 11.5;
zygomatic breadth, 18; mastoid breadth, 16; interorbital breadth,
4.7; alveolar length of upper molar series, 5.5.

Remarks.—The type locality is at the extreme upper edge of the
Upper Sonoran Zone and the upper limit of the range of the species.
This gopher is a small, pale, desert species of the Snake River Plains.
There is no indication of its intergradation with any of the surround-
ing forms, and its nearest relative is ocius of the upper Green River
Basin in Wyoming. The boundary of its range remains to be worked
out, and winter or early spring specimens are greatly needed to show
the winter pelage.

Specimens examined.—Total number, 42, as follows:

Idaho: Big Butte, 2; Big Lost River (near Sink), 3; Birch Creek (10 miles south
of Nicholia, 6,400 feet altitude), 21; Blackfoot, 11; Dubois, 1; Idaho Falls,
1; Sink of Birch Creek (5,100 feet), 3.

THOMOMYS PYGMAUS Merriam.
Pyeamy Pocket GOPHER.
(Pl. VII, fig. 11.)
Thomomys pygmxus Merriam, Proc. Biol. Soc. Washington, XIV, 115, July 19, 1901.

Type.—Collected at Montpelier Creek (exact locality about 10 miles
northeast of Montpelier, at 6,600 feet altitude, in open sagebrush of

110 NORTH AMERICAN FAUNA. [No. 39. -

Transition Zone), Bear County, Idaho, by Vernon Bailey, July 29,
1893. Type specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Southwestern Wyoming and southeastern Idaho
(Transition Zone) (fig. 10).

Characters.—Size smallest of the genus; color rich hazel-brown;
skull very slender and delicate with not very large audital bulle;
upper incisors incurved, distinctly grooved; ears small; mamme in 6
pairs, inguinal 2-2, abdominal 2-2, pectoral ‘2-2.

Color.-—Summer pelage: Upperparts rich hazel-brown, very uni-
formly distributed; dusky earpatches inconspicuous; nose plumbeous;
underparts dull ochraceous, rarely with any white markings; tail usu-
ally entirely buffy gray; feet whitish or buffy. Winter pelage: Paler,
more buffy brown. Young: Duller, more grayish and mare eae
colored belly.

Skull.—Very small, slender, and thin, the slight trace of temporal
ridges converging anteriorly; interparietal wide and oval; nasais
notched or emarginate at posterior tip; bullz rather small; auditory
meatus slender; interpterygoid fossa narrow and acute angled; basioc-
cipital narrow. Dentition very light; upper incisors incurved to a
slightly less degree than in ocius and idahoensis.

Measurements.—Type (3 ad.): Total length, 177; tail vertebre,
46; hind foot, 22. Topotype (¢ ad.): 165, 40, 20. Specimen from
Bear River (¢ ad.): 168, 50, 23. Skull (of type): Basal length, 28.4;
nasals, 10.4; zygomatic breath, 16.5; mastoid breadth, 14.5; interor-
bital breadth, 5; alveolar length of upper molar series, 6. .

Remarks.—This, the smallest of all known pocket gophers, seems —
not to be closely related to any of the neighboring forms nor to inter-
srade with them. It has some of the skull characters of fuscus, but
seems to be more nearly related to idahoensis and ocius, from which,
however, it is quite distinct. It occurs in the same localities but
apparently not on the same ground with the much larger bridgeri and
uinta, occupying the Transition Zone sagebrush ridges and mesas,
while they are restricted to the mellow soil and more fertile bottoms
of the stream valleys. A specimen in the Academy of Natural
Sciences of Philadelphia (No. 144), collected by J. K. Townsend in
1834, probably came from the Bear River region, but was later
labeled “Columbia River.”

Specimens examined.—Total number, 23, as follows:

Idaho: Montpelier Creek (12 miles northeast of Montpelier at 6,700 feet alti-
tude), 2

Fe Bear River and Bear River Divide (14 miles north of Evanston at
6,600—7,500 feet), 12; Big Piney (6,400 feet), 1; Big Sandy, 1; Fossil (6,600
feet), 1; Lone Tree (5 miles west, on Henry Fork, 7,400 feet), 2; Merna (on
Horse Creek, 7,800-8,000 feet), 3; Surveyors Park (12 miles northeast of
Pinedale, 8,000 feet), 1, .

1915.) TILOMOMYS FOSSOR GROUP. 141

Thomomys fossor Group.
THOMOMYS FOSSOR ALLEN.

CoLORADO POCKET GOPHER.
(P1. I, fig. 8; Pl. VII, fig. 13; text fig. 4.)

Thomomys fossor Allen, Bul. Am. Mus. Nat. Hist. V, 51, April 28, 1893.

Type.—Collected at Florida, La Plata County, Colorado, altitude
7,200 feet, by Charles P. Rowley, June 25, 1892. Type specimen in
Am. Mus. Nat. Hist.

Distribution.—Mountains of western Colorado, extreme southern
Wyoming, northern New Mexico, eastern and some ase and
northwestern Arizona (fig. 8).

Characters.—Size medium, hind foot averaging about 29 mm.;
color dull and dark brown; ears large; skull long, low, and narrow,
with rostrum especially slender in profile, as in the talpoides group;
mamme in 5 pairs, inguinal 2-2, pectoral 3-3.

Color—Summer pelage: Upperparts dull dark brown with some-
times a rich chestnut tone; ear and postauricular patch black; nose
and face dusky; underparts buffy or ochraceous; part of feet and tip
of tail usually whitish; chin usually, and spot on breast sometimes,
white. Winter pelage: Duller, more grayish or drab. Young: In
summer paler and more buffy than adults.

Skull.—Long and slender, with long rostrum, and anteriorally con-
verging temporal ridges; nasals narrow, with generally rounded pos-
terior tips; interparietal triangular; bulle large and full, basi-
occipital narrowed between them. Dentition very light. —

Measurements.—Average of 4 female topotypes: Total length, 221;
tail vertebre, 63; hind foot, 29. A large male measures 220, 68, 32.
Skull (of type, ¢ ad.): Basal length, 37; nasals, 15; zygomatic breadth,
22; mastoid breadth, 20; alveolar length of upper molar series, 7.5.
Skull of adult male from Lake City, Colo.:1 34, 13.5, 21, 18.7, 7.

Remarks.—Vhis is a wide-ranging boreal species with long, soft fur,
which even in midsummer does not become so short and harsh as in
low-country species. At high altitudes the long coat seems not to be
entirely lost during the brief summer.

As might be expected there is some slight variation in specimens
_ from different mountain ranges, some of which are entirely isolated by
low country, but nowhere is there enough variation to warrant
further subdivision. In such a variable group it is a great satisfac-
tion to find an occasional species that holds its distinctive characters
Over a wide area.

Specumens examined.—Total number, 183, as follows:

Arizona: Bright Angel Spring eibab Plateau), 3; DeMotte Park (Kaibab
Plateau), 3.

No. 48190, U. S. Nat. Mus.

Pr? NORTH AMERICAN FAUNA. [No. 39. ”

Colorado: Arapahoe Pass (Rabbit Ear Mountains), 5; Black Hawk, 2; Baxter
Pass (Book Plateau), 2; Boulder (5 miles west, at 5,600 feet altitude), 7;
Cascade, 1; Cochetope Pass (3 miles east), 5; Colorado City, 1; Colorado
Springs (23 miles north, at 6,000 feet), 1; Colorado Springs (east of Palmer
Park), 1; Como, 1; Coulter, 3; Crested Butte, 4; Culebra Canyon (Costilla
County), 1; Elk Head Mountains, 2; Elkhorn, 1; Estes Park, 6; Florida, 5;
Golden, 2; Hahns Peak, 2; Hayden, 1; Lake City, 2; Longs Peak, 3; Meeker,
3; Montgomery, 3; Nederland, 4; Pagosa Springs, 3; Pagosa Peak, 1; Pearl,
1; Saguache (23 miles northwest), 1; Sapinero, 2; Silverton, 2; Uncom-
pahgre Plateau, 2; Teller County Divide, 1; White River Plateau, 1.

New Mexico: Chusca Mountains, 3; Costilla Pass, 5; Costilla River, 3; Gallinas
Mountains (Rio Arriba County), 7; Halls Peak, 1; Hondo Canyon, 1; Hope-
well, 6; Horse Lake, 6; Jemez Mountains (head of Santa Clara Creek), 5;
Moreno Valley, 2; Mount Taylor, 1; Pecos Baldy, 4; Red River (Taos
County), 4; Taos Mountains, 4; Tres Piedras, 1; Tusas River, 1; Twining, 12;
Whites Peak, 1; Willis, 4.

Utah: Beaver Mountains, 9; Buckskin Valley, 1; Fish Lake, 4; La Sal Moun-
tains, 1; Panguich Lake, 1; Parawan Mountains, 4.

Wyoming: Bridgers Peak (8,800 feet altitude), 3; Medicine Bow Mountains
(10,200 feet), 1.

THOMOMYS BRIDGERI Merriam.

Fort BrimGerR PocKET GOPHER.
(Pl. VIL, fig. 15.)
Thomomys bridgeri Merriam, Proc. Biol. Soc. Washington, XIV, 113, July 19, 1901.

Type.—Collected at Fort Bridger, Wyoming (exact locality, Har-
vey’s Ranch on Smiths Fork, 6 miles southwest of Old Fort Bridger),
by Vernon Bailey, May 27, 1890. Type specimen in U. S. Nat.
Mus., Merriam collection. ,

Distribution.—Southwestern Wyoming and southeastern Idaho
(Transition Zone) (fig. 5).

Characters.—Size large; ears large and prominent; color dull dark
brown; skull heavily ridged and angular, with deeply emarginate
nasals; mammee in 5 pairs, inguinal 2-2, pectoral 3-3.

Color—Summer pelage: Upperparts rich warm brown, nearest
to cinnamon-brown of Ridgway; nose and face dusky brown; large
ear patch black; underparts dark buffy or dull ochraceous, occa-
sionally with white patch on chin; feet partly gray or mottled with
gray; toes whitish; tail brownish gray, paler below and occasionally
with white tips. Winter pelage: Upperparts duller and darker, more
nearly Prout’s brown of Ridgway; nose plumbeous, ear patch black;
underparts washed with buffy over hght plumbeous, with almost a
lavender effect. Young more grayish.

Skull.—Large, angular, and heavily ridged in adults; nasals long
and deeply emarginate at posterior tips; temporal ridges parallel
or shghtly converging in middle; interparietal triangular; audital
bullz long, narrow, and wide apart, with wide shaft of basioccipital

1915. ] THOMOMYS FOSSOR GROUP. 113

between; auditory meatus moderately large. Dentition relatively
light; incisors very long and slightly protruding.

Measurements—Type (6 ad.): Total length, 237; tail vertebre,
71; hind foot, 34. Topotype (9 ad.): 227, 68, 31. Average of 10
adult males: 244, 72, 33.2. Average of 10 adult females: 233, 71,
31.5. Har: From crown, 8. Skull (of type): Basal length, 39;
nasals, 16; zygomatic brondal 24.5; mastoid breadth, 22; interor-
bital pad, 7; alveolar length of upper molar series, 8.

Remarks. ae big gopher is abundant in Pale valleys in
the Transition Zone, but does not inhabit the arid upland. It
is a strongly marked, outlying form of the fossor group, and its
nearest relative is winta. It has a lower zonal range, however, and
seems to be entirely distinct. More thorough collecting may show
that it grades into winta higher up the streams, but at present the
two are best treated as distinct species.

At the type locality females were found normally to have 10
mamme and 5 embryos, as contrasted with the 14 mamme and 7
embryos in the little ocius of the dry sagebrush mesas close by.

Specimens examined.—Total number, 74, as follows:

Idaho: Montpelier, 18; Montpelier Creek (6,000 feet altitude), 1

Wyoming: Bear River (14 miles north of Evanston, 6,600-6,800 feet altitude),
5; Border, 1; Cokeville (6,400 feet), 8; Fort Bridger, 24; Henry’s Fork (5
miles west of Lonetree), 1; Lonetree (7,400 feet), 3; Lonetree (4 miles south),
2; Mountain View, 11.

THOMOMYS UINTA Merriam.
Uinta Pocket GOPHER.
(P1. VIL, fig. 14.) 3
Thomomys uinta Merriam, Proc. Biol. Soc. Washington, XIV, 112, July 19, 1901.

Type.—Collected in Uinta Mountains (10,000 feet altitude, north
base of Gilbert Peak), Summit County, Utah, by Vernon Bailey,
June 6, 1890. Type specimen in U.S. Nat. Mus., Biological Survey
collection.

Distribution.—Western Wyoming, southeastern Idaho, and northern
Utah (fig. 5).

Characters.—About the size of fossor and practically indistinguish-
able in external characters; skull shorter and wider, with shorter
rostrum and heavier dentition; mamme in 5 pairs, inguinal 2-2,
pectoral 3-3.

Color.—Summer pelage: eee dull dark brown, with dusky
nose and face and blackish ear and ear patch; mdlewuntts buffy or
ochraceous, with occasionally a white patch on chin; feet and tip
of tail usually whitish. Late fall and winter pelage: Duller and more
grayish. Young, paler, more buffy than adults.

98121°—15——8

114 NORTH AMERICAN FAUNA. [ No. 39.

Skull—Heavier and wider than in fossor, with heavier dentition,
smaller bulle, a more nearly five-angled interparietal, and wider
nasals with deeply emarginate instead of rounded posterior tips.

Measurements.—Average of type and topotype (¢ ad.): Total
length, 226; tail vertebre, 70; hind foot, 31. Topotype (¢ ad.): 211,
64, 28. Skull (of type): Basal length, 35; nasals, 13; zygomatic
breadth, 23; mastoid breadth, 21; alveolar length of upper molar
series, 8.

Remarks.—While very similar to its nearest relative, fossor, in
external characters, winta has such pronounced skull characters and
is so widely separated that it is best treated as a full species. It also
has the same boreal, mountain habitat and practically the same
environment and habits as fossor. Specimens from the mountains
of western Wyoming show some local variation but not sufficient
to be recognized as a subspecies. There seems to be no evidence
of intergradation with the smaller and smaller-eared fuscus group,
although they come close together in the region south of Jackson
Hole, Wyo.

Specimens examined.—Total number 110, as follows:

Idaho: Albion, 5; Blackfoot (hills east of town), 1; Bridge, 1; Inkom, 12; Irwin
(10 miles southeast), 7; Malade, 6; Pocatello, 4; Preuss Mountains, 5; Shelley,
3; Swan Lake, 4.

Utah: Kelton, (mountains 7 miles north), 3; Ogden, 6; Park City, 3; Parley
Canyon, 1; Uinta Mountains (9,000-10, 000 feet alintde) 6.

Wyoming: ion 7; Afton (10 miles southeast, in Salt River Mountains), 6;
Afton (10 miles north on Salt River), 7; Black Rock Creek (head, in Shoshone
Mountains), 5; Kendall (12 miles north), 6; Merna, 6; Needle Mountain
(10,000 feet altitude), 4; Stanley (3 miles west), 1; Thayne, 1. |

THOMOMYS QUADRATUS QUADRATUS Merriam.

DatLtEs Pocket GOPHER.
(Pl. VIL, fig. 8.)

Thomomys quadratus Merriam, Proc. Biol. Soc. Washington, XI, 214, July 15, 1897.

Type.—Collected at The Dalles, Oregon, by Clark P. Streator,
November 2, 1893. Type specimen in U. S. Nat. Mus., Biological
Survey collection.

Distribution.—Plains of eastern and central Oregon, northeastern
California, and northwestern Nevada (fig. 5).

Charatan —Smaller than winta; rather brighter colored; ears small;
skull relatively short and wide, with wide, truncate, posterior tip a
nasals; mamme in 5 pairs, inguinal 2-2, pectoral 3-3.

Color.—Summer pelage: Upperparts light russet; ear patch black;
nose dark plumbeous; underparts washed with dark buff; tail
brownish above, except at tip; feet whitish. Winter pelage: Duller
and grayer. Young, dull and rather dark.

1915.] THOMOMYS FOSSOR GROUP. 115

Skull——Relatively short and rectangular, with parallel lateral
ridges, and rectangular zygomatic arches; nasals short and _ pos-
teriorly truncate; pterygoids V-shaped; interparietal small, rounded,
triangular, or cordate; bulle rather small.

Measurements.—Average of 3 male topotypes: Total length, 210;

tail vertebree, 64; hind foot, 27. Average of 5 female topotypes:

195, 62, 26.4. Skull (of type): Basal length, 36; nasals, 13.5; zygo-
matic breadth, 24; mastoid breadth, 19; alveolar length of upper
molar series, 8.

Remarks.—This is a wide-ranging, variable form of the fossor
sroup, occupying the Upper Sonoran plains of eastern and central
Oregon, northeastern California, and northern Nevada, and extending
higher up in some of the narrow desert ranges.

Specimens exanvned.—Total number, 159, as follows:

California: Bieber, 7; Brownell, 1; Dry Creek (at 4,450 feet altitude), 1; Eagle
Lake, 2; Goose Lake, 1; Goose Lake Meadows (4,800 feet), 2; Lake City, 1;
Lassen Creek, 7; Madeline Divide, 1; Madeline Plains, 6; Pete’s Valley, 1;
Secret Valley, 1; Sugar Hill (Modoc County, 5,000 feet), 1; Susanville
(4 miles south), 2; Warner Mountains (head of Parker Creek, 7,300 feet), 16;
Warner Mountains (Parker Creek, 5,500 feet), 13; Warner Peak (east slope,
8,700 feet), 3.

Nevada: Badger, 1.

Oregon: Adel, 4; Crooked River, 3; Fremont, 4; Harney, 2; Ironside (4,000
feet altitude), 11; Lake Alvord, 1; Lakeview, 4; Matoleus River, 2; Merrill, 2;
Mount Warner, 1; Shirk, 17; Silver Lake, 6; The Dalles, 21; Tumtum Lake,
1; Wapinitia, 1.

Washington: Rockland, 12.

THOMOMYS QUADRATUS FISHER! Merriam.

FisHer Pocket GOPHER.
(Pl. VII, fig. 16; text fig. 3.)

Thomomys fuscus fisher’ Merriam, Proc. Biol. Soc. Washington, XIV, 111, July 19,

no Oris

Type.—Collected at Beckwith, Sierra Valley, Plumas County,
California, by W. K. Fisher, August 3,1900. Type specimen in U.S.
Nat. Mus., Biological Survey collection.

Distribution.—Northern, central, and western Nevada; west in
California to Sierra Valley and Mono Lake (fig. 6).

Characters—Shehtly smaller than quadratus, shorter-tailed, and
paler; smaller than clusius, about the same color but with narrower
skull and smaller bulle; much larger than idahoensis with relatively
smaller bull; claws moderately stout; mamme in 5 pairs, inguinal
2-2, pectoral 3-3.

Color.—Summer pelage: Upperparts buffy gray, lightly washed
with dull russet over back; ear patch blackish; nose plumbeous;
underparts buffy; chin white; feet whitish; tail whitish below, gray

116 NORTH AMERICAN FAUNA. [No.39. -

above usually to the tip. Winter pelage: Slightly grayer. Young,
more buffy.

Skull.—More like that of quadratus than of fuscus or monticola;
lateral ridges parallel in adults; nasals truncate posteriorly; inter-
parietal broadly triangular. Dentition lighter, and incisors less
abruptly decurved than in quadratus, fuscus, or monticola.

Measurements.—Average of 3 topotypes (¢ ad.): Total length,
191; tail vertebrae, 58; hind foot, 25.3. Average of 3 adult females:
194, 58, 25. Skull (of type, ¢ ad.): Basal length, 31; nasals, 11;
zygomatic breadth, 20; mastoid breadth, 17.5; alveolar length of
upper molar series, 6.

Remarks.—The subspecies fishert 1s typical only along the western
part of its range. In central Nevada it is darker and perhaps grades
into wnta. In northwestern Nevada it grades imperceptibly into
quadratus, the only form with which it seems to be closely connected.
It is quite distinct from monticola, with which it occurs at the type
locality, and from which it differs in much smaller ears and paler
color. 3

Specimens excamined.—Total number, 97, as follows:

California: Beckwith, 1; Casa Diablo, 6; Mono Lake, 8; Sierra Valley, 8.

Nevada: Arc Dome, 2; Big Creek 6; Bull Run, 2; Cottonwood Range, 14;
Eureka, 3; Monitor Mountains, 2; Mount Siegel, 2; Mount Sugar, 6; Pine
Forest Range, 4; Reese River Valley (50 miles south of Austin), 9; Reno, 6;
Ruby Lake, 3; Ruby Mountains, 2; Silver Creek (north of Austin), 5; Sum-
mit Lake, 3; Verdi, 3; Wells, 2.

Thomomys douglasi Group.
THOMOMYS DOUGLASI DOUGLASI (Ricuarpsoy).

Dovuaias PockEeT GoPHER; CoLUMBIA SAND Rar.
(Pl. II, fig. 10; Pl. VIII, fig. 5.)
Geomys douglasit Richardson, Fauna Boreali-Americana, I, 200, 1829.

Geomys fuliginosus Schinz, Syn. Mamm., II, 136, 1845. ‘“‘Habitat ad fluvium

Columbia.’”’
Thomomys douglasii Allen, Bul. Am. Mus. Nat. Hist., V, 66, April 28, 1893.

Type.—Collected ‘‘near the mouth of the Columbia,” at Fort
Vancouver, Washington, by David Douglas, probably m 1825. Mr.
Oldfield Thomas writes that it is not now in the British Museum
collection.

Distribution.—Known only from type locality (fig. 5).

Characters.—Size medium; claws stout; ears medium with rounded
tips; color nearly uniform dull hazel without dark ear patch; skull
long and slender and flat on top; incisors abruptly decurved and
distinctly grooved; mamme in 4 pairs, inguinal 2—2, pectoral 2-2.

Oolor.—(June specimens from type locality): Upperparts uniform
dull hazel, slightly paler on sides; underparts more ochraceous, with

1915.} THOMOMYS DOUGLASI GROUP. 1 Ba bee

a white spot on breast in all of the topotypes; nose gray; feet and
tail soiled whitish. Winter and young pelages not seen.

Skull.—Long and narrow, somewhat of the monticola type, but
with heavier rostrum, wider interorbital constriction, and with
zygomatic arches conspicuously widest at anterior angle; inter-
parietal very small and transversely oval; bulle small and narrow;
basioccipital triangular; pterygoids low and U-shaped. )

Measurements.—Average of 3 topotypes (¢ ad.): Total length,
215; tail vertebre, 64; hind foot, 30. Average of 4 females: 200, 58,
28.6. Skull (of topotype, 3 ad.):1 Basal length, 36; nasals, 14; zygo-
matic breadth, 22.5; interorbital breadth, 7; mastoid breath, 19;
alveolar length of upper molar series, 7.6. |

Remarks.—The well-developed ears, number and position of
mamme, general tone of coloration, and long narrow skull at first
suggest douglasi as the type of the monticola group, but the heavier
claws, more rounded ear tips, heavier rostrum, and wider interorbital
region seem to be group characters shared only by douglasi, oregonus,
yelmensis, melanops, limosus, and niger. This group of isolated
forms may somewhere connect with mazama, which seems to be its
nearest relative and neighbor, but for convenience it may be
termed the douglasi group. With the smaller-eared, slenderer-clawed
fuscus group, it seems less closely connected. From quadratus
and columbianus it differs widely in external as well as cranial
characters.

Specimens examined.—Total number, 7, as follows:

‘Washington: Vancouver (6 or 7 miles northeast, at Fourth Plain), 7.?

THOMOMYS DOUGLASI OREGONUS Merriam.
OREGON POCKET GOPHER.
(P1. VIII, fig. 8.)

Thomomys douglasi oregonus Merriam, Proc. Biol. Soc. Washington, XTV, 115, July 19,
1901.

Type.—Collected at Ely, near Oregon City, Oregon (exact locality in
Ely’s orchard), by Clark P. Streator, October 24, 1893. Type speci-
men in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Known only from type locality (fig. 5).

Characters.—Size about as in douglasi; ears smaller, but of the same
rounded form; color brighter hazel; skull relatively shorter and
wider, with distinctly triangular interparietal; mammz in 4 pairs,
inguinal 2—2, pectoral 2-2.

Color.—October specimens in thin summer pelage: Upperparts
clear bright hazel, with dusky nose and cheeks and blackish ear patch;

1 No. 65929, U. S. Nat. Mus. 2Collected by Clark P. Streator, June 15-16, 1894.

118 NORTH AMERICAN FAUNA. _ [No. 39. -

underparts paler, more ochraceous; feet and tail soiled whitish.
Young of the year, paler, more yellowish. A trace of long, rough fur
on the rumps of several breeding females is duller and darker, more
like the June pelage of douglasv.

Skull.—Relatively shorter and wider than in douglasi, with more-
spreading and more nearly parallel zygomatic arches, triangular inter-
parietal, larger bullz, V-shaped instead of U-shaped interpterygoid
fossa, and heavier molars; nasals narrow, with generally double
rounded posterior tips.

Measurements.—Average of 5 topotypes (¢ ad.): Total length,
216; tail vertebra, 67; hind foot, 29.5. Type (¢ ad.): 220, 70, 30.
Average of 5 adult females: 210, 66, 28.2. Skull (of type): Basal
length, 36; nasals, 14; zygomatic breadth, 23.7; interorbital breadth,
6.5; mastoid breadth, 19.4; alveolar length of upper molar series, 7.6.

Remarks.—The relationship between douglasi and oregonus is
not very close and the question of continuity of range has not been
worked out. The Columbia River probably separates their ranges,
but they are near enough in characters to allow subspecific rank to
show their relationship.

Specumens examiuned.—Twenty-eight, from type locality.

THOMOMYS DOUGLASI YELMENSIS Mergiam.

YELM PockEeT GOPHER.
(Pl. VIII, fig. 11.)
Thomomys douglasi yelmensis Merriam, Proc. Biol. Soc. Washington, XIII, 21, Jan.
al L399:

Type.—Collected at Tenino, Yelm Prairie, Thurston County,
Washington, by Clark P. Streator, October 24, 1891. Type specimen
in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Prairies around south end of Puget Sound, Wash.
(Cie 5))s

Characters.—Very similar to douglasi, but with more-pointed ears,
duller and darker coloration, and conspicuous black ear patches;
zygomatic arches not widest at anterior angle.

Color.—In faded October pelage: Upperparts dull hazel; nose and
sides of face dusky; ears and large ear patches blackish; underparts
buffy with irregular white patches; two specimens with white spots
on breast and one with white spots on side; feet and tail white or
soiled whitish. Winter pelage (just appearing on heads in October
specimens) dark umber on crown.

Skull.—Long and low like that of douglasi; slightly concave inter-
orbitally; zygomatic arches parallel or widest posteriorly; inter-
pterygoid fossa more nearly V-shaped; nasals narrower and more
deeply emarginate at posterior tips.

1915. ] THOMOMYS DOUGLASI GROUP. 119

Measurements.—Type (¢ ad.): Total length, 222; tail vertebrae,
68; hind foot, 32. Skull (of type): Basal length, 35.5; nasals, 15;
zygomatic breadth, 22.7; interorbital breadth, 6.6; mastoid breadth,
18; alveolar length of upper molar series, 7.7.

Remarks.—This is a well-marked form of the douglasi group
probably not occurring beyond the margins of the Puget Sound
prairies. The heavily timbered areas of western Washington and
Oregon seem to exclude pocket gophers, but the scattered colonies
of related forms strongly suggest a time when the valley prairies
were more extensive in area, and the range of the gophers more or
less continuous from Puget Sound to the Umpqua Valley. Isolated
colonies have become more or less differentiated in characters. Even
those from Roy, Spanway, and Steilacoom, Wash., differ to some
extent from the type series from Tenino, but not sufficiently for sep-
aration.

Specimens examined.—Total number, 44, as follows:

Washington: Roy, 19; Spanway, 6: Steilacoom, 16; Tenino, 3.

THOMOMYS DOUGLASI MELANOPS Merriam.
BiLacK-HEADED POCKET GOPHER.
(Pl. VIII, fig. 6.)
Thomomys melanops Merriam, Proc. Biol. Soc. Washington, XIII, 21, Jan. 31, 1899.

Type.—Collected at timberline at head of Soleduck River, Olympic
Mountains, Clallam County, Washington, by C. Hart Merriam and
Vernon Bailey, August 28, 1897. Type specimen in U.S. Nat. Mus.,
Biological Survey collection.

Mstribution.—Olympic Mountains, Wash. (fig. 5).

Characters.—About the size and color of yelmensis, but with more
black on nose and head; ears small and pointed; skull long, with
upper outline more arched than in yelmensis or douglasi; pterygoids
small and wide apart.

Color—August pelage: Upperparts dark russet, with dusky ‘nose
and face, and large black ear patches; underparts paler, more ochra-
ceous, usually with white chin, wrists, and lining of cheek pouches;
fore feet pure white; hind feet and tip of tail whitish.

Skull—bLong and well arched; lower outline of rostrum con-
stricted; nasals long and deeply emarginate; premaxille wide; inter-
parietal very small and oval or circular; bullae narrow, and basioc-
cipital wide between them; pterygoids high and wide apart.

Measurements —Type (¢ ad.): Total length, 206; tail vertebre, 63;
hind foot, 27. Topotype (@ ad.): 202, 58, 27. There are no adult
males from the type locality and those from Happy Lake, Wash., are
not correctly measured. Skull (of type): Basal length, 33; nasals, 13;
zygomatic breadth, 20; mastoid breadth, 16; interorbital breadth,

120 NORTH AMERICAN FAUNA. [NO. 39. °

6.4; alveolar length of upper molar series, 7.3. Skull of adult male
from Happy Lake: 34.6; 14.3; 22; 17; 6.3; 7.

Remarks.—Only females and one young male of this form have
been examined from the type locality, and in these the cranial
characters are not perfectly shown. Specimens of adult male and
female from Happy Lake, near the type locality, in the collection of
the Field Museum, indicate a closer relationship with yelmensis and
douglas than was formerly supposed. ‘The relationship of the form
seems best indicated by considering it a subspecies of douglasi,
although at present direct geographic connection is improbable.

Specumens examined.—Total number, 14, as follows:

Washington (Olympic Mountains): Happy Lake, 9; Soleduck River (head), 5.3

THOMOMYS LIMOSUS Merriam.
WHITE SALMON PocKET GOPHER.
(PI. VIII, fig. 12.)

Thomomys limosus Merriam, Proc. Biol. Soc. Washington, XIV, 116, July 19, 1901.

Type.—Collected at White Salmon, gorge of the Columbia, Klick-
itat County, Washington, by J. Alden Loring, June 26, 1897. Type
specimen in U.S. Nat. Mus., Biological Survey collection.

Distribution.—Known. only from type locality (fig. 5).

Characters.—Size and proportions of douglasi, but darker colored
and with blackish ear patch; skull shorter and wider.

Color.—June pelage: Upperparts dull chestnut, with plumbeous
nose and blackish ear patches; underparts slightly paler, more
ochraceous; in about half the specimens a white spot on breast; feet —
and tail soiled whitish. Young: Half-grown specimens very similar
to adults, but shghtly paler, and belly more buffy.

Skull.—Relatively short and wide with more arched dorsal outline
and wide-spreading zvygomata; nasals with emarginate, spreading,
posterior tips, terminating approximately even with tips of pre-
maxille; bullae low and narrow, and anterior shaft of basioccipital
correspondingly wide between them; pterygoids small, with narrow.
fossa.

Measurements.—Type (¢ ad.): Total length, 224; tail vertebre,
68; hind foot, 30. Average of 3 topotypes (¢ ad.): 224, 69, 29.3.
Topotype (¢ ad.): 216, 66, 27. Skull (of type): Basal length, 34.5;
nasals, 14; zygomatic breadth, 23.5; mastoid breadth, 20; interorbital
breadth, 6.2; alveolar length of upper molar series, 8.

Remarks.—While so strikingly different from douglasi as to require
specific rank, this form is included in the douglasi group because of
its prominent, rounded ear, heavy claws, and general skull characters

1 Two adult and two immature females, and one immature male.

1915.] THOMOMYS MONTICOLA GROUP. 121

It is in all probability an isolated form but evidently of ancestral
connection with douglasi.

Specimens examined.—Six skins and seven skulls, from type
locality.
THOMOMYS NIGER Merriam.

Brack Pocket GOPHER.
(Pl. VIII, fig. 7.)
Thomomys niger Merriam, Proc. Biol. Soc. Washington, XIV, 117, July 19, 1901.

Type.—Collected at Seaton, near mouth of Umpqua River,
Douglas County, Oregon, by J. Ellis McLellan, October 6, 1894.
Type specimen in U. S. Nat. Mus., Biological Survey collection.

Distribution.—Coast region of west-central Oregon (fig. 5). 3

Characters.—About the size of douglas, to which it shows nearest
relationship in its heavy claws and rounded ear; skull heavier,
shorter; rostrum depressed; upper incisors curved inward; color
black; mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color.—Upperparts uniform glossy black with purple and green
iridescence; underparts duller and more plumbeous; feet and distal
portion of tail white. Young, sooty black.

Skull.—Shorter, wider, and heavier than in douglasi, with heavier
rostrum; nasals posteriorly sulcate, with the anterior half abruptly
widened, and the posterior tips sharply emarginate; imterparietal
small; bull fuller and more rounded than in douglasi; pterygoids
thick and wide apart. Dentition heavier.

Measurements.—Type (¢ ad.): Total length, 225; tail vertebre,
81; hind foot, 30. Female topotype: 214, 71, 30. Skull (of type):
Basal length, 33.5, nasals, 14; zygomatic breadth, 22.5; mastoid
breadth, 18; interorbital breadth, 6; alveolar length of upper molar
series, 8.

Remarks.—This is probably a dichromatic species of which as yet
only specimens of the black phase have been secured. The cranial
characters place it with douglasi in a group of related but not eel
connected forms.

Specumens examined.—Total number, 22, as follows:

Oregon: Mapleton, 8; Mercer, 7; Scottsburg (2 miles east), 1 (albino);? Seaton, 6.

Thomomys monticola Group.
THOMOMYS MONTICOLA MONTICOLA ALLEN
CALIFORNIA MOUNTAIN POCKET GOPHER.
(PIII, figs. 9, 9’; Pl. VIII, fig. 1.)
Thomomys monticolus Allen, Bul. Am. Mus. Nat. Hist., V, 48, Apr. 28, 1893.

Type.—Collected on-Mount Tallac, California, at 7,500 feet altitude,
by W.W. Price, August 8, 1892. Typespecimen in Am. Mus. Nat. Hist.

1Collected June 26, 1897, by J. Alden Loring. Two are half-grown, the others adults.
2 Without skull and not certainly identified. EB:

122 NORTH AMERICAN FAUNA. [No. 39. -

Mstributeon.—Sierra Nevada of California and Nevada, from Lassen
Peak south to Mammoth Pass (fig. 5).

Characters.—Size small; feet and claws very slender; ears large,
thin, well formed, and pointed; skull long and slender, with anteriorly
converging lateral ridges; colors dull hazel above, buffy below; mam-
mee in 4 pairs, inguinal 2—2, pectoral 2-2. ;

Color.— Winter pelage: Upperparts dull hazel with blackish ear
patch and plumbeous nose; underparts and feet dull buffy; tail whit-
ish throughout or dusky above at base. Summer pelage: Slightly
clearer, brighter tawny. Young, somewhat paler, more yellowish
than adults.

Skull.—Long, slender, and low; rostrum slender, especially in pro-
file, abruptly arched anterior to molars; zygomata very slender ; inter-
parietal usually transversely oval, wider than long; nasals long, slen-
der, and slightly emarginate or doubly rounded at posterior tips,
reaching near tips of premaxille; bulle very small; lateral pits of
palate deep. Dentition light; upper incisors abruptly decurved at
right angles to axis of skull, not protruding beyond tip of nasals, and
distinctly grooved. | |

Measurements.—Average of 5 adult males from Emerald Bay, near
type locality: Total length, 212; tail vertebre, 70; hind foot, 27.4.
Average of 5 adult females: 209, 67, 26.4. Skull (of type, from original
description): Basilar length, 34; greatest breadth, 22; interorbital
breadth, 6; nasals, 14. Skull (of topotype, ¢ad.):! Basal length, 32;
nasals, 12.5; zygomatic breadth, 20; mastoid breadth, 17; interor-
bital breadth, 6.2; alveolar length of upper molar series, 7. |

Remarks.—While most nearly related to the fuscus and fossor
groups, monticola and the other subspecies, mazama, pinetorum, nasicus,
and helleri, form a well-defined group. The typical subspecies appears
to overlap the range of fisheri in Sierra Valley and actually to meet
the ranges of other surrounding forms. It is mainly a Canadian Zone
species, but locally extends into the Hudsonian and to a lesser extent
down into the Transition Zone.

Specimens excamined.—Total number, 360, as follows:

California: American River (head of South Fork), 2; Aspen Meadows, 2; Big
Trees, 3; Bloods, 1; Blue Canyon, 8; Blue Lake, 3; Buck’s Ranch, 2; Burney
(12 miles west), 1; Calaveras Big Trees, 3; Carberry Ranch, 8; Cisco (Placer
County), 25; Clouds Rest (near, meadow above Little Yosemite), 2; Donner,
21; Donner Lake, 3; Echo, 16; Emerald Bay, 24; Fallen Leaf Lake, 9; Gard-
ners, 1; Gates Creek, 1; Greenville (8 miles west), 2; Hat Creek (head), 1;
Heather Lake (Eldorado County), 1; Hermit Valley, 2; Hope Valley, 7; Inde-
pendence Lake, 18; Independence Lake (pass west of), 1; Lassen Peak (north
slope), 9; Lincoln Creek (Sierra County), 1; Little Yosemite, 2; Long Valley
(Willow Ranch), 1; McCloud (12 miles northeast), 1; McKinneys, 13; Marklee-
ville, 11; Milford, 4; Mill Creek (Mount Lassen), 2: Mokelumne River (head),
2: Mono Pass, 3; Mount Conness, 4; Mount Dana, 8; Mount Lyell, 1; Mount

1No. 100274, U. S. Nat. Mus., from Camp Agassiz, on Mount Tallac.

1915. ] THOMOMYS MONTICOLA GROUP. Ae

Shasta, 26; Mount Tallac, 36; Mount Unicorn, 3; Pacific Creek, 1; Pine City

(or Mammoth), 3; Pine Creek, 1; Pyramid Peak, 1; Robbins Creek (Lassen

County), 1; San Joaquin River (head), 1; Sierra Valley, 9; Silver Lake, 8;

Slippery Ford, 1; Sonora Pass, 4; Tallac, 1; Tenaya Lake, 2; Tuolumne

Meadows, 20; Warner Creek (Mount Lassen), 1; West Walker River, 2.
Nevada: Carson, 1; Glenbrook, 7; Sugar Loaf (Douglas County), 2.

THOMOMYS MONTICOLA MAZAMA Merriam.
MazaMA PocKET GOPHER.
(P1. VIII, fig. 2.)
Thomomys mazama Merriam, Proc. Biol. Soc. Washington, XI, 214, July 15, 1897.

Type.—Collected at Anna Creek (near Crater Lake), Mount
Mazama, Klamath County, Oregon, at 6,000 feet altitude, by
Edward A. Preble, September 3, 1896. Type specimen in U.S. Nat.
Mus., Biological Survey collection.

Mstribution.—Cascade and Siskiyou Mountains, Oreg., south in
California to the Trinity Mountains (fig. 5).

Characters—About the size of monticola, but darker and richer
colored; ears nearly as large, and bulle larger and more rounded;
rostrum shorter, with less slender nasals; zygomatic arches more
abruptly spreading; mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color—Summer pelage: Upperparts bright russet-brown; ear
patch blackish; nose plumbeous; underparts rich buff or ochraceous;
feet and tail whitish, tail usually gray above at base. Winter pelage
unknown. Young slightly paler than adults.
~ Skull—Not so long and slender as in monticola; nasals sharply
emarginate posteriorly and more spreading anteriorly; antérior base
of zygoma thin and prolonged against premaxilla. Dentition slightly
heavier than in monticola.

Measurements —Average of 4 topotypes (gad.): Total. length,
209; tail vertebra, 66; hind foot, 28. Average of 4 adult females:
202, 66, 28.7. Skull (of type, gad.): Basal length, 35; nasals, 13;
zygomatic breadth, 21; mastoid breadth, 17; interorbital breadth,
6.5; alveolar length of upper molar series, 7.5.

Remarks.—From monticola and pinetorum, mazama is readily dis-
tinguished by its shorter, wider, emarginate nasals and larger bull;
from the douglasi group, by smaller size, slenderer skull, lighter den-
tition, V-shaped instead of U-shaped pterygoids, and deeper lateral
pits of palate; from quadratus, by larger ears, slenderer skull, and
emarginate nasals. It occupies the Canadian Zone and to some
~ extent the Transition also.

Specumens examined.—Total number, 294, as follows:

California: Bear Creek (head of, Trinity County, at 6,400 feet altitude), 62; Bes-
wick (6 miles southwest, 6,000 feet), 1; Castle Lake (Siskiyou County,
5,400 feet), 8; Coffee Creek (north fork, Trinity County, 4,500 feet), 15;
Grizzly Creek (Trinity County, 6,000 feet), 2; Jackson Lake (Siskiyou
County), 13; Picard (Elgin Ranch), 1; Rush Creek (head of, Siskiyou County,

124 NORTH AMERICAN FAUNA. [NO. 39. -

6,400 feet), 63; Salmon Creek Divide (Siskiyou County), 4; Salmon River
(south fork, 5,000 feet), 5; Scott River (6 miles northwest of Callahans), 8;
Siskiyou Mountains (6,000-7,000 feet on White Mountain and Craggy Peak
and in Studhorse Canyon), 22; Taylor Fork of Salmon River (5,000 feet), 2;
Trinity Mountains (west end, 5,700 feet), 5.

Oregon: Anna Creek (Mount Mazama), 3; Ashland Peak, 2; Crater Lake (Mount
Mazama), 5; Diamond Lake, 7; Fort Klamath, 13; Grizzly Peak (near Ash-
land), 1; McKenzie Bridge, 19; Mount Hood (west slope, 6,000 feet), 5;
Mount Hood (Summit House at southern base), 5; Prospect, 1; Siskiyou
Mountains, 2; Sisters (north base), 3; Three Sisters (town), 17.

THOMOMYS MONTICOLA PINETORUM Merriam.
YELLOW PINE PocKEeT GOPHER.
(P1. VII, fig. 3.)

Thomomys monticola pinetorum Merriam, N. Am. Fauna No. 16, 97, Oct. 28, 1899.
Thomomys monticola premaxillaris Grinnell, Univ. of Cal., Pub. Zool., XII, 312, Nov.

21, 1914. Collected on South Yolla Bolly Mountain, at 7,500 feet altitude, by

G. Ferris, Aug. 6, 1913. Type specimen in Mus. Vert. Zool., Univ. of California.

Type.—Collected at Sisson, west base of Mount Shasta, California,
by R. T. Fisher, September 4, 1898. Type specimen in U. 8S. Nat.
Mus., Biological Survey collection.

Mstribution.—Mountains along west side of Sacramento Valley,
Cal., from Sisson south to South Yolla Bolly Mountain (fig. 5).

Characters.—Very similar to monticola, but upperparts brighter,
more golden brown; nose and cheeks conspicuously gray.

Color.—Summer pelage: Upperparts yellow-hazel, such as is seen
in yellow-pine bark, but never in color keys; ear patch blackish;
nose andecheeks conspicuously plumbeous or grayish, often extend-.
ing back to or beyond ears; underparts buffy; tail and feet gray or
whitish. Winter pelage: Shghtly duller than in summer. Young,
paler than adults. :

Skull.—Similar to that of monticola, but with nasals more con-
spicuously emarginate at posterior tips, and falling 1 to 2 mm. short
of tips of premaxille; interparietal smaller and more rounded, and
lateral ridges more nearly parallel in adults.

Measurements.—Type (gad.): Total length, 210; tail vertebra,
76; hind foot, 28. Average of 5 topotypes (¢ ad.): 200, 73, 27.6.
Skull (of type, ¢ ad.): Basal length, 33; nasals, 13; zygomatic
breadth, 20; mastoid breadth, 18; interorbital breadth, 6; alveolar
length of upper molar series, 7.

Remarks.—In series of specimens from South Yolla Bolly Moun-
tain ! and from the head of Grindstone Creek, still farther south, the

1 Thomomys monticola premaxillaris Grinnell differs from pinetorum in a slight accentuation of the same
characters that separate that form from monticola. The premaxille extend 2 mm. back of the nasals in
the type of pinetorum, 24 mm. in the type of premazillaris, while in monticola they are approximately even
with it. The nasals in premazillaris are decidedly narrower and more deeply emarginate than in monticola
but very slightly more so than in pinetorum. The types of premazillaris and pinetorum are indistinguish-
able in coloration. To recognize premazillaris would seem to me to be splitting one rather poor subspecies
in two.

1915.] THOMOMYS MONTICOLA GROUP. 125

characters of pinetorum are slightly accentuated, but not sufficiently
for further subdivision. Specimens from Wagon Camp are to some
extent intermediate between pinetorum and monticola, while from
higher up Mount Shasta they are typical of monticola. Others from
half a mile west of Sisson distinctly approach mazama. Apparently
pinetorum is a Transition Zone form extending from Sisson south
through the mountains along the west side of the Sacramento Valley.

Specomens examined. =Total number, 54, as follows:
California: Grindstone Creek (head), 5; Sisson, 9; South Yolla Bolly Mountain, 40.

THOMOMYS MONTICOLA NASICUS Merriam.

DESCHUTES POCKET GOPHER.
(PL. VILL, fig. 4.)
Thomomys nasicus Merriam, Proc. Biol. Soc. Washington, XI, 216, July 16, 1897.

Type.—Collected at Farewell Bend, Deschutes River, Crook
County, Oregon (west of Prineville), by Edward A. Preble, August 4,
1896. Type specimen in U. S. Nat. Mus., Biological Survey collec-
tion.

_ Distribution.—West-central Oregon (east of the Cascades), from
Farewell Bend, Deschutes River, south to the Yamsey Mountains
(fig. 5).

Characters.—Size and proportions of monticola; ears large and con-
spicuous; color slightly lighter and brighter; skull long and narrow,
with long spreading nasals.

Color—Summer pelage: Upperparts bright yellowish hazel with
plumbeous nose and ear patches; underparts rich buff; feet, most of
tail and chin usually whitish. Winter pelage: Duller hazel. Young,
paler and duller colored.

Skull.—Long, with narrow braincase and long rostrum; nasals long
and conspicuously widened anteriorly, almost spoon-shaped in some
individuals, sharply emarginate at posterior tip; interparietal small
and transversely oval; bulle rather small.

Measurements.—Type (& ad.): Total length, 214; tail vertebre, 69;
hind foot, 27. Skull (of type): Basal length, 34.7; nasals, 15.4;
zygomatic breadth, 21.5; mastoid breadth, 16.8; alveolar length of
upper molar series, 7.

Remarks.—The subspecies nasicus seems to be a well-marked form
of the monticola group, with skull characters placing it nearer monti-
cola (from which it is widely separated geographically), than to its
nearest neighbor, mazama, while in color it is very close to pinetorum,
with which it also agrees in a Transition Zone range. Its relationship
can best be shown by placing it as a subspecies of monticola.

Specvmens examined.—Total number, 49, as follows:

Oregon: Bend, 12; Deschutes River (Fareweli Bend), 2; Deschutes River
(mouth of Davis Creek), 8; Fort Klamath, 1; Lapine, 12; Little Meadows, 1;
Paulina Lake, 6; Pengra, 2. Yamsey Mountains, 9.

126 NORTH AMERICAN FAUNA. [NO. 39. .

THOMOMYS MONTICOLA HELLERI Ex niort.
HELLER PocKET GOPHER.

Thomomys helleri Elliot, Field Columb. Mus., zool. ser. III, 165, 1903.

Type.—Collected at Goldbeach, mouth of Rogue River, Oregon,
by E. Heller, in 1901. No type designated. Type series in Field
Mus. Nat. Hist.

Distribution.—Coast region of southwestern Oregon (fig. 5).

Characters.—About the size of mazama, but with darker richer color- —
ation and slenderer skull. eae

Color.—October pelage: Upperparts dull chestnut or mars brown;
sides and underparts becoming ochraceous; ear patches intense black;
nose and face blackish, rarely a trace of white on lips; feet mottled or
whitish; tip of tail usually white. Young of the year, duller colored.

Skull.—Similar to that of mazama but with narrower rostrum and
nasals, smaller bulls, and shorter pterygoids, with wider intervening
fossa. :

Measurements.—Average of 4 male topotypes: Total length, 203;
tail vertebrae, 55; hind foot, 29. Average of 4 female typotypes: 195,
57, 27.2. Skull (of topotype ¢ ad.):* Basal length, 34; nasals, 14.3;
zygomatic breadth, 22; mastoid breadth, 18.6; alveolar length of
upper molar series, 7.5.

Remarks.—These gophers are abundant on the flats on both sides
of the mouth of Rogue River but are not generally distributed on the
coast nor back in the timbered interior. They form a partially iso-
lated colony with their nearest relatives mazama on the open moun-
tain ridges of the Coast Ranges and Cascades. Their group relation-—
ship is so clearly with mazama and monticola that it seems best to
place them as a subspecies of monticola.

Specimens ecamined.—Total number, 19, as follows:

Oregon: Goldbeach, 11; Wedderburn (north side of river), 8.

Thomomys fuscus Group.
THOMOMYS FUSCUS FUSCUS Merriam.

Brown PocKET GOPHER.
(P1. VIII, figs. 13, 14; text fig. 4.)

Thomomys clusius fuscus Merriam, N. Am. Fauna No. 5, 69, July, 30, 1891.
Type.—Collected on mountains at head of Big Lost River, Idaho,
at about 8,000 feet altitude, by B. H. Dutcher, September 23, 1890.
Type specimen in U. S. Nat. Mus., Biological Survey collection.
Distribution.—Southeastern British Columbia, greater part of
northern and central Idaho and western Montana, northwestern
Wyoming, and parts of eastern Washington and Oregon (fig. 5).

1No. 166949, U. S. Nat. Mus.

1915.] THOMOMYS FUSCUS GROUP. 1a

Characters.—Size rather small; feet slender; ears small and pointed;
color light brown; skull slender, with large bulle; dentition light;
mamme in 4 pairs, inguinal 2-2, pectoral 2-2.

Color.—Summer pelage: Upperparts light brownish or dull walnut-
brown, not so dark as in fossor nor so bright as in monticola or mazama;
ear patch black or blackish; nose plumbeous; underparts buffy; feet
and tail soiled whitish or. buffy; tail usually gray above at base.
Winter pelage: Slightly duller than summer. Young, paler than
adults.

Skull—tLight and slender with short rostrum and anteriorly con-
verging lateral ridges; interparietal wide and oval; nasals short and
truncate posteriorly, reaching nearly to tips of premaxille; bulle
full and rounded; pterygoids long and narrowly V-shaped; skulls
of males and females very similar. Dentition light; upper incisors
abruptly decurved and distinctly grooved.

Measurements.—Average of 3 topotypes (¢ ad): Total length,
203; tail vertebre, 70; hind foot, 27. Average of 5 adult females:
205, 70, 27. Skull (of type, 2 ad.): Basal length, 33; nasals, 12.4;
zygomatic breadth, 20; mastoid breadth, 17; mterorbital breadth,
6; alveolar length of upper molar series, 7.

Remarks.—This form is fairly typical in central Idaho, western
Montana, and in the Blue Mountain region of Oregon. In northern
Idaho, eastern Washington, and southern British Columbia it becomes
noticeably darker, but retains more nearly the skull characters of
fuscus. West of the Columbia River it varies greatly with almost
every locality, but these local forms can be placed under fuscus better
than with quadratus or myops, both of which they suggest.

Specumens examined.—Votal number, 373, as follows:

British Columbia: Ashcroft, 1; Cascade, 6; Ducks, 5; Glacier, 1; Kamloops, 2;

Midway, 6; Monashee Ridge (Gold Range, at 4,000 feet altitude, 5; at 3,620

feet, 4), 9; Myers Creek, 1; Nelson, 9; Okanogan, 10; Pend Oreille River,

11; Salmon River, 2; Shuswap, 20; Skagit Valley, 1; Trail, 1; Wards

Ferry, 1.

Idaho: Ashton, 2; Big Lost River (head), 2; Blackfoot (west of river), 1; Blue
Lake, 12; Coeur d’Alene, 4; Freedom (near mouth of South Fork Salmon
River), 1; Hoodoo Valley, 1; Lemhi Indian Agency, 2; Lerdo, 1; Lost River
Mountains, 5; Midvale, 7; Mission, 1; Mount Carlton, 3; New Meadows, 3;

}

Salmon River Mountains, 11; Sawtooth Lake, 3; Seven Devils Mountains, 2;
South Fork Ranch, 1; Teton Canyon, 1; Van Wyck, 1.

Montana: Beartooth Mountains, 3; Benton (west of), 1; Corvallis, 2; Flathead
Lake, 4; Fort Ellis, 2; Little Bitterroot Creek, 1; Lo Lo, 1; Midvale, 2; Ross
Fork, 1; St. Marys Lake, 3; Stevensville, 2; Summit (west of Blackfoot
Station), 6; Thompson Falls, 8; Tobacco Plains, 10.

Oregon: Antelope, 2; Anthony, 14; Bingham Prairie, 1; Blue Mountains (10 miles
north of Harney), 1; Elgin, 1; Huntington, 1; Wallowa Mountains, 8.

128 NORTH AMERICAN FAUNA. [No. 39.

Washington: Blue Creek (13 miles east of Walla Walla), 3; Chelan, 2; Cheney,
1; Colfax, 1; Columbia County (25 miles southeast of Dayton and Humpeg
Falls), 2; Colville, 6; Davenport, 1; Douglas, 2; Easton, 9; Fort Spokane, 1;
Garfield, 1; Loon Lake, 2; Mabton, 3; Marcus, 5; Marshall, 11; Natches
River, 1; North Yakima, 22; Orondo, 1; Pullman, 11; Rock Lake, 2; Spo-
kane Bridge, 3; Spokane Falls, 15; Waterville, 1; Wenatchee, 8.

Wyoming: Black Mountains, 4; Moran, 4; Pahaska, 4; Pahaska Tepee. 12;
Teton Mountains, 6; Teton Pass, 4; Yellowstone Lake, 1.

THOMOMYS FUSCUS SATURATUS Batrey,

Ca@urR D’ALENE POCKET GOPHER.
(P1. VIII, fig. 16.)

Thomomys fuscus saturatus Bailey, Proc. Biol. Soc. Washington, XXVII, 117, July
10, 1914. ;

Type.—Collected at Silver (near Saltese) in the western corner of
Missoula County, Montana, at 4,300 feet altitude, in the Coeur d’ Alene
Mountains, by Clark P. Streator, June 20, 1891. Type specimen in
U.S. Nat. Mus., Biological Survey collection.

Distribution.—Higher parts of the Coeur d’Alene Mountains in
Idaho and Montana (fig. 5).

Characters.—Considerably larger than fuscus, with darker, richer
coloration and less tendency to white markings below; skull long and
high, with relatively narrower braincase and wider nasals.

Color.—Summer pelage: Upperparts dark rich hazel, becoming yel-
lowish on sides, and this shading into a buffy wash on underparts;
ear patches black; nose plumbeous; feet and tail buffy gray; rarely
a trace of white on chin or throat. Only June and August specimens
seen. The young are a shade paler than adults. |

Skull.—Long and narrow with high, truncate occiput; lateral
ridges prominent and nearly parallel; nasals widened anteriorly and
emarginate posteriorly; premaxille very narrow posteriorly; bulle
wide and flattened; basioccipital relatively wide between bulle.

Measurements.—Type (8 ad.): Total length, 225; tail vertebre,
77; hind foot, 30. Average of 3 adult males: 216, 72, 29. Average
of 5 females: 211, 75, 28.7. Skull (of type): Basal length, 33.2;
nasals, 13.7; zygomatic breadth, 21.4; mastoid breadth, 17; alveolar
length of upper molar series, 7.

Remarks.—This seems to be a well-marked but rather local form
of the fuscus group occupying the higher hemlock-covered part of the
Cour d’Alene Mountains. Specimens from Prospect Creek, lower
down on the east slope, show intermediate characters approaching
fuscus while others from Thompson Falls, at the east base of the
mountains are almost typical of fuscus.

Specumens excamined.—Total number, 26, as follows:

Idaho: Mullan, 3.
Montana: Prospect Creek, 6; Silver, 16; Thompson Pass, 1.

1915.] THOMOMYS FUSCUS GROUP. 129

THOMOMYS FUSCUS LORINGI Battey.

ALBERTA PocKET GOPHER.
(PL. VIII, fig. 9.)
Thomomys fuscus loringi Bailey, Proc. Biol. Soc. Washington, X XVII, 118, July 10,
1914.

Type.—Collected at South Edmonton, Alberta (exact locality not
given), by J. Alden Loring, September 23, 1894. Type specimen in
U.S. Nat. Mus., Biological Survey collection.

Distribution.—Known only from Edmonton and Moose Mountain,
Alberta (fig. 5).

Characters.—Similar to fuscus, but shghtly larger and duller colored,
with relatively heavier dentition, and small, circular interparietal;
ears small and pointed.

Color.— Winter pelage (September 23): Upperparts dull russet-
brown, becoming rich buff on sides; nose slaty gray; small ear patch
black; underparts rich buffy over plumbeous; chin and small spot
on breast white; feet soiled whitish; tail pale buffy. Swmmer
pelage (in two July 2 specimens from Moose Mountains): Similar but
slightly brighter russet.

Skull.—Long and slender, with narrow braincase and small, nearly
quadrate zygomatic arches, and almost circular interparietal; incisors

less abruptly decurved than in fuscus; molars heavier; bulle about the

same; basioccipital somewhat triangular; nasals narrow and trun-
cate at posterior tips.

Measurements.—Type (¢ ad.): Total length, 199; tail vertebre,
47; hind foot, 26.5. Skull (of type): Basal length, 34; nasals, 12.8;
zygomatic breadth, 20.5; mastoid breadth, 18; interorbital breadth,
6; alveolar length of upper molar series, 8.

Remarks.—Although represented by only one specimen from the type
locality, which is well out on the plains, this is evidently a form of the
mountain species fuscus from farther south and west. Two skins in
the Victoria Memorial Museum from Moose Mountain are apparently
the same, while specimens from St. Marys Lake, Glacier Park, Mont.,
show a slight tendency toward the characters represented by this
form, but are much nearer typical fuscus. Specimens from Shuswap,
British Columbia, much farther west, show none of the characters of
this form. There are no specimens of gophers available from the
mountains of western Alberta, between Moose Mountain and the
Montana line, but in this strip of country no collections have been
made, and probably gophers occur along the eastern slope of the
mountains from the United States boundary north to the Edmonton
region.

Loring collected specimens of talpoides about Edmonton, but these
large, dark-colored, large-footed gophers are quite distinct from the

98121°—15—_9

130 NORTH AMERICAN FAUNA. ENO. 39,

present little brown form. There were no notes to indicate whether
the two species occupy the same or separate areas.

In the type specimen the upper molars are abnormally close to-
gether at the anterior base, but this is probably not characteristic of
the species. Unfortunately there are no skulls for the Moose Moun-
tain specimens, but the skins match the type in color. They are
both breeding females collected July 2 by W. Spreadborough and are
particularly interesting in showing fully developed sets of 12 mamme
each; inguinal 2—2, abdominal 2-2, pectoral 2-2. This is the formula
for the talpoides and not for the fuscus group. The status of the form
is at present very uncertain and will be until more collecting is done
in the type region.

Specumens ecamined.—Total number, 3, as follows: |

Alberta: South Edmonton, 1; Moose Mountain, 2 (skins only).

THOMOMYS FUSCUS MYOPS Merriam
LirrLE-HEADED PocKET GOPHER.
(Pl. VIII, fig. 15.) .

Thomomys myops Merriam, Proc. Biol. Soc. Washington, XIV, 112, July 19, 1901.

Type.—Collected at Conconully, Okanogan County, Washington, by
J. Alden Loring, September 11, 1897. Type specimen in U. S. Nat.
Mus., Biological Survey collection.

Distributton.—Known only from type locality (fig. 5).

Characters.—Size smaller than fuscus but with very similar colora-
tion and the same minute ears; skull small; tips of premaxille even
with posterior tips of nasals.

Color.—September pelage: Upperparts light brownish or dull ad nti
brown; ear patch blackish; nose plumbeous; underparts and base of
tail batty ; feet and tip of onl whitish; chin usually white. Young,
paler, more grayish.

Skull.—Smali, but of about the form and proportions of that of
fuscus; nasals and premaxille terminating posteriorly on an approxi-
mately even line; interparictal transversely elongated; bulle small;
anterior shaft of basioccipital relatively wide; pterygoids wide apart,
with edges flattened at base.

Measurements.—Average of 2 topotypes (¢ ad.): Total length,
182; tail vertebre, 57; hind foot, 25. Average of 5 females: 186, 59,
24.4. Skull (of type, @ ad.): Basal length, 31.8; nasals, 12.5; zygo-
matic breadth, 20; mastoid breadth, 17.5; alveolar length of upper
molar series, 7.5.

Remarks.—Specimens from the type locality, Conconully, on Con-
conully Creek, a western affluent of the Okanogan River, are very uni-
form in characters and so peculiar that they were originally described
with full specific rank. They evidently belong to the fuscus group,

1915.] THOMOMYS FUSCUS GROUP. 3H

however, which shows great variation in that general region, and
probably mark only a local valley form partly cut off from the general
range of the species.

Specimens examined.—Seven, from type locality.

THOMOMYS HESPERUS Merriam.
West Coast Pocket GOPHER.
(Pl. VIII, fig. 10.)
Thomomys hesperus Merriam, Proc. Biol. Soc. Washington, XIV, 116, July 19, 1901.

Type.—Collected at Tillamook, Tillamook County, Oregon, by J.
Ellis McLellan, November 9, 1894. Type specimen in U. S. Nat.
Mus., Biological Survey collection.

Distribution.—Coast region of northwestern Oregon (fig. 5).

Characters.—Size very small, with the small, pointed ears of the
fuscus group; color dark rich auburn; skull small, short, and rounded.

Color.—Upperparts bright rich auburn, brighter than in helleri or
melanops, darker and richer than in mazama; nose and cheeks dusky;
large ear patch black; underparts lighter, more ochraceous; lining of
cheek pouches white; feet and tip of tail whitish.

Skull.—Short; brainease rather wide; rostrum short and wide;
posterior tip of nasals emarginate; bullae very small, short, and
rounded; basioccipital short and wide; pterygoids low and wide
apart; molars very small but app ey of about the same propor-
tions as those of fuscus.

Measurements.—Type (¢@ ad.): Total length, 175; tail vertebrae,
54; hind foot, 24. Old male from Chintimini Moaneaine: 210, 60, 27.
Shull (of oe Basal length, 27.3; nasals, 10.7; zygomatic breadth,
18; mastoid breadth, 15; interorbital breadth, 6; alveolar length
of upper molar series, 6. Skull of old male from Chintimini Moun-
tains: 30, 12.4, 20, 16.5, 6, 7.

Remarks.—This is evidently a little, dark form of the fuscus group,
which seems to be isolated below the mouth of the Columbia River.
In color it strongly resembles melanops of the Olympic Mountains
and hellera from farther south, but the tiny ears and general skull char-
acters seem to place it in the fuscus group. Until a good series of
specimens of adult males is obtained its true position can not be fully
determined. An old male from the top of Chintimini Mountains
(near Corvallis) seems to be the same, but may not be wholly typical.

Specomens examined.—Total number, 6, as follows:

Oregon: Chintimini Mountains (at 4,000 feet altitude), 1 old male; Tillamook, 5
(no adult males).

Fie:

PuaTeE II.
[Natural size; allfrom U.S. Nat. Mus., Biological Survey collection.]

Thomomys bulbworus, 3, Beaverton, Oreg. (No. 57337.)

Thomomys townsend townsendi, 3, Nampa, Idaho. (No. 181196.)

Thomomys bottx botte, 3, Point Reyes, Cal. (No. 134989.)

Thomomys perpallidus aureus, 3, topotype, Bluff, Utah. (No. 57166.)

Thomomys talpoides rufescens, &, near Ft. Clark, N. Dak. (No. 161381.)

Thomomys talpoides clusius, 8, topotype, Bridger Pass, Wyo. (No. 25534.)

Thomomys idahoensis, 3, type, Birch Creek, Idaho. (No. 30900.)

Thomomys fossor, &, Lake City, Colo. (No. 48190.)

Thomomys monticola monticola, 3, topotype, Tallac, Cal. (No. 100660.)
Dorsal and lateral view.

Thomomys douglasi douglasi, 3, topotype, Vancouver, Wash. (No. 65928.)

. Thomomys bottx leucodon, 3, Ashland, Oreg. (No. 203671.) Lateral view.
. Thomomys ocius, 3, type, Fort Bridger, Wyo. (No. 25586.)

Thomomys umbrinus umbrinus, 3, Boca del Monte, Vera Cruz. (No. 64093.)
122 .

North American Fauna No. 39, U.S. Dept. Agr. Biological Survey. PLATE Il.

B2035-122

SKULLS OF THOMOMYS.

. bulbivorus. . T. t. rufescens. . T. d. douglasi.
. t. townsendi. . T. t. clusius. . T. b. leucodon.
. b. botte. . T. idahoensis. 2. T. ocius.
. p. aureus. . T. fossor. 3. T. u. umbrinus.
; h and 91, T. m. monticola.
Se 110)

Fic.

COON nmoark WwW

PratE IT:
[Natural size; all from U.S. Nat. Mus., Biological Survey collection.]

Thomomys bulbivorus, ¢, Salem, Oreg. (No. 57328.)

Thomomys townsend: townsendi, 3, type, Nampa, Idaho. (No. 181196.)
Thomomys townsend: nevadensis, 3, type, Austin, Nev. (No. 32413.)
Thomomys bottx bottz2, ¢, Point Reyes, Cal. (No. 134989.)

Thomomys bottx leucodon, 3. Ashland, Oreg. (No. 203671.)

Thomomys bottx laticeps, 3, topotype, Arcata, Cal. (No. 58454.)

Thomomys botte angularis, 3, type, Los Banos, Cal. (No. 58123.)
Thomomys bottx pallescens, 3, topotype, Grapeland, Cal. (No. 94013.)
Thomomysaltivallis, 3 , topotype, San Bernardino Mountains, Cal. (No. 89954.)

an Fauna No. 39, U.S. Dept. Agr. Biologi rvey. PLATE Ill.

B2036-122
SKULLS OF THOMOMYS.
. bulbivorus. 4, T. b. botte. 7. T. b. angularis.

1p AR
2. T. t. townsendi. 5. T. b. leucodon. 8. T. b. pallescens.
3. T. t. nevadensis. 6. T. b. laticeps. . T. altivallis.

Fic.

PLATE<TV.
[Natural size; allfrom U.S. Nat. Mus., Biological Survey collection.]

Thomomys bottx pascalis, 8 , type, Fresno, Cal. (No. 44702.)

Thomomys botte minor, 3, type, Fort Bragg, Cal. (No. 146463.)

Thomomys botte navus, &, type, Red Bluff, Cal. (No. 57791.)

Thomomys fulvus texensis, & , type, Davis Mountains, Tex. (No. 31408.)

Thomomys fulvus pervagus, 3, type, Espanola, N. Mex. (No. 58293.)

Thomomys perpallidus apache, 3, type, La Jara Lake, N. Mex. (No. 135366.)

Thomomys sinalox, 3, type, Altata, Sinaloa. (No. 96745.)

Thomomys bottex alticolus, 3, topotype, Victoria Mountains, Lower California.
(No. 74208. )

Thomomys bottx russeolus, 3 , type, San Angel, Lower California. (No. 139920.)

North American Fauna No. 39, U.S. Dept. Agr.

Biological Survey. PLATE IV.

B2037-122
SKULLS OF THOMOMYS.

4. T. f. texensis. T. sinaloee.
5. T. f. pervagus,

T. b. alticolus.
6. T. p. apache. T. b. russeolus.

- pascalis.
. Toinor.
. navus,

Q

| Oo eel

PLATE? Vi
[Natural size; all from U.S. Nat. Mus., Biological Survey collection, except fig. 2.]

Thomomys perpallidus albatus, 3, topotype, Fort Yuma, Cal. (No. 99528.)

Thomomys perpallidus perpallidus, 3, topotype, Palm Springs, Cal. (No. 4,
collection of Dr. William Bebb.)

Thomomys perpallidus perpes, 3, type, Lone Pine, Cal. (No. 32503.)

Thomomys operarius, &, type, Keeler, Cal. (No. 32466.)

Thomomys perpallidus aureus, 3, topotype, Bluff, Utah. (No. 57155.)

Thomomys perpallidus canus, &, type, Deep Hole, Nev. (No. 78365.)

. Thomomys latirostris, 3, type, Painted Desert, Ariz. (No. 24914.)

Thomomys cabezone, 3, type, Cabezon, Cal. (No. 53989.)

. Thomomys alpinus awahnee, 2, type, Yosemite Valley, Cal. (No. 133076.)
. Thomomys botte mewa, 3, type, Raymond, Cal. (No. 133183.)
. Thomomys botte anitxe, 3, topotype, Santa Anita, Lower California. (No.

146831.)

2. Thomomys magdalenx, 3, type, Magdalena Island, Lower California. (No.

146832.)

3. Thomomys alpinus alpinus, 3, type, Mount Whitney, Cal. (No. 42512.)
. Thomomys jacinteus, 3, San Jacinto Mountains (Tahquitz Valley), Cal. (No.

126469. )

5. Thomomys neglectus, 3, type, San Antonio Mountains, Cal. (No. 137869.)
. Thomomys bottxe nigricans, 3, topotype, Witch Creek, Cal. (No. 148685.)

North American Fauna No. 39, U. S. Dept. Agr. Biological Survey. PLATE V.

\ es
"]

ene:

eBqt) (oe

/

el
f

a
oy

pee7gh/

SKULLS OF THOMOMYS.

. T. p. albatus. 6. T. p. canus. 2. T. magdalene.
. T. p. perpallidus. T. latirostris. 3. T. a. alpinus.
3. T. p. perpes. 8. T. cabezone. . T. jacinteus.
. T. p. operarius. T. a. awahnee. 5. T. neglectus.
. T. p. aureus. T. b. mewa. 3. T. b. nigricans.
T. b. anite.

Fic.

PA Vile
{Natural size; all from U.S. Nat. Mus., Biological Survey collection.]

Thomomys fulous fulvus, 3, Flagstaff, Ariz. (No. 169603.)

Thomomys cervinus, &, topotype, Phoenix, Ariz. (No. 23707.)

Thomomys fulvus toltecus, 8, typical, Colonia Diaz, Chihuahua. (No. 132468.)
Thomomys mearnsi, 3, type, Animas Valley, N. Mex. (No. 157008.)
Thomomys fulvus intermedius, 3, Fort Huachuca, Ariz. (No. 46334.)
Thomomys fulvus desertorum, 3, type, Mud Spring, Ariz. (No. 6082.)
Thomomys baileyi, 3, type, Sierra Blanca, Tex. (No. 25159.)

Thomomys lachuguilla, 3, type, El Paso, Tex. (No. 110336.)

Thomomys perditus, 3, type, Lampazos, Nuevo Leon. (No. 32998.)
Thomomys goldmani, 8, type, Mapimi, Durango. (No. 58075.)

. Thomomys nelsoni, 3, type, Parral, Chihuahua. (No. 96451.)

2. Thomomys sheldomi, 3, type, Santa Teresa, Tepic. (No. 90819.) |
. Thomomys umbrinus umbrinus, &, Boca Del Monte, Vera Cruz. (No. 64093.)
. Thomomys umbrinus peregrinus, 2, type, Salazar, Mexico. (No. 50130.)

5. Thomomys umbrinus orizabe, 2, type, Mount Orizaba, Puebla. (No. 53616.)
. Thomomys atrovarius, 3, typical, Mazatlan, Sinaloa. (No. 96750.)

North American Fauna No. 39, U. S. Dept. Agr.

Biological Survey. PLATE VI.

SKULLS OF THOMOMYS.

. T: f. fulvus.
2. T. cervinus.
. T. f. toltecus.
. T. mearnsi.
5. T. f. intermedius.

. f. desertorum.
. baileyi.

. lachuguilla.

. perditus.

. goldmani.

. nelsoni.

. Sheldoni.

'. u. umbrinus.
. u. peregrinus.
.u. orizabe.

‘. atrovarius.

Fig. 1.

eo

Pruate VII.
{Natural size; a!l from U. 8. Nat. Mus., Biological Survey collection.]

Thomomys talpoides rufescens, 3, near Fort Clark, N. Dak. (No. 161381.)
Thomomys talpoides bullatus, 3, type, Powderville, Mont. (No. 55159.)
Thomomys talpoides nebulosus, &, type, Sand Creek Canyon, Wyo. (No.
202495.)
Thomomys talpoides agrestis, 2 , type, Medano Ranch, Colo. (No. 150725.)
Thomomys talpordes clusius, &, topotype, Bridgers Pass, Wyo. (No. 25534.)
Thomomys talpoides caryi, 2 , type, Bighorn Mountains, Wyo. (No. 168946.)
Thomomys talpoides pryori, 3, type, Pryor Mountains, Mont. (No. 66469.)
Thomomys quadratus quadratus, 3, type, The Dalles, Oreg. (No. 31117.)
Thomomys ocius, 3, type, Fort Bridger, Wyo. (No. 25586.)
Thomomys idahoensis, 3, type, Birch Creek, Idaho. (No. 30900.)

. Thomomys pygmxus, 3, type, Montpelier Creek, Idaho. (No. 55271.)
2. Thomomys columbianus, 3, type, Touchet, Wash. (No. 57134.)

3. Thomomys fossor, 3, Lake City, Colo. (No. 48190.)

. Thomomys uinta, 3, type, Uinta Mountains, Utah. (No. 30051.)

5. Thomomys bridgeri, 3, type, Fort Bridger, Wyo. (No. 25592.)

16.

Thomomys quadratus fisheri, 3, type, Sierra Valley, Cal. (No. 101238.)

North American Fauna No. 39, U.S. Dept. Agr. Biological Survey. PLATE VII.

B2040-122

SKULLS OF THOMOMYS.

T. t. caryi. . T. columbianus.
T. t. pryori. 3. T.fossor.
T. q. quadratus. . T. uinta.

. ocius. . T. bridgeri.

. idahoensis. . TL. q. fisheri.

. pygmeeus.

T. t. rufescens.
. T. t. bullatus.
. T. t. nebulosus.
. T. t. agrestis.
yD. te clusius:

er
Sonne

Fic.

Puate VIII.
{Natural size; all from U.S. Nat. Mus., Biological Survey collection.]

Thomomys monticola monticola, 3, topotype, Tallac, Cal. (No. 100660.)

Thomomys monticola mazama, 3, type, Anna Creek, Oreg. (No. 80502.)

Thomomys monticola pinetorum, &, type, Sisson, Cal. (No. 95152.)

Thomomys monticola nasicus, 3, type, Farewell Bend, Deschutes River, Oreg.
(No. 79815.)

Thomomys douglasi douglasi, 3 , topotype, Vancouver, Wash. (No. 65928.)

Thomomys douglast melanops, 2, type, Olympic Mountains, Wash. (No.
90630.)

Thomomys niger, 3, type, Seaton, Oreg. (No. 69407.)

Thomomys douglasi oregonus, 3, type, Oregon City, Oreg. (No. 56939.)

Thomomys fuscus loringi, &, type, Edmonton, Alberta. (No. 68746.)

Thomomys hesperus, immature ° , type. Tillamook, Oreg. (No. 69825.)

. Thomomys douglasi yelmensis, 3, type, Tenino, Wash. (No. 43777.)

. Thomomys limosus, 3, type, White Salmon, Wash. (No. 89724.)

. Thomomys fuscus fuscus, &, Lemhi, Idaho. (No. 31219.)

. Thomomys fuscus fuscus, 2, type, head of Big Lost River, Idaho. (No.

31671.)

. Thomomys fuscus myops, 2, type, Conconully, Wash. (No. 91066.)
. Thomomys fuscus saturatus, 3, type, Silver, Mont. (No. 40885.)

North American Fauna No. 39, U.S. Dept. Agr. Biological Survey. PLATE VIII.

B2041-122

SKULLS OF THOMOMYS.

. T. m. monticola. 6. T. d. melanops. 12. T.limosus.
. T. m. mazama. ie een ger: 13a Dore tuUSCUus soy
3. T.m. pinetorum. 8. T. d. oregonus. 14. T.f.fuscus, @.
. T.m. nasicus. 9. T. f. loringi. 15. T. f. myops.
. T. d. douglasi. 10. T. hesperus. 16. T. f. saturatus.
11. T. d. yelmensis.

ieee te}

ke

5s

sap"

TNE EX.

[New names in bold=face type; synonyms in italics.]}

ING

Abnormai pelage conditions, 20.
Acknowledgments, 31.
agrestis, Thomomys talpoides, 105-105.
albatus, Thomomys perpallidus, 70.
Albinism, 20.
alpinus, Thomomys alpinus, 63-64.
alticolus, Thomomys botte, 60.
altivallis, Thomomys, 62-63.
angularis, Thomomys botte, 53.
anitze, Thomomys botte, 59.
apache, Thomomys perpallidus, 75-76.
aphrastus, Thomomys, 56.
Ascomys borealis, 97.

bulbivorus, 40.

talpoides, 97.
atrogriseus, Thomomys nevadensis, 42.
atrovarius, Thomomys, 95-96.
aureus, Thomomys perpallidus, 74-75.
awahnee, Thomomys alpinus, 64-65.

B.

baileyi, Thomomys, 87-88.
borealis, Ascomys, 97.
Geomys, 96, 100.
Pseudostoma, 100.
Saccophorus, 97.
bottz, Oryctomys (Saccophorus), 45.
Thomomys botte, 45-46.
Breeding habits, 11.
bridgeri, Thomomys, 112-113.
bulbivorum, Diplostoma, 40, 45.
Pseudosioma, 40.

bulbivorus, Ascomys, 40.

Geomys, 40.

Thomomys, 40-42.

Thomomys, 45. é
bullatus, Thomomys talpoides, 101-102.

Burrows, 8.
Cc.

cabezonz, Thomomys, 76-77.

Camas rat, 40-42.

canus, Thomomys perpallidus, 73-74.
caryi, Thomomys talpoides, 103-104.
cervinus, Thomomys, 79.

Characters, cranial, 16.

general, 14.

generic, 32.
chrysonotus, Thomomys perpallidus, 71.
clusius, Thomomys talpoides, 100-101.
columbianus, Thomomys, 106-107.
Cranial characters, 16.
Cratogeomys, 7.
Cricetus talpoides, 96. -

98121°—15——1]1

‘D.

Damage by pocket gophets, 10.
Dentition, 17. :
desertorum, Thomomys fulvus, 83-84.
Destruction of pocket gophers, 14.
diaboli, Thomomys botte, 51-52.
Diplostoma bulbivorum, 40, 45.
Disposition of pocket gophers, 13.
Distribution of pocket gophers, 7, 22.
douglasi, Thomomys douglasi, 116-117.
douglasii, Geomys, 116.

Thomomys, 116.

E.

Enemies of pocket gophers, 13.
EH:

fisheri, Thomomys quadratus, 115-116.
Food of pocket gophers, 10.

pocket gophers as, 13.
fossor, Thomomys, 111-112.
fuliginosus, Geomys, 116.
jfulvus, Geomys, 80.

Thomomys fulvus, 80-82.
fuscus, Thomomys fuscus, 126-128.

G.

General characters, 14.
Generic characters, 32.
Geomys, 7.

borealis, 96, 100.

bulbivorus, 40.

bursarius, 100.

douglasii, 116.

fuliginosus, 116.

fulvus, 80.

oregonensis, 100.

talpoides, 96.

(Thomomys) talpoides, 97.

townsendii, 42, 43.

umbrinus, 89.

unisulcatus, 97.
goldmani, Thomomys, 94-95.
Gopher. See Pocket gopher.
Group relationships, 32.

18k

Habits, breeding, 11.

food, 10.

general, 8.

winter, 11.
Heavy rostrum groups, 32.
helleri, Thomomys monticola, 126.
hesperus, Thomomys, 131.
Heterogeomys, 7.

133

134 NORTH AMERICAN FAUNA.

Ile

idahoensis, Thomomys, 108-109.
infrapallidus, Thomomys bottz, 55-56.
intermedius, Thomomys fulvus, 84-85.

J.
jacinteus, Thomomys, 66-67.

K.
Key to species, 36-40.
L.

lachuguilla, Thomomys, 88-89.
laticeps, Thomomys botte, 46-47.
latirostris, Thomomys, 78-79.
leucodon, Thomomys botte, 47-49.
limosus, Thomomys, 120-121.

List of species, 34-36.

loringi, Thomomys fuscus, 129-130.

M.
Macrogeomys, 7.
magdalenze, Thomomys, 61-62.
Map (genus), 9.
Maps (species) 23, 24, 25, 26, 27, 28.
martirensis, Thomomys, 67-68.
Material examined, 30.
mazama, Thomomys monticola, 123-124.
mearnsi, Thomomys, 87.
Megascapheus (subgenus), 40.
Melanism, 21.
melanops, Thomomys douglasi, 119-120.
mewa, Thomomys botte, 50.
minor, Thomomys botte, 51.
Molt, 18.
monticola, Thomomys monticola, 121-123.
monticolus, Thomomys, 121.
myops, Thomomys fuscus, 130-131.

N.

nasicus, Thomomys monticola, 125.
Natural enemies of pocket gophers, 13.
navus, Thomomys botte, 49-50.
nebulosus, Thomomys talpoides, 102-103.
neglectus, Thomomys, 65-66.

nelsoni, Thomomys, 92-93. “
nevadensis, Thomomys townsendi, 44-45.
niger, Thomomys, 121.

nigricans, Thomomys bottz, 56-58.

oO.

ocius, Thomomys, 107-108.

operarius, Thomomys, 77-78.

oregonus, Thomomys douglasi, 117-118.
orizabe, Thomomys umbrinus, 90-91.
Orthogeomys, 7.

Oryctomys (Saccophorus) bottx, 45.

1.

pallescens, Thomomys botte, 55.
Pappogeomys, 7.
pascalis, Thomomys botte, 54.
Pelage, 15.

conditions, abnormal, 20.
perditus, Thomomys, 95.
peregrinus, Thomomys umbrinus, 91-92.
perpallidus, Thomomys perpallidus, 68-69.
perpes, Thomomys perpallidus, 72-73.

pervagus, Thomomys fulvus, 82-83.

pinetorum, Thomomys monticola, 124-125.

Platygeomys, 7.

Pocket gopher. Sce also Thomomys.
Alberta, 129-130.
Bighorn, 103-104.
black, 121.
black-and-brown, 95-96.
black-headed, 119-120.
Black Hills, 102-103.
brown, 126-128.
Cabezon, 76-77.
California, 45-46.
California Mountain, 121-123.
camas, 40-42.

Cape San Lucas, 59.
Carrizo Plain, 55-56.
Coeur d’ Alene, 128.
Colorado, 111-112.
Columbia, 106-107.
Coues, 100-101.
Dakota, 98-99.
Dalles, 114-115.
Davis Mcuntain, 85.
Deschutes, 125.
desert, 83-84.
Diablo, 51-52.
digger pine, 50.
Douglas, 116-117.
Espanola, 82-83.
fawn-colored, 79.
Fisher, 115-116.
Fort Bridger, 112-113.
Fresno, 54.
fulvous, 80-82.
Goldman, 94-95. .
Grapeland, 55.
gray, 73-74.
Green River, 107-108.
Heller, 126.
Humboldt Bay, 46-47.
Idaho, 108-109.
Jicarilla, 75-76.
Lachuguilla, 88-89.
La Puerta, 58-59.
little gray, 95.
little-headed, 130-131.
Lone Pine, 72-73.
Los Banos, 53.
Magdalena, 61-62.
Mazama, 123-124.
Mearns, 87.
Mendocino, 51.
Mexican, 91-92.
mountain-top, 84-85.
-Mount Whitney, 63-64.
Nelson, 92-93.
Nevada, 44-45.
Oregon, 117-118.
Orizaba, 90-91.
Owens Lake, 77-78.
Painted Desert, 78-79.
Palm Springs, 68-69.
~ Phoenix, 79.
prairie, 98-99.
Pryor Mountain, 104-105.
pygmy, 109-110.
red, 49-50.

{ NO. 39.

1915.1

Pocket gopher, sage, 101-102.
San Angel, 60-61.
San Bernardino Mountain, 62-63.
San Gabriel, 65-66.
San Jacinto, 66-67.
San Luis, 105-106.
San Pedro’ Martir, 67-68.
Saskatchewan, 96-98.
Sheldon, 93-94.
Sierra Blanca, 87-88.
Sierra Laguna, 60.
Sinaloa, 80.
Southern, 89-90.
Stephens, 56-58.
Toltec, 86.
Townsend, 42-43.
Uinta, 113-114.
West Coast, 131.
white, 70.
White Salmon, 120-121.
white-toothed, 47-49.
yellow, 74-75.
yellew-backed, 71.
yellow pine, 124-125.
Yelm, 118-119.
Yosemite, 64-65.
Pockets, 15.
premarillaris, Thomomys monticola, 124.
pryori, Thomomys talpoides, 104-105.
Pseudostoma borealis, 100.
bulbivorum, 40.
talpoides, 97.
puerte, Thomomys botte, 58-59.
pygmeus, Thomomys, 109-110.

Q.

quadratus, Thomomys quadratus, 114-115.

R.

rufescens, Thomomys, 98-99.
russeolus, Thomomys bottz, 60-61.

Ss.

Saccophorus (subgenus), 45.

borealis, 97.

bottz, 45.

talpoides, 96.
Sand rat, Columbia, 116-117.
saturatus, Thomomys fuscus, 128.
scapterus, Thomomys, 72.
Sexual variation, 17.
sheldoni, Thomomys, 93-94.
sinaloe, Thomomys, 80.
Slender rostrum groups, 33.
Species, list, 34-36.

maps, 23, 24, 25, 26, 27, 28.

T.
talpoides, Ascomys, 97.

Cricetus, 96.

Geomys, 96, 97.

Pseudostoma, 97.

Saccophorus, 96.

Thomomys talpoides, 96-98.
texensis, Thomomys fulvus, 85. ~
Thomomys, 32. See also Pocket gopher.

agrestis, 105-106.

INDEX.

Thomomys albatus, 70.

alpinus, 63-64.
alticolus, 69.
altivallis, 62-63.
angularis, 53.
anite, 59.
apache, 75-76.
aphrastus, 56.
atrogriseus, 42.
atrovarius, 95-96.
aureus, 74-75.
awahnee, 64-65.
baileyi, 87-88.
bottz, 45-46.
bridgeri, 112-113.
bulbivorus, 40-42.
bulbivorus, 45.
bullatus, 101-102.
cabezone, 76-77.
canus, 73-74.
caryi, 103-104.
cervinus, 79.
chrysonotus, 71.
clusius, 100-101.

columbianus, 106-107.

desertorum, 83-84.
diaboli, 51-52.
douglasi, 116-117.
fisheri, 115-116.
fossor, 111-112.
fulvus, 80-82.
fuscus, 126-128.
goldmani, 94-95.
helleri, 126.
hesperus, 131.
idahoensis, 108-109.
infrapallidus, 55-56.
intermedius, 84-85.
jacinteus, 66-67.
lachuguilla, 88-89.
laticeps, 46-47.
latirostris, 78-79.
leucodon, 47-49.
limosus, 120-121.
loringi, 129-130.
magdalene, 61-62.
martirensis, 67-68.
mazama, 123-124.
mearnsi, 87.
melanops, 119-120.
mewa, 50.

minor, 51.
monticola, 121-123.
moniicolus, 121.
myops, 130-131.
nasicus, 125.
navus, 49-50.
nebulosus, 102-103.
neglectus, 65-66.
nelsoni, 92-93.
nevadensis, 44-45.
niger, 121.
nigricans, 56-58.
ocius, 107-108.
operarius, 77-78.
oregonus, 117-118.
orizabee, 90-91.

135

136

Thomomys pallescens, 55.

pascalis, 54.
perditus, 95.
peregrinus, 91-92.
perpallidus, 68-69.
perpes, 72-73.
pervagus, 82-83.

pinetorum, 124-125.

premaxillaris, 124.
pryori, 104-105.
puerte, 58-59.
pygmeeus, 109-110.

quadratus, 114-115.

NORTH AMERICAN FAUNA.

Thomomys umbrinus, 89-90.
yelmensis, 118-119.
toltecus, Thomomys fulvus, 86.

townsendi, Thomomys townsendi, 42-43.

townsendii, Geomys, 42, 43.
Type localities, list, 34-36.

U.

uinta, Thomomys, 113-114.

umbrinus, Geomys, 89.
Thomomys umbrinus, 89-90.

unisulcatus, Geomys, 97.

rufescens, 98-99. V..
russeolus, 60-61. Variation, 22.

saturatus, 128. sexual, 17.

scapterus, 72. Ww.
sheldont/3-9%: Winter habits, 11.

sinaloz, 80. -

talpoides, 96-98.
texensis, 85.
toltecus, 86.
townsendi, 42-43.
uinta, 113-114.

yelmensis, Thomomys douglasi, 118-119.

Ze
Zygogeomys, 7.

[NO. 39. *

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MERICAN FAUNA —
No 40. 3. oe a8

= : = [Actual date of publication, June 20, 1916] ae

‘A SYSTEMATIC ACCOUNT OFTHE
o. PRAIRIE-DOGS oe

BY

ue N. HOLLISTER

s _ ASSISTANT CURATOR OF MAMMALS, UNITED STATES © Poe ae a - s

“NATIONAL MUSEUM : SEES ee
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bes DEPARTMENT OF AGRICULTURE
BUREAU OF BIOLOGICAL SURVEY

HENRY W. HENSHAW, Chief —

NORTH AMERICAN FAUNA
No. 40

[Actual date of publication, June 20, 1916]

A SYSTEMATIC ACCOUNT OF THE
PRAIRIE-DOGS

BY

N. HOLLISTER

ASSISTANT CURATOR OF MAMMATS, UNITED STATES
NATIONAL MUSEUM

WASHINGTON
GOVERNMENT PRINTING OFFICE
1916

LEITER’ OF TRANS ia

U. 8. DEPARTMENT OF AGRICULTURE,
BurREAU OF BIOLOGICAL SURVEY,
Washington, D. C., November 16, 1915.

Sm: I have the honor to transmit herewith for publication as
North American Fauna No. 40, a systematic account of the prairie-
dogs of North America, by Mr. N. Hollister, Assistant Curator of
Mammals, of the United States National Museum. This review of
the group is based largely upon material in the collection of the
Biological Survey. Prairie-dogs are distributed over a large part of
the Great Plains and Rocky Mountain regions. Their colonies often
number thousands of individuals, and their destruction of grasses
and other forage plants makes them of considerable economic im-
portance. Drastic measures are frequently necessary to prevent
their destroying crops of grain and hay. The Biological Survey is
engaged in exterminating these rodents in national forests and on
the public domain. The definite information in this report in regard
to the several species and their distribution, as indicated by maps,
will materially aid in efforts, National or State, to control or exter-
minate them.

Respectfully,
Henry W. HEnsHAwW,
Chief, Biological Survey.
Hon. Davin F. Houston,
Secretary of Agriculture.
2

CONTENTS.

LE Eis css aR ee a Ae eg Oat ae ee
Memiouion. habits,and economic status...-..........5.0. 020.2202 0....-
emmMmme aa, Somer a8 ee ae oh Le ee la
SE ee er ee oe Pe Le ee es
ee GUITAR ee rs. OS at ae fel ec Uk oh Pee ees.
ema FeNoOwledOMments. ... 52 2 Lie eee ee ee Se ete eee

Seen rnre cers Ss OO ee So ek oe bw ase ek
List of species and subspecies with type localities. .............---------

SPE EM UUCOCLOSSHMOMNYR 2 30.55 .'. Se PS S 2 eS seeds een
Dummer momeMar aMMECITeTIT S's) 700) es) se oe oo Se

pes

je
DOOho OS oo) = S ower ¢

fed fe!

ILLUSTRAZIONG

PLATES.
[Following page 36.]

Puats I. Skulls (dorsal views) of Cynomys ludovicianus ludovicianus, C. 1. arizonensis,
and C. mexicanus, and (posterior views) of C. 1. ludovicianus and C. 1.
arizonensis.

II. Skulls (ventral views) of Cynomys ludovicianus ludovicianus, C. l. ari-
zonensis, and C. mexicanus, and mandibles of C. 1. ludovicianus and C.
mexicanus.

III. Skulls (dorsal views) of Cynomys leucurus, C. parvidens, C. gunnisoni
gunnisoni, and C. g. zuniensis.

IV. Skulls (ventral views) of Cynomys leucurus, C. parvidens, C. gunnisoni

gunnisoni, and C. g. zuniensis.

V. Skulls (lateral views) of Cynomys ludovicianus ludovicianus, C. 1. arizo-
nensis, C. mexicanus, C. parvidens, C. gunnisoni gunnisoni, and C. g.
zuniensis.

VI. Skulls (dorsal and ventral views) of two individuals of Cynomys ludo-
vicianus ludovicianus from the same locality. ,

VII. Skulls (various views) of Cynomys ludovicianus ludovicianus, C. mexicanus,
C. gunnisoni gunnisoni, C. parvidens, and C. leucurus.

TEXT FIGURES.

Page.
Fig. 1. Distribution of subgenus Cynomys (Cynomys ludovicianus ludovicianus,
C. l. arizonensis, and C. mexicanus).:...2. 222s eee ee 13
2. Distribution of subgenus Leucocrossuromys (Cynomys leucurus, C. par-
videns, C. gunnisoni gunnisont, and C. g. zuniensis)-_ 22-225 --246---- 24
2

No, 40. NORTH AMERICAN FAUNA, June, 1916,

A SYSTEMATIC ACCOUNT OF THE
PRAIRIE-DOGS.

By N. HouuisteEr.

INTRODUCTION

The prairie-dog is a true ground squirrel, or spermophile, and its
common designation ‘‘dog”’ is most unfortunate from the standpoint
of exact science. The name is now so generally used, however, that
any attempt to substitute one more appropriate would be futile.
The animals may be conveniently divided into two general classes—
the black-tailed prairie-dogs (subgenus Cynomys, three forms of
two species) and the white-tailed prairie-dogs (subgenus Lewco-
crossuromys, four forms of three species).

DISTRIBUTION, HABITS, AND ECONOMIC RELATIONS.

Early explorers of the Western States were much impressed by
the peculiar habits of the prairie-dog, one of the most abundant and
conspicuous mammals over much of what was then ‘‘Louisiana”’
and the region of the upper Missouri River. Their journals and
narratives contain numerous references to this spermophile under
such names as prairie-dog, petit chien, wishtonwish, barking squirrel,
and prairie squirrel, together with long accounts of the ‘‘ dog-towns”’
and the social habits and notes of alarm of the animals. As usual
in the case of a new and interesting animal, these accounts are partly
true and partly erroneous, having been drawn from actual obser-
vation supplemented by stories of travelers, settlers, and Indians.

Prairie-dogs are distributed over a large part of the Great Plains
and Rocky Mountain regions of the United States and southward
into northern Mexico. This actual area is relatively small, however;
the animals are found in no other region. Though the several forms
are constantly extending their ranges into new valleys and pushing
out into new pastures on the plains, the animal has penetrated into
only a small area of the Great Basin, and seems sharply restricted by
its specialized nature from occupying any great part of the humid
eastern plains, the northern prairies of Canada, the Tropics, or the
immense stretches beyond the Rockies.

5

6 NORTH AMERICAN FAUNA. [No. 40. .

The habits of the black-tailed species and, to a slightly less degree,
those of the more mountain-loving white-tailed forms, are decidedly
social, The great size of the ‘‘dog-towns”’ on the plains has indeed
been one of the principal subjects of comment by early travelers.
The villages sometimes extend for many miles and the number of
individuals inhabiting a big town is enormous. While in certain
localities the white-tailed species are almost as, gregarious as the
black-tailed forms, in others their dens are much more scattered
and the animals live in solitary families. These differences in habit
are due mainly to variations in local habitat. In certain parts of
the mountainous ranges of Cynomys gunnisoni and of C. leucurus,
the country is broken and otherwise unsuited to sustain large num-
bers of the animals, and the creatures necessarily live much as do
certain rock squirrels of the genus Citellus, in isolated pairs or widely
separated families.

The prairie-dog is not strictly a hibernating animal except at high
altitudes or in the more northern regions where the great depth
of snow and the extreme severity of winter make hibernation a
virtual necessity. In the more southerly parts of the Great Plains
there is actually no period of hibernation. In the central Plains
States hibernation is partial and irregular, and even in the extreme
north the animals are not infrequently seen out on fine winter days,
though they hibernate for long periods during severe weather. In
the higher mountainous habitat of the white-tailed forms the villages
are often covered for many weeks with a great depth of snow, and
activity on the part of the prairie-dogs is, therefore, out of the
question. In such regions the animals are rarely seen after November
or they may retire even by middle October. In the higher parts of
the New Mexican range prairie-dogs cease to appear above ground
by approximately the first of December, although, if conditions are
favorable, they are out on fine days throughout the winter. In the
Green River Basin, Wyoming, Vernon Bailey saw Cynomys leucurus
eating sage-brush tips on snow a foot deep in zero (Fahrenheit)
weather, and also after a night when the temperature had fallen to
22° below zero.

The burrows vary greatly according to local necessities. In some
regions the mounds heaped up around entrances to dens are enormous,
while in others they are smaller or almost wanting. Much rainfall
necessitates high mounds, constant care, and incessant work by the
industrious animals until the safety of the home is insured. The
mounds in such cases are often especially well made and the earth
is very hard packed. Each double armful of wet dirt which is
pushed into position by the animal is rammed and packed into a firm
condition by repeated drives with the nose; the body, in curved

1916.] INTRODUCTION. 7

position, with shoulders bent, acts as a powerful machine to drive the
hammerlike muzzle into the earth.

While at times virtually omnivorous, the prairie-dog finds its
principal food in grasses and other green plants. Roots and stems are
eaten as well as the tender leaves. In times of scant vegetation, or
during a drought, the villages are almost destitute of plant life, so
closely is every living bit cropped.

Prairie-dogs are unquestionably responsible for great annual
damage to crops and pasturage. In certain areas the destruction
amounts to virtually the entire forage. Crops of grain and culti-
vated hay are often entirely ruined unless drastic preventive measures
are taken. In other out-of-the-way places the animals do not inter-
fere in the least with the operations of man.

In captivity prairie-dogs eat almost any food, and are especially
fond of cakes and fruit; watermelon also is relished.

NATURAL ENEMIES.

Among the principal natural enemies of the prairie-dog are the
badger, coyote, black-footed ferret, eagle, rough-legged hawk, and
raven. Prairie-dog meat furnishes much of the food of individuals
of these species that live in close proximity to the villages. Many
absurd stories of the joint occupation of dens by prairie-dogs, rattle-
snakes, and burrowing owls have been written, but careful observers
have found that whatever the relation between these denizens of the
plain may be—and the creatures are often found together in the
villages—it is anything but advantageous to the prairie-dog, and
large numbers of its young are destroyed by the unwelcome visitors.

NOMENCLATURE.

Several systematic mammalogists early bestowed technical names
on the prairie-dog, in some cases basing both names and descrip-
tions wholly upon published accounts in explorers’ narratives. The
resulting confusion in both zoology and nomenclature was con-
siderable. Systematists supposed the newly discovered prairie-dog
to be a species of marmot, and the first technical name properly pro-
posed for it, Arctomys ludovicianus, the Louisiana marmot, was given
by George Ord in the second American edition of Guthrie’s Geogra-
phy, 1815. Ord’s description was drawn from a stuffed specimen
of the prairie-dog in Peale’s Museum, and his account of the habits
of the animal was taken chiefly from the history of Lewis and Clark’s
expedition, with supplemental observations from the narratives of
Pike and Brackenridge. The specimen examined by Ord in Peale’s
Museum was certainly the skin collected by Lewis and Clark, and

8 NORTH AMERICAN FAUNA. [No. 40.

later, 1823, more fully described by Say.t' This was doubtless the
first specimen of the prairie-dog to reach any museum, and Lewis
and Clark are probably entitled to the credit for the actual discovery
of the species.

Rafinesque, in 1817, proposed the new genus Cynomys for the
‘“barking squirrel’’ of Lewis and Clark, calling the species Cynomys
socials. At the same time he provisionally named the ‘‘petit chien”’
of these authors Cynomys grisea, not knowing that it too was a ‘‘bark-
ing squirrel.’”’ Warden, in his Historical Account of the United
States, 1819, named the species Monax missouriensis, basing his de-
scription on information obtained in the narratives of late explorers,
particularly Maj. Pike; and Harlan, 1825, based still another name,
Arctomys latrans, on the barking squirrel of Lewis and Clark. These
names have now been fixed in the synonymy of one species, but there
was considerable confusion regarding the nominal species and their
nomenclature up to the time of publication of Baird’s Mammals of
North America, 1857. In this work two’ species of Cynomys were
recognized, one black-tailed and one white-tailed. Baird, who in 1855
had first described his Spermophilus gunmisona, a species of the white-
tailed group of prairie-dogs, from Colorado, still, in 1857, entertained
considerable doubt as to whether his new species was distinct from a
still more recently discovered white-tailed prairie-dog from Wyoming.
To this latter form he thought the older name Arctomys columbianus
of Ord, based on the ‘‘burrowing squirrel”’ of Lewis and Clark, might
be applied, should further investigation establish its validity.

The next authoritative work on the group was Allen’s Monograph
of the Sciuride, 1877. In this work, as in his previous list of Ameri-
can Sciuride,? Dr. Allen recognized two species of prairie-dogs, the
ludovicianus of Ord and a white-tailed species to which he applied the
name Cynomys columbianus (Ord), treating gunnisons of Baird as a
synonym. As later pointed out by Dr. Merriam,’ the name colum-
bianus really belongs to a spermophile of the genus Cutellus, and
Baird’s name gunnisoni is the earliest valid name for any member of
the group of white-tailed prairie-dogs. Dr. Merriam had been able
the previous year to substantiate Baird’s suspicions of many years
before regarding the existence of two distinct species of white-tailed
prairie-dogs, and had named the Wyoming form Cynomys leucurus.
The same year Dr. Mearns described a southwestern form of the
ludovicianus type as Cynomys arizonensis, and mn 1892 Merriam
added @. mexicanus to the list of known species of the genus. The
Sevier River valley white-tailed prairie-dog remained unknown, so
far as systematists were concerned, until it was described and named
by Dr. J. A. Allen in 1905.

1 Long’s Exped. Rocky Mts., I, pp. 451-452, 1823. 3 North Amer. Fauna No. 5, pp. 39-42, 1891.
2 Proc. Boston Soc. Nat. Hist., XVI, p. 294, 1874. '

1916.] INTRODUCTION. 9

In the present paper two subgenera of Cynomys are recognized.
These are equivalent to the black-tailed and the white-tailed groups.
One new subspecies is described and one currently recognized name
is placed in synonymy, leaving the number of recognized forms at
seven, the same as in the last (1912) general list of North American
mammals.!

PELAGES AND MOLTS.

In all prairie-dogs, except Cynomys mexicanus, there are two well-
marked annual molts over the entire animal, except on the tail, where
there is only a single renewal each year. The two coats are the harsh,
thin, summer pelage and the soft, heavily underfurred winter pelage.
In southern States, where the spring molt takes place much earlier
than in the North, the two renewals are conspicuous. Farther north
and at greater altitudes, where the season is shorter, the summer coat
is so little worn and the fall renewal so subtle that the autumnal
change 1s difficult to detect.

In general, the spring molt begins on the pectoral region and axil-
le, and the entire underparts from front to back are completely
renewed before any real molt is visible above. On the upperparts,
the head and shoulders, with occasional irregular areas on the back,
renew first, and the renewal progresses posteriorly. Considerable
wear is apparent over the anterior half of the back before the tail is
in fresh coat. It is quite common to find museum specimens in full
summer coat except for the tail, which retains the old pelage of the
previous winter. In late summer or early autumn, at about the time
when the tail is at last in full fresh pelage, the winter coat begins to
appear on the extreme posterior parts of the body. The renewal of
the underfur progresses anteriorly in a regular definite area, and the
fresh, soft winter coat gradually covers the whole body. The
autumnal renewal thus exactly reverses the order of progression in
spring. The whole process is much as if there was a continuous
summer change of hair which started at the nose in spring, worked
backward-to the tail, reversed, and in fall traveled forward to the
nose again. The tail, midway in the route of this process, receives
only one coat annually.

There is, naturally, a considerable variation in the season of
renewal according to latitude or altitude. Breeding females are
always slow to acquire the fresh coat, renewal being greatly retarded
by lack of excess vitality. Young animals of the first year do not
molt or renew at the same dates as do adult males and nonbreeding
females which have lived over one winter. The molt of Cynomys
Mexicanus appears to be much more complex than in the more north-
ern forms and is almost continuous. Some individuals of this spe-
cies show parts of three distinct pelages.

1 Miller, G. S., jr., Bull. 79, U. S. Nat. Mus., pp. 294-295, 19i2.

10 NORTH AMERICAN FAUNA. [No. 40. |
MATERIAL AND ACKNOWLEDGMENTS.

This revision of the prairie-dogs is based upon a study of 876 speci-
mens, almost all of which are well-prepared skins and skulls.1. A few
skeletons and odd skulls also have been examined. ‘The liberal policy
of American museums has made it possible to assemble practically
all the study material of the group preserved in American collections,
including the types and type series. Thanks for these courtesies are
especially due Dr. J. A. Allen, American Museum of Natural History;
Mr. Samuel Henshaw and Mr. Outram Bangs, Museum of Comparative
Zoology; Mr. Edward R. Warren, Colorado Springs, Colo.; and Mr.
Charles B. Cory and Mr. W. H. Osgood, Field Museum of Natural
History. Mr. Warren’s private collection of Colorado specimens of
Cynomys leucurus and C. gunnisoni has been of especial value and
help. In the lists of specimens examined the source is indicated of
all material from localities not represented in the United States Na-
tional Museum collections.

This review is published as a North American Fauna with the per-
mission of the Secretary of the Smithsonian Institution.

Genus CYNOMYS Rafinesque.

1817. Cynomys Rafinesque, Amer. Monthly Mag., II, p. 45.

1819. Monax Warden, Stat., Pol., and Hist. Acc. U.S., I, p. 226. Type, Monax
missouriensis Warden es Poni ludovicianus).

1827. Cynomis Lesson, Manuel de Mamm., p. 244 (pro Cynomys Rafinesque).

1894. Cynomomus Omar, Science, XXIII, p. 103 (pro Cynomys Rafinesque).

1899. Mamcynomiscus Herrera, Sin. Vulg. Cient. Vert. Mexicanus, p. 22. (pro
Cynomys Rafinesque).

Type species.—Cynomys socialis Rafinesque (=Arctomys ludo-
viciana Ord).

Diagnosis.—Like Citellus, but with cheek teeth relatively much
larger; maxillary tooth rows strongly convergent posteriorly; molari-
form teeth with protocone much more hypsodont; and crown of last
upper molar with well-developed mesostyle and accompanying
additional transverse ridge.

General characters.—Size as in larger species of Citellus or gres ter;
form stout; tail short, rarely more than one-fourth the total length;
ears short, not extending beyond fur of winter pelage; cheek pouches
moderate. Wrist and heel heavily furred; a tuft of hair in center of

1 The material examined has been assembled from collections as follows:

U.S. National Museum, Biological Survey collection... 2... < <22 a2 csijain- ome = eee ee eerie let 465
U.S. National Museum propeti.cis 2s 6. socio cone wiciciernin elm ww 2 micleye wn eset ae ee ete a 221
American Museum of Natural History, New York, N. Y....-.--.------.--- ad cRatG a CS EE, 81
Museum of Comparative Zoology, Cambridge, Mass.2. .... 5/0... 6-2 -4+ - 252 4 eee eee = 47
Collection of E. R.. Warren; Colorado Springs, Colo...----.-.2 =. <)- 222 ese - ene eln = = ie ee eee ole 44
Field Museum of Natural History, Chicago, Tb. 02.21.02 2a. .02 22 at wc e eee on cle ee eee eee etal 18

Total number of'specimens:.'.. . 02. es as os eas oe ae oo ad oe ae nate eee ao eters 876

1916.] GENUS CYNOMYS. dial

- palm of foot usually extending forward to phalangeal tubercles;
manus with five distinct claws, the claw of pollex well developed,
subequal to claw of outer (fifth) toe. Mamme 8 to 12. Skull
rather broad and heavily built; sinciput high and rounded; squamosal
arm of zygoma widely spreading; occipital crest well developed;
sagittal crest moderate anteriorly, but well developed posteriorly
in old adults; antorbital foramen subtriangular, with strongly devel-
oped tubercular process. Molar series strongly convergent posteri-
orly; teeth very large and greatly expanded laterally; first premolar
large, sometimes nearly equal to second; crown pattern of molars
essentially as in true Citellus, but teeth with higher crowns and
deeper grooves; molars and premolars with protocone much more
hypsodont; m? with additional transverse ridge extending (from
mesostyle) across outer half of center of crown; upper incisors some-
times with very indistinct groove along inner face.

Color pattern.—Simple; body unmarked, not sharply bicolor.
_ Upperparts reddish brown, buffy, or grayish, finely lined with darker
or lighter hair-tips; underparts paler, clear buff or pale cinnamon. |
More or less well-marked dark areas above and below eye; tail con-
spicuously tipped with black or white.

Geographic distributionn—Rocky Mountain and Great Plains
regions of the United States and northern Mexico. From the Milk
River near the Canadian boundary in northern Montana south to
northern San Luis Potosi, Mexico; east to the Missouri River in North
Dakota and to about the ninety-seventh meridian in Nebraska and
Oklahoma; west to the Rocky Mountains in Montana, to the valleys
of central Utah, and to the border of the Grand Canyon on the
Hualpai Indian Reservation, Arizona (figs. 1 and 2).

Remarks.—The prairie-dog is a true ground squirrel, orspermophile; -
there is no real evidence of the frequently suggested close relationship
between Cynomys and Marmota.

In an article by Prof. K. A. Satunin in the Mitteilungen des
Kaukasischen Museums, volume IV, pages 175 to 193, 1909, the con-
clusion is reached that Colobotis Brandt, currently recognized as a
subgenus of Citellus, is a synonym of Cynomys.. Through the kind-
ness of Mr. Oldfield Thomas, I have been able to examine, from the
collection of the British Museum, skulls of the type species of Colo-
botis—Citellus fulvus (Lichtenstein). It is at once apparent that
Cynomys is even less closely related to Colobotis than to Citellus proper,
and that it is unquestionably an error to use the generic name Cyno-
mys for any of the Old World spermophiles. Colobotis is exceedingly
like true Citellus and differs generically from Cynomys in the same
characteristics, especially in the less complicated last upper molar, lack
of the striking internal hypsodontism of the molariform teeth, and in
the nearly parallel-sided palate.

12 NORTH AMERICAN FAUNA. [No. 40.

List of Species and Subspecies with Type Localities.
Subgenus CYNOMYS.

Cynomys ludovicianus ludovicianus (Ord)........-.---- Upper Missouri River.
ludovicianus arizonensis Mearns. ........--..-.----- Point of Mountain, Arizona.
menicanus Meriamis: 220). 025. je. b50ase Se eee La Ventura, Coahuila, Mexico.

Subgenus LEUCOCROSSUROMYS.

Cynomys leucurus Mermamys.2 2225255 28o= = So gees Fort Bridger, Wyoming.
Darwiden’s AAWCW SSS ace en < te ne nee a ee ee Buckskin Valley, Utah.
gunanisoni guunisont (Baird)_ 22.23.25. 22 eI. - Cochetopa Pass, Colorado.
QUAmISON? ZuNniensis NObIS..:...228h2s- s52- Eee ee Wingate, New Mexico.

Key to Species and Subspecies.

a.! Tail tipped with black; jugal bone heavy, thickened, the outer face at angle of
ascending branch presenting a broad triangular surface (Cynomys).
6.1 Black on tail covering most of distal half; tail longer; audital bulle large and
greatly inflated; posterior border of inflected angle of mandible nearly atright
angle 'toaxis of Jaw. siisets songee eet woss es eee C. mexicanus (p. 21).
6.2 Black on tail confined to distal third; tail shorter; audital bulle smaller and less
inflated; posterior border of inflected angle of mandible at angle of about
45° to axis of jaw. :
c.1 Coloration above in fresh summer pelage dark pinkish cinnamon; audital
bulle smaller; anterior face of maxillary root of zygoma deeply emerginate,
C.1. ludovicianus (p. 14).
c.2 Coloration above in fresh summer pelage more vinaceous-cinnamon; audital
bulle larger; superior surface of maxillary root of zygoma very broad, not
deeply emarginate anteriorly...............----- C. 1. arizonensis (p. 19).
a.2 Tail tipped and bordered with white; jugal bone weak, thin, and flat, the outer
surface at angle of ascending branch only very slightly thickened, the margin
rounded, not distinctly triangular (Leucocrossuromys).
6.1 Terminal half of tail white, without dark center.
c.1 Size larger; coloration in summer buffy or grayish (not reddish or rich cinna-

mon); skull with less interorbital breadth............-. C. leucurus (p. 24).
c.2 Size smaller; coloration in summer reddish or rich cinnamon (not buffy or
grayish); skull with greater interorbital breadth. ..... C. parvidens (p. 27).

6.2 Terminal half of tail with gray center, bordered and tipped with white.
c.1 Size smaller, with smaller hind foot; coloration more buffy and blackish, less

pinkish cinnamon and ochraceous................- C. g. gunnisoni (p. 29).
c.2 Size larger, with larger hind foot; coloration more pinkish cinnamon and
ochraceous, less buffy and blackish..............-- C. g. zuniensis (p. 32),

Subgenus CYNOMYS Rafinesque.

Characters.—Size large; tail comparatively long, averaging more
than one-fifth the total length, and tipped with black; mamme 8.
Skull: Angular and massive; general shape of occipital region, viewed
from behind, oval; jugal bone heavy, thickened, the outer surface
at angle of ascending branch very broad, triangular, with inferior
vertex produced far downward; maxillary root of zygoma corre-
spondingly strengthened, the shelf and suprajugal arm much thick-
ened. Teeth larger, more expanded laterally, than in species of the
white-tailed group.

1916.] SUBGENUS CYNOMYS. 13

~S & ai CF
a iit ae ae

B2119-129

Fic. 1.—Distribution of subgenus Cynomys: 1, Cynomys ludovicianus ludovicianus; 2, C. 1. arizonensis
8, C. mexicanus.

14 NORTH AMERICAN FAUNA. [No. 40.

CYNOMYS LUDOVICIANUS (Orp).

(Synonymy under subspecies.]

Characters.—Differs from Cynomys mexicanus in more reddish col-
oration and shorter tail; black area on terminal portion of tail much
more restricted; skull with smaller audital bulle and less developed
triangular plate of jugal; teeth larger.

Subspecies—Two subspecies of Cynomys ludovicianus are here
recognized, the typical form, occupying a large part of the Great
Piains region from Montana to Texas; and C. l. arizonensis, the more
arid southwest, in New Mexico, Arizona, and adjacent portions of
Mexico and southwestern Texas.

CYNOMYS LUDOVICIANUS LUDOVICIANUS (Orp).
BuackK-TAILeD Prartrie-Doa.
(Pl. I, figs. 1, 4; Pl. II, figs. 1, 4; Pl. V, fig. 1; Pl. VI; Pl. VII, figs. 1, 7, 8.)

1815. Arctomys ludoviciana Ord, Guthrie’s Geogr., 2d Amer. ed., II, pp. 292, 302.

1817. Cynomys socialis Rafinesque, Amer. Monthly Mag., II, p. 45, November.
(Plains of the Missouri.)

1817. Cynomys ? grisea Rafinesque, Amer. Monthly Mag., II, p.45, November. (“On
the Missouri.’’)

1819. Monax missouriensis Warden, Stat., Pol., and Hist. Acc. U.S8., 1, p. 226. (‘‘The
Missouri country.’’ Based largely on the account of ‘‘Major Pike, in his expedi-
tion through Louisiana.’’)

1822. Arctomys missourtensis Desmarest, Mammalogie, pt. 2, p. 329.

1825. Arctomys ludoviciani Harlan, Fauna Amer., p. 305.

1825. Arctomys latrans Harlan, Fauna Amer., p. 306. (Plains of. the Missouri.)

1827. Spermophilus ludovicianus Lesson, Manuel de Mamm., p. 244.

1827. Cynomis socialis Lesson, Manuel de Mamm., p. 244 (syn.).

1827. Spermophilus griseus Lesson, Manuel de Mamm., p. 245.

1827. Cynomis griseus Lesson, Manuel de Mamm., p. 245 (syn.).

1827. Cynomys griseus Lesson, Manuel de Mamm., p. 260.

1829. Arctomys ludovicianus Cuvier, Régne Animal, I, p. 197.

1829. Alrctomys] missuriensis Fischer, Syn. Mamm., p. 345.

1829. A[rctomys] griseus Fischer, Syn. Mamm., p. 345.

1829. Arctomys (Spermophilus? ) ludovicianus Richardson, Fauna Boreali-Amer., pt. 1,
p. 154.

1829. C[ynomys] cinereus Richardson, Fauna Boreali-Amer., pt. 1, p. 155 (pro grisea
Rafinesque, 1817).

1845. Cynomys cinereus Schinz, Syn. Mamm., IT, p. 64 (syn.).

1855. Monax missuriensis Giebel, Die Saugethiere, p. 630 (syn.).

1857. Cynomys ludovicianus Baird, Gen. Rep. North Amer. Mamm., pp. xxv, 331.

1905. Cynomys pyrrotrichus Elliot, Proc. Biol. Soc. Washington, XVIII, p. 139,
April 18. (White Horse Spring, Woods County, Oklahoma; type in Field Mus.
Nat. Hist., Chicago.)

Type locality—Upper Missouri River (‘‘vicinity of the Missouri,
and throughout the greater part of Louisiana’’).

Geographic distribution.—Great Plains region of western United
States, south from near the Canadian border in Montana to west-

1916.] CYNOMYS LUDOVICIANUS LUDOVICIANUS. 15

central Texas (Mason County to eastern Pecos Valley); east to about
the ninety-seventh meridian in Nebraska, Kansas, and Oklahoma;
west to the Rocky Mountains in central Montana, Wyoming, and
Colorado, and in extreme eastern New Mexico. Chiefly Upper
Sonoran, but also ranging into Transition and Lower Sonoran Zones.
Introduced colonies exist, or have been reported as formerly existing,
in Sac County and at Burlington, Iowa; near Monroe, Louisiana; at
Seneca, South Carolina; and on Nantucket Island, Massachusetts. '

* Characters.—Skull with superior surface of maxillary root of
zygoma bordering premaxillary and frontal bones narrow, sharply
emarginate anteriorly; audital bulle comparatively small. Colora-
tion in fresh summer coat less vinaceous-cinnamon than in Cynomys
ludovicianus arizonensis.

Color.—Adult in full fresh summer pelage: Upperparts from nose to
between eyes, between ears, nape, and over entire body above, dark
pinkish cinnamon, finely lined with black and buff. Individual hairs
black at base, followed by buffy white, then cinnamon, with sub-
terminal band of buff, and, in unworn condition, narrow tip of
blackish. Mixed with these are numerous wholly black and half-
black hairs, rather longer than the ordinary pelage, the chief cause
of black streaking. Upper lip, sides of nose, and eye ring buff or
buffy white; whiskers black; cheeks and sides:of head buffy or
vinaceous-cinnamon, with numerous blackish hairs; arms above,
sides of body, and legs above pale ochraceous-cinnamon; feet buffy.
Tail above for about two-thirds its length like back, below paler
vinaceous-cinnamon; terminal third chiefly black or blackish brown
above and below, with extreme tip of lighter blackish brown. Under-
parts of body from chin to near tail whitish or buffy white. Nails
blackish, tipped with lighthorn. Irishazel. Adult in full fresh winter
pelage: Peldge much thicker, softer, and longer than in the summer
coat; less ochraceous or pinkish cinnamon, more buff and gray.
Upperparts grayish cinnamon (mixed pale cinnamon, whitish, and
black). Individual hairs intense black at bases, then pale buff, with
subterminal band of cinnamon and tip of almost pure white. Mixed
throughout this pelage, as in the summer coat, are numerous long
overlying hairs of black. Forehead with considerable blackish; tail
asinsummer. Underparts dark buff or pale cinnamon, the hairs black
at bases, whitish midway, and broadly tipped with pale cinnamon-
buff. Juvenile pelage: Upperparts clear ochraceous-cinnamon, with
admixture of a few white and blackish hairs; underparts whitish or
pale buffy yellow; tail clear ochraceous-cinnamon, broadly penciled
with black; bases of many of the black pencil hairs pure white.

Both the summer and the winter coats rapidly fade and wear, and
the resulting variations in color are considerable. Specimens are
frequently greatly affected by the color of the soil, and numerous

16 NORTH AMERICAN FAUNA. | [No. 40.

examples are considerably reddened or darkened by stain, either on
the upper and lower body or uniformly throughout the pelage.
Specimens in complete fresh summer coat are rather uncommon as
the vernal molt is so slow that the forward parts, renewed first, are
already considerably worn before the tail, last to renew, is in full
fresh coat.

Molt.—The spring molt and renewal are very conspicuous but the
fall molt, especially in northern localities, is often obscure. There
is considerable variation in the dates of renewal but the followifig
schedule is about the average for northern and central Great Plains
States: Late March and early Aprilskins are worn, pale, and faded, but
already show some signs of molt and renewal on the underparts, where
the first new hair appears on the pectoral region and axille. May
specimens have completely renewed below and the renewal has com-
menced on the head, shoulders, and in irregular patches on the back.
June specimens are almost completely in fresh coat, but still show, as
a rule, some of the old pelage on the rump and tail. July and early
August skins are usually completely renewed, except for a few in
which the tails above are still in the winter coat. The hair of the
body is short and harsh, and rapidly becomes broken, worn, and
faded. Late August adults show considerable progress in the fall
renewal; the winter underfur and long soft hair is appearing on the
lower back, and the animal looks generally lighter colored. The
tails are fully renewed. November specimens are in complete new
fur, and the hair on the tails still appears fresh and unworn after its
single annual renewal; all traces of the summer coat have dis-
appeared. In December skins, the coat has thickened, grown longer,
and appears considerably lighter in color. This is in part due to
fading, but is also caused by the general effect of — long white
underfur showing through the reddish hair tips.

Skull and teeth.—The skull of Cynomys ludovicianus ludovicianus
differs from that of C. 1. arizonensis in its distinctly smaller audital
bullz and narrow, deeply emarginate anterior face of maxillary root
of zygoma. It may readily be distinguished from skulls of C. leucurus
(whose range meets that of C. ludovicianus in parts of Wyoming) by
its shghtly larger size, smaller bulle, more notched anterior face of
maxillary root of zygoma, less flattened zygoma (muscle attachment
reaching farther forward, nearly or quite to plane of first tooth), gen-
erally heavier mandible, ighter postorbital processes, and small audi-
tory meatus. The incisors vary in color from white ‘to deep yellow in
specimens from one locality. The variation in size and shape of skull
among specimens of the same sex and age from one locality is remark-
able (see Pl. VI).

- Measurements.—Averages of 13 adult males from South Dakota:
Total length, 388 (360-415); tail vertebrae, 86 (75-98); hind foot,

1916.] CYNOMYS LUDOVICIANUS LUDOVICIANUS. ye

62 (61-83). Skull: Condylobasal length, 59.8 (57.2-62.6); zygomatic
breadth, 45 (42.9-48.1); mastoid breadth, 28.5 (27.5-31); length of
nasals, 23.8 (22-25); length of mandible, 45.9 (43-49.4); maxillary
tooth row, alveoli, 16.6 (16-17.5). Females average very slightly
-smaller than males, but the difference in size is, as a rule, barely per-
ceptible. For detailed measurements of specimens, see page 34.

Weight.—The weights of adult examples of C. l. ludovicianus are
recorded in collectors’ field catalogues as follows: Alma, Nebraska,
October 21, 3, 3 lbs., 9, 2 lbs. 4 oz.; Tilyou’s Ranch, Montana,
October 1, ?, 2 lbs. 3 oz.

Type specumen.—Ord’s description of this prairie-dog was drawn
from a stuffed skin in Peale’s Museum, Philadelphia, supple-
mented by notes from the narrative of Lewis and Clark. This
specimen was without doubt the identical skin which later on was
more fully described by Say.! It has long since been lost sight of.

Remarks.—As explained in the account of the pelage changes, there
is considerable seasonal color variation in the common prairie-dog,
but a comparison of specimens in exactly the same state of pelage
from all parts of the animal’s range proves the absence of any con-
stant geographical differences. There is likewise a remarkable
uniformity in size, the averages and extremes of long series from
northern States being virtually identical with measurements of
specimens from Texas. The alleged form from Oklahoma, described
by Dr. Elliot as Cynomys pyrrotrichus, is not subspecifically distin-
guishable from C. l. ludovicianus. Specimens from this region are
commonly stained by the red soil, and sometimes so naturally as
almost to defy detection as a stain. Numerous examples, however,
show the telltale red soil on the toes and feet, and a careful micro-
scopic examination of the hairs often shows the particles composing
the stain, which can be perceptibly lightened by a moment’s washing
with alcohol or benzine. There are, furthermore, absolutely nor-
mally colored specimens from various parts of the area included in
the distribution of the ‘‘red-haired’’ prairie-dogs, and all the young
specimens are indistinguishable from juvenile examples from the
Dakotas and other parts of the range of the species. Reddish skins,
in all respects like the specimens of ‘‘pyrrotrichus’’ from Oklahoma,
are not infrequently found in other parts of the prairie-dog’s range
where soil conditions are favorable for producing such a stain.

On the basis of skull characters the subspecies ludovicianus is typ-
ical over the northern parts of its general distribution, in Montana,
the Dakotas, and Wyoming. The ‘‘area of intergradation”’ with C. 1,

1 See Guthrie’s Geog., 2d Amer. ed., p. 303, under ‘“‘Columbia Marmot,’’ where Ord states that ‘‘A stuffed

skin of the Louisiana Marmot is in the Museum of Mr. Peale.’’ Also, Say, in Long’s Exped.-to Rocky Mis.,
I, p. 452, 1823.

20651°—16——2

18 NORTH AMERICAN FAUNA. [No. 40.

arizonensis is large, and skulls from Nebraska, Kansas, Oklahoma,
and Texas are sometimes difficult to distinguish. As here mapped
the range of (. 1. ludovicianus includes the entire area where typical
examples most frequently occur. The characters separating ari-
zonensis are slight and are only average. For example, in an exami-
nation of 120 adult skulls from Montana, the Dakotas, Arizona,
and Chihuahua, about 80 per cent can readily be placed with their
proper form; comparing 115 skulls from Wyoming, *Nebraska, and
New Mexico, about 75 per cent are easily determinable; 31 skulls
from Colorado and extreme western Texas (west of Pecos River) still
show about 75 per cent readily distinguishable; while of 56 skulls
from Kansas and Texas east of the Pecos, there are just about 50 per
cent which can be differentiated, and the characters are relatively
hard to distinguish satisfactorily.

An albinistic specimen, entirely white, from the Central Park
Menagerie, New York, has been examined in the collection of the
American Museum of Natural History.

Specimens examined.—Total number, 290, as follows:

Colorado: Boulder County, 1; Colorado Springs, 1;! Denver, 1; Don Carlos, 1;
Fremont County, 1;? Larimer County, 4;3 Loveland, 3; Monon (Baca County),
2;1 Olney (Otero County), 4; Pueblo, 1; Rockvale, 2;? Soda Springs, 2;4
Springfield, 2.1

Kansas: Arkansas River, 1; Banner, 8; Cairo, 2; Coyote Station, 1;* Fort Hays,
8;4 Garden City, 2;? Garden Plain, 1; Hoxie, 1; Long Island, 3;3 Pendennis
(Lane County), 1; Republican River, 1; Trego County, 6.

Massachusetts: Nantucket, 2 (introduced).

Montana: Billings, 1; Boxelder Creek, 1; Calf Creek, 1;3 Craig (10 miles
northwest of), 1; Darnall’s Ranch, 1; Fort Assinniboine, 5; Fort Custer, 3;
Glendive, 6; Great Falls, 2; Milk River (near mouth), 2; Newland, 2; Pom-
pey’s Pillar, Yellowstone River, 1; Shelby Junction, 4; Teton River, 2;
Tilyou’s Ranch, 1.

Nebraska: Alma, 2; Birdwood Creek, 3; Columbus (Platte County), 1; Ken-
nedy, 3; Platte River, 4.

New Mexico: Chico Springs, 1; Koehler Junction, 2; Pecos, 1; Tompkin’s
Lake, 2.

North Dakota: Glenullin, 2; Little Missouri River, 1; Medora, 7.

Oklahoma: Beaver River, 4;?,:3 Chattanooga, 1; Mount Scott post office, 3;
Neutral Strip, 5;?> Ponca Agency, 2; White Horse Spring, 5 (including type
of ‘‘pyrrotrichus”’).?

South Dakota: Armour, 1; Buffalo Gap, 8; Cheyenne River (Custer County),
2;3 Corral Draw, 3;3 Edgemont, 2; Fort Pierre, 6; Fort Pierre to Badlands,
15 (skulls); Fort Randall, 11; Phinney (Cheyenne River), 1; Pine Ridge
Indian Reservation, 1;2 Rapid City, 10; Upper Missouri River, 3 (skulls and
skeletons).

Texas: Brazos River, 1; Colorado, 1; Fort Chadbourne, 1;* Henrietta, 4; Lips-
comb, 1; Llano Estacado, 1; Mason, 9; Monahans, 1; Red River, 2; Stanton,
2; Texline, 1; Vernon, 11; Wichita Falls, 3.

1 Collection E. R. Warren. 8 Collection Amer. Mus. Nat. Hist.
2 Collection Field Mus. Nat. Hist. 4 Collection Mus. Comp. Zool.

|
)
|

1916.] CYNOMYS LUDOVICIANUS ARIZONENSIS. 19
J

Wyoming: Arvada (Powder River Valley), 2; Camp Carling (=Fort Russell, near
Cheyenne), 2; Cheyenne, 6; Deer Creek, 1; Douglas, 4; Fort Fetterman, 2;
Ishawooa, 4; Newcastle, 4; Pine Bluff, 2; Pole Creek, 7; Sage Creek, 11.1

CYNOMYS LUDOVICIANUS ARIZONENSIS Mzarns.
ARIZONA PRAIRIE-Doa.
(Pl. I, figs. 2, 5; Pl. Il, fig. 2; Pl. V, fig. 2.)

1890. Cynomys arizonensis Mearns, Bull. Amer. Mus. Nat. Hist., II, no. 4, p. 305,
February 21.

1892. Clynomys] ludovicianus arizonensis Merriam, Proc. Biol. Soc. Washington, VII,
p. 158, July.

Type locality—Point of Mountain, near Willcox, Cochise County,
Arizona.

Geographic distribution.—Southeastern Arizona, southern and
central New Mexico, southwestern Texas, and adjacent portions of
Sonora and Chihuahua, Mexico. North to San Pedro and Santa
Rosa, New Mexico; east to the Pecos Valley; west to Huachuca,
Arizona; south to San Diego and Casas Grandes, Chihuahua, and to
Presidio County, Texas.

Characters —Like Cynomys ludovicianus ludomcianus but averaging
very slightly larger, and brighter colored; skull with larger audital
bullz, and with superior surface of maxillary root of zygoma bordering
premaxillary and frontal bones very broad, not deeply emarginate,
the zygoma as viewed from above not branching at sharp angle from
rostrum.

Color —AlImost precisely as in the typical subspecies, but averaging
slightly brighter vinaceous-cinnamon in fresh summer pelage.

There is a single wholly melanistic specimen from south of the
Guadalupe Mountains, in Texas, in the National Museum collection.

Molt.—Specimens showing excessive molt are more common than
in the northern subspecies, and molting examples frequently exhibit
strange patterns of coloration. There are frequent mixtures of the
faded, left-over winter coat and the fresh reddish summer coat.
Excessive wear before any renewal sometimes leaves large parts of
the animal entirely black, the hair having been worn away com-
pletely except for the black underfur.. Specimens in the much-worn
pelage of iate summer, before any evidence of fall renewal, frequently
present a curious appearance: the color is pale ochraceous or yel-
lowish buff and the hairs appear brittle and dead, often matted and
twisted as if scorched by excessive heat.

January and February skins are in the long, soft, faded winter
pelage. In the lower levels the spring molt occurs earlier than in the
more northern race, and the animals are usually in fuil fresh summer
coat, except for the tail, between the first part and middle of May.
The short, stiff, summer coat rapidly wears and is much changed

1 Collection Amer. Mus. Nat. Hist.

20 NORTH AMERICAN FAUNA. [No. 40.

before the hair of the tail has renewed. At higher altitudes the
pelage is not completely renewed until the middle of July. When
the summer pelage is acquired this late the animals have molted
again and renewed into full fall pelage by the first of October, while
specimens from the lower levels, which acquired the summer pelage
much earlier, are sometimes as late as November in attaining the
same condition. |

Skull and teeth—The skull of C.l. arizonensis differs from that
of C. 1. ludovicianus in the very broad surface of the superior maxillary
arm of the zygoma, bordering the ascending branch of the premaxilla
and the frontal bone, which in the typical examples is nearly twice as
wide as in C. 1. ludovicianus, less sharply emarginate anteriorly, and
rather more convex on the outer surface along the jugal. As a
result the maxillary arm of the zygoma appears to leave the rostrum
at a much less sharp angle than in the northern form. The audital
bulle average considerably larger than in typical ludovicianus.
Teeth essentially as in the northern race.

Measurements.—Averages of 19 adult males from southeastern
Arizona: Total length, 388 (350-412); tail vertebra, 89 (78-100);
hind foot, 62 (57-65). Skull: Condylobasal length, 60 (56.6-64);
zygomatic breadth, 44 (41-47.2); mastoid breadth, 27.3 (26.1—-29.5);
length of nasals, 24 (22.3-25.4); length of mandible, 45.2 (42.8-
47.2); maxillary tooth row, alveoli, 16.2 (15.5-17). Females average
slightly smaller. For detailed measurements of specimens, see
page 35.

Weight.—Three male specimens from Altuda, Texas, July 30, have
the following weights recorded in collector’s field catalogue: 2 lbs. »
2 oz., 2 Ibs. 6 oz., and 2 lbs. 8 oz.

Type specimens.—There are two cotypes, in the American Museum
of Natural History, New York: Skin (No. 2509, ? ad.), from Point
of Mountain, near Willcox, Arizona, April 9, 1885; and skin and
skull (No. 2183, g ad., teeth considerably worn), from Dragoon
Summit, Arizona, May 3, 1885; both collected by Dr. Edgar A.
Mearns. Dr. Mearns later! restricted the type locality to Point of
Mountain, and listed No. 12162 as the type specimen. Two female
specimens collected on the same date bear this skin number of the
American Museum. One has a skull numbered 12162, but the other,
without a skull number, bears a red type label, like No. 2185 from
Dragoon Summit.

Remarks.—This is a slightly characterized form, barely recogniza-
ble except for average skull characters. Remarkable as it may seem,
there are specimens from near the type region which can be matched
in every particular with examples of typical ludovicianus from Mon-
tana. Nevertheless, as mentioned under the preceding form, about

1 Mearns, E. A., Bull. 56, U. S. Nat. Mus., p. 239, 1907.

————————— ll —

1916.] CYNOMYS MEXICANUS. 21

80 per cent of the skulls examined from the two extremes are readily
determinable. The very slight average increase in size and the
slightly more vinaceous summer coat are of such trivial degree that
alone they would be valueless as characters for subspecific separa-
tion. Throughout southeastern Arizona and over most of the New
Mexican range of the form the skull characters are well marked, but
in western Texas such a strong tendency toward C. l. ludovicianus
appears as to make identification of single specimens or unsatisfac-
tory series virtually impossible. I have therefore considered the
range of the form restricted to the region from the Pecos Valley
south and west, where 75 per cent or more of the skulls are deter-
minable. The area of intergradation with the subspecies ludovi-
cianus is so extensive (from Texas to Nebraska) that it is really
larger than the range of either subspecies in its typical form.
Specimens examined.—Total number, 184, as follows:

Arizona: Bonita, 1; Dos Cabesos, 1; Dragoon Summit, 1 (cotype);! Fort Hua-
chuca, 10; Huachuca Plains, 4; 2» Point of Mountain, 4 (including cotype);!
San Pedro River, Mexican Boundary Line, 20; * Sulphur Spring Valley, 2; 1
Willcox, 13.

Chihuahua: Colonia Juarez, 5; Juarez, 2; 13 Sierra en Media, 1.

New Mexico: Animas Valley (Grant County), 14; Cactus Flat (20 miles north
of Cliff), 4; Capitan Mountains, 3; Cliff, 1; Cloverdale, 3; Dog Spring (Grant
County), 9; Faywood, 2; Gila (Grant County), 1; Hachita, 2; Jornada del
Muerto, 1; Lake Valley, 3; Lone Mountain, 1; Manzano Mountains (east foot-
hills), 2; Organ City, 1; Playas Valley, 1; Queen, 3; Roswell, 2; San Luis
Springs (=Lang’s ranch, Grant County), 26 (including 17 odd skulls); San
Pedro, 3; Santa Rosa, 7; Silver City, 5.

Texas: Alpine, 1; Altuda, 3; Belen (El Paso County), 4; El Paso, 4; Fort Davis,
1; Guadalupe Mountains (near), 1; Limpia Mountains (=Davis Moun-
tains), 1; Pecos River, 1; Presidio County, 2; Sheffield, 3; Sierra Blanca, 5

CYNOMYS MEXICANUS Merriam.
MEXICAN PrarriE-Doe.
(ie ness Pet ties, 53° Pl. V; fig. 3;-Pl: VI, fig*2.)
1892. Cynomys mexicarus Merriam, Proc. Biol. Soc. Washington, VII, p. 157, July.
Type locality. La Ventura, Coahuila, Mexico.
Geographic distribution.—Southeastern Coahuila and northern San
Luis Potosi, Mexico. North to Saltillo; south to Vanegas.
Characters.—Size of Cynomys ludovicianus, but with longer tail;
coloration above in summer coat less reddish, more grayish and
vinaceous-buff, and much more heavily lined with black; coloration
in winter still more hoary; black on terminal portion of tail much
more extensive; teeth smaller.
1 Collection Amer. Mus. Nat. Hist.
2 Collection Mus. Comp. Zool.
3 Collection Field Mus. Nat. Hist.

4 Dr. Mearns tells me that a few of these specimens were actually collected on the Mexican side of the
boundary line, in Sonora.

22 NORTH AMERICAN FAUNA. [No. 40.

Color.—Adult in fresh summer pelage: Upperparts as in @. ludo-
vicianus, but less reddish and much more grayish and vinaceous-buff;
entirely black hairs more numerous, giving a much more grizzled
general effect. Cinnamon areas less intense, more vinaceous. Tail
above like rump for half its length; terminal half chiefly intense black,
the black. extending forward on sides of tail, leaving a lighter area
_ midway; hairs of sides of pencil tipped with whitish, bases of all
the pencil hairs almost pure white. Adult in fresh winter pelage:
General appearance decidedly hoary or vinaceous-buff, the hairs
with broad subterminal bands of whitish, which mix in the general
effect with the narrow black tips and cinnamon undercolor. Indi-
vidual hairs narrowly black at base, then broadly white and nar-
rowly cinnamon, with subterminal band of buff and tip of blackish.
The general effect is a glossy marbled vinaceous-buff, making the
coat of this species by far the handsomest of all the prairie-dogs.
Juvemle specimens are much paler than those of the corresponding
age in C. ludovicianus, and rapidly change into a pelage like the
fresh autumnal coat of adults. |

Molt.—The molt of C. mexicanus is almost continuous and ex-
ceedingly complex, and without series of specimens collected in all
the summer months it can not be worked out satisfactorily. With
only March, April, July, and August skins before me I am inclined to
believe that there are three complete renewals of the pelage of the
body annually. Specimens collected in March have all renewed
below, but are chiefly in the worn winter coat above, except in small
areas forward, or even over the greater part of the head, shoulders,

and withers, where renewal into a pelage much like the fresh winter —

coat has commenced. April skins from Saltillo are in a much-worn
condition. The real summer pelage is certainly retained only a
short time, as specimens collected at La Ventura from July 19 to
August 5 exhibit every condition from what appears to be a worn,
faded, left-over winter coat; through renewal into a short, stiff sum-
mer coat, without underfur; and the progress of the heavy winter
pelage with a dense underfur over the posterior half of the back.
Several specimens show all three of these pelages in definite areas on
the body. Breeding females, as usual, are slow to renew, and the
extreme state of wear is common until after the nursing season is
over.

Skull and teeth.—Skull broad and angular, with wide-spreading
zygomatic arches and sharply defined processes; in general effect
more like skulls of C. 1. ludovicianus from Montana and the Dakotas
than like those of C. 1. arizonensis; upper maxillary root of zygoma
narrow along contact with ascending premaxillary bones and along
ascending arm of jugal; face of upper incisors bright yellow. Differs
from skulls of both ludovicianus and arizonensis in its much larger,

1916.] SUBGENUS LEUCOCROSSUROMYS. 03

greatly inflated audital bulle; teeth smaller; nasals broad and trun-
cate posteriorly; triangular plate of jugal especially well developed
and greatly produced at downward point; posterior border of in-
flected angle of mandible nearly at right angle with axis of jaw (in
ludovicianus at an angle of about 45° with axis); the indistinct
grooves along inner half of face of upper incisors usually more
noticeable than in other forms.

Measurements.—Averages of 8 adult males from La Ventura,
Coahuila: Total length, 416 (390-430); tail vertebra, 102 (89-115);
hind foot, 63 (59-68.5). Skull: Condylobasal ieee 59.3 (58.4-
60.5); zygomatic breadth, 44.6 (43-45.6); mastoid breadth, 28.6
(27.9-29.5) ; length of nasals, 23 (22.3-23.8) ; length of mandible, 43.8
(41.5-45.8) ; maxillary tooth row, alveoli, 15.9 (15.4-16.3). Females
average slightly smaller. For detailed measurements of specimens,
see page 30.

Type specimen.—No. 28433 U. S. National Museum, Biological
Survey collection. pee male, skin and skull (teeth moderately
worn). Collected March 24, 1891, by C. P. Streator; original number
625. Except for the underparts, head, nape, and shoulders, which
are in process of renewal, the specimen is in the worn, left-over winter
pelage.

Remarks.—This is a well-marked species, easily distinguishable ex-
ternally and by skull characters from the other forms of the genus.
It is no more closely related to Cynomys ludovicianus arizonensis than
to typical ludovicianus, and the southern subspecies arizonensis can
not be considered as intermediate in the true sense between it and
ludovicianus. In the excessive size of the audital bulle, mezicanus
is the extreme of the arizonensis type, but in other characteristics it
more closely resembles Judovicianus, and is separated from both by a
number of constant differences. It is completely isolated in its
distribution from other forms of Cynomys and occupies only a
limited range.

Specimens examined.—Total number, 42, as follows:

Coahuila: La Ventura, 37 (including type); Saltillo, 5.}

Subgenus LEUCOCROSSUROMYS nov.

Characters.—Size slightly smaller than in the subgenus Cynomys,
general appearance more spermophile-like; tail short, averaging less
than one-fifth the total length, and tipped with white; mamme
normally 10, rarely 12. Skull: General shape of occipital region,
viewed from behind, elliptical-oval; jugal bone weak, thin, and flat,
the outer surface at angle of ascending branch only very slightly
thickened, the margin rounded, not triangular; maxillary root

1 Collection Mus. Comp. Zool.

94 NORTH AMERICAN FAUNA. [No. 40.

of zygoma correspondingly weak, the shelf and suprajugal arm not
especially thickened. Teeth smaller than in the subgenus Cynomys,
not so much expanded laterally.

Type species. —Cynomys gunnisoni (Baird).

Remarks.—The mountain-inhabiting group of prairie-dogs com-
monly known as the “white-tailed” species form a compact group
clearly separated from the typical subgenus. It includes three well-

B2120—129

¥iG. 2.—Distribution of subgenus Leucocrossuromys-: 1, Cynomys leucurus; 2, C. parvidens; 3, C. gunnisoni
gunnisoni; 4, C.g. zuniensis.

marked species, leucurus, parvidens, and gunnisont. In habits these
are distinctly spermophile-like and in coloration they are peculiar.
All have sharply marked areas of color above the eye and on the
cheek, and have the tail tipped or bordered with white.

CYNOMYS LEUCURUS Merriam.
WHITE-TAILED PRatIRiz-Dog.
(PLAIII, fig. 1; PL TV, figs ds Pa Vi tise om)

1874. Cynomys columbianus Allen, Proc. Boston Soc. Nat. Hist., XVI, p. 294 (part;
not Arctomys columbianus Ord, 1815).

1890. Cynomys leucurus Merriam, North Amer. Fauna No. 3, p. 59, September 11;
North Amer. Fauna No. 4, p. 33, October 8.

1916.] CYNOMYS LEUCURUS. 25

1898. Cynomys lewisii Allen, Bull. Amer. Mus. Nat. Hist., X, p. 456, November 10
(not Arctomys lewisii Audubon and Bachman, 1854).
1904. Cynomys lewist Trouessart, Cat. Mamm., suppl., p. 342.

Type locality —F ort Bridger, Sweetwater County, Wyoming.

Geographic distribution.—Irregular areas in the mountainous parts
of Montana, Wyoming, Utah, and Colorado. South from the Bighorn
Basin, in southern Montana, across central and southwestern Wy-
oming into western Colorado and northeastern Utah; east to the Lara-
mie Mountains, Wyoming, and into North Park, Colorado; south into
the lower Gunnison Valley; west a few miles across the Bear River
Divide into extreme northern Utah and, farther south, into the Green
River valley. Chiefly Transition Zone.

Characters.—Size, largest of the subgenus, only slightly smaller than
in Cynomys ludovicianus, but tail much shorter; general coloration
less reddish, more grayish buff than in C. parvidens; differs from C.
gunnisom in having terminal half of tail entirely white; supraorbital
spot and cheek dark brown.

Color.— Adult in fresh summer pelage: General color of upper parts
yellowish buff, streaked with blackish. Nose yellowish buff, un-
marked; spot above eye and large area on cheek dark blackish brown;
ears pale cmnamon. Top of head to center of tail uniformly mixed
pale cmnamon-buff or yellowish buff and blackish; the individual
hairs black at bases, then hight gray, then pale cinnamon, with sub-
terminal band of buff and, in unworn condition, tip of blackish.
Limbs, feet, and underparts clear buffy; nails blackish, tipped with
light horn. ‘Tail white, the hairs of proximal half above with bands
of blackish, below pale cinnamon; distal half clear white. Adult in
fresh winter pelage: Decidedly darker than in summer; more blackish
above, especially posteriorly; the buff tints richer in tone and the
dark areas on head considerably spread out, less sharply defined.
There is a heavy underfur of grayish white, but the bases of all the .
hairs are clear black; sides of nose, chin, and throat white; nape and
flanks lighter, more grayish, than back and rump. Juvenile pelage:
Above grayish brown; below paler grayish; supraorbital spot of
brownish black sharply marked. Postjuvenile pelage: Specimens in
first fresh autumnal coat darker than adults, more reddish, and hairs
of upperparts more heavily tipped with blackish.

Adults in the faded, left-over winter coat are often very yellowish
above, with little or no black streaking from the hair tips. Many
specimens in various stages of molt and renewal present strange com-
binations of color. Examples deeply stained by color from the soil
are frequently noted.

Molt.—The two annual molts are greatly affected by altitude, with
a consequent variation of a month or more in near-by localities. In
Wyoming the molt is approximately as follows: Specimens taken

26 NORTH AMERICAN FAUNA. [No. 40. .

before May 10 are still in the old winter coat, with little evidence of
molt. Skins collected from May 20 to June 1 have renewed over
most of the underparts and somewhat on the head and shoulders.
Numerous examples taken from June 1 to 10 are all in fresh coat
except on the lower rump and tail. Skins collected July 15 to 30 are
in full summer coat. By August 10 there is much evidence of wear
over the forward half of body, and by early September the fall renewal
has commenced. As in the case of C. ludovicianus, this progresses
forward, and by September 25 to October 1 is complete.

Skull—tLarger than in C. parvidens or C. gunnisoni; otherwise
much like skull of parvidens but with narrower interorbital region.
Differs further from skull of gunnisoni in the less broadly spreading
maxillary arm of zygoma, large and flat mastoids, larger audital
bulle, and less rounded inferior rim of angle of ascending branch
of jugal; occiput viewed from behind broader and flatter. 7

Measurements.—Averages of 13 adult males from Wyoming: Total
length, 358 (340-370) ; tail vertebrae, 57 (44-60) ; hind foot, 62 (60-65).
Skull: Condylobasal length, 58.7 (56-61.3); zygomatic breadth, 43.8
(41.7-45.4); mastoid breadth, 29 (27.4-30); length of nasals, 22
(20.7-23.1); length of mandible, 43.1 (41.6-44.9); maxillary tooth
row, alveoli, 15.7 (15.1-16). For detailed measurements of speci-
mens see page 35.

Type specumen.—No. 186472, U. S. National Museum. Adult
female, skin and skull (teeth moderately worn). Collected Septem-
ber 15, 1888, by Vernon Bailey; original number 224 (Merriam
collection, skin 4668, skull 5319). In worn summer coat anteriorly
and in fresh winter coat posteriorly. |

Remarks.—In 1898, Dr. J. A. Allen! proposed that the name
Arctomys lewisii of Audubon and Bachman be revived for this species.
The specimen on which Arctomys lewisit was founded was recorded as
_ No. 461 of the Zoological Society of London collection. In reply to
my inquiry as to the possible existence of this type specimen in the
British Museum, Mr. Oldfield Thomas wrote, under date of January
6, 1915, that the specimen is still preserved, is numbered 55.12.24.144
of the Museum register, but that it is a species of Marmota, and not of
Cynomys as supposed by Dr. Allen.

The ranges of Cynomys leucurus and C. gunnisoni are not known
actually to meet, but the ranges of C. lewcurus and C. ludovicianus
do meet, and for a short distance overlap, at several points in Montana
and Wyoming. Both species are constantly enlarging their ranges,
pushing out into new valleys, and consequently frequently find
themselves in close proximity. Individuals of one species sometimes
wander into villages of the other. In the Bighorn Basin, in Montana,
Bailey found the species living in close proximity but always apart,

1 Bull. Amer. Mus. Nat. Hist., X, p. 456, 1898.

1916] - ; CYNOMYS PARVIDENS. 97

although there was no natural barrier. Cary found them intermin-
gling in colonies near Ishawooa, Wyoming, and Bailey captured a
specimen of leucurus in a “‘town”’ of ludovicianus in Clark’s Fork
valley, Montana.

Except for what appears to be a slight average increase in size in
the extreme southern portions of its range, the specimens of C.
_leucurus that have been assembled present a remarkably uniform
appearance, both in coloration and in skull characters. There has
been considerable misunderstanding in times past as to the western
limit of distribution of this species. Many writers believed that
its range extended to the Plains of the Columbia, and this mistake
is one of the principal reasons for past confusion in nomenclature.
As a matter of fact the animal is entirely unknown in Idaho, and
crosses the Bear River Divide to the Wyoming-Utah boundary at
only one place, so far as known. Regarding this western limit of
distribution, Dr. Merriam has given me an interesting note. He tells
of tracing the species, in 1913 and 1914, across the Utah-Wyoming
boundary not far from Evanston, westward into Utah for a distance
of six or seven miles where it stops abruptly.

Specimens examined.—Total number, 107, as follows:

Colorado: Beaver Creek to bridge over White River (Rio Blanco County), 2;!,2
Big Beaver Creek, 2;1 Buford, 1; Canadian Creek, 2; Coyote Basin, 1;!
Craig to Kelly’s (Moffat County), 4;1 Crawford, 1;1 Douglas Spring, 1;!
Escalante Hills, 1; Grand Junction, 17; Hayden, 1; Hebron, 1; Hell Creek
(Jackson County), 1;! Meeker, 4; North Park, 1; Sand Creek to Snake River,
21,2

Montana: Clark’s Fork, 1; Sage Creek, 2.

Utah: Uinta Mountains, 1;3 Uncompahgre Indian Reservation, 3.?

Wyoming: Bear Creek, 1; Bighorn Basin, 1; Big Piney, 1; Big Sandy, 1; Brid-
ger’s Pass, 1; Cheyenne (west of), 2; Cumberland, 2; Deer Creek, 2; Dubois,
1; Fontenelle, 1; Fort Bridger, 10 (including type); Fort Steele, 2; Fossil,
1; Garrett, 2; Green River, 2; Independent Rock (25 miles southwest of),
1; Ishawooa, 1; Lander, 3; Laramie Mountains, 2; Lost Cabin, 1; Medicine
Bow Mountains, 4; New Fork of Green River, 6; Otto, 1;2 Pole Creek, 1;
Shirley Mountains, 1; Spring Creek, 1; Sweetwater, 1; Woods Post Office, 5.

CYNOMYS PARVIDENS Aten.
Utan PrarrikE-Doe.
(PI ibe tic, 2-:Pl, IV, fie. 2; Pl. V, fig. 4; Pl. VII, fig: 4:)
1905. Cynomys parvidens Allen, Science Bull., Mus. Brooklyn Inst. Arts and Sci.,

I, no, 5, p. 119; March 31.

Type locality — Buckskin Valley, Iron County, Utah.
Geographic distribution.—Mountain valleys of central Utah in the
Sevier River region; south from Nephi to Iron and Garfield Counties.

1Collection E.R. Warren. ?Collection Amer. Mus. Nat. Hist. %Collection Mus. Comp. Zool.

28 NORTH AMERICAN FAUNA, [No. 40.

Characters.—In general like Cynomys leucurus but smaller and
with color of upperparts in summer and winter decidedly reddish or
rich cinnamon, less buffy or grayish; tail with terminal half white,
as in deucurus. Skull as in leucurus, but smaller, with greater in-
terorbital breadth.

Color.—Adult in late summer: Entire upperparts of head and body
cinnamon or dark buffy cmnamon, with very little admixture of buff
or blackish hairs; the individual hairs blackish at bases, then narrowly
pale buff, broadly cinnamon, and tipped with dark brown. Spot of
black above eye very sharply marked; cheek patch of brown distinct;
sides of nose pale buff; eye ring, ears, and limbs cinnamon; feet dark
buff; lips and chin whitish; underparts buffy or cinnamon. ‘Tail for
half its length above concolor with rump; below cinnamon; terminal
half all around white, without markings. Adult in early winter pelage:
In general like summer coat but with dense underfur which is black
at base, then light buff; upperparts rich cinnamon, the long hairs with
broad bands of black and buff; sides of body yellowish buff, distinctly
marked from color of back, and streaked by the bands of black on
the longer overlying hairs. _

Molt. —Owing to lack of Lapnacel collected at certain seasons, the
molt of C. parvidens is not well known. A single skin collected ; in
the Sevier National Forest, May 19, is still entirely in the old winter
coat. Specimens collected in late August and early September are
mostly in rather worn summer coat, with little indication of molt.
The tails are in freshly renewed pelage. Some specimens, taken the
first week in September, are well advanced in renewal of winter pelage, —
with full underfur and long, fresh coat. Evidently the transition from
summer to winter coat is gradual and obscure, without a distinct
molt, much as in the animals of the northern part of the ranges of
C. ludovicianus and C. leucurus. <A series of skins collected in the
Sevier National Forest, October 27 to November 3, are in full win-
ter coat. )

Skull and teeth.—Skull much hike that of C. leucurus, but smaller
and with greater interorbital breadth; postorbital processes less ab-
ruptly projecting. The teeth are not noticeably smaller.

Measurements.—Averages of 8 adult males from Utah: Total length,
338 (305-360); tail vertebra, 43 (80-57); hind foot, 59 (55-61).
Skull: Condylobasal length, 55.9 (53-57.9); zygomatic breadth, 42.2
(38.3-44.5); mastoid breadth, 28.6 (26.7-29.8); length of nasals, 21.8
(20.5-22.4); length of mandible, 42.3 (40.3-43.7); maxillary tooth
row, alveoli, 15 (14.8-15.4). For detailed measurements of specimens
see page 35.

Type specimen.—No. 28738, American Museum of Natural History,
New York. Adult not sexed, but probably a male; skin and skull

1916.] CYNOMYS GUNNISONI GUNNISONI. 29

(teeth moderately worn). Collected August 25, 1904, by George P.
Engelhardt, on the Brooklyn Institute of Arts and Sciences expedi-
tion of 1904 (Museum of Brooklyn Institute No. 437). Skin in good
condition, but not made up into a modern study specimen. It is in
worn summer pelage, but with freshly renewed coat on the tail.

Remarks.—The relationship of this distinct species with Oynomys
leucurus rather than with C. gunnisoni is clear. No evidence of inter-
gradation with leucurus is found in the material of each form which I
have examined, and every specimen of C. parvidens can instantly be
distinguished by its marked reddish coloration. The superficial re-
semblance to certain specimens of (. g. zuniensis is striking, but
the color of the tail and general skull characters are diagnostic.
Specimens of this prairie-dog from many parts of its range and
collected at all seasons, particularly early summer and late fall, are
much needed.

Specimens examined.—Total number, 46, as follows:

Utah: Buckskin Valley, 18 (including the type! and paratype); Sevier National
Forest, 28.

CYNOMYS GUNNISONIT (Barrp).

[Synonymy under subspecies. ]

Characters.—Smaller than Cynomys leucurus; larger than C. parvi-
dens; general coloration darker; tail gray, with white tip and white
border on the terminal half (in leucurus and parvidens the terminal
half of tail is entirely white). Skull smaller than in leucurus, larger
than in parvidens; differs from both in more broadly spreading maxil-
lary arm of zygoma; mastoids smaller and more obliquely placed,
rather than in general occipital plane; audital bulle smailer; occiput
viewed from behind higher and less broadened; inferior rim of angle
of ascending branch of jugal averaging less pointed, more rounded,
with little trace of special lateral flattened surface.

Subs pecies.—T wo subspecies of Cynomys gunnisoni are here recog-
nized. The typical race is confined to parts of Colorado and northern
New Mexico, while a paler desert form, C. g. zwniensis, ranges from
central New Mexico and southwestern Colorado to the San Francisco
Mountain region and to the Hualpai Indian Reservation, Arizona.

CYNOMYS GUNNISONI GUNNISONI (Barrp).
GUNNISON PRAtIrRIE-Doc.

(Pl. III, fic. 3; Pl. IV, fig. 3; Pl. V, fig. 5; Pl. VIL, figs. 3, 6.)

1855. Spermophilus gunnisoni Baird, Proc. Acad. Nat. Sci. Philadelphia, VIT, p. 334,
April. il

1857. Cynomys gunnisoni Baird, Gen. Rep. North Amer. Mamm., pp. XXV, 335.

1877. Cynomys gunnisoni Allen, Proc. Boston Soc. Nat. Hist., XVI, p. 294 (syn.).

1 Collection Amer. Mus. Nat. Hist.

380 NORTH AMERICAN FAUNA. [No. 40.

Type locality.—Cochetopa Pass, Saguache County, Colorado.

Geographic distribution.—Rocky Mountain region of central and
central-southern Colorado and northern New Mexico. North into
South Park, Colorado; east to El Paso, Fremont, and Huerfano
Counties, Colorado; south into the Sangre de Cristo and Jemez Moun-
tains, New Mexico; west to western Gunnison and Hinsdale Coun-
ties, Colorado, and to western Rio Arriba County, New Mexico.
Chiefly Transition Zone, but also in Cie Sonoran and lower parts
of Canadian Zone.

Characters.—Shightly smaller than Cynomys gunnisoni zuniensis,
with smaller hind foot; coloration more buffy and blackish, less pink-
ish cinnamon and ochraceous. General coloration much as in C.
leucurus, but averaging darker, and with only tip and border of ter-
minal half of tail white.

Color.—Adult wn Fresh summer pelage: General coloration, except
for tail, very much as in C. leucurus, but darker; averaging more
drab, less yellowish, and the cheeks less sharply marked by dark
patch. Tail for proximal half above concolor with back; distal half
dark gray, bordered and tipped with white, and with subterminal
band of blackish; underside of tail pale cinnamon, blending into
white at tip, and edged with white. The fresh spring coat, which is
glossy and much intermixed with blackish, rapidly turns a dull
brown, the jet black hair tips having worn away or faded. Winter
pelage, except for the tail, much like that of leucurus.

Molt.—There is a variation of about six weeks in the time of molt,
according to altitude, but the average dates in Colorado are about as
follows: Spring specimens are in the left-over winter coat or in various —
degrees of molt up to June 10 or July 1, when the pelage, with the
exception of the tail, is entirely renewed. A fews pecimens have com-
pletely renewed the pelage on the tail as early as July 15, but numer-
ous examples collected as late as August 10, still have the tail clothed
in the old winter hair. The fresh coat rapidly loses its luster, and
during the latter half of August the pelage becomes much worn over
the anterior half of the body while the posterior half shows consider-
able influx of new hair. The molt is usually complete and the animal
in full fresh coat again by September 15. Mid-October skins are.
in full long coat, with a dense underfur.

Skull.—Averaging smaller and of less robust build than skull of
C.g. zuniensis. Readily distinguishable from skulls of C. leucurus
by its considerably smaller size (see also specific skull characters,
page 29).

Measurements.—Averages of 12 adult males from Saguache County
and Fort Garland, Colorado: Total length, 340 (809-365); tail verte-
bree, 53 (39-65); hind foot, 56 (52-59). Skull: Condylobasal length,

1916.] CYNOMYS GUNNISONI GUNNISONI, 31

54.8 (52.4-56.5); zygomatic breadth, 42.5 (40.7-44.2); mastoid
breadth, 27 (25.7-28.2); length of nasals, 20.5 (18.9-22.3); length of
mandible, 41.9 (40.9-43.3); maxillary tooth row, alveoli, 14.5 (14-15).
Females average slightly smaller. For detailed measurements of
specimens see page 35.

Weight.—An adult male collected September 6 in the Jemez Moun-
tains, New Mexico, weighed 14 pounds.
* Type specimen.—No. 1636, U.S. National Museum. Skull of young
adult, not sexed (teeth little worn). Collected in September, 1853,
by F. Kreutzfeldt; original number 22. The museum catalogue
shows that the skin (No. 501) was formerly preserved in the collec-
tion, but it has been lost for many years.

Remarks.—So far as known the ranges of Cynomys gunnisoni and
O. leucurus do not actually meet at any point, although, as stated by
Cary,? they are separated by only a very narrow strip in the Cimarron
region. Though so much alike in general characters the two species
seem perfectly distinct, with no evidence of intergradation. While
chiefly a Transition Zone form, gunnisoni is common and at home in
the Upper Sonoran valleys within its range, as throughout suitable
places in the upper Rio Grande and San Luis Valleys, Colorado.
The range of gunnisoni nearly if not quite meets that of the black-
tailed prairie-dog at several points in the eastern foothill region of
Colorado. Prof. Lantz noted a colony of gunnisoni in South Park,
Colorado, in which black and partly black individuals were common,
a dozen or more melanistic specimens being seen.

This dark form of C. gunnisoni intergrades directly with the paler
race, next described, of the more arid parts of New Mexico and Ari-
zona, but the two subspecies are well marked and the area of intergra-
dation is small.

Specimens examined.—Total number, 111, as follows:

Colorado: Antonito, 2; Cascade, 1; Castrel’s Ranch (Park County), 2;3 Cochetopa
Pass, 30 (including type skull); Colorado City (northwest of), 3; Colorado
Springs (west of), 1; Divide (Teller County), 2;* Florissant, 2;+ Fort Garland,
20 (including some older skins labeled ‘‘Fort Massachusetts”); Fremont
County, 2;> Head of Rio Grande, 1;° Pike’s Peak, 3; Poncha Pass, 2;* Querida,
2;* Saguache, 4; Salida, 5;* ® Southfork, 2; Tarryall Creek (Park County), 1;+
Twin Creek, 1;3 Twin Lakes, 1; Wagon Wheel Gap, 12.%: +6

New Mexico: Costilla Pass, 3; Coyote Creek, 2; Gallinas Mountains, 1; Jemez
Mountains, 1; La Jara Lake, 1; Moreno Valley, 2; Tres Piedras, 2.

1 See Lyon and Osgood, Bull. 62, U. S. Nat. Mus., pp. 161-162, 1909.
2 North Amer. Fauna No. 33, p. 95, 1911.

3 Collection Mus. Comp. Zool.

4 Collection E. R. Warren.

5 Collection Field Mus. Nat. Hist.

6 Collection Amer. Mus. Nat. Hist.

32 NORTH AMERICAN FAUNA, [No. 40.

CYNOMYS GUNNISONI ZUNIENSIS sussp. nov.
ZuNi Prarrie-Doe.
(Pl. IIT, fig. 4; Pl. IV, fig. 4; Pl. V, fig. 6.)

Type locality Wingate, McKinley County, New Mexico.

Geographic distribution.—Southwestern Colorado, extreme south-
eastern Utah, northwestern and west-central New Mexico, and north-
central Arizona. North in western Colorado to Montrose County;’
northeast in New Mexico in the Rio Grande Valley to Espanola and
east to Pecos and the Manzano Mountains; south on the west side of
the Rio Grande Valley to Sierra and Socorro Counties, New Mexico;
west in central Arizona to Prescott and the Hualpai Indian Reserva-
tion. Chiefly Upper Sonoran, but also in Transition Zone.

Characters.—Slightly larger than Cynomys gunnisont gunnisoni,
with larger hind foot and heavier skull. Coloration less buff and
blackish, more pinkish cinnamon and ochraceous. |

Color.—Adult wm fresh summer pelage: General color of upper-
parts pale cinnamon, heavily streaked with lighter cinnamon-buff
and blackish; individual hairs blackish at bases, then narrowly buff,
then broadly cinnamon, with subterminal band of buff, and tip of
blackish. Nose, lips, eye ring, and ears, buff; between eyes, between
ears, and over nape darker brown, mixed with blackish; large area of
blackish brown on cheek, and a smaller, sharply marked, blackish
brown spot above eye; feet, limbs, and underparts pale cinnamon-buff.
Basal half of tail above concolor with back; distal half mixed gray
and white, bordered and tipped with clear white. The black tips
to the hairs of the back rapidly wear away and the pelage becomes
a uniform dull cinnamon, mixed with light buff. The distinctive
markings of the head become more obscure. Adult in fresh full winter
pelage: General color more grayish and buff, less cinnamon, than in
summer, with greater admixture of blackish hairs on head and more
of white and gray on body. General color of upperparts dark cream-
buff, heavily mixed with pale buff and blackish, giving a decided
grizzled appearance; underfur narrowly black, then broadly whitish;
the long hairs with bands of black, pale cinnamon, and white, and
narrow tips of black. Top of head and cheeks heavily intermixed with
blackish, the spot over eye sometimes intense black; sides of nose,
eye ring, ears, limbs, and sides of body rich cream-buff; lips and throat
paler; underparts pale cinnamon. Tail above mixed gray and white,
with a narrow sub-border of dark gray and border of clear white;
below clear pale cinnamon, terminal half with faint sub-border of
grayish brown and border and tip of creamy white. Young examples
in the fresh postjuvenile pelage are more vinaceous or pinkish cin-
namon than the corresponding pelage of C. g. gunnisoni.

1916.] CYNOMYS GUNNISONI ZUNIENSIS. 33

Molt.—The molt of C. g. zwniensis is much more conspicuous than
in the northern subspecies. May specimens are frequently in a
very ragged state, with very little indication of renewal. Speci-
mens in complete summer coat, including the tail, are rarely taken
before July 1. The summer coat frequently is so rapidly worn and
bleached as to change the entire appearance of the animal before
there are any traces of the fall pelage. The new pelage of autumn
first appears in adults over the posterior half of the body about
September 1. No specimens in full winter coat have been seen
which were killed earlier than November 1.

Skull—As in C. g. gunnison but averaging larger and more
heavily built.

Measurements.—Type specimen: Total length, 363; tail vertebre,
53; hind foot, 60. Skull: Condylobasal length, 57.9; zygomatic
breadth, 43.8; mastoid breadth, 29.1; length of nasals, 22.4; length
of mandible, 43; maxillary tooth row, alveoli, 15.6. Averages of 9
specimens from Arizona: Total length, 355 (330-373); tail verte-
bre, 62 (54-68); hind foot, 60 (57.5-62.5). Skull: Condylobasal
length, 55.8 (51.9-57.8); zygomatic breadth, 43.5 (39.7-45.5); mas-
toid- breadth, 27.4 (25.7-28.6); length of nasals, 21.9 (20.6—-23.5);
length of mandible, 42.7 (40.3-44.2); maxillary tooth row, alveoli,
14.5 (14.2-14.8). For detailed measurements of specimens see
page 35.

Weight—An adult male collected at Acoma, New Mexico, Sep-
tember 28, weighed 2 pounds.

Type specuomen.—No. 137555, U. S. National Museum, Biological
Survey collection. Adult male, skin and skull (teeth considerably
worn). Collected at Wingate, New Mexico, June 26, 1905, by N.
Hollister; original number 2374. Except on the tail the skin is in
fresh summer pelage.

Remarks.—This well-marked-color subspecies of Cynomys gun-
nisoni occupies the arid southwestern part of the range of the species
and is the common prairie-dog of large parts of New Mexico and Ari-
zona. Material from southwestern Colorado is not plentiful, but the
specimens examined indicate that the range of the pate form extends
into that portion of the State. Specimens are wanting from the
near-by section of Utah, where this form undoubtedly occurs.
Specimens from Flagstaff, Arizona, and the vicinity of San Francisco
Mountain in general are slightly darker, more drab, than skins from
eastern Arizona and western and central New Mexico, but on the
whole the form is very uniform in coloration throughout its range.
While in the main an Upper Sonoran form, this prairie-dog is gener-
ally distributed in the Transition Zone parks of numerous mountain

20651°—16——3

34 NORTH AMERICAN FAUNA, [No. 40.

ranges. In the Bear Spring Mountains, New Mexico, in 1905, I saw

one nearly pure-white animal of this species. Bailey reports finding

a mixed colony of individuals of this form and of C. ludovicianus

near Pecos, New Mexico, and he captured specimens of each form

at San Pedro, but the ranges of the two species rarely overlap.
Specomens ecamined.—Total number, 96, as follows:

Arizona: Agua Fria, 4;' Aubrey Valley, 1; Clarks Valley, Mogollon Mountains,
2;1 Flagstaff, 7; Keams Canyon, 1; Kendrick Park, 4; Little Spring, 6;
Mormon Lake, 5; San Francisco Mountain, 7; Spaulding’s Ranch, Agua
Fria Valley, 1;! Springerville, 18, Winslow, 2.

Colorado: Bedrock, 2;? Cortez, 1;1 Coventry, 1.?

New Mexico: Acoma, 1; Albuquerque, 7; Chusca Mountains, 2; Espanola, 2;
Fairview, 2; Fruitland, 1; Liberty, 1; Magdalena, 3; Manzano Mountains,
1; Ojo Caliente, 2; Pecos, 3; Rio Alamosa, 3; San Augustine Plain, 2; San
Mateo Mountains (Socorro County), 1; San Pedro, 1; San Rafael, 1; Thoreau,
2; Wingate, 3; Zuni Mountains, 1.

TABLES OF MEASUREMENTS.

Adult male specimens of Cynomys.

: Skin. Skull.
z
pe lalé q |2 me eae
Species and locality. a | Be eo] Bailes ea salen on aalar
qd.) 2 | | 2 |2h) 8s) saiee
o ra P [og [oe Peel ce alee
= oH a an Ores = =
C. ludovicianus ludovicianus.
Montana:
Shelby Junction.......-..- 66690 | 375 72 55 | 59.1 | 44.7 | 45.5 | 16.2 | Moderately worn. ’
WD) OF eee stare Soca . 66691 374 85 60 | 59.0 | 44.4 | 45.3 | 15.9 Do.
Darnall’s Ranch.........-- 169447 410 95 65 | 62.5 | 46.6 | 47.7 | 16.9 Do.
North Dakota: Medora....-.... 161314 | 405 96 64 | 61.4 | 45.5 | 45.9 | 16.9 | Much worn.
South Dakota:
HOT ERICEeLe ase cee eee eee 191426 373 76 62 | 61.0 | 45.2 | 45.8 ; 16.8 | Considerably worn.
ButialovGapesece. oo. eee 191428 384 90 63 | 60.8 | 43.8 | 46.4 | 16.8 | Moderately worn.
IB) Opes ars eres 191429 360 84 61 | 59.5 | 42.9 | 45.0 | 16.6 Do.
Wyoming:
Ishawooa......-..-2.---- -..| 169493 396 93 64 | 60.6 | 45.2 | 45.8 | 17.0 Do.
INieWeAaStleS. 22.55 se5ssceese 65894 | 390 79 62 | 60.9 | 44.2 | 46.7 | 16.8 Do.
Nebraska:
Keennediysseease sen seen | 191427 392 78 57 | 61.4 | 44.8 | 46.1 | 16.8 Do.
ND) Oe et oe sey eons 18678 390 92 63 | 59.2 | 45.7 | 44.8 | 16.5 Do.
PATIN Bee see ca ncis ee oe ee 110770 409 89 65 |} 62.1 |.47.0 | 48.1 | 17.2 Do.
Colorado:
Woveland.. 2242.2. tascus. o~ 69521 370 82 62 | 60.2 | 45.5 | 46.0 | 17.1 | Considerably worn.
OMGY 2 5225 sos comes scarce 69952 379 83 61 | 59.6 | 44.1 | 45.5 | 17.0 | Moderately worn.
DD ORs ee eee -.| 70020 355 82 61 | 57.2 | 43.1 | 43.9 | 16.9 Do.
Kansas: Pendennis. ....... eel eS 762ml Sil 81 62 | 60.3 | 47.2 | 46.2 | 16.9 | Considerably worn.
Oklahoma:
IMO UMTS COlbeeee nese os seer 132440 | 375] 115 63 | 61.5 | 44.2 | 47.2 | 17.3 } Moderately worn.
ID) Oe Ne eae oe ee ee | 132441 376 104 63 | 59.6 | 44.8 | 45.9 | 17.0 Do.
Texas
WiGETI OTS ey sents eee te ee 64783 | 365 82 61 | 60.6 | 44.4 | 46.4 | 16.8 | Considerably worn.
1D) ORE, Sees pee eee ye | 64784 413 | 93 60 | 63.6 | 48.1] 47.3 | 16.6 Do.
Henrietta........:. -.! 64780! 386! 87 66 | 62.0 | 45.3 | 48.4! 17.0 | Moderately worn.

1 Collection Amer. Mus. Nat. Hist. 2 Collection E. R. Warren.

1916.] TABLES OF MEASUREMENTS. 35

Adult male specimens of Cynomys—Continued.

: Skin. Skull.
5
2
eee % lz Slige er
Species and locality. = to | 2 es re . | ae | Condition of molar
a = S 2g = © Ho teeth.
& © a } St | 8c Wat Sy
3 ras 2 = Sceito. sae = lire
o oS ue ue} omy) ope iS wo
3 Ss 4 R=! =| = a seo
= = ~ joo et) N a le
C. ludovicianus arizonensis.
Texas
IMIG Uda seine es a. le 23183 | 398 | 96] 64] 61.5 | 45.0] 46.0 | 16.7 | Moderately worn.
OSS6 Doon eGo eee 23184 400 92 62 | 60.8 | 46.2 | 47.5 | 16.7 Do.
IDO. Cakeensee eee 23396 404 89 67 | 61.7 | 46.4 | 47.4 | 16.8 | Much worn.
New Mexico:
Siliverd@ityiee ss asces as. - 51279 | 380 79 61 | 60.9 | 47.0 | 45.0 | 16.1 Do.
DogiSprings S52. 2:2...- 20380 412 94 66 | 62.2 | 44.3 | 45.7 | 16.9 Do.
Animas Valleys. is--2<2.-: 58920 | 395 83 64 | 59.9 | 44.5 | 44.2 | 16.0 Do.
Chihuahua: Sierra en Media...! 99362 | 400 80 65 | 62.9 | 45.0 | 46.0 | 16.6 Do.
Arizona:
Honteluachucass: +... 34029 | 412 97 60 | 61.8 | 45.2 | 46.8 | 16.6 | Considerably worn
ID) OM tee ee. ee 34030 | 410} 100 64 | 62.8 | 45.0 | 44.9 | 16.1 | Much worn,
ID) Oa Pte eae ee siecocsic cit 34031 400 93 62 | 62.3 | 44.1 | 45.9 | 16.5 Do.
C. mexicanus.
Coahuila:
MaaVienturass hens s 1 26423 419 107 63 | 59.2 | 45.6 | 43.9 | 16.3 Moderately worn.
ID) OMe cee 26426 430 106 64 | 59.9 | 45.0 | 48.9 | 15.5 Do.
DOM eee Se 26428 | 415 | 103 61 | 59.5 | 44.7 | 44.0 | 15.9 Do.
ID) OP ees i See 26429 415 102 59 | 58.5 | 43.0 2.8 | 1528 Do.
MIB) Co Fapae Napa ee a 79546 415 115 64 | 58.7 | 44.4 | 45.8 | 15.4 Do.
C. leucurus.
Wyoming:
SH AW0 Oa ee see 169336 370 60 60 | 60.3 | 45.4 | 44.3 | 15.6 | Much worn.
Spring Creek... 22.2.2. ..- 56002 | 363 60 60 | 58.7 | 43.6 | 43.0 | 15.4 | Moderately worn.
iHontenellerss es -2--.. /.- 147187 | 370 48 62 | 61.3 | 45.0 | 44.9 | 16.0 | Much worn.
Cuamiberlandes 22-22) 22.2 179515 | 368 60 63 | 59.9 | 44.4 | 44.6 |] 15.9 Do.
HOLS teeleme cs le 171457 367 52 61 | 59.2 | 44.9 | 44.1} 15.3 Do.
Colorado:
Escalante Hills...-......:. 148145 | 382 65 67 | 59.4 | 44.8 | 45.8 | 16.1 | Moderately worn.
Grand Junction... 5... f- 54111 | 361 55 64 | 59.9 | 44.7 | 44.0 | 15.6 | Considerably worn.
C. parvidens.
Utah:
Pucslin Walleye... io. 158985 | 337 57 60 | 54.5 | 40.7 | 42.2 | 14.9 | Moderately worn.
eC B UE ee 158992 326 57 59 | 54.1 | 40.7 | 41.8 | 14.8 ; Do.
Peers eee fice Sse 158993 305 52 57 | 53.0 | 38.3 | 40.3 | 14.8 | Little worn.
Sevier National Forest 206323 | 335 31 55 | 57.9 | 43.9 | 42.2 | 15.0 » Moderately worn.
Ose acco uae 206337 354 49 60) 1-57.16 21-4352) |) 42-91) 1522 Do.
C. gunnisoni gunnisoni.
Colorado:
pockeons TEES. ae eee 35845 | 343 52 53 | 53.8 | 40.7 | 41.1 | 14.7 | Moderately worn.
Smee ye sin coo 35849 | 323 Al 52 | 52.4 | 41.5 | 40.9 | 14.0 | Considerably worn.
Do Scant 35854 | 330 45 56 | 54.9 | 42.6 | 42.5 | 14.7 | Moderately worn.
13) One 35855 | 332 45 56. 1-54.29 | 42:5 | 42.5) 14:6 Do
HTD) Oar ea 1 35856 331 2 55 | 54.4 |. 42.7] 41.4 | 15.0 | C onsiderably worn.
Mew Mexico: La Jara Lake....| 134908 | 328 68 54 | 53.4 | 41.3 | 40.0 | 14.8 Do.
C. gunnisoni zuniensis.
New Mexico:
MN ONGAUM ES: a 221055 cece aces 160752 | 340 60 57 | 53.8 | 41.1 | 41.6 | 14.5 | Considerably worn.
WWalnt Pate cate eee fe. 1137555 | 363 53 60 | 57.9 | 43.8 | 43.0 | 15.6 | Much worn.
PAICOMMA see eee sa: 148290 | 360 60 62 | 55.9 | 48.5 | 44.5 | 15.4 | Moderately worn.
Mao d alen ase ore a. 167574 | 350 60 61 | 54.7 | 43.7 | 42.0 | 15.0 | Much worn.
Ojowaltentes.= =. 0222 4-.--. 167021 | 350 49 60 | 57.6 | 43.0 | 42.2 | 15.1 | Considerably worn.
Arizona: j
Springervalleses. 3. 5 h..52 5. 24653 | 355 63 60 | 55.2 | 48.4 | 42.8 | 14.7 | Moderately worn.
WW ORE ee ee eee 24654 357 7 60 | 55.0 | 48.2 | 41.7] 14.€ Do.
IRAE S Catlepere eye nae ae 201638 | 356 64 59 | 56.4 | 44.0 | 42.7 | 14.3 ; Much worn.
Kendrick Park... .-.--.... 202942 373 68 60 | 57.8 | 44.5 | 44.2 | 14.8 0.
MALLE SDE Ge s 5. Shee cL 202556 | 363 54 58 | 55.9 | 45.0 | 42.9 | 14.4 | Considerably worn.

3 6 NORTH AMERICAN FAUNA, [No. 40.
Averages of adult male specimens of Cynomys.
aie Skin, Skull.
Ets!
=
- a . xe Lom ° ko) Pp
; Se eee Eesti aus
Species and locality. i Be 2 = Bales) ecs Bai Sas
Hun | o 6 | Sho | so ec é Taya Re
Sais S st ||| Ss cleave! coaa| seca coal eral ayes
ae] is OS |Oo ! om x | 3 tS | tah
Lame emer eSecie = | 2 |e toe
7, = a eS N =) Z Sp ty
C. ludovicianus ludovicianus.
IMOn tana terce asec nce ee necro ee 6 | 383 84 61 | 60.4 } 45.1 | 28.0 | 22.8 | 45.8} 16.4
INOTGheD Akola ce s..o eee see ce eerie ee 1 405 96 64 | 61.4 | 45.5 | 28.1 | 23.1 | 45.9] 16.9
South Dakotase-sess- ee oe eee 131388 86 62 | 59.8 | 45.0 | 28.5 | 23.8 | 45.9] 16.6
Way ONIN SS 258 5< cretion Sat nercceeke 4 | 393 86 63 | 60.7 | 45.3 | 28.3 | 24.0 | 46.2] 16.8
ING DEAS Kans: Bnctacee eee ers ere 3 397 86 62 | 60.9 | 45.8 | 28.9 | 23.4 | 46.3 16.8
ColoradO nes ae seee eee eee acne 4] 364 82 62 | 58.9 | 44.1 | 27.8 | 23.1 | 45.1 16.9
KANSAS a aoce nse tee ek soar oie meee 10 380 88 624-597 | 4457 20S p23 |k45..5 16.5
Okishomawk es ..-.42 Sceeeeee case eee 2 376 110 63 | 60.6 | 44.5 | 28.3 | 23.5 | 46.6 W574
MOXAS Ait oe cence so eeecinee ne nee Sosa T2\=°392 88 62 | 61.5 | 45.5 | 29.0 | 24.5 | 46.9] 16.7
C. ludovicianus arizonensis.
MORAGH ee aamcje ele see oie Boece nee eee 7 | +395 88 64 | 60.3 | 44.7 | 27.7 | 24.3 | 46.5] 16.6
INOW MGXICOl 2. acces Saee eele Sate eee Ls 381 84 62 | 60.1 | 44.3 | 28.0 | 23.8 | 45.0 16.2
IATIZ ONAEe aot estes ee eee cere see ser 19 388 89 62 | 60.0 | 44.0 | 27.3 | 24.0 | 45.2 16.2
Chihviahwaec cas semecer ete eae sceneries 1 | 400 80 65 | 62.9 | 45.0 | 29.1 | 23.5 | 46.0 | 16.6
C. mexicanus.
Coahttilaces..chie sscstee see cc ae cee 8} 416} 102 63 | 59.3-| 44.6 ; 28.6 | 23.0 | 43.8 | 15.9
C. leucurus.
WyOMlin eo ).6: cece ona tose ee 13.| 358 | — 57 | 62 | 58.7 (4328 | Qos o2NO RA Stel e527
Colorad Oss seen sree ore ee ee eee ee 6 | 368 56 63 | 60.0 | 44.6 | 29.8 | 22.6 | 45.0} 15.6
C. parvidens.
Gee era ee es ey ee 8 338 43 59 | 55.9 | 42.2 | 28.6 | 21.8 | 42.3 15.0
C. gunnisoni gunnisoni.
COlOrad Dee ae eee nee ae eee 12 340 53 56 | 54.8 | 42.5 | 27.0 | 29.5 | 41.9 14.5
INewANhlexiCO asso. ce sects = coce seems i 328 68 54 | 53.4 | 41.3 | 25.8 | 21.5 | 40.0 14.8
C. gunnisoni zuniensis.
New Mexico and eastern Arizona.....--- 17 | 344 60 58). 55.00], 41.9: | 2764) a2 on |e425.3 14.8
Arizona: San Francisco Mountain....... 8) 355 62 60> |) 55./8 4355 de27e 4s QI OAS Ay 14S
PLrArE- 1,

Figs. land 4. Cynomys ludovicianus ludovicianus (Ord).

(No. 191411, U. S. Nat. Mus.)

Figs. 2 and 5. Cynomys ludovicianus arizonensis Mearns.

(No. 35873, U..S.-Nat: Mus.)

Arizona.

(About natural size.)

Fig. 3. Cynomys mexicanus Merriam.
79550, U. S. Nat. Mus., Biological Survey collection.)

3 ad.

La Ventura, Coahuila, Mexico.

g ad. Glendive, Montana.

g ad. San Pedro River,

(No.

North American Fauna No. 40, U. S. Dept. Agr. Biological Survey.

PLATE Il.

B2112—129
SKULLS OF CYNOMYS.

1, 4. C. 1. ludovicianus,
2, 5. C. 1. arizonensis.
&. C. mexicanus,

Piate II,
(About natural size.)

Fig. 1. Cynomys ludovicianus ludovicianus (Ord). ¢& ad. Glendive, Montana.
(No. 191411, U. S. Nat. Mus.)

Fig. 2. Cynomys ludovicianus arizonensis Mearns. og ad. San Pedro River, Arizona.
(No. 35873, U. 8. Nat. Mus.)

Fig. 3. Cynomys mexicanus Merriam. ¢ ad. La Ventura, Coahuila, Mexico. (No.
79550, U. S. Nat. Mus., Biological Survey collection.)

Fig. 4. Cynomys ludovicianus ludovicianus (Ord). @ ad. Glendive, Montana. (No.
191407, U. S. Nat. Mus.) : .

Fig. 5. Cynomys mexicanus Merriam. @ ad. La Ventura, Coahuila, Mexico. (No.
79554, U. S. Nat. Mus., Biological Survey collection.)

North American Fauna No. 40, U.S. Dept. Agr. Biological Survey.

B2113—129

SKULLS OF CYNOMYS.

1, 4. C. 1. ludovicianus.
2. C. 1. arizonensis.
3, 5. C. mexicanus,

Puate IIT.

(About natural size.)

Fig. 1. Cynomys leucurus Merriam. $ ad. Cumberland, Wyoming. (No.
179515, U. 8. Nat. Mus., Biological Survey collection.)

Fig. 2. Cynomys parvidens Allen. 9 ad. Buckskin Valley, Utah. (No. 158994,
U.S. Nat. Mus., Biological Survey collection.)

Fig. 3. Cynomys gunnisoni gunnisoni (Baird). & ad. Saguache, Colorado. (No.
176695, U. S. Nat. Mus., Biological Survey collection.)

Fig. 4. Cynomys gunnisoni zuniensis Hollister. ¢ ad. Wingate, New Mexico.
(Type, No. 187555, U. 8. Nat. Mus., Biological Survey collection.)

North American Fauna No. 40, U.S. Dept. Agr. Biological Survey.

PLATE Ill.

a
q

B2114—129
SKULLS OF CYNOMYS.

1. C. leucurus,. . guNnNisoni.
2. C. parvidens,

3 g
4. C. g. zuniensis.

PLATE LY.

(About natural size.)

Fig. 1. Cynomys leucurus Merriam. § ad. Cumberland, Wyoming. (No.
179515, U. S. Nat. Mus., Biological Survey collection.)

Fig. 2. Cynomys parvidens Allen. @ ad. Buckskin Valley, Utah. (No. 158994,
U.S. Nat. Mus., Biological Survey collection.)

Fig. 3. Cynomys gunnisoni gunnisoni (Baird). ¢ ad. Saguache, Colorado. (No.
176695, U. S. Nat. Mus., Biologicait Survey collection.)

Fig. 4. Cynomys gunnisoni zuniensis Hollister. g ad. Wingate, New Mexico.
(Type, No. 187555, U. S. Nat. Mus., Biological Survey collection.)

North American Fauna No. 40, U.S. Dept. Agr. Biological Survey. PLATE IV.

B2115—129

SKULLS OF CYNOMYS.

1. C. leucurus. 3. C. g. gunnisoni.
2. C. parvidens, 4. C. g. zuniensis.

PLATE V.
(About natural size.)

Fig 1. Cynomys ludovicianus ludovicianus (Ord). ¢ ad. Glendive, Montana. (No.
191411, U. 8. Nat. Mus., Merriam collection, original number, 4454.)

Fig. 2. Cynomys ludovicianus arizonensis Mearns. ad. San Pedro River, Arizona.
(No. 35873, U.S. Nat. Mus.)

Fig. 3. Cynomys mexicanus Merriam. ¢ ad. La Ventura, Coahuila, Mexico. No.
79550, U. S. Nat. Mus., Biological Survey collection.)

Fig. 4. Cynomys parvidens Allen. @ ad. Buckskin Valley, Utah. (No. 158994,
U.S. Nat. Mus., Biological Survey collection.)

Fig. 5. Cynomys gunnisont gunnisoni (Baird). ¢& ad. Saguache, Colorado. (No.
176695, U. S. Nat. Mus., Biological Survey collection.)

Fig. 6. Cynomys gunnisoni zuniensis Hollister. g ad. Wingate, New Mexico.
(Type, No. 1387555, U. S. Nat. Mus., Biological Suvery collection.)

PLATE V.

Biological Survey.

North American Fauna No. 40, U. S. Dept. Agr.

"SISTOLUNZ

6t1-91ItE

i
Fs)

Toh ty

TUOSTUUNS

3 °O

‘suoprAred *O ‘fF
"SAWONAQ 4O STINUS

‘BSNUBOTXOUL *(—) “§

*‘sISUOUOZT

Iv

I

-

\

7

G

‘SNUBIDIAOPHE “TO

|

PLATE -VI.
(About natural size.)

Figs. 1 and 2. Cynomys ludovicianus ludovicianus (Ord). & ad. Mason, Texas.
(No. 191388, U. S. Nat. Mus.)

Figs. 3 and 4. Cynomys ludovicianus ludovicianus (Ord). & ad. Mason, Texas.
(No. 191389, U. S. Nat. Mus.)
Selected skulls of same sex and age from one locality; to show the great individual

variation frequently noted in skulls of Cynomys.

North American Fauna No. 40, U. S. Dept. Agr. Biological Survey. PLATE VI.

SKULLS OF CYNOMYS LUDOVICIANUS LUDOVICIANUS.

PLATE Ville
(About natural size.)

Fig. 1. Cynomys ludovicianus ludovicianus (Ord), left mandibular ramus. 9? ad.
Glendive, Montana. (No. 191407, U.S. Nat. Mus.)
Fig. 2. Cynomys mexicanus Merriam, left mandibular ramus. @ ad. La Ventura,
Coahuila, Mexico. (No. 79554, U. 8S. Nat. Mus., Biological Survey collection.)
Fig. 3. Cynomys gunnisoni gunnisoni (Baird), left mandibularramus. gad. Sagua-
che, Colorado. (No. 176695, U. S. Nat. Mus., Biological Survey collection.)

Fig. 4. Cynomys parvidens Allen. @ ad. Buckskin Valley, Utah. (No. 158994,
U.S. Nat. Mus., Biological Survey collection.)

Fig. 5. Cynomys leucurus Merriam. ¢ ad. Cumberland, Wyoming. (No. 179515,
U.S. Nat. Mus., Biological Survey collection.) .

Fig. 6. Cynomys gunnisoni gunnisoni (Baird). ¢ ad. Saguache, Colorado. (No.
176695, U. S. Nat. Mus., Biological Survey collection.)

Figs. 7 and 8. Cynomys ludovicianus ludovicianus (Ord). @ juv., showing milk pre-
molars and unworn crowns of molars. Ishawooa, Wyoming. (No. 169674,
U.S. Nat. Mus., Biological Survey collection.)

North American Fauna No. 40, U. S. Dept. Agr. Biological Survey PLATE VII.

B2118-129
SKULLS OF CYNOMYS.

C. parvidens.
C. leucurus.

1, 7, 8. C. 1. ludovicianus, 4,
2. C. mexicanus. De
3, 6. C. g. gunnisoni,

20651°—16——-4

LANG

ax

{New names in bold=face type; synonyms in italics.]

Acknowledgments, 10.
Arctomys latrans, 8, 14.
columbianus, 8.

griseus, 14.

lewisii, 26.

ludoviciana, 10, 14.

ludoviciani, 14.

ludovicianus, 7, 14.

missouriensis, 14.

missuriensis, 14.
arizonensis, Cynomys, 8, 19-21.

cinereus, Cynomys, 14.
Citellus, 8, 10, 11.
fulvus, 11.
Colobotis, 11.
columbianus, Arctomys, 8.
Cynomys, 8, 24.
Cynomis, 10.
griseus, 14.
socialis, 14.
Cynomomus, 10.
Cynomys (genus), 5, 8, 10, 11.
(subgenus), 12-23.
Cynomys arizonensis, 8, 19-21.
cinereus, 14.
columbianus, 8, 24.
grisea, 8, 14.
griseus, 14,
gunnisoni, 6, 10, 24, 29-31.
gunnisonii, 29.
leucurus, 6, 8, 10, 24-27.
lewisi, 25.
lewisii, 25
ludovicianus, 8, 10, 14-19.
mexicanus, 8, 21-23.
parvidens, 27-29.
pyrrotrichus, 14, 17, 18.
socialis, 8, 10, 14.
zuniensis, 32-34.

Distribution, 5.

Food, 7.
grisea, Cynomys, 8, 14.
griseus, Arctomys, 14.
Cynomis, 14.
Cynomys, 14.
Spermophilus, 14.
gunnisoni, Cynomys, 6, 10, 24, 29.
Cynomys gunnisoni, 29-31.
Spermophilus, 8, 29.
gunnisonii, Cynomys, 29.

Habits, 6.

©

latrans, Arctomys, 8, 14.

lewisi, Cynomys, 25.

lewisti, Arctomys, 26.
Cynomys, 25.

Leucocrossuromys (subgenus), 5, 23-34.

leucurus, Cynomys, 6, 8, 10, 24-27.

ludoviciana, Arctomys, 10, 14.
ludoviciani, Arctomys, 14.
ludovicianus, Arctomys, 7, 14.
Cynomys, 8, 10, 14.
Cynomys ludovicianus, 14-19.
Spermophilus, 14.

Mamcynomiscus, 10.

Marmot, Columbia, 17.
Louisiana, 7, 17.

Marmota, 11.

Measurements, tables of, 34-36.

mexicanus, Cynomys, 8, 21-23.

missouriensis, Arctomys, 14.
Monaz, 8, 10, 14.

missuriensis, Arctomys, 14.
Monaz, 14.

Molt, 9.

Monar, 10.
missouriensis, 8, 10, 14.
missuriensis, 14.

Natural enemies, 7.
Nomenclature, 7-9.

parvidens, Cynomys, 27-29.
Pelage, 9.

Petit chien, 5, 8.
Prairie-dog, Arizona, 19-21.
black-tailed, 14-19.

Gunnison, 29-31.
Mexican, 21-23.
Utah, 27-29.
white-tailed, 24-27.
Zuni, 32-34.
pyrrotrichus, Cynomys, 14, 17, 18.

socialis, Cynomis, 14.
Cynomys, 8, 10, 14.
Specimens examined, 10.
Spermophilus griseus, 14.
gunnisoni, 8, 29.
ludovicianus, 14.
Squirrel, barking, 5, 8.
burrowing, 8.
prairie, 5.
Subgenera, 9.

Tables of measurements, 34-36.

Wishtonwish, 5.

zuniensis, Cynomys gunnisoni, 32-34.

37

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