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MUS. COMP. Z00L.
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HARVARD
UNIVERSITY

TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VOLUME X

PRINTED FROM THE

W. W. Wuitney PUBLICATION ENDOWMENT

SAN DIEGO, CALIFORNIA

PRINTED FOR THE SOCIETY
1943-1946

COMMITTEE ON PUBLICATION

josHua L. Barty
Cari L. Husss

LAURENCE M. KLAUBER

MUS. COMP. Z00L.
LIBRARY

MAY 16 194

HARVARD
RY VGLUME X

On the Generic Relationships of Certain Californian Xerophile
Snails. By S. Stillman Berry. Published December 30, 1943... 1-24
New Mollusks from the Round Mountain Silt (Temblor)
Miocene of California. By A. Myra Keen. Published
December. sO WSS eek ce eh ee ewe PA TL Pe AP ee ah eee, 2 25-60
Growth in the Western Blue-Tailed Skink. By Thomas L.
Rodgers and Viola H. Memmler. Published December 30, :
JIC Be Saag Oe OE Oe a enc SAD Oke at Bean SEO Ns COE Ab ME on ee pe ae 61-68
A New Snake of the Genus Sonora from Lower California,

Mexico. By Laurence M. Klauber. Published December 30,

4S taper omer A Slater eet he WEA eA Re Mol eer Sea a: eatin, Mee cere AI 69-70
A Desert Subspecies of the Snake Tantilla Eiseni. By Laurence

M. Klauber. Published December 30, 1943.20.00... 71-74
The Coral King Snakes of the Pacific Coast. By Laurence M.

Klauber. Published December 30, 1943............c..-.cclccscecse------ 73282
The Subspecies of the Rubber Snake, Charina. By Laurence

M. Klauber. Published December 30, 1943 .....000000000000000000000.-. 83-90

The Sidewinder, Crotalus Cerastes, with Description of a New
Subspecies. By Laurence M. Klauber. Published August 18,

[US rah Re =e 5 Ape ede coms ale seat ee OR PLR NOS a eee 91-126
New Occurrences of Fossil Tapir in Southern California. By
Ghestec’ Stock Publishedw August 18,1944 602 127-130

A New Race of Kangaroo Rat from the Argus Mountains,
California. By Laurence M. Huey. Published March 9, 1945..131-132
The Geckos of the Genus Coleonyx with Descriptions of New
Subspecies. By Laurence M. Klauber. Published March 9,
SEIS) pe tne ak eee aa eR mee eee Boe |S ead ea Mande» a ae 133-216
The Transverse Volcanic Biotic Province of Central Mexico and
its Relationship to Adjacent Provinces. By Robert T. Moore.
ublishedeAmpust 39 \O45%.. cen ee So 217-236
Preliminary Studies on the Black-Throated Sparrows of Baja
California, Mexico. By A. J. van Rossem. Published August
3 ee 1S PE SEI ie oe epee nl re AS ER ek 78 a Orr Me 237-244 '
The Pocket Gophers of Baja California, Mexico, with Descrip-
tions of Nine New Forms. By Laurence M. Huey. Published

eo te 245.268
The Chuckwallas, Genus Sauromalus. By Charles E. Shaw.
Biplistica gauiocsty hw lO4p Sel ei ee 269-306

A New Wood Rat, Genus Neotoma, from the Viscaino Desert
Region of Baja California, Mexico. By Laurence M. Huey.
Pmplismecyastetisty ot. AOA). to eal el 307-310

The Glossy Snake, Arizona, with Descriptions of New Sub-
species. By Laurence M. Klauber. Published March 29, 1946..311-398

Data and Field Notes on the Desert Tortoise. By Chapman
Keane: Published March 295 1946.2 Soon eee cctctacseee ane 399-402

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TRANSACTIONS

OESREIE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vot. X, No. 1, pp. 1-24, plates 1-2, figs. 1-8, map
YN

yan 14 1944

Nt 1p RAB

ON THE GENERIC RELATIONSHIPS OF
CERTAIN CALIFORNIAN XEROPHILE SNAILS

S. STILLMAN BERRY
Redlands, California

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY

DECEMBER 30, 1943

Ko : es
24,949 “gan 14 1944

LisRAks

120° 119° 18° 117° 116° 115° 114°

Distributional map showing areas inhabited by various genera of xerophile
helicoid snails in southern California.

RECTANGLE. Known localities occupied by species of Mohavelix (micrometalleus).

CircLes. Known localities occupied by species of Sonorelix: 1—avawatzica, 2—rix-
fordi, 3—borregoensis, 4—borregoensis ora.

Dots. Approximate areas occupied by species either shown in the present paper to
belong to Sonorelix or the shell-characters of which indicate a reasonable probability that
their affiliation here will eventually be established.

BROKEN LINE. Approximate western limit of area occupied by species of the group
Eremarionta according to present information.

ON THE GENERIC RELATIONSHIPS OF

CERTAIN CALIFORNIAN XEROPHILE SNAILS
BY

S. STILLMAN BERRY
Redlands, California

The snails of the Californian deserts have received increasing
attention from students in late years, but they are still poorly understood
and their systematics remain even to this date in a distinctly chaotic
condition. Since they form a somewhat inconvenient, not to say
inaccessible, group for thoroughgoing study, they have unhappily escaped
this desideratum altogether, and have meanwhile suffered rather severely
at the hands of the skimmer and dilettante. It is unfortunate that the
most recent author to deal with them (Pilsbry 1939) was able to perform
very little of his work at first hand, with the result that the system he
adopted is confusedly full of unnatural groupings if not outright errors,
in the end advancing but little from the general chaos that has all along
prevailed.

I have had evidence for a number of years that the generic taxonomy
of these snails has been quite inadequately worked out and particularly
that the group Eremarionta is not nearly so all-inclusive in the fauna
as has much too hastily been assumed. The implied changes appeared
so radical, however, that I have not only delayed publication of my
findings, but have even committed the tacit error of continuing my own
published treatment of divers species in the perfunctory traditional
sense, hoping (and expecting) that opportunity would arise for the
collection of such additional data regarding critical forms as might
enable me to fortify my initial findings and thus in the end perform a
more convincing service. Some fresh material has indeed come to hand,
but I do not have very frequent opportunity to reach the good collecting
grounds, while of late years the interposition of many extraneous duties
has further impeded me. I have now reached the conclusion that no
good purpose can be served by further withholding publication of at
least a part of my notes, incomplete and imperfect though they may be,
and however short they fall of the more broadly fortified treatment
which I have had in mind.

Under all the circumstances there need be little wonder that the
taxonomic vicissitudes of Californian desert snails have been many.

4 SAN DrgeGo Society oF NATURAL HIstory

This has been inevitable from the circumstance that the distances to be
covered are vast, living animals in good condition not too easy to come
by, and any fortuitous treatment of the comparatively infrequent
specimens collected has usually been more a reflection of the latest
discoveries among the more or less similar snails of neighboring regions
than an autochthonic study. It was thus not unnatural that the first
of the group to be described was referred, along with most other large
American snails, to the (properly European) genus Helix (Yates 1890).
A little later, when a few more kinds were brought to light, they, like
so many other west American helicoids, found lodgement in the
(tropical American) genus Epiphragmophora (Pilsbry 1898). Still
later, the spectacular discoveries of Ashmun and Ferriss in Arizona.
with the elucidation at the hands of Pilsbry (1900) of the remarkable
anatomical peculiarities of the important genus Sonorella, resulted in
the early transference thither, on the uncertain basis of shell characters
alone, of all the snails then known to be endemic in the Mohave and
Colorado Deserts (Bartsch 1904), a treatment which has not only
proved premature but far too sweeping. The peculiar characteristics of
Sonorella which mostly concern us here lie in the remarkably simplified
reproductive apparatus, which upon dissection reveals no trace of the
specialized dart-sac and mucus-glands that characterize the superficially
similar coastal genera, no diverticulum or branch of the spermatothecal
duct, and a very great reduction or complete obsolescence of the
“flagellum” or epiphallic caecum, while the penial sac contains a well-
developed verge. The spermatotheca is characteristically quite small.

The dissection by Pilsbry (1907) of the newly described Epiphrag-
mophora (Micrarionta) hutsoni G. H. Clapp, from near Quartzsite,
Arizona, proved a jolt to complacency, for although this species has a
Sonorella-like shell, none of the anticipated features were revealed, but
instead a well-developed dart-sac and mucus-glands of such character as
to ally it with the coastal Micrarionta-group and not at all with Sonorella.
Pilsbry at this point conservatively remarked (op. cit., p. 139), “How
many of the supposed Sonorellas of southeastern California may really
prove to belong to Micrarionta is problematic, but perhaps all those
with the embryonic sculpture like E. hutsoni will eventually be removed
from Sonorella.” His insight soon found confirmation in the new
desertorum (Pilsbry and Ferriss 1908) in a paper which elevated
Micrarionta to independent generic rank, and in wolcottiana, the large

snail so conspicuous in the neighborhood of Palm Springs (Pilsbry

BERRY—CALIFORNIAN XMEROPHILE SNAILS 5

1918). Since then the tendency has exclusively been to regard all
the Sonorella-like snails found west of the Colorado River and those
to the east of that stream which appeared closely similar to hutsoni as
species of Micrarionta (cf. especially Berry 1922, Pilsbry 1939). The
subgenus Eremarionta was meanwhile proposed by Pilsbry (1913: 382)
with M. desertorum as type, the assumption being that it would likely
be found to include most or all of the desert forms.

Several years ago (Berry 1930), I showed reason for the removal
of one Californian desert snail, originally described as a Sonorella, to
Helminthoglypta (H. fisheri). In the present paper, anatomical
evidence is set forth that at least four more divergent and geographically
scattered species cannot possibly be retained in Micrarionta, but are
definitely akin to Sonorella, though showing certain peculiarities of
their own.

I must here acknowledge my great obligation to Prof. Edmund C.
Jaeger of Riverside Junior College, the late Fred M. Reed of Riverside,
Dr. L. G. Ingles of Chico State College, and Mr. C. C. Searl of Hemet
for essential aid in the task of securing living material, and likewise to
Mr. A. E. Burns of Oakland, Mr. Allyn G. Smith of Berkeley, and
Prof. Jaeger for the excellent photographs used in illustration, most of
which were made by Mr. Burns.

All anatomical drawings were done by the author with the aid of
a camera lucida, but, owing to certain difficulties of a mechanical nature,
as well as those inherent in the material, their pretension is to approxi-
mate accuracy only, rather than to absolute exactness in every detail.
The following abbreviations have been used :

ABBREVIATIONS USED IN TEXT FIGURES
c.d.—common duct of spermatotheca and diverticulum

di.—diverticulum
e.c.—epiphallic caecum or “flagellum”
ep.—epiphallus
ov.—oviduct

p-—pPpenis

p.c.—penial chamber
r.—penis retractor
s.d.—duct of spermatotheca
sp.—spermatotheca
v.—vagina

v.b.—basal part of verge
v.d.—vas deferens

v-p.—papilla of verge

6 San Dreco Society oF NATURAL HISTORY

Genus SONORELLA Pilsbry 1900
(Pilsbry 1900:556; 1939:267)

Type: Sonorella hachitana (Dall 1895)

This genus is characterized by the extreme simplification of the female side
of the hermaphrodite system. All of the amatory organs so conspicuous in
Micrarionta and Helminthoglypta are here entirely absent and there is no
spermatothecal diverticulum. The epiphallic caecum is very small or obsolete.
The penis contains a verge, which shows great diversity of form and often attains
a high degree of specialization. The shell is simply helicoid, generally lacking
in very remarkable features, and frequently rendering difficult or impossible the
proper allocation of unknown species without dissection.

Mohavelix, new subgenus

(Derived from Ind. Mohave, name of desert and Indian tribe, plus Gr.
helix, a spiral, hence a snail shell).

Type: Micrarionta (Eremarionta) micrometalleus Berry 1930

Shell small, thin, subdiscoid, with a wide perspective umbilicus, the
periostracum crudely granular-papillose throughout, except for the embryonic
sculpture of numerous rather sharply cut, elongate, pointed papillae, arranged
in forward-descending series and sometimes confluent into nearly solid decurrent
lines.

Hermaphrodite system with epiphallus weakly differentiated, but equipped
with a minute adnate caecum; penis very robust, in length about equal to the
similarly robust vagina, abruptly and definitively set off from the epiphallus, its
apical portion strongly bulbously expanded around the heavy, cylindrical base
of the relatively large, elongate, conical papilla or verge. Retractor inserted on

distal part of epiphallus.

Sonorella (Mohavelix) micrometalleus (Berry 1930)
Plate 1, figures 1-3

1930. Micrarionta (Eremarionta) micrometalleus Berry—Ann. Mag. Nat.
Flist., ser. 10; 6:189; fies. 3, 4.

1931. Micrarionta {Helminthoglypta?} micrometalleus Willett—Naut., 44
(4): 125 (brief note).

1939. Micrarionta micrometalleus Pilsbry—N. A. Land Moll., 263, fig. 137.

Descriptive Notes—The female system is simple, without accessory organs.
The vagina is both robust and moderately long. The spermatothecal duct is
very long, slender, and simple, entering the vagina a little distad from the level
of emergence of the vas. I can find no trace of a diverticulum.

The male system is simple proximally but has the distal portions more
strongly specialized. The epiphallus appears but weakly differentiated from the

BERRY—CALIFORNIAN XEROPHILE SNAILS 7

vas externally, its enlargement at this point slight; it has a very inconspicuous,
minute, conical caecum, which is closely adnate to the distal end of the vas and
bound to it by the thin enclosing membranes and the apparent involvement of
its apex in a branch of the small muscle or connective which attaches the elbow
of the vas to the base of the vagina. The penis is about as long as the vagina,
thin-walled, robust, abruptly set off from the epiphallus; its proximal third is
strongly bulbously swollen around a thick cylindrical core, whence abruptly
arises a thin-walled, elongate-conic verge nearly half as long as the remaining
portion of the penis. The retractor is inserted on the epiphallus a little above
its junction with the penis.

Remarks.—It is tantalizing to have no better anatomical material of a snail
as important from the standpoint of the relationships and distribution of our
southwestern snails as is this one, but the few living mature examples sent to

Fig. 1 ‘ Pig. 2

Sonorella (Mohavelix) micrometalleus (Berry).

Fig. 1. Anterior part of hermaphrodite system of holotype.
Fig. 2. Proximal portion of penis, slit open to expose the verge.

me in the original collection failed to expand well in drowning or to pull
satisfactorily afterward, and the long-hoped-for opportunity to pay a personal
visit to the locality to observe the creature first-hand and collect a fresh supply
has never materialized.

In its depressed form, perspective umbilicus, and rather horny texture of
the shell, this species is more suggestive of a pygmy Sonorella ferrissi Pilsbry
than any other species of the genus seen by me. From this, nevertheless, it
differs radically in its embryonic sculpture and in the structure of the penis
and verge. The embryonic sculpture is curiously similar to that of Eremarionta,
though cruder and coarser, no Sonorella with which I have chanced to compare

8 San Disco SociETY OF NATURAL History

it being at all closely comparable in this particular. If now correctly aligned,
it must be one of the smallest species so far known in Sonorella.

It is interesting to note that, as Californian snails go, micrometalleus
appears to be a pretty deep delver. This phase of its natural history is, of
course, entirely in harmony with its Sonorelline affinities. The necessity now
shown for the elimination of this species from Eremarionta pushes the supposed
range of the latter group quite out of Kern County and a considerable distance
to the eastward, but hurdles Sonorella across a wide intervening area to establish
this important genus for the first time as a true member of the Californian
fauna, even though as a considerably modified offshoot.

Sonorelix, new genus

(Derived from the Sp. Sonora, the Mexican State which gives its name
to the Sonoran Desert and Life Zone, + Gr. helix, anything which assumes a
spiral shape, hence the genus Helix, and has particular reference to the inter-
mediate position between Sonorella and the more elaborately specialized helicine
snails which the anatomical characters appear to indicate).

Type: Micrarionta (Eremarionta) borregoensis Berry 1929

Sonorelline snails having a pale, waxy-porcellaneous, umbilicate, helicoid
shell, ornamented by a peripheral brown band of varying prominence.
Embryonic shell sculptured by a variably developed sub-retiform papillation.

Hermaphrodite system lacking a dart-sac, mucus-glands, or other accessory
amatory organs on the female side; vagina very long; spermatothecal duct long,
the often enormously large spermatotheca lodged beneath the albumen gland in
the species investigated; diverticulum of spermatothecal duct robust, of
moderate length, the portion of the duct distad to the junction with it excessively
short. Epiphallus short, stout, bearing a well-developed caecum; penis abruptly
set off and enlarged from the epiphallus, thence narrowing either suddenly or
more regularly to the aperture, concentrically ridged or folded within, and
provided with a short conical verge; retractor inserted on epiphallus.

Remarks.—It will be noted that the group now for the first time given sys-
tematic recognition differs radically from Eremarionta in the complete absence of
dart-sac and mucus-gland, while the embryonic sculpture is likewise somewhat
peculiar, lacking the sharpness and geometric regularity of papillation seen in
most species of the latter genus. Compared with Sonorella, noteworthy similarity
is found in the simplification intrinsic in the absence of accessory amatory
organs, and in the presence of a well-developed verge, but the new group diverges
from this genus so definitely in the possession of a large epiphallic caecum, in
the presence of a diverticulum, and in the great size of the spermatotheca itself,
that unless the definition of Sonorella is greatly amplified, there appears no
escape from the establishment of an independent category. As there are rather
valid objections to either course, it becomes a matter of personal choice which
arrangement any particular student will choose to adopt, and I shall therefore

BERRY—CALIFORNIAN XEROPHILE SNAILS 9

find no quarrel with anyone who may prefer a less analytic treatment than that
here proposed.

Other described species which appear to share to a considerable extent in
the characteristic sculpturing of the embryonic whorls noted in the type-species
of Sonorelix are ora (Willett) , carrizoensis (Willett) , rixfordi (Pilsbry) , aetotis
(Berry), depressispira (Berry), harperi (Bryant) , orcutti (orcuttiana' Bartsch),
avawatzica (Berry), baileyi (Bartsch), eremita (Pilsbry), and melanopylon
(Berry). Of these the first, as I shall shortly endeavor to show, very definitely
belongs here, being an immediate ally of borregoensis, and rixfordi likewise
possesses essentially the same type of reproductive system. The relationships of
avawatzica also lie near. Most or all of the others I believe-are likewise likely
to belong to Sonorelix, but it would probably be both premature and unwise to
be too dogmatic upon this point until the anatomy of each form can be made
known in its own right.

The species thus indicated as belonging to this group are all. desert-dwelling
creatures, exhibiting a markedly discontinuous distributional range, which extends
from southern Inyo County to the Mexican border. The more northern of the
known localities are all of relatively small extent and considerably isolated from
one another. More extensive are a fairly large area south and west of
Twentynine Palms in southern San Bernardino and north-central Riverside
Counties, and another occupying the hilly eastern portion of San Diego County
and the extreme western edge of Imperial County. No snails certainly referable
to Sonorelix are yet known from the immediate vicinity of the Colorado River
or from any point to the eastward of that stream in Arizona, but as so many
of our xerophile snails are doubtless still to be discovered, while many of those
known merely from the shells remain to be investigated anatomically, it is
reasonable to anticipate that from time to time material extensions of the range
as here outlined are almost certainly to be anticipated (see map).

In its general ecology, Sonorelix appears to be more like Eremarionta than
the usual Sonorella. It is found under loose slabs of rock, among rubbish and
in rock slides, but dead and bleached shells are usually, unlike most Sonorella,
to be found scattered in some abundance on the surface, and, where taken alive,
it has not turned out to be a very deep delver. Two characteristic sites where
species of the new genus have been found are shown on Plate 2.

Sonorelix borregoensis (Berry 1929)
Plate 1, figures 4-6

1929. Micrarionta (Eremarionta) borregoensis Berry,—Naut., 43 (2) :39.

1929. Micrarionta reedi Willett,—Bull. Sou. Cal. Ac. Sc., 28 (2): 17, 19, pl.
6, lower figs.

1939. Micrarionta borregoensis Pilsbry—_N. Am. Land Moll., p. 256, fig. 132.

1Should this species prove with more complete knowledge to be either a Sonorella
or a Sonorelix, the original spelling will stand, since orcutti was not still born but be-
comes preoccupied only by reason of its transference to Micrarionta.

10 San Dieco Society of NaturAL History

Descriptive Notes—The shell is of fair size and thickness, in form
depressed to very low conic. The whorls are usually about 5 in number, convex,
quite regularly and rapidly enlarging, the suture distinct. The last whorl is
moderately descending parietally. The aperture is rounded and quite oblique.
The peristome is but little thickened, everted scarcely at all above, more so
below, and there is a moderate reflex at the columella over the margin of the
umbilicus. The umbilicus is wide and permeable to the apex, its diameter
being contained usually a little over 7 times in that of the shell. Spiral sculpture
is wanting. The embryonic shell is at first smooth, then concentrically wrinkled
for a fraction of a whorl, after which it becomes covered with numerous heavy

——

fel
Fig. 3. Sonorelix borregoensis (Berry).

Anterior part of hermaphrodite system and spermatotheca of holotype. The apparent
narrowing of the upper part of the vagina is due to a twist in the organ. The form and
position of the verge are shown as though in optical section by dotted line.

microscopic papillations, which only occasionally exhibit traces of an arrange-
ment in decurrent slanting lines, and are for the most part elongate, coarse,
and irregular, sometimes fusing or anastomosing to give somewhat the effect
of an ill-defined network.

The female system is simple, without traces of mucus-glands or dart-sac.
The vagina is oval in section, moderately robust, and very long. The
spermatotheca i is enormously large, ovate-spherical, and lodged under the apical
portion of the albumen gland; its duct is very long, entering at the base of the
oviduct, and branches rectangularly a very short distance proximad to form a
stout, sradually tapering diverticulum of moderate length, which is for the most

BERRY—CALIFORNIAN XEROPHILE SNAILS 11

part considerably wider than is the spermatothecal duct above the junction.

The male system comprises a slender vas deferens, which increases in girth
in its last quarter to enter the very robust, acutely elbowed, steadily enlarging
epiphallus. The latter bears a rather robust, tapering caecum not quite so long
as itself. The retractor muscle is thick and broadly inserted on the elbow of
the epiphallus. The penis is perhaps a little shorter than the vagina, but very
large, robustly conical, and abruptly set off and enlarged from the epiphallus,
thence tapering regularly to its termination. The proximal end of the penis
sometimes shows a few weak concentric ridges. The penial chamber is ample,
especially in the enlarged proximal part, and is closely concentrically folded or
ridged within. Apically it contains a short, stout, weakly annulate, conical
verge, which opens on a nipple-like prominence through transverse, rather thick
lips.

Measurements—The following table gives the more essential caliper
measurements, in millimeters, of twelve mature shells in the Berry Collection.

Coll. Max. Min. Diam. No.
Lot No. Diam. Diam. Alt. Umbilicus W horls
6914 Paratype 2307) 20.4 11B33 2 5.2
6914 Paratype 23.4 19.4 12.6 3.0 23
6915 Paratype 22.4 18.7 12.6 3.3 5.0
6915 Paratype 22.3 19.0 13.0 BE2. Dee.
6914 Paratype 223 18.5 12.4 3.0 D2
6914 Paratype 22.1 18.4 12.6 Bel 5.0
6913 Holotype 22.0 18.2 12a DS) 4.7
6914 Paratype 20.7 17.3 11.0 2.8 4.7
7053 Coyote Mt. 21.4 18.3 123 2.8 Dell
7053 Coyote Mt. PD Nef 18.0 WI 3.4 Bea)
7053 Coyote Mt. 19.2 16.5 EL 2.6 ahye
7053 Coyote Mt. 17.6 Dal 10.6 2.0 a2

Type Material—Holotype, Cat. No. 6913, Berry Collection. Paratypes

are in the collections of the San Diego Society of Natural History, the Academy
of Natural Sciences of Philadelphia, and the late Fred M. Reed, as well as
Cat. Nos. 6914 and 6915, Berry Collection.

Type Locality—Palm Canyon, Borrego Valley, San Diego County, Cali-
fornia; 2 live mature, and many dead and for the most part bleached mature and
immature shells; collectors, Fred M. Reed, April 4, 1929, and L. M. Klauber,
April 14, 1929.

Material Examined. —

No. W here
Spec. Locality Collector Deposited
2 Near Thousand Palms,
Borrego Valley C. A. Dodd, 1928 Berry Coll. 6917
1 Palm Canyon, Borrego Valley F.M. Reed, Apr. 4, 1929 Berry Coll. 6913
13 Palm Canyon, Borrego Valley F. M. Reed, Apr. 4, 1929 Berry Coll. 6914
3 Palm Canyon, Borrego Valley F.M. Reed, Apr. 4, 1929 Reed Coll. 2456
13 Palm Canyon, Borrego Valley F.M.Reed,Apr.4,1929 =... ee eee
4 Palm Canyon, Borrego Valley LL.M. Klauber, Apr. 14,1929 — Berry Coll. 6915
4 Palm Canyon, Borrego Valley L.M.Klauber, Apr. 14,1929 — S. D. Soc. Nat. Hist.
2 Palm Canyon, Borrego Valley L.M. Klauber, Apr. 14,1929 — Ac. Nat. Sc. Phila.
13 E. slope Coyote Mt.,

N.W. of Clark’s Lake

E. C. Jaeger, Dec. 23, 1929

Berry Coll. 7053

12 San DrgEco SoctEty oF NATURAL HISTORY

Geographic Range—Mountain slopes surrounding the upper end of
Borrego Valley from Coyote Mt. to the San Ysidro Mts. and possibly further,

all in northeastern San Diego County, California.

Remarks.—This is one of the finest of our desert snails. When Mr. Reed
brought me the fruits of his 1929 collecting trip to the Borrego Valley, he was
disappointed to learn that I had already seen several lots of this species and
had the description in hand. That material, however, consisted of empty shells
only, and as Reed generously left a good part of his own shells with me,
including one fine living individual, the latter was appointed to serve as holotype.
It subsequently turned out that Reed shared his remaining shells with George
Willett, with the unfortunate result that we both published descriptions almost
simultaneously, for although my manuscript went to the publishers in July, it
was not printed until October, by which time Willett’s paper, unknown to me,
had also gone to press. As only a brief preliminary diagnosis was published in
1929, I have taken advantage of the present opportunity to insert from my notes
not only an anatomical account, but a more complete description of the shell.

The Santa Rosa Mountains, to the north of the range of borregoensis,
are inhabited only by species of Eremarionta, so far as at present known. Its
closest relative, as well as its immediate neighbor to the south, is the following
form.

Sonorelix borregoensis ora (Willett 1929)

1929 Micrarionta ora Willett,—Bull. So. Cal. Ac. Sc., 28 (2): 17, 19, pl. 6,
upper figs.

1937. Maucrarionta harperi, (pars) Willett,—Naut., 50 (4) :122.
1939. Micrarionta ora Pilsbry—N. Am. Land Moll., 251, 254, 255, fig. 127a.

Descriptive Notes—The shell is not very different from that of borregoensis
s.s., but it averages somewhat smaller, and the umbilicus is narrower.

In color the living animal is in somber contrast to the light brown or cream
of the shell, the dorsum (main body and eyes) being black, with an edge of
Deep Mouse Gray. The sole is Deep Mouse Gray with an inner paler zone
between Deep Olive Gray and Dark Olive Gray.

The hermaphrodite system is essentially as I have described for borregoensis
s.s.., but the vagina is rather more robust, and the terminal part of the sperma-
tothecal duct even shorter and thicker, apparently entering below the oviduct,
and with the diverticulum appreciably shorter and stouter. The penis is more
bulbously swollen apically and its distal portion more evenly cylindric, but of
the same general character otherwise. It contains a rather longer, conical,
annulate verge, projecting from a somewhat bulbous base. The wall of the
penis is quite thick, but the strong concentric folds of the interior are evident
as weak annulations externally. The diverticulum shows a thin-walled sacculated
expansion or dilatation near its midway point, the significance of which is not
for the moment apparent.

BERRY—CALIFORNIAN XMEROPHILE SNAILS 13

Measurements.—The following are measurements, in millimeters, of three
mature shells.

Coll. Max. Min. Diam. No.
Lot No. Diam. Diam. Alt. Umbilicus Whorls
6916 Berry Coll. 20.9 17.0 12.2 2.6 5.0
6916 Berry Coll. 19.4 16.6 11.2 2.3 5.0
6916 Berry Coll. 18.7 15.5 10.0 pe) 4.8

Material Examined.— 4 mature and 1 immature individuals from the west
end of Sentenac Canyon, San Diego County, California; all taken alive by
Clyde C. Searl, Jan. 6, 1929.

Remarks —I received my examples from Sentenac Canyon in the living
state early in 1929 and wrote out a description of them as a new species, but
withheld it from publication in favor of borregoensis. Since then I have let it
lie because of the publication of ora Willett and its apparent similarity thereto,
and further because I had the hope of obtaining more complete material of all

Fig. 4. Sonorelex borregoensis ora (Willett).

Anterior part of hermaphrodite system of specimen 6916a from Sentenac Canyon.
The form and position of the verge are shown by dotted line as though in optical section.

these forms before attempting to establish their various relationships. Lacking
material from the type-locality of ora, I can at this juncture only assume the
correctness of Willett’s own considered opinion (1937:122) that the snails of
Sentenac Canyon belong to the same race, and point out that, if this be so,
the relationships of ora to borregoensis rather than to harperi, as the latter is at
present understood, become clearly manifest. The rather smaller average size
of ora and its somewhat narrower umbilicus are in fact the only shell characters
I find to separate the two, and I have examples from at least one locality well
within the range of borregoensis s.s., but not discussed in the present paper,
which are considerably smaller than ora. The reproductive anatomy of the
Sentenac Canyon ora proved, as anticipated, to be essentially similar to that of
borregoensis. There are, however, various small differences to be noted, and
these, unless and until they can be shown to lie within the normal range of

14 San DrisEco Society oF NAturRAL History

the typical race, appear sufficient to justify the retention of ora as a rather
weakly marked subspecies of borregoensis. In range the two races appear to be
closely adjacent. If actually contiguous, as I anticipate will prove to be the
case, their characters are such that the point of actual passage of one form to
the other will probably be found in practice quite imperceptible.

I can at present discover little support for Willett’s affiliation of ora with
harperi (1937:122), and it would appear from the subsequent remarks of
Pilsbry (1939:254) that Willett himself shortly abandoned the opinion as

untenable.

The somber coloring of the animal is much like that of Eremarionta.

Sonorelix rixfordi (Pilsbry 1919)
Plate 1, figures 7-9

1919. Micrarionta rixfordi Pilsbry,—Naut., 33 (2) :53.

1930. Micrarionta rixfordi Willett,—Bull. So. Cal. Ac. Sc., 29 (1) :15.

1939. Micrarionta rixfordi Willett,—id., 38 (1) :15.

1939. Micrarionta rixfordi Pilsbry—N. Am. Land Moll., 257, 258, 261, 263,
figure 133.

Descriptive Notes —The female system lacks a dart apparatus. The vagina
is very long, slightly increasing in amplitude near the exit. The spermatothecal
duct is provided with a robust and fairly long diverticulum, considerably greater
in caliber than the duct with which it unites, the portion of the duct below the
union being more robust than above and exceedingly short.

The male system includes a short robust epiphallus, and there is a well-

}
ii
Sk

Imm

ees ALLEL i
>

a
oa
ron)

Fig. 5
Sonorelix rixfordi (Pilsbry).
Fig. 5. Anterior part of hermaphrodite system of specimen 6441a from near type-
locality, west of Twentynine Palms.

Fig. 6. Distal portion of epiphallus and proximal portion of penis of same specimen
slit open to expose the verge.

BERRY—CALIFORNIAN XEROPHILE SNAILS 15

developed, sometimes strongly coiled epiphallic caecum nearly as long as the
epiphallus itself. The penis is robust, not very thick-walled, little more than
half as long as the vagina, and abruptly enlarged and set off from the epiphallus;
it tapers more rapidly at first than in the distal more evenly cylindrical portion.
The penial chamber is ample, especially proximally, and its wall is provided with
many strong concentric ridges or folds which to some extent are apparent
externally. Its apical portion contains a moderately firm, somewhat flattened,
bluntly conical, finely concentrically ridged verge, which extends down the
penial chamber for roughly a fifth of its length; the basal part of the verge is
only moderately swollen. The retractor is strong, inserted about midway of the

epiphallus.

Material Examined—The first listed is from the original lot, and those
from 8-10 miles west of Twentynine Palms are approximate topotypes.

No. W here
Spec. Locality Collector Deposited
1 10 m. W. of Twentynine
Palms, San Bernardino Co. Emmet Rixford, Mar., 1919 Berry Coll. 5143
3 Cliff 8-10 m. W. of Twenty-
nine Palms, San Bernar-
dino Co. E. C. Jaeger, Dec. 28, 1927 Berry Coll. 6441
3 Large cn. in Little San Ber-
nardino Mts., 4 m. W. of
Twentynine Palms, San
Bernardino Co. S. S. Berry, Nov. 12, 1928 Berry Coll. 6722
7 On Pinto Basin Road, 8 m.
S. of Twentynine Palms, E. P. and E. M. Chace,

Riverside Co., Jan. 31, 1936 Berry Coll. 8421
2 2m. S. of Keys Ranch, E. P. and E. M. Chace,

Riverside Co. Jan. 31, 1936 Berry Coll. 8419
2 Live Oak Tank, E. P. and E. M. Chace,

Riverside Co. Jan. 31, 1936 Berry Coll. 8420
2 7 m. down cr. from Quail _ E. P. and E. M. Chace,

Spr., San Bernardino Co. Jan. 31, 1936 Berry Coll. 8418

Geographic Range——Northern outliers and canyons of the Little San
Bernardino Mts. on both sides of the boundary between San Bernardino and
Riverside Counties west and south of Twentynine Palms, and extending south-
eastward to the Eagle and perhaps the Orocopia Mts., should aetotis Berry
prove indefensible as a segregate, all in the southern Mohave Desert region of
California.

Remarks.—It is interesting to find the affinity suggested by the similarity
in embryonic sculpturing of the shells of rixfordi and borregoensis entirely borne
out by the anatomy of the reproductive system. The anatomical characters
described are much as has been shown for borregoensis, but the extremely long
vagina, comparatively short penis, and lack of a heavy thickened base to the
verge appear to be somewhat special features. Unfortunately my only available
preparation is in a badly damaged state and I can give no information respecting
the spermatotheca, but that portion of its duct in front of the entrance to the
diverticulum is thick and exceedingly short, as in borregoensis.

Willett (1930:15) unites my aetotis and depressispira with rixfordi, and

16 SAN Disco Society oF NAturAL History

I am not prepared successfully to confute him. Nevertheless, although aetotis,
in particular, appears appreciably closer to rixfordi with our present information
than I thought to be the case when I described it, the more extensive material
of both forms now available to me still continues to show certain slight
differences, real or imaginary though they be, and therefore, until the animals
can be investigated, it may be just as well to continue an abeyant recognition of
all three forms.

Sonorelix avawatzica (Berry 1930)
Plate 1, figures 10-13

1930. Micrarionta (Eremarionta) avawatzica Berry,—Ann. Mag. Nat. Hist.,
ser. 10, 6:190, figures 5-8.

1939. Micrarionta avawatzica Pilsbry—N. Am. Land Moll., 245, 260, 262,
263, figure 134 (after Berry).

Descriptive Notes—The living animal is practically black on the dorsum
around the head and eyes (Dark Mouse Gray in one example), paling to nearly
Mouse Gray on the upper surface of the foot, or sometimes a little lighter as
in the paler specimen mentioned. The sole is Mouse Gray with a fairly wide

border of Deep Mouse Gray.

The female system shows no trace of a dart-sac, mucus-glands, or other

Sonorelix avawatzica (Berry).

Fig. 7. Distal portion of hermaphrodite system and spermatotheca of holotype from
south of Cave Springs, Avawatz Mountains.

Fig. 8. Epiphallus and proximal portion of penis of paratype 6885b, with the out-
line of the verge dotted in as though in optical section.

BERRY—CALIFORNIAN XEROPHILE SNAILS 17

specialized amatory organs. The vagina is of rather robust caliber throughout,
but only moderately long. The spermatotheca is of moderate size, ovate-globular,
its duct very long and branched in T-shaped fashion perhaps twice as high on
the oviduct as in the species of the group hitherto described, the diverticulum
being fairly long, slender, and appreciably heavier than the duct at its point
of entrance.

The male system has a rather short and evenly calibrated vas deferens
which becomes a trifle more robust distally. The epiphallus is short and of
moderate thickness, its tapering caecum well developed and nearly as long as
itself. The penis is fairly long, and sharply set off from the epiphallus by a
moderate but abrupt bulbous expansion at this point, whence it diminishes with
only a very gentle taper. The enlarged part of the penis shows a few weak
concentric ridges apically, and contains a conical, thin-walled, subterminally
perforated verge, arising from a heavily thickened base, and terminating in a
softer papilla-like portion, its aperture lip-like. The inner wall of the penis is
closely concentrically ridged.

Geographic Range—The species is still unrecorded except from the
immediate vicinity of the type locality in the Avawatz Mountains in northern
San Bernardino County, but I have seen a little imperfect material from one or
two more distant localities which leads to the belief that it will eventually be
found to inhabit rather an extended area.

Remarks.—Pilsbry (1939:261, 263) comments concerning this species that
it is very close to M. rixfordi, “but differs by the projecting apex and slightly
smaller umbilicus....but all of these forms [rixfordi, baileyi, eremita, avawatzica |
are very much alike, and further collections in the vicinity of Resting Springs
may perhaps show that our distinctions are rather fine-drawn.” When I
described the shell of avawatzica it seemed to me a pretty distinct thing as the
shells of desert snails go. Of those mentioned, baileyi is very incompletely
known, the recently published eremita seeming the nearest approach to it that
I know; but with respect to rixfordi there need be no sort of confusion, for as
compared with this species the peculiar shell characters of avawatzica now find
themselves well fortified by the detailed anatomical features here described,
especially the near equality attained by the vagina and penis, the more proximal
position of the diverticulum, the much longer common duct of the spermato-
theca, and the differently shaped verge.

San Dreco Society oF Natura History

LITERATURE CITED

BarTSCH, P.

1904.

Notes on the genus Sonorella, with descriptions of new species.
Smithsonian Miscellaneous Collections, 47:187-200, pl. 28-33, Oct.
1904.

Berry, S. S.

19272.

1929.

1930.

1930a.

Notes on the Mollusca of the Colorado Desert—I. Proceedings
Academy Natural Sciences Philadelphia, 74:69-100, map, text figs.
1-5, pl. 8-10, 1922.

Three new snails from the hills of California. Nautilus, 43 (2):
39-40, Oct. 1929.

Snail notes from the California desert. Nautilus, 43 (3) :73-75,
Jan. 1930.

New helicoid snails from the Mohave Desert—IV. Annals and
Magazine Natural History, ser. 10, 6:187-192, text figs. 1-8, Aug.
1930.

Piussry, H. A.

1897-98. A classified catalogue of American land shells, with localities.

1900.

1907.

1913:

1918.

1919:
1939.

Nautilus, 11:45-48, 59-60, 71-72, 83-84, 93-96, 105-108, 117-120,
127-132, 138-144, Aug. 1897 to Apr. 1898. Reprinted, with cor-
rections, as A classified catalogue with localities of the land shells

of America north of Mexico, pp. 1-35, April 1898.

Sonorella, a new genus of Helices. Proceedings Academy Natural
Sciences Philadelphia, 52:556-560, pl. 21, 1900.

On the soft anatomy of E. (Micrarionta) hutsoni. Nautilus, 20
(12) :138-139, pl. 9, figs. 5-8, Apr. 1907.

Notes upon some Lower Californian Helices. Proceedings Aca-

demy Natural Sciences Philadelphia, 65:380-393, text figs. 1-3, pl.

15-16, July 1913.

On the generic position of Sonorella wolcottiana Bartsch. Proceed-
ceedings Academy Natural Sciences Philadelphia, 70:139-140, text
fig. 1, 1918.

A new Californian Micrarionta. Nautilus, 33 (2) :53, Oct. 1919.

Land Mollusca of North America (north of Mexico), Vol. 1,
Part 1, 1939.

BERRY—CALIFORNIAN XMEROPHILE SNAILS 19

Pitspry, H. A., and Ferriss, J. H.
1908. A new Micrarionta from Arizona. Nautilus, 21 (12): 134-136,
pl. 11, figs. 6-10, Apr. 1908.
WILLETT, G.

1929. Descriptions of two new land shells from southern California.
Bulletin Southern California Academy Sciences, 28:17-19, pl. 6,
Oct. 1929.

1930. Notes on Micrarionta rixfordi Pilbsry. Bulletin Southern California
Academy Sciences, 29: 15-16, June 1930.

1931. Two new helicoids from the Mohave Desert, California. Nautilus,
44 (4) :123-125, pl. 7, figs. 3-4, Apr. 1931.

1937. Micrariontas of the southwestern Colorado Desert. Nautilus, 50
(4) :122-125, Apr. 1937.

1939. Micrariontas of desert ranges bordering the east side of Coachella
Valley and Salton Sink, California. Bulletin Southern California
Academy Sciences, 38 (1) :14-16, pl. 2, Jan.-Apr. 1939.

YATES 1G:
1890. A new variety of Helix. Nautilus, 4 (6) :63, Oct. 1890.

20 San DigGo Society OF NATURAL History

PLATES

Figs. 1-3. Sonorella (Mohavelix) micrometalleus (Berry). Shell of holotype
from El Paso Mts., Kern County, California; x ca. 3.

Figs. 4-6. Sonorelix borregoensis (Berry). Shell of holotype from Palm
Canyon, San Ysidro Mts., San Diego County, California;
slightly enlarged.

Figs. 7-9. Sonorelix rixfordi (Pilsbry). Shell of specimen from original lot,
Berry Coll. No. 5143, from 10 miles west of Twentynine Palms,
San Bernardino County, California; x ca. 1/2.

Figs. 10-12. Sonorelix avawatzica (Berry). Shell of holotype from 5 miles
south of Cave Spring, Avawatz Mts., San Bernardino County,
California; x ca. 11.

Fig. 13. Sonorelix avawatzica (Berry). Apical whorls of holotype; x ca. 9.

BERRY—CALIFORNIAN XEROPHILE SNAILS

2p San Disco Society oF NATURAL HIstory

PEATE?

Fig. 1. Rocky slope just south of highway about 10 miles west of Twenty-
nine Palms, San Bernardino County, California. The approximate type-locality
of Sonorelix rixfordi (Pilsbry). (Photograph by S. S. Berry, Nov. 10, 1928).

Fig. 2. Rocky point west of road in the pass at junction of Barstow and
Silver Lake roads, 5 miles south of Cave Spring, Avawatz Mountains,
San Bernardino County, California; the type locality of Sonorelix avawatzica
(Berry). The holotype was taken in the small rock-slide just behind the youth

standing near the center of the picture. (Photograph by S. S. Berry, March 1,
1929).

BERRY—CALIFORNIAN XEROPHILE SNAILS

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TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VotuME X, No. 2, pp. 25-60, plates 3-4, figs. 1-5

Zoolteay

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\ JAN 141944
Bee LLL)

NEW MOLLUSKS
FROM THE ROUND MOUNTAIN
SILT (TEMBLOR) MIOCENE OF CALIFORNIA

A. Myra KEEN

Stanford University

SAN DIEGO, CALIFORNIA
~ PRINTED FOR THE SOCIETY

DECEMBER 30, 1943

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2 MILES

GEOLOGY BY D.BIRCH

Fig. 1. Geology of the Kern River area near Round Mountain. The
course of Kern River is marked by the band of Quaternary alluvium near
the center of the map.

NEW MOLLUSKS
FROM THE ROUND MOUNTAIN
SILT (TEMBLOR) MIOCENE OF CALIFORNIA’
| BY
A. Myra KEEN

Stanford University

ABSTRACT

Nineteen new species of mollusks are described and one new pelecypod
subgenus proposed—Bornia (Temblornia). All material discussed comes from
outcrops along Kern River, in the Round Mountain silt member of the Temblor
formation. Study of the fauna indicates that it is of lower to middle Miocene
age and is probably a correlative of some part of the Alum Bluff formation of
Florida. In the discussion of comparative material, a homonym, Typhis (Tali-
typhis) costaricensis Olsson, 1942, not Olsson, 1922, is renamed T. (T.) olssont.

ACKNOWLEDGMENTS

As indicated in the body of the paper, I am greatly indebted to Mr.
Donald C. Birch and Mr. Robert T. White for specimens, maps, and
advice. Professor Hubert G. Schenck has read and criticised the type-
script and has suggested many improvements. In addition I have
received aid in various ways from Professor Konrad Krauskopf, Miss
Elizabeth A. Watson, Miss Lois T. Martin, and Mr. John H. Beach.
For any errors of interpretation which may have crept in, however, J
alone am responsible.

Funds for the preparation of illustrations have generously been
made available by the Research Committee of Stanford University.

SUMMARY OF THE STRATIGRAPHY OF THE KERN RIVER AREA

When new species of mollusks representing genera not previously
recorded in the Pacific Slope Tertiary were detected in samples submitted
by Donald C. Birch from collecting localities along Kern River, the
potential signficance of the discovery for inter-regional correlation was
at once obvious, and further investigation was begun. Detailed maps

and sections were prepared by Birch and by Robert T. White which,

1An abstract of a portion of this paper has appeared under the title “New Typhis
from the California Miocene’, Bull. Geol. Soc. America, vol. 50, no. 12, pt. 2, December
1, 1939, p. 1972.

GENERALIZED COLUMNAR SECTION
EAST SIDE or tHe SAN JOAQUIN VALLEY

FRUITVALE OILFIELD KEAN RIVER OILFIELD KERN GORGE

Y Y
HES

FOSSIL
LOCALITIES
2/2/.2122 LS SU

BARKER'S RANCH
“HORIZON”

RECENT

"“B" ZONE

BUTTON-B8ED

EQUIVALENT

R.M._ KLEINPELL

“A” ZONE
GR/T BED
A, 8, AND
ZONES REFER
TO FAUNAS OF
FM. ANDERSON

—

M/ICRO-FAUNAL STAGES

AFTER

PRIOGENIE SPIE Ss Ore Ee NIE
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VAQUEROS

OLIGOCENE

“KREYENHAGEN’?

N
EOCENE MH oMENGINE-IONE”

MEASURE

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MN Minas age

YS GEOLOGY BY DONALD C BIRCH

at
Say)

Fig. 2. Columnar sections at three points in the Kern River area.

KEEN—MOoLLUSKS FROM RoUND Moun rain SILT 29

through their generous cooperation, are published here.

The geology of an area about five miles wide north and south of
Kern River, which here flows westward, is shown in figure 1. This region
is about twelve miles northeast of Bakersfield, Kern County, California,
and is included in the northwest corner of Caliente Quadrangle (U. S.
Geol. Survey Topographic sheet). Fossils occur at a number of localities,
the three most important of which, for the present study, are Sharktooth
Hill, at the upper left of the figure; the classic “Barker’s Ranch” (LSJU
loc. 2641) near the center; and LSJU loc. 2121 at the right center.
The Round Mountain silt member of the Temblor formation is the
principal cartographic unit in the area; less widespread are the Olcese
sand and the Freeman-Jewett silt. Additional members of the Temblor
underlie the area but outcrop only beyond the borders of this map.

Columnar sections at three points from west to east through the
Bakersfield-Kern River district, in figure 2, show the relationships and
relative thicknesses of all members and formations in the region. Age-
assignments shown at the left represent what might be called the
standard classification for California; some revisions are suggested in
figure 5. The Barker’s Ranch “horizon” of early writers is near the top
of the right-hand section in figure 2. It includes the upper part of the
Olcese sand and the major part of the Round Mountain silt; LSJU
locality no. 2641 is in the sand of the lower part and LSJU loc. 2121
in the sandy silt in the upper part of this “horizon.” In stratigraphic
order below the Round Mountain are (1) the Olcese sand, (2) the
Freeman and Jewett silts, often indistinguishable and grading down-
ward locally into (3) the Pyramid Hill sand or “grit zone.” All of
these members are considered to form a part of the Temblor formation.
The underlying Vedder sand is generally correlated with the Vaqueros
formation. Above the Round Mountain silt the Fruitvale shale occurs
in some parts of the district. Beds of presumed Santa Margarita age,
the “Fruitvale sand” of some workers, overlie the Round Mountain.
The complete section shown in this figure is, it must be emphasized,
based upon subsurface as well as surface observations. A more extensive
review of subsurface structure is given by Ferguson.”

2Ferguson, Glenn C., “Correlation of oil field formations on east side San Joaquin
Valley”, State of California, Division of Mines, Bull. 118, pt. 2, Pre-print, pp. 239-246
1 chart, 2 plates, Aug., 1941.

30 SAN Drgsco Society OF NATURAL HIstTory

Tue Rounp Mountain SILT?

Figure 3 is a detailed columnar section of a part of the Round
Mountain silt at Stanford University locality 2121. Although fossils
occur in lenses throughout the section, they are exposed best in bands
at three levels. Species of mollusks occurring in some abundance at
each of these levels are cited in the column at the right. The longest
list is from a collection made by the writer in the siltstone twenty feet
above the base of the measured section. A mineralogical analysis of the
silt, made by Konrad Krauskopf, is summarized in the column
“Lithology.” Krauskopf comments, further :

The heavy mineral assemblage is notable for its limited variety, for the
scarcity of magnetite, and for the relative abundance of a peculiar type of
sphene. The light mineral assemblage is notable for the abundance of clear,
unaltered glass and for the relative scarcity of quartz. For so fine a rock the
sorting is surprisingly good. The glass fragments are angular and irregularly
shaped, but in general they do not have the shard-like form of tuffs. . . . The
limited variety and the abundance of such easily altered minerals as horn-
blende and biotite probably imply that the material has not been carried far
from its source.

Altogether, specimens of some 77 species of mollusks were found
at the single locality LSJU no. 2121. Compared with Miocene faunules
elsewhere, such at the Bowden fauna (Jamaica) or the Alum Bluff
(Florida) where 700 to 800 species have been recorded, this may seem
an unimpressive number. For the Miocene of California, however, it
represents unusual abundance. Preservation of the material and the
relative wealth of small forms are particularly noteworthy.

To extract the more minute specimens from the loosely-consolidated
silt, an adaptation of micropaleontologic technique was used. A block
of matrix was brought to the laboratory, where it was gently crumbled
by hand. Larger specimens, as they appeared, were removed with forceps
or needles. The remaining fragmentary material was then sifted, dry, in
a 16-mesh screen, through which the small shells passed without injury;
no water was used in screening, for moisture disintegrated the chalky
shell-material. Fragile specimens were coated with a dilute solution of

3The name “Round Mountain silt” (as also “Olcese sand”) was proposed by Alex.
Diepenbrock, “Mt. Poso oil field”, California Oil Fields, vol. 19, no. 2, 1933, p. 14, pl. 2.
The type locality is at the west edge of the Mt. Poso oil field, northeast of Bakersfield,
Kern Co., in Ohio Oil Co. well no. “Glide” 1, sec. 13, T. 27 S., R. 27 E. If this locality
cannot be considered fixed as type locality by monotypy it may be regarded as here
designated.

KEEN—MOLLUSKS FROM ROUND Mounrain SILT

DETAILED COLUMNAR

SECTION OF

ROUND MOUNTAIN (MIOCENE) SILT
Pinos 2ielaNeAR KEAN GORGE,
KERN CO., CALIFORNIA

LITHOLOGY

Gray siltstone

aoo0o000

ae Concretionary, calcareous, silty
9.910 O07

sand; fossils in lower 2 feet

Greenish-gray siltstone
Fossiliferous sandy siltstone

Gray to buff platy sandy siltstone
Fossiliferous sandy siltstone

Gray to greenish-gray, buff-
weathering platy sandy siltstone

Size analysis
Median
0.06 mn.

Smallest
0.004 mm.

Largest
0.02 mm.

Mineral analysis
Reavy mineral fraction:
Abundant—-hornblende; biotite
Common--epidote; sphene
Present--zircon; magnetite;
muscovite
Rare--augite; zoisite
Light mineral fraction:
Abundant--clear angular unaltered
glass fragments; plagioclase;
aggregate-structured brownish
«rains of weathered feldspar,
chlorite, etc.
Present--quarts; potash feldspar
forthoclase and microcline);
chlorite

Gray siltstone, thin-bedded, platy,
fossiliferous

IN: FEET

S.C ASE

Interbedded pinkish-gray sandy silt-
stone and fine buff silty sand

,.] Gray to buff friable fine sand

=] Interbedded eenish-gray pu
muds tone afd eray afiteeone

FOSSIES

Chione latilaminosa Anderson and ‘ertin
Turricula ochsneri (Anderson ané Martin),
etc.
Anadara osmonti (Dall)
Bruclarkia seniculata (Conrad)
Dentalium petricola Dall
Ficus (Trophosycon) ocoyana (Conrad)
Neverita andersoni Clark
etc.

Anadara osmonti (Dall)

Chione latilaminosa Anderson and !artin

Dosinia marsaritana ‘Jiedey

Mactra sectoris Anderson and Martin

Nucula (Snnucule) n. s>.

Nuculana ochsneri (Anderson & !artin)

Transennella joaouinensis And. & Martin

Yoldia temblorensis Anderson and tertin

Cancellaria dalliana Ancerson

Cylichna ramonensis Clark

Dentalium conradi Dall

Ficus (Tronhosycon) ocoyana (Conrad)

Mangelia kernensis Anderson and Martin

Melanella ne. sp.

Mitrella n. sp.

Nassarius arnoldi (Anderson)

Neverita andersoni Clark

Odostomia (Evalea) andersoni Bartsch

Phos dumbleana Anderson in Hanna

Pyramidella cooperi Anderson and Martin

Terebra cooperi Anderson

Trophon kernensis Anderson

Turbonille n. sp.

Turricula ochsneri (Anderson *% 'artin),
etc.

MEASURED BY R.T.WHITE

Fig. 3. Analysis of the Round Mountain silt at Stanford University

locality PAVE

gum arabic. To avoid the possibility of later recrystallization of the
gum, with consequent breakage of type specimens, the gum arabic was
removed from all specimens selected for illustration and was replaced
either with a dilute solution of cellulose acetate in acetone or with
bakelite varnish.

32 San Disco Society OF NATURAL History

PALEONTOLOGICAL RELATIONSHIPS

The genera of mollusks present in the Round Mountain silt are
tropical American, such an assemblage as may be found at almost any
collecting station off either the West or the East Coast of Central
America or in fossil deposits in those regions from Miocene time onward.
One gastropod is especially noteworthy in lending added emphasis to

WASHINGTON
SAN FRANCISCO

ANS)

Qer

A=TYPE LOCALITY OF TEMBLOR
B= KERN RIVER DISTRICT

C = RECENT,PACIFIC COAST

D = BOWDEN, JAMAICA

E = GATUN, PANAMA

F =CHIPOLA, FLORIDA

G =DOMINICAN REPUBLIC

H = MANZANILLA, TRINIDAD

EOCAEINIES “Ad- WAG
TMPAUS CTA LEP ENS) OGEURS

Fig. 4. Localities of occurence of the subgenus Talityphis.

the accepted hypothesis that a marine connection existed during the
Miocene between the California coast and the Gulf of Mexico. This,
a new species, is a member of a genus, T’yphis, which is geographically
well-distributed but, paradoxically, seldom-encountered; the species can
be allocated further to the subgenus Talityphis, also of rare occurrence,
which is restricted geographically to the tropical American region.

KEEN—MOLLUSKS FROM ROUND Mountain SILT 33

Distribution records of the subgenus Talityphis, as culled from the
literature and as observed in several museum collections in the United
States, are plotted in figure 4. Talityphis is represented in the Recent
fauna by two species; in the Panamic marine faunal province, as shown
by the letter “C”, localities of collection fall in a limited area near the
tip of Lower California; although the subgenus has been considered
extinct in the Caribbean, specimens collected among beach sand by
A. A. Olsson at Monte Cristi, San Domingo, confirm the existence of
the genotype species in the Recent fauna of the Antillean area. The
species T. alatus Sowerby and the variety T. alatus obesus Gabb were
established at several localities there during the Tertiary. The earliest
record for the subgenus is in the Chipola formation (usually considered
early Miocene) of Florida. A second species, T. (T.) pterinus Gardner,
occurred also in Florida, in the late Miocene Shoal River formation.
T. alatus and obesus have been reported from the Bowden—“D” of fig.
4—(middle Miocene) of Jamaica; from the Miocene of the Dominican
Republic, i.e., Santo Domingo (without stratigraphic allocation )—“G”;
from the Gatun (middle Miocene) at Toro Cay, Panama—“E”; and
from the Manzanilla (middle Miocene) of Trinidad—“H”. The first
recognition of the subgenus in the Tertiary of the Pacific Slope of North
America is now reported.

In the Recent fauna Typhis, sensu lato, is restricted to the neritic
zone, at depths of from 6 to 60 fathoms. The related genus
Siphonochelus has been reported from depths to 400 fathoms. Dredging
records for the subgenus Talityphis range from 20 to 40 fathoms.
Finding it in a fine-grained well-sorted sandstone or silt, as in the Round
Mountain occurrence, would indicate that the deposit was laid down
some distance off shore in a warm-temperate or tropical sea.

Other genera represented at LSJU loc. 2121, although not as
restricted in geographic or bathymetric distribution, lend plausibility to
the conclusion that this is a shallow to moderately deep-water subtropical
assemblage, closely related to Caribbean and Panamic Miocene and
Recent faunules.

AGE AND CORRELATIONS

Figure 5 is a chart to show the stratigraphic and age relationships
of those formations in which Talityphis has been recorded and their
presumed correlation with formations in the Kern River area. Data for

34 SAN Disco Society OF NATURAL History

the Antillean-Caribbean region are from Senn’, with modifications.

Most of the genera present in the Round Mountain silt range
through the Tertiary; hence, their appearance here, even if new to
California, cannot be used as paleontological evidence of age. Only two
generic units may be said, with some degree of certainty, to have
originated in the Miocene strata of the tropical American region.
Talityphis, according to the literature, makes its first appearance in the

“SANTA MARGARITA”

MIOCENE

SAND

|
| (OR FRUITVALE) | [SARMATIAN}
|
T

PORT AU
FRUITVALE SHALE TORTONIAN

SHOAL RIVER S PRINCE

=
|| GURABO
| MANZANILL A} HELVETIAN
| CERCADO ROUND MOUNTAIN

+ SILT
3

MIDDLE
MIOCENE

OAK GROVE

oe
CHIPOLA BURDIGALIAN

MIOCENE

OLCESE SAND

TEMBLOR

FREEMAN-JEWETT AQUITANIAN
SILTS

PYRAMID HILL
SAND

OLIGOCENE

CHATTIAN

VEDDER SAND RUPELIAN

MIDDLE
OLIGOCENE

TALITYPHIS ALATUS OR VARIETIES RECORDED
T. PTERINUS
, NSP.

Fig. 5. Tentative correlation of certain Antillean-Caribbean formations
with members of the Temblor formation in the Kern River, California, area.

Chipola (lower Miocene) of Florida’, which may, therefore, be a
correlative of the Round Mountain silt. The new subgenus Temblornia
is, so far as can be ascertained, represented by two species only, one in
the Round Mountain silt and one, undescribed, in the basal beds of the
lower Gatun. The Round Mountain silt would appear, on the basis of
this molluscan evidence, to fall somewhere between the Gatun and the

4Senn, A. “Paleogene of Barbados and its bearing on history and structure of
Antilean-Caribbean region”, American Assoc. Petrol. Geol., Bull., vol. 24, 1940, pp. 1548-
1610, 4 text figs., chart.

>An undescribed species from Colombia, collected in strata of possible Oligocene age,
may be a candidate for allocation to Talityphis.

KEEN—MOLLUSKS FROM ROUND Mountain SILT 35

Chipola, as shown in figure 5.°

Correlation of either the Caribbean or the Californian basins of
deposition with the standard European section must as yet be by indirect
evidence. The compilation by Senn (figure 5) attributes Burdigalian
age to the Chipola formation, Helvetian to the lower Gatun and
correlatives, and Tortonian to the Bowden. Evidence against Tortonian
age for the Round Mountain silt is to be found in the occurrence of a
fossil sperm whale, Aulophyseter morrice: Kellogg,’ in the beds at
Sharktooth Hill. This whale Kellogg considers comparable to those in
the Helvetian of Europe. As the Sharktooth Hill beds are at the very
top of the Round Mountain silt, the Round Mountain cannot be
Tortonian; the fact that this vertebrate fossil occurs in the uppermost
strata would argue, rather, in agreement with other evidence cited, that
the Round Mountain is equivalent to or older than the Helvetian. There
remains a possibility that it is Burdigalian and, even more, that the
1,600 feet of fossiliferous strata of the Temblor formation below the
Round Mountain silt member are Burdigalian or older. The provisional
correlation of these Temblor members and of subjacent and superjacent
beds in the Kern River district, as derived from a study of the contained
molluscan fauna is, then, shown in figure 5. An attempt to harmonize
this correlation with those derived from studies of other phyla, such as
Foraminifera, is beyond the scope of this report.

6For a summary of foraminiferal evidence of age, which is not altogether in accord
with the correlations suggested here, the reader should consult R. M. Kleinpell’s “Miocene
Stratigraphy of California” (American Assoc. Petrol. Geol.: Tulsa, Okla., 1938, 450 pp.,
10 figs., 19 tables, 22 plates). Kleinpell would allocate the Round Mountain silt to the
upper part of his Saucesian microfaunal Stage. The Saucesian, according to Schenck (oral
communication), is considered to be of Oligocene age. A correlation chart showing
relationships of certain microfaunal Stages of California to European Stages and to
Antillean-Caribbean formations is given on p. 14 of Schenck, H. G., and T. S. Childs, Jr.,
“Significance of Lepidocyclina (Lepidocyclina) californica, new species, in the Vaqueros
formation (Tertiary), California”, Stanford Univ. Publ. Univ. Ser., Geol. Sci., vol. 3,
no. 2, July 7, 1942, 59 pp., 4 pls.).

7Kellogg, Remington, “Study of the skull of a fossil sperm-whale from the Temblor
Miocene of southern California”, Carnegie Inst. of Washington, Publ. no. 346, 1927, pp.
1-23, pls. 1-9.

SAN Disco Society oF NATURAL HIsTory

MOLLUSCAN SPECIES COLLECTED AT
STANFORD UNIveErRSITY Loc. 2121

A=Abundant (more than 50 specimens)
C=Common (25 to 50 specimens)

I= Infrequent (5 to 25 specimens)
R=Rare (5 specimens or less)

PELECYPODA
Aequipecten andersoni barkerianus? (Arnold)... C
Anadatazosmontis (Dally. is ec seet cee te eee eth res meee C
Bornia (Temblornia) triangulata (Anderson and Martin) ........ I
Chione (Lirophora) latilaminosa Anderson and Martin............ S
Chione (Chione) temblorensis (Anderson) .........-....222.---2.---------. GS
Donasd nisp ts sock on de Scher we Bee Oe ast ae R
Dosinia (Dosinidia) margaritana Wiedey................--..-.--.-----. E
Jsuiciniscamenucarin: spss.-ia = eee re eee eee R
Macoma (Rexithaerus) copelandi Wiedey.................-..-..-.-.--.-.-- I
Mactra sectoris Anderson and Martin.................0--222---------e GS
Milthansanctaeertcis: (Arnold =. 8s ee eee ee I
Nuculay(Ennucula)sbirchin sp. ee et ee (G
Nuculana ochsneri (Anderson and Martin) ............---..----2.0-2-------- G
Pandora (Kennerlyia) acutirosttata Clark! eae ee R
Taras buwaldana (Anderson and Martin) ................----2--.-----0---- R
ellina-spact <li carctata\(Contad oe ee ee ee ee ii
‘vellinan(Arcopacia)) ecoyanal@ontad=0 =e nn eee I
Tellina (Peronidia) oldroydi Wiedey............... fe joe eee ana I
Tellina wilsoni Anderson and Martin... .........0..0..20002-20-0-------- I
Transennella joaquinensis Anderson and Martin... A
Yoldia ioregoma Shumate ees eee ee I
Yoldia temblorensis Anderson and Martin.,.........00--- A
GASTROPODA
Acteon: boulderana Etheringtons== =. 2s ee I
“Amphissa” posunculensis Anderson and Martin... R
Anachisswatsonae isp ere eee Sheet sro eee R
BalGis;*conchnita sie spe tes eee ne epee ce Cc
Brucatka¥ceniculatan (Conrad) oe ee I
Bulla cantuaensis Anderson and Martin.................-.---------- R
G@alliostomazct.- ©. diabloensiss Clark == ee eee R
Calyptraea!filosacGabbs:. 2 eee eg ee ee ee I
Cancellaria barkeri (Anderson in Hanna) ..............-.-2----2-------------- I
Cancellariacondon: Anderson..+.....0.23 ee R
Cancellatia dalliana Anderson.......20020226.2.220 oe A
Cancellaria- pactiiea, “Andetson:.2 == ee I
Gancellaria ramonensis* Clark... 42 ee. Be R
@ancellariassmmplex Anderson ee eee ee R

Chrysallida rotundomontatial fs cpa eas eee R

KEEN—MOoLLUSKS FROM ROUND Mountain SILT

Conus owenianusy Andersons)... 25028. Se Ea A
Grepidulasrostralisn Conrad: s. = 22 fea ea R
@vlichna (2) loismattinae i. Sp... cs1-5-2tt-.-..... Pee Se Rees > A
@ylichnatemblorensisy iis spot oe ek a lM se! R
Epitontum,ct-/E./indiasorum® (Carpenter) =... ne I
Eulima gabbiana (Anderson and Martin)... R
perminoscalagdutianiivn.;spescnn ee ee ee Re a I
Berminosealanwhitetstcsp: 2 ses cae as ee R
Ficus: (@irophosycon) ocoyana (Conrad) 22... 228 oe G
Fetestur ety on ornate Soto Sateen kee cence she ace paeet rennet I
Mangelia kernensis (Anderson and Martin)... A
Megasnrcula howei,Idanna and Plertletn a2 = 2 0..2.2ct te I
Mitrelllag (Wiicrella)anichuela: ms Sp. oie. 2 eee ee I
INMionilaopsisyelectilisitis Spies. cnc tte. ces en ot ape scene R
Nassarius antiselli (Anderson and Martin) ............-.-...-.-2-222-.--.-. R
INassarius arnoldir (Anderson) =e 2a ee A
INassariusiblaker (Anderson and Martin) =. 2.-2024.22--62--c-ces00--- R
Nassarius ocoyanus (Anderson and Martin) ................-.--.---------- R
INatica posunculavidanna‘and Plertlein’...0 2-28. et R
IWeverttagandensonic Clarkes tate ee. oe eis iets ee oh A
INeverttayeallosa,Gabpie eee oe ee eg I
@dostomia’ (Evalea): andersoni Bartsch: 2.20 ee I
@Olivellasischnonstie spo eee eo ee eee Ue Bh ee R
Phos (Antillophos) dumbleana Anderson in Hanna.................... I
Pyramidella cooperi Anderson and Martin..............2-.2-.-.2:0----0---- A
Sinumscopulosuim (Comtad) =o aes nse e it eee econ ees eee R
Syemolapseaned ixatives pre 1 ome ee ee eS Soe aha ena R
iieimostomal (Meinostoma’ yelensinrsp). ce. .228 et 2s. ee R
Werebra- cooper < Atider soit x tastes fats tae as ste taes A
Mirephonskernensisy Anderson: t.25 22s et kh eect I
Murbonilla<(Pyrgiscus) bravoensis n. sp... 522-2. 2e I
Turbonilla (Pyrgolampros) mariposa n. sp.........-.-.-.---------------- R
Turricula buwaldana (Anderson and Martin) .....................-.-.-. R
Turricula ochsneri (Anderson and Martin) ..............-.-.--------------- I
aMiemeulavpiercer (Arnold 2 Si. eB oss hiedee A
Njuceitellatocoyanay Conrad e-eh ROP SN I
Biv piss ( Walicyphis)\Wlampadans Sp... 2.22. eee ee R
Wolvulellaselumanasp: 25 81g ES A R
SCAPHOPODA
Dentalhtumeconradie Dall a0 2 ee ee A

Herrera peter cen aM al es eek ee eae coe acs ea cata cne Senet C

38 SAN Deco Society OF NATURAL History

STANFORD UNIVERSITY COLLECTING LOCALITIES

LSJU loc. no. 2121: California, Kern Co., Caliente Quadrangle,
near center of southwest quarter of sec. 6, T. 29 S., R. 30 E., Mount
Diablo B. L. and M., in small gully close to terrace contact. Strati-
graphic horizon: lowermost part of Round Mountain silt. Collectors:
Donald C. Birch, 1938; Robert T. White, 1939; Lois T. Martin and
Elizabeth A. Watson, 1940; A. Myra Keen, 1939.

LSJU loc. no. 2641: California, Kern Co., Bakersfield or Caliente
Quadrangles, in sec. 5, T. 29 S., R. 29 E., M. D. B. and M., 1,000 feet
south and 600 feet west of the northeast corner of the section, on the
side of a gully, approximately 250 feet north of where a small gully
flows onto the alluvium of Kern River. Stratigraphic horizon: basal
Round Mountain silt or uppermost Olcese sand.* Collector, Robert T.
White.

DESCRIPTION OF SPECIES

The paleontological descriptions which follow are arranged alpha-
betically by genera and species, under the major divisions of “Pele-
cypoda” and “Gastropoda.”

All holotypes are deposited in the Stanford University Paleonto-
logical Type Collection. Available paratypes have been distributed to
the California Academy of Sciences, the University of California, the
United States National Museum, and San Diego Society of Natural
History. ;

Unless otherwise specified, the material was collected by the
writer.

PELECYPODA

Genus BORNIA Philippi, 1836
Type (by subsequent designation, Stoliczka, 1870) : Bornia corbuloides Philippi,
1836.
Subgenus TEMBLORNIA Keen, n. subg.
Type: Donax triangulata Anderson and Martin, 1914.

Description —Resembling Bornia in outline, with radial sculpture on

8Comment by White on this stratigraphic allocation: “Fossils at LSJU loc. 2641
range through a stratigraphic section totalling 120 feet in thickness. This unit is pre-
dominantly a fine silty sand and sandy silt which may represent a sandy facies of the
lower Round Mountain silt as exposed at Round Mountain. The sand content, on the
other hand, might justify interpretation of the unit as the uppermost portion of the Olcese
sand.”

KEEN—MOLLUSKS FROM ROUND Mountain SILT 39

anterior and posterior slopes; differing from Bornia, sensu stricto, in the structure
of the hinge; resilifer small and shallow, ventral margin of hinge plate entire,
not bisected as in Bornia, s. s., by the insertion of the resilium; hinge teeth well
developed, consisting of two cardinals and a posterior lateral in the left valve,
two cardinals in the right.

Discussion—Temblornia is represented by one species in California. An
undescribed and closely similar species has been detected in the lower Gatun
formation of the Panama Canal Zone. There is a striking resemblance,
especially as regards the hinge, between Temblornia and the New Zealand
Tertiary group Semeloidea Bartrum and Powell (genotype, S. donaciformis
Bartrum and Powell, 1928, Trans. N. Z. Inst., vol. 59, p. 158, pl. 29, figs.
49-50). In Semeloidea, however, the lower margin of the hinge plate is
angulate, not smoothly arched; the posterior lateral tooth of the left valve is
longer and heavier, and the posterior cardinal is more curved; exteriorly, the
radial corrugations are fewer and markedly heavier. Finlay has suggested
(Trans. N. Z. Inst., vol. 61, 1930, p. 255) that Semeloidea, which was proposed
as a genus of the Semelidae, should be regarded as a subgenus of Bornia. Thus
it is apparently coordinate in rank with Temblornia. A revision of the
numerous taxonomic units of the Erycinidae is much needed.

Bornia (Temblornia) triangulata (Anderson and Martin), 1914

Donax triangulata Anderson and Martin. Proc. Calif. Acad. Sci., ser. 4, vol.
4, p. 63, pl. 3, fig. 9.
Plate 3, figs. 6, 7

Discussion—Examination of the holotype (California Academy of
Sciences Paleo. Type Coll. no. 130) shows that this is not a Donax, but
rather a representative of a new subgenus of Bornia in which the hinge is strongly
developed. The shell is of shining porcellaneous texture with six to eight faintly
incised grooves at the anterior and posterior ends. The grooves crenulate the
inner margin slightly.

Hy potypes.—Stanford Univ. Paleo. Type Coll. nos. 7523, 7523-a.

Dimensions —No. 7523, height 4.3, length 6.5 mm.

GeNus DONAX Linnaeus, 1758

Type (by subsequent designation, Schumacher, 1817) : Donax rugosa Linnaeus,
1758.

Donax n. sp.

Plate 3, fig. 8

Although the species assigned by Anderson and Martin to Donax turns
out to be a Bornia, the genus Donax is present in the Round Mountain Silt.
A single broken specimen was obtained which is illustrated here. It is too
fragmentary to warrant bestowal of a specific name.

Hypotype—LSJU Paleo. Type Coll. no. 7524, from LSJU loc. 2121.

Dimensions.—Height (incomplete) 3.6, length (incomplete) 5.5 mm.

40 SAN Digeco Society oF NATURAL HIstory

Genus DOSINIA Scopoli, 1777

Type (by subsequent designation, Herrmannsenn, 1847): Chama dosin Adan-
son, 1757, = Artemis africana Hanley, 1843, ?=Arthemis africana Gray,
1838.

Subgenus Dosinip1a Dall, 1902
Type (by original designation) : Venus concentrica Born, 1778.

Dosinia (Dosinidia) margaritana Wiedey, 1928

Trans. San Diego Soc. Nat. Hist., vol. 5, no. 10, p. 145, pl. 18, figs. 1-3. Type
locality: Near La Panza, eastern San Luis Obispo Co., California;
Vaqueros formation.

Plate 3, figs. 3-5

So abundant are young specimens of Dosinia margaritana in the Round
Mountain silt that they might easily be mistaken for adults of some minute
pelecypod. At the same horizon occur adult Dosinia with well-preserved beaks
which, by their outline and the spacing of concentric sculpture, leave no room
for doubt of the identity of the small shells. As compared with the adult, the
juvenile Dosinia has a much less massive hinge plate; the hinge teeth are
relatively more widely spaced and the anterior cardinal and lateral teeth more
conspicuous.

Hypotypes—LSJU nos. 7525, 7525-a, 7525-b, from LSJU loc. 2121.

Dimensions.—No. 7525, height 3.6, length 3.8 mm.; nos. 7525-a, 7525-b,
height 2.5, length 2.7 mm.

Genus LUCINISCA Dall, 1901
Type (by original designation) : Lucina nassula Conrad, 1846.

Lucinisca menuda Keen, n. sp.
Plate3, figs..155 16

Shell small, nearly circular in outline; sculpture of radial and concentric
riblets which intersect as beads, more closely spaced on posterior slope than on
central and anterior parts of shell (somewhat obscured in holotype by an
incrustation of sand grains which could not be removed) ; beaks nearly central,
dorsal margin sloping downward at a low angle; posterior margin somewhat
truncate, joining dorsal at an angle of about fifty degrees; ventral and anterior
margins evenly rounded; margins crenulated by the ends of the radial ribs;
muscle scars subequal, pallial line entire; hinge of left valve strong, with two
anterior lateral teeth, two subequal cardinal teeth, and a double posterior lateral
tooth or clasper; right valve not available.

Holotype.—A left valve, Stanford Univ. Paleo. Type Coll. no. 7526,
collected by R. T. White from LSJU loc. 2121.

Dimensions —Height 6.7, length 6.5 mm.

Discussion—From the West American Recent species Lucinisca nuttalli
(Conrad) this is distinguished by the truncate posterior margin and the more

KEEN—MOoLLUSKS FROM ROUND Mountain SILT 41

attenuate anterior margin. Possibly L. menuda may be the Lucinisca aff. L.
nuttalli (Conrad) recorded by Loel and Corey (Univ. Calif. Publ. Bull. Dept.
Geol. Sci., vol. 22 ,1932, p. 210); attempts to locate the specimens mentioned by
Loel and Corey have not been successful.

Several species of Lucinisca are reported in the Chipola and Bowden forma-
tions of the Caribbean area.

The species name menuda is from the Spanish, meaning “minute.”

Genus NUCULA Lamarck, 1799
Type (by monotypy): Arca nucleus Linnaeus, 1758.

Subgenus ENNUCULA Iredale, 1931
Type (by original designation) : Nucula obliqua Lamarck, 1819.

Nucula (Ennucula) birchi Keen, n. sp.

Nucula (Ennucula) n. sp. Schenck and Keen, 1940. Calif. Fossils for the Field

Geologist, p. 10, pl. 4, figs. 6-7.

Plate 3, figs. 9-12

Shell small, solid, ovate, beaks low, opisthogyrate; lunule and escutcheon
not marked; sculpture of incremental lines only; posterior end truncate, anterior
produced and rounded, base arched; interior nacreous, adductor and auxiliary
muscular impressions present, faint (not shown in figures), basal margin
smooth; hinge strong, chondrophore oblique, anterior teeth 15, posterior 6.

Holotype.—Stanford Univ. Paleo. Type Coll. no. 7527; paratypes nos.
7527-a, 7527-b; hypotype (specimen figured by Schenck and Keen) no. 7320.

Dimensions.—Holotype, length 6.5, height 5.2, semi-thickness 1.5 mm.;
paratype 7527-a, length 6.9, height 5.2; paratype 7527-b, length 7.0, height 5.0,
thickness 3.0 mm.

Discussion—From the only other described West American Miocene
Nucula, N. washingtonensis Weaver (Univ. Washington Publ. Geol., vol. 1,
no. 1, 1916, p. 34, pl. 3, figs. 27-29), this is distinguished by its smaller
diameter, its more ovate outline, and its lack of impressed “lunule” (ie.,
escutcheon). Nucula postangulata Clark (Univ. Calif. Publ. Bull. Dept. Geol.,
vol. 11, 1918, p. 122, pl. 13, figs. 2,5) of the Oligocene San Ramon formation,
California, is less attenuated anteriorly and has an impressed escutcheon,
according to the original description. From Nucula taphria Dall (Trans.
Wagner Free Inst. Sci., vol. 3, pt. 4, 1898, p. 576, pl. 32, fig. 14) of the
Florida Miocene N. birchi differs by its smoother exterior and its low beaks.

Named in honor of Donald C. Birch, whose careful collecting and mapping
paved the way for the present study.

Genus TRANSENNELLA Dall, 1883
Type (by monotypy) : Cytherea (Transennella) conradina Dall, 1883.

Transennella joaquinensis Anderson and Martin, 1914

Proc. Calif. Acad. Sci., ser. 4, vol. 4, p. 60, pl. 3, figs. 6 a-c.
Plate 3, figs. 1, 2

Interiors of two specimens are figured here to show the hinge, inadequately

42 SAN DigEco SociETY oF NATURAL HIstTory

illustrated by Anderson and Martin.

In common with most West American species assigned to Transennella,
the interior shell margin does not exhibit the striations characteristic of the
genotype, which is from the Recent fauna of Florida. Woodring? suggests that
some or all West American species should be excluded from Transennella.
Research to settle this point is beyond the scope of the present paper; such
material in the Stanford University collection as has been scrutinized does not
entirely confirm Woodring’s observation. For the time being, therefore,
joaquinensis is retained in Transennella.

Hypotypes.—Stanford Univ. Paleo. Type Coll. nos. 7528, 7528-a.

Dimensions —No. 7528, height 4.9, length 5.5; no. 7528-a, height 3.1,
length 3.5 mm.

GASTROPODA

Genus ACTEON Montfort, 1810
Type (by monotypy): Acteon tornatilis (Gmelin) =Bulla tornatilis Linnaeus,
1758.

Acteon boulderana Etherington, 1931
Univ. Calif. Publ. Bull. Dept. Geol. Sci., vol. 20, no. 5, p. 113, pl. 14, fig. 9.
From the Astoria (Miocene) of south-west Washington.
Plater4; dig. 122

Hypotype—Stanford Univ. Paleo. Type Coll. no. 7529, from LSJU loc.
2641, R. T. White collector.

Dimensions —Height 18.0, diameter 9.3 mm.

Discussion—Although Etherington cited specimens of boulderana from
the Kern River area, no illustrations of California specimens have been published,
nor has the occurrence received mention by other authors. To authenticate the
record, therefore, a figure is included here.

Genus ANACHIS H. and A. Adams, 1853

Type (by subsequent designation, Tate in Woodward, 1875): Columbella
scalarina Sowerby, 1832.

Anachis watsonae Keen, n. sp.
Plate 4, figs. 1, 2

Shell moderately small and stout, whorls of spire almost flat; anterior canal
short, pillar wide; inner lip smooth, outer lip broken in only specimen available;
sculpture apparently of axial ribs only on spire (8 ribs on antepenultimate whorl
in holotype); body whorl smooth, with 8 incised spiral grooves on pillar and
base.

Holotype—Stanford Univ. Paleo. Type Coll. no. 7530, from LSJU loc.
22

9In Woodring, W. P., R. Stewart, and R. W. Richards, “The geology of the
Kettleman Hills oil ftield, California,’ U. S. Geol. Survey Prof. Paper 195, “1940”
(published in 1941), p. 95.

KEEN—MOLLUSKS FROM RouUND MounraIn SILT 43

Dimensions —Height 9.7, diameter, 4.3 mm.

Discussion—Only one broken specimen is available. On this the cortical
layer is lacking on the spire; however, the persistent axial ribs and the general
outline of the shell make the diagnosis as a new species of Anachis plausible.
No closely similar species appears in the literature on the Miocene of the
Caribbean area or in the California Tertiary record. Numerous species of
Anachis have been described from the Panamic marine province.

Named for Miss Elizabeth Watson, in appreciation of assistance in
collecting material from the Round Mountain silt.

Genus BALCIS Leach in Gray, 1847

Type (by monotypy): Balcis montagui Leach in Gray=Strombiformis albus
Da Costa, 1778.

Balcis conchita Keen, n. sp.
Plate 4, fig. 5

Shell elongate-conic, slender, straight, whorls flattened, appressed at the
summits; periphery of the last whorl rounded, base long, well rounded; aperture
large, oval; posterior angle acute; outer lip rounded; inner lip short, stout,
somewhat curved, reflected and appressed to the base; parietal wall covered with
a callus.

Holotype.—Stanford Univ. Paleo. Type Coll. no 7538, from LSJU loc.
2121.

Dimensions—Height 6.4 (estimated), diameter 1.5, height of aperture
1.3 mm.

Discussion —-The nearest Recent West American relative of this species
is Balcis rutila (Carpenter) (Suppl. Rept., British Assoc. Adv. Sci. for 1863,
1864, p. 659, figured, as Melanella, by Pach (Proc. Ws 5: Nate Mus. vol-
53, 1917, pl. 35, figs. 2, 3, 6), which is slightly larger and proportionately
widen None of the Caribbean species in the literature seems to be close to
B. conchita.

Winckworth (Jour. Conch., vol. 20, no. 1, May, 1934, pp. 12-13) has
ereaenrne nemre Wiclarciin astordoabial validity, as the genotype is
virtually unidentifiable. The next available synonym is Balcis, which is here
tentatively accepted.

The specific name conchita is derived from the Spanish word for “little

shell.”

Genus CHRYSALLIDA Carpenter, 1856

Type (by subsequent designation, Dall & Bartsch, 1904): Chrysallida
communis Carpenter, 1856, (not C. B. Adams, 1852) = Odostomia (Chry-
sallida) torrita Dall and Bartsch, 1909.

Chrysallida rotundomontana Keen, n. sp.
Plate 4, fig. 28
Shell small, ovate-conic; nuclear whorls not preserved on holotype; post-
nuclear whorls flattened, strongly contracted at sutures and slightly shouldered

44 SAN Disco SoctEty oF NATURAL HIStory

at summits, marked by about 16 curved axial ribs which cross four strong spiral
cords with nodose intersections; periphery of last whorl marked by a spiral
cord not crossed by axial sculpture; base nearly smooth, with about 8 shallow
spiral grooves which become fainter toward columella; aperture oval, posterior
angle acute; columella stout, reflected anteriorly, with a strong fold at its
insertion.

Holotype—LSJU Paleo. Type Coll. no. 7531, from LSJU loc. 2121.

Dimensions —Height 2.8, diameter 1.4 mm.

Discussion —In the Recent fauna of the West Coast Odostomia (Chry-
sallida) pulcia Dall and Bartsch, 1909, is the closest relative. It is slightly
smaller than C. rotundomontana, with more numerous axial ribs. From
Odostomia (Chrysallida) melanoides (Conrad) as figured by Martin, 1904,
Maryland Geol. Survey, Miocene, p. 220, pl. 54, fig. 1, C. rotundomontana
differs in being smaller, proportionately broader, and less conspicuously
sculptured on the base.

The name rotundomontana is the Latinization of “Round Mountain.”

GeNus CYLICHNA Loven, 1846

Type (by subsequent designation, Herrmannsenn, 1852): Bulla cylindracea
Pennant, 1777.

Cylichna? loismartinae Keen, n. sp.
Plate 4, figs. 16, 18

Shell small, cylindrical, posterior end rounded; spire concealed, apex slightly
depressed and bounded by a low ridge; aperture as long as shell, anterior end
dilated, posterior end narrowed toward apex; columella with a low, oblique
basal fold; sculpture of faint incised grooves, more apparent at anterior end, and
incremental lines.

Holotype—Stanford Univ. Paleo. Type Coll. no. 7532, from LSJU loc.
ZAG

Dimensions.—Height 3.4, diameter 1.8 mm.

Discussion—From Cylichna temblorensis this species is readily separable
by its greater width, its smaller size, and its posteriorly narrowed aperture. At
first glance the outline would suggest Cylichnella, but all six specimens available
lacked the second columellar fold of that genus. The species is therefore
tentatively allocated to Cylichna until such time as a revision of the Scaphand-
ridae makes possible a more precise assignment. No other Miocene or Recent
species in the Caribbean or West American area seem comparable.

Named in honor of Miss Lois T. Martin, who assisted in collecting material
from the Round Mountain silt.

Cylichna temblorensis Keen, n. sp.
Plate 4, figs. 13, 14

Shell small, cylindrical, posterior end truncated; spire immersed, apex
concave; aperture as long as shell, anterior end dilated, posterior end somewhat
elevated above remainder of body whorl; columella bearing an obscure, oblique

KEEN—MOoLLUSKS FROM ROUND MounrtAIN SILT 45

basal fold; sculpture of occasional faint grooves and incremental lines.

Holotype—Stanford Univ. Paleo. Type Coll. no. 7533, from LSJU loc.
2NZAe

Dimensions.—Height 4.0, diameter 1.6 mm.

Discussion—From Cylichna ramonensis Clark, 1918 (Univ. Calif. Publ.
Bull. Dept. Geol., vol. 11, p. 188, pl. 20, fig. 7) this species is distinguished by
its smaller size and more truncate posterior end; from C. aula Woodring of the
Bowden Miocene (Carnegie Inst. Publ. 385, 1928, p. 129, pl. 2, fig. 19) by its
greater height and the lack of a grove at the apex. From all Chipola (Florida,
Miocene) species temblorensis is distinguished by its proportionately greater
height and the shallower umbilical groove, and from the West American Recent
“C. alba Brown” of authors it is distinguished by its consistently smaller size
and its almost complete absence of spiral sculpture.

Genus EULIMA Risso, 1826

Type (by subsequent designation, Herrmannsenn, 1846): Turbo subulatus
Donovan=Strombiformis glaber Da Costa, 1778.

Eulima gabbiana (Anderson and Martin)

Eulimella gabbiana Anderson and Martin, 1914. Proc. Calif. Acad. Sci., ser.
4, vol. 4, p. 68, pl. 7, fig. 20. Near Barker’s Ranch, Kern Co., Miocene.
Plate 4, fig. 6

Hypotype.—Stanford Univ. Paleo. Type Coll. no. 7543, from LSJU loc.
2641, collected by R. T. White.

Dimensions.—Height 9.5, diameter 1.9, height of aperture 2.3 mm.
Discussion.—This species was, on the basis of the original figure, allocated

to Melanella by Bartsch, 1917 (Proc. U. S. Nat. Mus., vol. 53, p. 316). Study
of the incomplete holotype (Calif. Acad. Sci. Paleo. Type Coll. no. 143), a
specimen with only the last 214 whorls, and comparison with material from the
type locality show this to be an Eulima'®. This genus is distinguished from
Melanella of authors (Balcis Leach of this paper), by the attenuated outline and
narrow, elongate aperture. A virtually complete specimen from the type locality
is figured here.

No Recent West American species of Eulima approaches E. gabbiana
closely in size and proportions; the nearest is E. californica (Bartsch) (Proc.
U. S. Nat. Mus., vol. 53, 1917, p. 340, pl. 45, fig. 5) which, however, is larger
and proportionately more slender. E. migrans Conrad, 1846, as figured by
Martin, 1904 (Maryland Geol. Survey, Miocene, p. 217, pl. 53, fig. 12), from
the St. Mary’s Miocene of Maryland is, if Martin’s dimensions are correct,
considerably more slender. Martin’s illustration shows a wider, shorter shell,
with a wider aperture and a more impressed suture.

10“Strombiformis” of authors, not of Da Costa as designated by Harris, 1894. For a
discussion of nomenclature in this family, see Winckworth, Jour. of Conch., vol. 20, no. 1,

May, 1934, pp. 12-13.

46 SAN Disco Society oF NATURAL History

GENus FERMINOSCALA Dall, 1908
Type (by original designation): Epitonium (Ferminoscala) ferminianum Dall,
1908.

Ferminoscala durhami Keen, n. sp.

Plate 4, fig. 31

Shell small, conical, whorls rounded, sutures impressed; sculpture of six
raised spiral threads about equally spaced, crossed by weak axial threads more
apparent on spire than on later whorls; aperture round, inner lip reflected,
outer lip broken on both specimens observed, probably thin; basal disk with
spiral threads only.

Holotype-—Stanford Univ. Paleo. Type Coll. no. 7534, from LSJU loc.
DOANE

Dimensions.—Height (incomplete) 8.7, diameter 3.0 mm.

Discussion—Compared with F. whitei, described below, this shell has a
wider apical angle and lacks the heavy axial sculpture. It corresponds to
F. pseudoleroyi (Maury) (Bulletins of Amer. Paleo., vol. 10, no. 42, 1925, p.
394) from Jamaica, whereas F. white: corresponds to F. spathe Woodring.
From F. pseudoleroy: it differs in the greater apical angle and in the narrowness
of its spiral ribs. It resembles F. prunicola (Martin) (Maryland Geol. Survey,
Miocene, 1904, p. 214, pl. 53, fig. 6) but has fewer and weaker spiral ribs.

As Woodring remarks (Carnegie Inst. Publ. 385, 1928, p. 402), “There
is no reason to suppose that Ferminoscala bears any direct relation to Acrilla
H. and A. Adams’, under which it is often subsumed.

This species is named for Dr. J. Wyatt Durham in recognition of his
his work upon West American Tertiary Epitoniidae.

Ferminoscala whitei Keen, n. sp.
Plate 4, figs. 32, 33

Shell of medium size, slender, with eight whorls sculptured with about
25 to 30 retractive axial lamellae per whorl and 6 high spiral threads, producing
deep rectangular pits; in addition with microscopic axial and spiral sculpture,
especially in pits; sutures impressed, somewhat channeled; basal disk with four
spiral threads; basal lip bent backward, forming a narrow and obscure fasciole.

Holotype-——Stanford Univ. Paleo. Type Coll. no. 7535, from LSJU loc.
2121, collected by R. T. White.
Dimensions.—Height 15.5, diameter 6, height of aperture 4.6 mm.

Discussion—This species is similar to F. spathe Woodring (Carnegie
Inst. Publ. 385, 1928, p. 403, pl. 32, fig. 5) from the Bowden and to
F. reticulata (Martin), 1904 (Maryland Geol. Survey, Miocene, p. 214, pl.
53, fig. 5) from the Miocene of Maryland, but is wider, with more impressed
sutures than either.

Named for R. T. White, the collector, in appreciation of advice on
problems of stratigraphy, preparation of a columnar section of the Round
Mountain silt, and collection of specimens.

KEEN—MOLLUSKS FROM ROUND MounraIN SILT 47

Genus HASTULA H. and A. Adams, 1853

Type (by subsequent designation, Cossmann, 1896): Buccinum strigilatum
Linnaeus, 1758.

Hastula gnomon Keen, n. sp.
Plate 4, fig. 11

Shell of medium size, slender, whorls flat; sculpture of narrow axial ribs
which disappear on anterior part of later whorls, numbering (on holotype) 17
to 20 ribs per whorl; surface except for the puckered axial ribbing smooth and
shining; outer lip of aperture evidently thin (broken on all specimens
examined) ; parietal wall with a thin callus, columella bearing an obscure basal
fold, siphonal fasciole with a faint median groove.

Holotype—Stanford Univ. Paleo. Type Coll. no. 7536, from LSJU loc.
2121, Donald Birch collector.

Dimensions.—Height 17.2, maximum diameter 4.6, height of aperture 4.6
mim.

Discussion—In the West American Recent fauna Hastula is represented
in the Panamic area by Terebra luctuosa Hinds, 1844 (Proc. Zool. Soc. London
for 1843, p. 157), which is larger, with more numerous axial plications.
Woodring has reported two species of Hastula from the Bowden formation
of Jamaica (H. jamaicensis Woodring, Carnegie Inst. Publ. 385, 1928, p. 143,
pl. 4, fig. 4, and H. homala Woodring, ibid., pl. 4, fig. 5). H. gnomon
resembles the former in size, the latter in outline but has less prominent axial
ribs than either. Of the four St. Mary’s and Calvert Miocene species described
by Martin (Maryland Geol. Surv. Miocene, 1904, pp. 143-144, pl. 40, figs.
10-14), Terebra (Hastula) patuxentia is nearest to gnomon; its axial sculpture
is more closely spaced, however, and the illustration shows spiral striations. No
Hastula have been reported from lower or middle Miocene deposits in the
Caribbean. Therefore, this record in California seems to be the earliest in the
Americas. The genus is recorded in the Eocene of Europe.

The word gnomon is a Latin noun, masculine gender; a gnomion is a
pin or pointer on a sun dial.

Genus MANGELIA Risso, 1826
Type (by subsequent designation, Herrmannsenn, 1852): Mangelia striolata
Risso—Murex attenuatus Montagu, 1803.
Subgenus CACODAPHNELLA Pilsbry and Lowe, 1932
Type (by original designation): Cacodaphnella delgada Pilsbry and Lowe,
1932.

Mangelia (Cacodaphnella?) kernensis (Anderson and Martin)

Mangilia kernensis Anderson and Martin, 1914. Proc. Calif. Acad. Sci., ser.
4, vol. 4, p. 94, pl. 7, figs. 6 a-b. Barker’s Ranch, Kern Co., Miocene.
Plate 4, fig. 21
Hypotype——Stanford Univ. Paleo. Type Coll. no. 7537, from LSJU loc.
Pale

48 San Disco SocitETy oF NATURAL HIstory

Discussion—The nuclear whorls of M. kernensis do not conform to the
pattern of any of the numerous genera of small sculptured turrids described by
Woodring (Carnegie Inst. Publ. 385, 1928, pp. 165-198). The first two whorls
are smooth, the next half-turn with 5 to 7 spiral threads crossed but not
cancellated by fine, almost microscopic, axial riblets. In the third or post-
nuclear whorl the first sinuous axial ribs appear, puckering the spiral ribs at
regular intervals. Axial ribs are more prominent than spiral in later whorls,
the spiral ribs crossing over axials as undulating threads. In Cacodaphnella,
as described by Pilsbry and Lowe (Proc. Acad. Nat. Sci. Philadelphia, vol.
84, p. 58, 1932), both axial and spiral sculpture appear in the nuclear whorls
though never as the heavy cancellations common in other groups of small
turrids such as Cryoturris Woodring (op. cit., p. 178). Although Pilsbry and
Lowe did not observe any other Recent West American species which they
could assimilate to Cacodaphnella, there seems to be a candidate in Mangelia
densilineata (Dall) (U.S. Nat. Mus. Bull. 112, p. 83, 1921). Cacodaphnella
would thus appear to be a turrid group comprised of three species, one Miocene
and two Recent, and restricted in distribution to the West Coast of North
America.

GENUS MITRELLA Risso, 1826

Type (by subsequent designation, Cox, 1927): Mitrella flaminea Risso, 1826
= Murex scriptus Linnaeus, 1758.

Mitrella (Mitrella) anchuela Keen, n. sp.
Plate 4, fig. 12

Shell small, stout, pillar constricted; whorls of spire slightly bulging;
aperture relatively wide; outer lip thin, set with five heavy denticles within the
aperture, attenuated anteriorly and curving into a notch-like narrow canal
(broken in holotype during preparation); inner lip smooth, reflected and
appressed to body whorl; sculpture of about five coarse threads on the pillar
and anterior portion of body whorl.

Holotype.—Stanford Univ. Paleo. Type Coll. no. 7539, from LSJU lcc.
212A.

Dimensions.—Length 5.1, diameter 2.7 mm.

Discussion —This is the shortest and stoutest of West American Mitrellas,
fossil or living. It is also distinguished from M. (Striomitrella) tenuilineata
(Clark) (Univ. Calif. Publ. Bull. Dept. Geol., vol. 11, 1918, p. 173, pl. 22,
figs. 2, 3) by its scanty spiral sculpture. Caribbean Miocene Mitrellas such as
M. lissa Woodring (Carnegie Inst. Publ. no. 385, 1928, p. 275, pl. 16, fig. 14)
or M. lepta Woodring (ibid., pl. 16, fig. 15) from the Bowden or M. communis
(Conrad) (figured as Columbella by Martin, Maryland Geol. Survey, Miocene,
1904, p. 199, pl. 50, figs. 5-7) are larger and more slender, lacking the peculiar
elongation of the outer lip.

The specific name anchuela is from the Spanish, meaning “somewhat

broad.”

KEEN—MOoLLUSKS FROM RouND Mountain SILT 49

Genus MONILIOPSIS Conrad, 1865
Type (by monotypy): Pleurotoma elaborata Conrad, 1833.

Moniliopsis electilis Keen, n. sp.
Plate 4, fig. 15

Shell small to medium in size, biconic, aperture less than half the length
of the shell; nuclear whorls not preserved on sole specimen obtained; post-
nuclear whorls rounded, suture impressed; surface cancellately sculptured by
axial ribs which bend to the right at the anal fasciole and spiral ribs of equivalent
strength, axial ribs 23 on last whorl, more pronounced below anal fasciole,
spiral ribs 6 to 8 per whorl, upper 3 ribs weaker than those below anal fasciole;
body whorl cancellately sculptured like spire; anal fasciole just above periphery
of whorl, marked by bending of axial ribs; pillar narrow, inner lip erased,
outer lip thin; aperture rounded above, prolonged into a canal below.

Holotype.—Stanford Univ. Paleo. Type Coll. no. 7540, from LSJU loc.
2121, R. T. White collector.

Dimensions.—Height (incomplete) 12.2, diameter 4.8, length of aperture
2.3 mm.

Discussion—No similar species occurs in the Miocene of West America
or the Caribbean area. Comparable Pliocene and Recent species of the West
Coast are Moniliopsis graciosana (Arnold) (Smithsonian Misc. Coll., vol. 50,
pt. 4, 1907, p. 430, pl. 54, fig. 18) from the Careaga formation (Pliocene) of
Santa Barbara County, California, which has coarser axial sculpture and is not
evenly cancellate, and M. incisa (Carpenter) (Suppl. Rept. British Assoc. Adv.
Sci. for 1863, pp. 603, 657, 1864), which has spiral sculpture only.

The name electilis is a Latin adjective signifying “dainty.”

Genus ODOSTOMIA Fleming, 1817

Type (by subsequent designation, Dall and Bartsch, 1904): Turbo plicatus
Montagu, 1803.

Subgenus Evatea A. Adams, 1860

Type (by subsequent designation, Dall and Bartsch, 1904): Evalea elegans
Adams, 1860.

Odostomia (Evalea) andersoni Bartsch, 1917

Eulimella californica Anderson and Martin, 1914. Proc. Calif. Acad. Sci., ser.

4, vol. 4, p. 67, pl. 7, figs. 19 a-c. Miocene, near Barker’s Ranch, Kern

Co. (Not Odostomia (Evalea) californica Dall and Bartsch, 1909).
Odostomia (Evalea) andersoni Bartsch. Proc. U. S. Nat. Museum, vol. 52,

no. 2193, May, 1917, p. 667.

Plate 4, figs. 17, 23

Two specimens of Odostomia andersoni are figured here. Fine spiral
striae on the surface of the shells, too minute to show in the photographic
illustraticns, indicate the correctness of the allocation to Evalea made by
Bartsch.

Hypotypes—Stanford Univ. Paleo. Type Coll. nos. 7541, 7541-a.

50 SAN DtgEco Society OF NATURAL HIstory

Dimensions —No. 7541, height 3.0, diameter 1.5; no. 7541-a, height 2.5,

diameter 1.4 mm.

GeNus OLIVELLA Swainson, 1831

Type (by subsequent designation, Dall, 1909): Olivella purpurata Swainson,
1831=Voluta dama Wood, 1828.

Olivella ischnon Keen, n. sp.
Plate 4, figs. 3, 4

Shell relatively small and slender; spire high, about three-fourths the length
of the aperture; sutural channel deep and wide; columellar folds weak, five in
number, near anterior end of inner lip.

Holotype-—Stanford Univ. Paleo. Type Coll. no. 7542, from LSJU loc.
2641, R. T. White collector.

Dimensions.—Height 9.4, diameter 4.5 mm.

Discussion—None of the Caribbean Miocene Olivellas seem to approach
this species closely in outline or in relative proportions. In the Recent West
American fauna Olivella pedroana Conrad (Pacific Railway Survey Reports,
vol. 5, 1855, pl. 6, fig. 51) is probably nearest, but it lacks the deeply channeled
suture of ischnon. From the San Ramon formation (Oligocene) of, California
Clark has described O. quadriplicata (Univ. Calif. Publ. Dept. Geol., vol. 11,
1918, p. 185, pl. 19, figs. 10, 17); this is distinguished from ischnon by its four
columellar plications and by its lower and wider spire.

The name ischnon is a Latin noun, neuter gender, signifying “a slender
thing.”

Genus SYRNOLA A. Adams, 1860
Type (by monotypy): Syrnola gracillima A. Adams, 1860

Syrnola scandix Keen, n. sp.
Plate 4, figs. 24, 29, 30

Shell of medium to small size, elongate-conic; nucleus heterostrophic, axis
at right angle to spire, about 1/; immersed in succeeding whorl; first post-nuclear
whorl rounded, later whorls nearly flat; suture deeply impressed, later whorls
appressed over sutural channel; sculpture of microscopic growth lines and spiral
striations, early whorls with a few incipient axial ribs; aperture oval, outer lip
thick in adult shell; columella with a single low fold.

Holotype.—Stanford Univ. Paleo. Type Coll. no. 7544; paratypes nos.
7544-a and 7544-b, from LSJU loc. 2121.

Dimensions.— Holotype, height 7.0, diameter 2.1, no. 7544-a, » height BOY
diameter 1.1; no. 7544-b, height 2.3, ejemmeter 0.7 mm.

Dineaan are genus S pela has not been reported in the Recent West
American fauna, and only one other Pacific Slope Miocene species is known,
Eulimella ochsneri Anderson and Martin (Proc. Calif. Acad. Sci., ser. 4, vol.
4, 1914, p. 66, pl. 7, figs. 23 a-b) which Bartsch (Proc. U. S. Nat. Mus. vol.
52, 1917, p. 639) correctly allocated to Syrnola; this species is larger and much
more deeply channeled at the sutures than scandix. Syrnola is represented in

KEEN—MOoLLUSKs FROM ROUND MOounraIn SILT Dil

the Caribbean Tertiary by such species as S. marylandica (Martin) (Maryland
Geol. Survey, Miocene, 1904( p. 221, pl. 54, fig. 6), which is proportionately
shorter with a more quadrate aperture than scandix.

Scandix is a Latin noun, feminine gender, meaning “stork’s bill.”

Genus TEINOSTOMA H. and A. Adams, 1853

Type (by subsequent designation, Cossmann, 1888): Teinostoma politum
A. Adams, 1853.

Teinostoma (Teinostoma?) lens Keen, n. sp.
Plate 4, figs. 7-9

Shell small, thin, circular, spire depressed and partially covered by suc-
ceeding whorls; outer lip, as viewed from above, arched forward between suture
and periphery; basal lip, as viewed from below, arched backward; umbilicus
obscured by a thin callus which covers about 1 of base and merges with the
thin parietal callus; sculpture of incremental lines only.

Holotype.—Stanford Univ. Paleo. Type Coll. no. 7545.

Dimensions.—Height 0.7 mm., diameter 1.8 mm.

Discussion —Teinostoma lens is intermediate in characters between the T.
nanum (Lea) and the T. calvertense Martin of the Maryland Miocene, as
figured by Martin, 1904, Maryland Geol. Survey, Miocene, pp. 263, 264, pl.
62, figs. 1-2, 3, resp.), being more depressed than the former and having a
heavier callus than the latter. It is smaller and more oblique than T. sublimatum
Dall (Proc. U. S. Nat. Mus., vol. 51, no. 2162, 1916, p. 520, pl. 88, figs. 7, 8)
from the Flint River (Georgia) “Oligocene.” It is also more depressed than
T. depressum (Gabb) figured by Pilsbry (Proc. Acad. Nat. Sci. Philadelphia,
vol. 73, 1921, pl. 37, fig. 2) from the Miocene of Santo Domingo. In the
Recent West American fauna the nearest relative appears to be T. sapiellum
Dall (Proc. U. S. Nat. Mus., vol. 56, no. 2295, 1919, p. 369), which is a
thinner shell with a more convex spire and fewer whorls.

The Latin name lens (“lentil”—masculine gender) is chosen in reference
to the lenticular shape of the shell.

Genus TURBONILLA Risso, 1826

Type (by subsequent designation, Dall and Bartsch, 1904): Turbonilla
plicatula Risso (not Brocchi) =T. typica Dall and Bartsch, 1904.

Subgenus Pyraiscus Philippi, 1841

Type (by subsequent designation, Dall and Bartsch, 1904): Melania rufa
Philippi, 1836.

Turbonilla (Pyrgiscus) bravoensis Keen, n. sp.
Plate 4, figs. 20, 26, 27
Shell small, elongate-conic; nuclear whorls 2, helicoid, axis at right angles
to spire, slightly immersed in succeeding whorl; post-nuclear whorls nearly flat,
narrowly tabulate at the summits, ornamented by 14 to 16 sinuous axial ribs

52 SAN DieEGo Society OF NATURAL HIstory

per whorl; intercostal spaces about 11/ times as wide as ribs, crossed by 7 to 8
spiral threads which divide the interspaces into a series of pits; suture impressed,
somewhat wavy; periphery of body whorl well rounded, without axial sculpture;
base nearly smooth (a few faint spiral grooves present); aperture quadrate,
outer lip rounded, columella strong, a little twisted.

Holotype—LSJU Paleo. Type Coll. no. 7546; paratype, no 7546-a.

Dimensions.—Holotype, height (incomplete) 4.1, diameter 1.3 mm.; para-
type no. 7546-a, height (incomplete) 3.0, diameter 1.0; no. 7546-b, height 1.8,
diameter 0.7 mm.

Discussion—Among Recent West American species, the nearest approach
to this seems to be Turbonilla (Pyrgiscus) virgo Dall and Bartsch (U. S. Nat.
Mus. Bull. 68, 1909, p. 93, pl. 8, fig. 4) which has, however, more axial ribs
per whorl and stronger basal sculpture than bravoensis. All of the Caribbean
Miocene species which have been figured appear to be larger and to have more
axial ribs per whorl (e.g., T. (P.) interrupta (Totten) as figured by Martin,
1904, Maryland Geol. Survey, Miocene, p. 224, pl. 54, figs. 13, 14 and T. (P.)
dominicensis Gabb from San Domingo, figured by Pilsbry, 1922, Proc. Acad.
Nat. Sci. Philadelphia, vol. 73, p. 391, pl. 36, fig. 3).

The name bravoensis is taken from the only place name which is shown
on the Caliente quadrangle near the collecting locality.

Subgenus PyRGOLAMPROS Sacco, 1892
Type (by original designation): Pyrgolampros mioperplicatus Sacco, 1892.

Turbonilla (Pyrgolampros) mariposa Keen, n. sp.
Plate’ 4; fies: 195,25

Shell small, elongate-conic; nuclear whorls 214, helicoid, turned at right
angles to spire, only slightly immersed; succeeding two whorls faintly sculptured;
remainder of shell with 12 to 14 irregularly sinuous axial ribs crossed by
numerous faint spiral grooves; intercostal spaces wider than the ribs; suture a
little channeled and wavy; aperture oblique-subquadrate, outer lip thin, joining
the twisted columella in a broad curve.

Holotype—LSJU Paleo. Type Coll. no. 7547; paratype, no. 7547-a.

Dimensions.—Holotype, height (incomplete) 3.1, diameter 1.1; paratype,
height 2.8, diameter 0.8 mm.

Discussion—No comparable Miocene species from the Caribbean area
have been described, as the Pyramidellidae of that region have not yet received
monographic treatment. Several West American Recent species have been
allocated to Pyrgolampros; the one most closely resembling T. (P.) mariposa is
T. (P.) halibrecta Dall and Bartsch, 1909 (U. S. Nat. Mus. Bull. 68, p. 65,
pl. 5, figs. 10, 10a), which, however, is larger and has more axial ribs on the
spire.

The name mariposa is from the Spanish, meaning “butterfly.”

GeNus TYPHIS Montfort, 1810

Type (by original designation): Purpura tubifer Bruguiére, 1792.

KEEN—MOoLLUSKS FROM RoUND Mountain SILT 53

Subgenus TALITYPHIS Jousseaume, 1882

Type (by original designation): Typhis expansus Sowerby, 1874 (PI. 3, fig.
20).

Typhis (Talityphis) lampada Keen, n. sp.
Plates; figs. 14,19; 23

Shell moderately large, spire somewhat low; four varices per whorl, each
prolonged at shoulder into a spine; tubules or hollow spines alternating with
varices at shoulders of whorls, closer to preceding varices and directed backward
away from aperture; varix at outer lip greatly expanded, a little foliated, joined
to preceding whorl by a broad rampart which is continuous with inner lip;
another rampart reinforces the outer edge of the varix, forming a sutural line
along the face of the varix; spiral sculpture of about six faint raised lines on
body whorl; aperture elliptical, large; anterior canal long, closed throughout.

Holotype—Stanford Univ. Paleo. Type Coll. no. 7548, collected by
Donald Birch; paratype no. 7549 collected by R. T. White; paratype, Calif.
Acad. Sci. Paleo. Type Coll. no. 7949, collected by Elizabeth Watson and
Lois Martin, all from LSJU loc. 2121. Three additional paratypes, Univ. Calif.
Paleo. Type Coll., from UC loc. 2715, Kern River area.

Dimensions —Holotype, height 24, diameter 16.5, aperture 8.2; paratype
no. 7949, height 22, diameter 16.2, aperture 7.3; paratype no. 7549, height 21.5,

diameter 16 (estimated), aperture 8.5 mm.

Discussion—The distribution of Talityphis has been commented upon
elsewhere in this paper. Here it is more appropriate to mention points of
difference which separate T. lampada from other members of the subgenus.
The genotype of Talityphis, Typhis expansus Sowerby (Proc. Zool. Soc. for
1873, p. 719, pl. 69, fig. 4, 1874) was described without locality. Specimens
collected by A. A. Olsson on the beach at Monte Cristi, San Domingo, compare
so closely with the original figure that one may assume the holotype to have
come either from San Domingo or from one of the neighboring islands of the
West Indies. Spiral sculpture of expansus is stronger and the aperture is
smaller than in lampada. The only other known Recent species of Talityphis
is Typhis latipennis Dall (Proc. U. S. Nat. Mus., vol. 56, no. 2295, 1919, p.
339, holotype figured by Maxwell Smith, Rock Shells, 1939, sp. 257, pl. 14, fig.
9), described from U. S. Bur. Fish. Sta. 2822, off La Paz, Lower California,
Mexico. The specimens figured on Plate 3, figs. 17, 21, 24, 25 of this paper
are virtual topotypes. I have compared these illustrations with the holotype of
latipennis (no. 96653, U. S. National Museum) and regard the specimens as
conspecific.

The earliest described Talityphis both geologically and chronologically is
Typhis alatus Sowerby, 1850 (Quart. Jour. Geol. Soc. London, vol. 6, p. 48,
pl. 10, fig. 4), from the Tertiary [ Miocene} of Santo Domingo. Typhis obesus
Gabb, 1873 (Trans. American Philos. Soc., n. s., vol. 15, p. 203, figured by
Pilsbry, 1922, Proc. Acad. Nat. Sci. Philadelphia, vol. 73, p. 354, pl. 28, figs.
5-6), from the Miocene of the Dominican Republic is usually regarded as a

54 San DigeGo Society oF NATURAL History

low-spired variety of alatus. A specimen of obesus figured by Olsson as Typhis
alatus (Bulletins of American Paleo., vol. 9, 1922, p. 304, pl. 13, fig. 15) from
Toro Cay, Panama (Gatun formation, Miocene), is refigured here for com-
parison. Like T. expansus it has a smaller aperture and more conspicuous spiral
sculpture than T. lampada. Typhis pterinus Gardner, which seems to be a
Talityphis (Florida Geol. Survey Bull. 14, p. 52, pl. 10, fig. 10), from the
Shoal River Miocene of Florida, has a higher spire than any other Talityphis.
A Pliocene species has recently been described from the West Coast of Costa
Rica!

The name lampada (a Latin noun, feminine gender, meaning torch or
lantern) was selected in indication of the light which this form throws upon
interregional correlation.

Genus VOLVULELLA Newton, 1891

Type (by subsequent designation, Bucquoy, Dautzenberg, and Dollfus, 1886) :
Volvula rostrata A. Adams, 1850.

Volvulella gluma Keen, n. sp.
Plate 4, fig. 10

Shell spindle-shaped, slender; aperture as long as shell, narrow, anterior
end dilated, posterior end extended; at apex a short spine concealing spire;
columella with an obscure fold; inner lip slightly reflected over a narrow
umbilical groove; sculpture of fine incised lines at both ends of shell, more
conspicuous on anterior end.

Holotype.-—Stanford Univ. Paleo. Type Coll. no. 7550, from LSJU loc.
2641, R. T. White collector.

Dimensions.—Height 3.4, diameter 1.4 mm.

Discussion—Volvulella (a substitute name for Volvula A. Adams, 1850,
not Gistel, 1848) has not previously been reported from the Tertiary in
California. From Recent West American species V. gluma is clearly distinct,
V. californica Dall (Proc. U. S. Nat. Mus., vol. 56, 1919, p. 298) being
larger and of greater diameter and V. cylindrica (Carpenter) (Suppl. Rept.
British Assoc. Advanc. Sci. for 1863, p. 647, 1864) having a shorter spine at
the apex. In the Caribbean Miocene several comparable species occur. V. oxytata
(Bush) as figured by Woodring (Carnegie Inst. Publ. 385, 1928, p. 125, pl. 2,
fig. 10) is proportionately wider than V. gluma; V. marylandica Martin (Mary-
land Geol. Survey, Miocene, 1904, p. 134, pl. 39, fig. 6) is larger, with fainter

11The description of this species, Typhis (Talityphis) costaricensis Olsson (Bulletins of
American Paleont., vol. 27, no. 106, p. 228, pl. 25, figs. 5, 8, December 25, 1942),
appeared too late for the information to be included in figure 4. The name being
preoccupied by T. linguiferus costaricensis Olsson, 1922, Dr. Olsson has graciously assigned
me the opportunity of renaming the homonym. It gives me pleasure to suggest that this
Pliocene species be known as Typhis (Talityphis) olssoni Keen, new name. The type
locality is the mouth of Quebrada Penitas, Burica Peninsula, Costa Rica, in beds of
probable Pliocene age; holotype, Pal. Research Inst., no. 4064. The species differs from
lampada in its more slender outline and its smaller aperture.

KEEN—MOoLLUSKS FROM RouUND Mountain SILT 55

sculpture; V. oxytata dodona (Gardner) of the Chipola Miocene, Fiorida
(U.S. Geol. Survey Prof. Paper 142F, 1937, p. 267, pl. 37, fig. 25) is,

likewise, larger than gluma.

The name gluma is a Latin noun, feminine gender, meaning “husk.”

EXPLANATION OF PLATES
PLATE 3
All specimens except those illustrated in figs. 13, 17, 18, 20, 21, 22, 24
and 25 are deposited in Stanford University Paleontological Type Collection.

Bigs. Transennella joaquinensis Anderson and Martin. Hypotype no.
7528; left valve. LSJU loc. 2121, Round Mountain Silt
(Miocene), Kern Co., California. * 4.3. P. 41.

Fig..2. Transennella joaquinensis Anderson and Martin. Hypotype no.
7o2e-asught valve: IESJU) loc. 2121; 4.1. P: 41.

Fig. 3. Dosinia (Dosinidia) margaritana Weidey. Hypotype no. 7525-a;
interior of right valve of juvenile specimen. LSJU loc. 2121.
7A: Pi 40:

Fig. 4. Dosinia (Dosinidia) margaritana Wiedey. Hypotype no. 7525-b;
exterior of left valve of juvenile specimen. LSJU loc. 2121.
7.0: P40:

Big-2: Dosinia (Dosinidia) margaritana Wiedey. Hypotype no. 7525;
interior of left valve of juvenile specimen. LSJU lcc. 2121.
x 6.9. P. 40.

Fig. 6. Bornia (Temblornia) triangulata (Anderson and Martin). Hypo-
type no. 7523; interior of left valve. LSJU loc. 2121. * 4.8.
P. 39.

Fig. 7. Bornia (Temblornia) triangulata (Anderson and Martin). Hypo-

type no. 7523-a; interior of defective right valve. LSJU loc.
JIE <3 SP? 39:

Fig. 8. Donax, n. sp. Hypotype no. 7524; interior of defective right valve.
ES|U loc 2121: 5.1. P. 39.

Fig. 9. Nucula (Ennucula) birchi Keen, n. sp. Paratype no. 7527-a; in-
terior of right valve. LSJU loc. 2121. X 4.3. P. 41.

Fig. 10. Nucula (Ennucula) birchi Keen, n. sp. Paratype no. 7527-b; dor-
saloviewaleo|Wiloe: 2121. xX 42. P41.

Big. 1 Nucula (Ennucula) birchi Keen, n. sp. Exterior of right valve of

same specimen as in fig. 9. 3.1. P. 41.

Fig. 12. Nucula (Ennucula) birchi Keen, n. sp. Holotype no. 7527; in-
terior of left valve. LSJU loc. 2121. X 4.3. P. 41.

56 San DigGo Society oF NATURAL History

PLATE 3 (CoNnTINUED)

Figs. 13,18, Typhis (Talityphis) alatus obesus Gabb. Hypotype, Paleontolo-
Ups, gical Research Inst., Ithaca, New York. Miocene, Gatun for-
mation, Toro Cay, Panama. Fig. 13, view from left side; fig.
18, apertural view; fig. 22, view of spire, showing arrange-

ment of varices and tubes. * 1.6. P. 54.

Figs. 14,19, Typhis (Talityphis) lampada Keen, n. sp. Holotype no. 7548.
LSJU loc. 2121. Fig. 14, from left side; fig. 19, from front;
fig. 23, from above. X 1.6. P. 53.

Figs. 15,16. Lucinisca menuda Keen, n. sp. Holotype no. 7526. LSJU loc.
2121. Fig. 15, interior of left valve, X 3.0; fig. 16, exterior
of same valve, X 3.5. P. 40.

Bigs l7. Typhis (Talityphis) latipennis Dall. Hypotype no. 7951
(C.A.S.) from Calif. Acad. Sci. loc. 27585, lat. 23° 02’
north, long. 109° 32’ west, dredged August, 1932, in 25
fathoms a few miles off shore at Gorda Point, San José del
Cabo Bay, about 7 miles northeast of Palmilla Point, Lower
California. Recent. X 1.1. P. 53.

Fig. 18. See fig. 13.
Figal 9: See fig. 14.
Fig. 20. Typhis (Talityphis) expansus Sowerby. Copy of original figure

(Proc. Zool. Soc. London for 1873, 1874, p. 719, pl. 59,
fig. 4); holotype probably in British Museum (Nat. Hist.).
Type locality unknown, probably West Indies. Dimensions
of holotype not recorded by Sowerby; magnification of this
figure probably X 1.5 to 2. P. 53.

Fig. 21. Typhis (Talityphis) latipennis Dall. Hypotype no. 7950
(C.A.S.) from Calif. Acad. Sci. Sta. 136-D-14 (Eastern
Pacific Zaca Expedition), dredged in 45 fathoms on Arena
Bank, lat. 23° 29’ 30” north, long. 109° 25’ west, off Lower
California. Recent. Specimen viewed from back. X 1.1. P.

Dae
Big: 22. See fig. 13.
Bign23: See fig. 14.
Fig. 24. Same specimen as fig. 21, viewed from above, showing arrange-

ment of varices and tubes.

Fig. 2: Same specimen as fig. 21, apertural view.

KEEN—MOoLLusks FROM ROUND Mountain SILT PLATE 3

KEEN—MOoLLUSKS FROM ROUND MOounraIN SILT 57)

PEAT E4.

All specimens illustrated on this Plate are deposited in the Stanford Uni-
versity Paleontological Type Collection.

Fig.l:

Fig. 2.
Fig. 3.

Bross
Fig. Ds

Bigel2:

Anachis watsonae Keen, n. sp. Holotype no. 7530. LSJU loc.
2121, Round Mountain silt (Miocene), Kern County, Cal-
ifornia. X 2.7. P. 42.

Back view of specimen shown in fig. 1.

Olivella ischnon Keen, n. sp. Holotype no. 7542. LSJU loc. 2121.
IIT EM PES) 0)

Back view of specimen shown in fig. 3.

Balcis conchita Keen, n. sp. Holotype no. 7538. LSJU loc. 2121.
Xx 4.2. P. 43.

Eulima gabbiana (Anderson and Martin). Hypotype no. 7543.
LSJU loc. 2641, “Barker’s Ranch”, Kern Co., California.
Round Mountain silt (possibly uppermost Olcese sand),
Miocene. X 3.3. P. 45.

Teinostoma (Teinostoma?) lens Keen, n. sp. Holotype no. 7545.
LSJU loc. 2121. Fig. 7, view from above; fig. 8, apertural
view; fig. 9, basal view. X 7.1. P. 51.

Volvulella gluma Keen, n. sp. Holotype no. 7550. LSJU loc.
26412 X 7.35-2.4.

Hastula gnomon Keen, n. sp. Holotype no. 7536. LSJU loc. 2121.
IO26: P47.

Mitrella (Mitrella) anchuela Keen, n. sp. Holotype no. 7539.
ES) loa 2121) XX 3:82 B. 48:

Figs. 13, 14. Cylichna temblorensis Keen, n. sp. Holotype no. 7533. LSJU loc.

Fig. 15.
Fig. 16.
Big. 17.

Fig. 18.
Fig. 19.

Fig. 20.

2121. Fig. 13, view from above; fig. 14, apertural view.
X 7.0. P. 44.

Moniliopsis electilis Keen, n. sp. Holotype no. 7540. LSJU loc.
212 X 255 P. 49:

Cylichna? loismartinae Keen, n. sp. Holotype no. 7532. LSJU
loc: 2121. X 7.3. -P: 44.

Odostomia (Evalea) andersoni Bartsch. Hypotype no. 7541-a.
LSJU loc. 2641. X 67. P. 49.

Back view of specimen shown in fig. 16.

Turbonilla (Pyrgolampros) mariposa Keen, n. sp. Holotype no.
7547. USI loc 2121. X70. P. 52.

Turbonilla (Pyrgiscus) bravoensis Keen, n. sp. Paratype no.
7546-b; nucleus and early whorls of spire. LSJU loc. 2121.
M14 DP

San Digeco Society oF NATURAL History
PLATE 4 (ConTINUED)

Mangelia (Cacodaphnella?) kernensis (Anderson and Martin).
Hypotype no. 7537; showing nuclear sculpture. LSJU loc.
2A2L AS -19:0, Po 47:

Acteon Honk ee Hypotype no. 7529. LSJU loc.
2641 X21 ps Pe:

Odostomia one andersoni Bartsch. Hypotype no. 7541.
LSJU loc. 2641. X 6.6. P. 49.

Syrnola scandix Keen, n. sp. Paratype no. 7544-a; showing early
whorls of spire. LSJU loc. 2121. X 6.9. P. 50.

Turbonilla (Pyrgolampros) mariposa Keen, n. sp. Paratype no.
7547-a; nucleus and early whorls. LSJU loc. 2121. * 14.0.
P52?

Turbonilla (Pyrgiscus) bravoensis Keen, n. sp. Holotype no.
DAO seo doe 212, oe ole

Turbonilla (Pyrgiscus) bravoensis Keen, n. sp. Paratype no.
7546-a; showing aperture. LSJU loc. 2121. X 7.2. P. 51.

Chrysallida rotundomontana Keen, n. sp. Holotype no. 7531.
ES|U toes 21215-X6.9:2P5.43,

Syrnola scandix Keen, n. sp. Holotype no. 7544. LSJU loc. 2121.
X70 PRO:

Syrnola scandix Keen, n. sp. Paratype no. 7544-b; showing nu-
clear whorls and early whorls of spire. LSJU loc. 2121.
X15.0. P. 50.

Ferminoscala durhami Keen, n. sp. Holotype no. 7534. LSJU loc.
2121. X 4.0. P. 46.

Ferminoscala whitei Keen, n. sp. Holotype no. 7535. LSJU loc.
PIO 3 Pa46:

Ferminoscala whitei Keen, n. sp. Enlargement of sculpture on
back of basal whorls; same specimen as fig. 32.

KEEN—MOLLuSKS FROM ROUND Mountain SILT

a
pant

TRANSACTIONS

O}F Wal

SAN DIEGO SOCIETY OF NATURAL HISTORY

Vo.uME X, No. 3, pp. 61-68, fig. 1

oi Comps
Ry) ra
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SgAN 251944

Ligra®*

GROWTH IN THE
WESTERN BLUE-TAILED SKINK

Tuomas L. RopGers AND Vi0oLA H. MEMMLER
Museum of Vertebrate Zoology, Berkeley, California

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY

DECEMBER 30, 1943

24.4918 “comp

* yan 25 1944

COMMITTEE ON PUBLICATION
U. S. Grant, IV, Chairman

L. M. KLauBer CLINTON G. AppotTt, Editor

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GROWTH IN THE
WESTERN BLUE-TAILED SKINK
BY

Tuomas L. RopGers and VioLta H. MEMMLER
Museum of Vertebrate Zoology, Berkeley, California

When sufficient material is available, studies of the distribution of
size groups yield data on growth of lizards that is nearly as reliable as
repeated capture and measurement of marked individuals in nature. In
the region where this study was conducted, and probably in any part of
the range of the Western Blue-tailed Skink (Eumeces skiltonianus), it
would be highly impractical to capture and mark a sufficiently large
number of these lizards to insure recapture of them on a scale that would
afford information about their growth. For this reason it was decided to
collect skinks from a restricted area throughout an entire year, with the
expectation that age groups could be detected in measurements of size
and traced from month to month, yielding information on rate of
growth. Such an undertaking required regular trips to the area and
many man hours of collecting (139). The senior author was present on
24 of the 36 collecting trips, but a total of 21 persons contributed to
the collecting of the 409 specimens used in the study. Prominent among
these were Walter W. Dalquest, whose deep interest in field activity
supplied the initiative that started the work and the drive that netted
145 of the specimens, and Sherburne F. Cook, Jr., who collected 52 of
the specimens and made special effort to get a much needed series of
specimens in the month of February. All the specimens were taken
within two miles of Bald Peak in the Berkeley Hills of extreme western
Contra Costa County, California, an area sufficiently restricted to
reduce to a negligible degree geographic variation represented in our
material.

In late winter and spring, during wet weather, skinks are found
with little difficulty under rocks. In summer, they are more active on
the surface and have practically unlimited cover in the form of grass
and cracked earth in which to hide, which makes them difficult to
capture at this time. In November, after the first rains, they are found
fairly commonly under rocks, but in the colder months of December
and January they are found less commonly in such places, probably
because they are deeper under the surface.

64 San Digco Socrety oF NATURAL History

The size measurement used was snout-vent length, and, of course,
it was first necessary to find whether or not there is a sexual difference
in this character. Specimens collected in April and May which did not
fall in the smallest age group were used for this test. Twenty-five
females collected in April averaged 63.3 + 0.98 mm. long, and 28 males
averaged 62.9 = 0.75 mm. The difference between the means is .4, and
the standard error of the difference is 1.23. Twenty females collected
in May averaged 64.9 + 1.0 mm. long, and 17 males averaged 65.5
+ 1.2 mm. The difference here is .6 and the standard error of the
difference is 1.56. Since the differences are small, are reversed in the
two sets of data, and are exceeded by the standard errors of the
differences, it can be safely assumed that there is no significant sexual
difference in this character. This circumstance makes it possible to
combine males and females for study of age groups.

Three clutches of eggs were laid in captivity on July 6, July 11,
and July 15. Another clutch was found in the field on July 21. Two
broods of young were represented among the August specimens. One
of these hatched in captivity on August 16 from eggs collected in the
field on July 15. The other brood was collected in the field on August
9, and may have been as much as one or two weeks old at the time.
It therefore seems safe to assume that young normally hatch in July

and August.

The specimens measured at hatching ranged from 24.7 to 26.3 mm.
Few specimens were collected in the next few months. The smallest
specimen taken in October was 37 mm. long. A group of small ones
collected in November ranged from 37 to 43 mm. long. The smallest
in December was 42 to 49 mm. In January, the smallest group ranged
from 41 to 49 mm. and in February it ranged from 38 to 49 mm. The
growth indicated by these specimens must be that of the youngest age
group, that is, those animals hatched in the preceding July and August.
Also, rapid growth is indicated in August, September and October, the
warm months of the fall season, and little growth in December, January,
and February, the coldest months in this region.

The specimens represented by the data for January and February
were collected at the end of the period of study, in 1943, while those
for March were collected at the beginning of the study, in 1942. It can
be noted (figure 1) that the smallest groups in March average smaller
than those in February. The young skinks hatched in the fall of 1942,

Ropcers & MEMMLER—GROWTH IN THE SKINK 65

must have experienced better growing conditions during their first six
months of life than those hatched in the fall of 1941. This period of
better growing conditions also appears to be reflected in the 20-month-
old skinks (see break in growth curve at this point in figure 1).

In March there seems to be no increase in size of the smallest
groups collected throughout the month; there is probably little growth

oecenoe’ 80

Alnr 30 anne

Figure 1. Circular scatter diagram showing the change in distribution and
grouping of data along the scale of snout-vent length (on the radii) at different
times of year, and thus the growth trend of the species. Each sector of the
circle represents one month. The light concentric circles mark distances, each
representing ten millimeters of snout-vent length, along the radii. Each dot
represents one individual. The data for a series of specimens collected on a
given day are plotted along the radius representing that date of collection. The
spiral lines represent growth trends, and the heavy outer circle represents the
approximate average snout-vent length for all specimens over two years old.

66 SAN Deco Society oF NATURAL History

in this month. In April, the specimens of the smallest groups average
about 41 mm. in length, in May about 45 mm., and in June about 50
mm. This indicates rapid growth in these warm spring months. It also
leads to the conclusion that one-year-old individuals average about 50
mm. long, a gain of about 25 mm. since hatching.

The measurements of the larger specimens (those over 55 mm.
long) collected on a given day comprise a group with a wide range
that usually may be subdivided into two groups. This is best seen in
the large mass of material taken in February, March, April, and May.
The smallest of these two groups averages about 62 mm. in February,
and must represent the 20-month-old individuals. The later history of
this group indicates that by the time an individual is two years old it has
reached approximately 65 mm., a gain of about 15 mm.; this age group
can not, from then on, be distinguished from the group representing
all older individuals.

The smallest breeding individuals are 61 and 62 mm. long, but
most of the breeding animals are 68 to 75 mm. long. This would suggest
that some individuals might breed at the time they are two years old,
but that usually they start breeding the third summer after they are
hatched, when they are three years old.

All specimens over two years of age are represented in the remaining
group that centers around the 68 mm. line in the graph. The number
of specimens represented in this group is roughly equal to those in the —
group that is approaching two years of age. It can be assumed from
this that the group on the graph represents an accumulation of animals
of at least three year classes. Since the accumulation in this group is
not great, it seems probable that the normal life span for individuals
once having attained breeding condition is five or six years.

The normal growth rate is thus determined to be 25 mm. the first
year, 15 mm. the second year, probably not more than 3 or 4 mm. the
third year. If growth continues after the third year, it probably does
not exceed 2 mm. the fourth or 1 mm. after that. In light of the great
amount of individual variation shown to exist in the well defined groups
of 8- to 1l-month-old individuals, it would be illogical to carry this
line of argument to the extreme of assuming that the 75 mm. individuals
are 10 years old or that the 82 and 84 mm. individuals, taken in other
parts of Contra Costa County, are 20 or more years old. These large
specimens could be individuals that by rare chance were permitted to

Ropcers & MEMMLER—GROWTH IN THE SKINK 67

live to be several years older than normal, but they are more probably
lizards that experienced unusually good growing conditions or that
inherited the ability to grow rapidly. Some specimens that are 70 to 72
mm. long appear to be older, by virtue of their heavy scarred heads,
than some of the largest ones.

In the light of our findings it seems incredible that Taylor (A
taxonomic study of the... genus Eumeces ... , Bull. Univ. Kansas,
23:67, 1935), could have detected 16 age groups, 5 of which fall
between our first two, by using measurements of the snout-vent length
of the specimens of skiltonianus available to him at the time of his study.
Taylor does not state how many specimens he used, how great an area
was represented in the “locality” from which they came, or how many
years were involved. However, his list of material examined indicates
that he did not have more than 50 specimens of skiltonianus taken in
May (of many years) from any combination of four adjoining counties.

Summary.—Y oung of Eumeces skiltonianus are hatched in the months ot
July and August. They average about 25 mm. long (snout-vent length) then
and grow to about 50 mm. by the time they are one year old. Most of their
growth takes place in the first three months of life and in the following April,
May, and June. These skinks grow to about 65 mm. by the time they are two
years old, and to about 68 mm. when they are three years old. Some may breed
when they are two years of age, but most of them breed at the end of their third
year. The normal life span for individuals once having attained breeding
condition is probably five or six years. The oldest individuals are probably not
more than nine years old. The results of this study make it impossible for the
authors to accept Taylor’s 16 age groups for skinks of this species, four of which
fall between our first two.

Lm 4

TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vo . X, No. 4, pp. 69-70 DECEMBER 30, 1943

A NEW SNAKE OF THE GENUS SONORA oo *! Comnp

j : Ps >.
on notogy 80D

FROM LOWER CALIFORNIA, MEXICO (“yay 0,"
AG IEK BY Nara 7
LAURENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

Some time ago I received from Mrs. Grifing Bancroft a Sonora
of the semiannulata group, which, as far as I know, is the first specimen
of this group to have been taken on the Pacific side of the mountains
on the peninsula of Lower California. Since this snake differs from the
known members of the genus, I take pleasure in naming it after Mrs.
Bancroft, a friend of more than forty years, who, with her husband, has
been the source of many fine reptile specimens collected in the course
of their oological and archaeological expeditions to Lower California,
Sonora, and the islands of the Gulf of California.

Sonora bancroftae sp. nov.

Type.—No. 35,077 in the collection of LMK, collected two miles east of
San Jorge,! Lower California, Mexico, by Mrs. Griffing Bancroft, April 10,
1942.

Diagnosis —A Sonora of the rounded-snout group, differing from mosauert
of the middle of the peninsula in having cross bands, whereas mosaueri is
unicolor; also, bancroftae has more ventrals and subcaudals. From semiannulata
semiannulata it differs in the color of both the background and bands.

Description of the Type—A female; length over-all 194 mm.; tail length
36 mm. The scale rows are 15-15-14; the dorsal scales are smooth, and with
single apical pits. The ventrals number 171 and the subcaudals 47; the latter
are divided, as is also the anal. The supralabials are 7-7, infralabials 8-8, loreals
probably? 1-1, preoculars probably 1-1, postoculars 2-2, temporals 1+2, 1+2.

1San Jorge is in the valley of the San Telmo River about twenty miles above its
mouth. It will be found on the American Geographical Society map of Baja California—
Norte, Provisional Edition, 1928, a few miles northeast of the intersection of Lat. 31° N.
and Long. 116° W.

2The type specimen died from an injury incident to its capture a day or so before it
reached me, in consequence of which the head is rather shrunken anteriorly.

70 SAN Dieco Society oF NaturAL History

The scale rows were counted at the points recommended by Stickel in his
summary of the genus.?

The pattern comprises a series of evenly-edged gray cross bands (34 on
the body, 8 on the tail) on a light-brown ground. The ground color (as
preserved in alcohol) is Orange-Cinnamon.* The cross bands are Deep Neutral
Gray. Below the color is Pinkish Buff. The dark bands are usually wider than
the interspaces; they are from 242 to 342 scales (end to end) wide, while the
spaces between are 2 to 3. The gray color comprises many fine punctations,
the ground color showing through to some extent. The gray bands fade out as
they reach the lowest lateral scale rows; however, they do not narrow laterally
as is usual in S. s. semiannulata, nor are the interspaces clouded with dark spots
laterally as is often the case in the latter subspecies. All dorsal scales are
somewhat lighter on the edges than in the centers. The ventrum is immaculate

buff.

The top of the head is dark. There is a narrow light crescent which engages
the eyes; behind this on the neck is the first dark ring, which likewise is
crescent-shaped.

Remarks —The species is known only from the type. It was captured when
a rock was overturned in seeking a missile to throw at a red diamond rattler.
There was a second specimen under the stone, but it was not collected.

This species is most closely related to S. s. semiannulata, with which it may
eventually be shown to intergrade, although, as far as I know, no semiannulata
has yet been collected in Lower California. The differences between bancroftae
and semiannulata are primarily in pattern. The yellow-brown ground color in
bancroftae is not like the cream or red suffusions between the dark rings usually
observed in live semiannulata, but supplants the true white ground color of the
latter. The gray rings in bancroftae are relatively wider and closer together than
the black rings of semiannulata and do not taper laterally as is the case in the
Arizona snakes.

3William H. Stickel: The Snakes of the Genus Sonora in the United States and
Lower California. Copeia, No. 4, pp. 182-190, 1938.

4Capitalized colors are those of Ridgway: Color Standards and Color Nomenclature,
1912.

ow eooay
LS, $4% TRANSACTIONS re a na)
SAN DIEGO SOCIETY OF NATURAL HISTORY
Vot. X, No. 5, pp. 71-74 DECEMBER 30, 1943

A DESERT SUBSPECIES OF THE SNAKE
TANTILLA EISENI
BY

LauRENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

The California Black-headed Snake, Tantilla eiseni Stejneger, 1896,
is found from southeastern Alameda County and Fresno south to San
Quintin, northern Lower California. Of this species I have accumulated
a moderately adequate series from San Diego County. Among these
are six from the desert side of the mountains, which are sufhciently
different from those on the coastal slope to warrant segregation as a
separate subspecies. I therefore describe this as

Tantilla eiseni transmontana subsp. nov.
DESERT BLACK-HEADED SNAKE

Type—No. 29,273 in the collection of LMK, collected on the road one
mile east of Yaqui Well, San Diego County, California, by Charles E. Shaw
and Cyrus S. Perkins, June 6, 1938, at 8:10 P. M. Five paratypes are available
from San Diego County and one from Riverside County.

Diagnosis —A desert slope subspecies of Tantilla eiseni characterized by
a higher number of ventral scutes, a shorter tail, and lighter color than the
cismontane form. T. e. transmontana has a higher ventral scale count than any
other known Tantilla.

Description of the Type-—Adult male. Length over-all (before shrinkage
in preservative) 302 mm.; tail length 70 mm. The scale rows are 1515-1);
all scales are smooth. The ventrals are 182; anal divided; there are 68
subcaudals, all divided. The plates on top of the head are normal. The nasals
are divided; there are no loreals. The supralabials are 7-7, and the infralabials
6-6. The preoculars are 1-1; postoculars 2-2. The temporals are 111, 1+1.

The following color description refers to the specimen as preserved in
alcohol: The top of the head is purplish-black, although the rostral and
internasals are lighter. On the sides the dark color touches only the tops of
the supralabials, except posteriorly where an extension of the dark area is
carried down across the last supralabial to a point behind and below the angle

72 SAN Deco Society oF NATURAL History

of the mouth. The dark color on top of the head reaches a distance two scales
behind the parietals. Here there is a transverse band very slightly lighter than
the ground color, no doubt the vestige of the usual Tantilla ring. On the under
surface of the head there are some faint punctations on the edges of the
infralabials; the third and fourth infralabials are heavily punctated where they
abut the genials. The body is cream-colored throughout, both dorsally and
ventrally. Fine brown punctations are faintly in evidence on the seven mid-
dorsal scale rows. A dark streak of the vertebral process shows through the
skin; there is a rather marked mid-dorsal groove.

Summary of Paratypes—Six paratypes are available, the data on them
being as follows:
Sub- Supra- — Infra-
Sex Ventrals caudals labials —_labials

LMK 2633 Yaqui Well M 175 62 6-7 TI,
LMK 2634 Yaqui Well M 184 65 Vaal 6-6
LMK 32419 Sentenac Canyon F 197 63 Ty 6-6
LMK 33760 Palm Springs F 195 66 HI) 6-6
LMK 33997 Sentenac Canyon F ~- — TY Tach
SDSNH 11260 La Puerta F 190 60 Tah WY

All localities are in transmontane San Diego County except Palm Springs,
which is in Riverside County. All specimens have 15 scale rows, one preocular,
two postoculars, and 1+1 temporals. None has loreals.

In these paratypes there is some variation in the amount and extent of
dorsal pigment. The pigmented dorsal scale rows vary from 7 to 13, the
punctations being most in evidence mid-dorsally. The punctations are light and
scattered; they are darkest in the Palm Springs specimen, which is the only one
that approaches, in depth of color, the lightest of the coastal snakes. The black
of the heads continues from two to three scale lengths posterior to the parietals,
and is carried back of and below the angle of the mouth, as Blanchard discovered
is characteristic of eiseni and differentiates this form from utahensis (Blanchard,
1938). The light neck ring is more or less in evidence, depending on the extent
of the dorsal punctations. Only in the Palm Springs specimen are there black
dots posterior to the light ring.

Range——No definite localities of this subspecies are known except those
where the type and paratypes were collected. Eventually it will no doubt be
found along the desert base of the mountains from San Gorgonio Pass in
Riverside County southeastward well into Lower California. While transmontana
may occur along the eastern base of the San Bernardino Mountains and in the
Little San Bernardinos, it has not been collected in either. Further north we
may expect to find utahensis; at least such is the case in the Panamints and in
the Kingston Range. Without doubt eiseni and transmontana intergrade through
San Gorgonio Pass in Riverside County and via some of the San Diego County
passes. Three specimens from Snow Creek, on the north slope of Mt. San
Jacinto, Riverside County, are considered intergrades, as they show characters of
both subspecies.

Remarks.—Transmontana has more ventral scutes and is a lighter colored
snake than eiseni. The change in color is produced both by there being fewer

KLAUBER—NeEw DeEsERT SNAKE 73

punctated dorsal scale rows and by the punctations of transmontana being
lighter and more widely spaced. This parallels the change which takes place
in the modification of Leptotyphlops humilis humilis into L. h. cahuilae in
passing from the coast to the desert; also in the worm snakes there is an
increase in the longitudinal dorsal scales corresponding to the increase of the
ventrals in Tantilla.

The following are comparative data on the ventrals of the two subspecies:

MALES FEMALES
Eiseni Transmontana — Eiseni Transmontana
Number of specimens 22 3 18 3
Mean 169.4 180.3 176.3 194.0
Interquartile range 1674-1714) V77AS1835 1173.9=178:8) 19 1/6-196:4:
Extreme range 164-175 175-184 169-182 190-197

Notwithstanding the few specimens of transmontana available, the differ-
ence between the means is found to be highly significant in both sexes.

With respect to the coloration it may be said that the lightest of the
coastal specimens is darker than the darkest of those from the desert, although
this is not true of the specimen from Palm Springs, which is an intergrade as
far as pattern is concerned. In eiseni eiseni all 15 rows are usually punctated,
although the side rows are lighter than the mid-dorsals. Occasionally the spots
are carried onto the edges of the ventrals. Transmontana, besides being lighter,
usually has at least two unmarked lateral rows on each side. The heads of the
coast specimens are also darker, and the underjaws are usually more heavily
punctated, although this is not always the case. The light collar is generally
more clearly outlined in the coastal specimens, but the difference is not infallible.
The general color divergences are apparent in the live specimens as well as the
preserved.

With regard to the tail length, using the methods which I have detailed
elsewhere (Klauber, 1943), although handicapped by the rather inadequate
transmontana material, I find the differences to be significant. Comparing snakes
converted to a standard over-all, length of 300 mm., the following are the mean
tail lengths in mm.:

Males Females

Eiseni 74 66
Transmontana 69 59
Coefficient of divergence, % Fig 12

I have little doubt that differences comparable to these will be authenticated
when additional desert specimens become available.

Three specimens from Snow Creek, on the north slope of Mt. San Jacinto,
are available. In color they are light, like transmontana, and their ventral scutes
fall within the transmontana range, the counts being 179, 180, and 183 (all
males). However, in tail-length proportionality they resemble the coastal sub-
species; I therefore consider them intergrades.

I wish to acknowledge the assistance of the following individuals

and museums in permitting the study of specimens in their collections:

Mr. J. R. Slevin, California Academy of Sciences; Mr. Thomas Rodgers,

74 SAN DrgeGo Society oF NatTurRAL History

Museum of Vertebrate Zoology, University of California; Mr. Charles
M. Bogert, American Museum of Natural History; Dr. Howard R.
Hill, Los Angeles Museum; Mr. Charles H. Lowe, Jr.; and Mr. Harold
T. Woodall.

BIBLIOGRAPHY

BLANCHARD, F. N.
1938. Snakes of the Genus Tantilla in the United States. Zool. Ser. Field
Mus. Nat. Hist., Vol. 20, No. 28, pp. 369-376.
KLAUBER, L. M
1943. Tail Length Differences in Snakes, with Notes on Sexual
Dimorphism and the Coefficient of Divergence. Bull. Zool. Soc.
San Diego, No. 18, pp. 1-64.
Smit, H. M.
1942. A Résumé of Mexican Snakes of the Genus Tantilla. Zoologica,
Vol. 27, No. 7, pp. 33-42.
STEJNEGER, L.

1896. Description of a New Species of Snake (Tantilla eiseni) from
California. Proc. U. S. Nat. Mus., Vol. 18, No. 1044, pp. 117-118.

TRANSACTIONS

OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY
Vot. X, No. 6, pp. 75-82 DECEMBER 30, 1943
A le
THE CORAL KING SNAKES ere
OF THE PACIFIC COAST ate ely

‘Re ¢ 34% BY LL

LAURENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

I have lately completed an examination of a number of Coral, or Mountain,
King Snakes from Washington, Oregon, California, and Lower California. I
find in California two pattern types which I believe should be recognized as
subspecies; one of these is characteristic of the Coast Range, the other of the
Sierra Nevada. Where these two mountain ranges merge, both to the north
(in the Siskiyous and Cascades) and southward (in the San Gabriel, San
Bernardino, and San Jacinto ranges), there are mixtures of the two patterns.
The snakes of the extreme southern extension of the range, into San Diego
County and Lower California, adhere rather closely to the coastal pattern.

The primary difference between the two patterns lies in the nature of the
triads comprising the dorsal rings; in the Coast Range form the two black
boundary rings of each triad are generally well separated so that the central
red ring crosses the dorsum, while in the Sierra pattern the two black boundary
rings usually coalesce dorsally, thus restricting the red marks to paired lateral
blotches, or the red may even be completely suppressed. It is true of few snakes
in either area that every triad follows one form or the other, but the criterion
of whether more or less than half of the black boundaries of the triads merge
dorsally, adheres substantially to a consistent geographical arrangement.

In Lower California two other snakes belonging to this group have been
described, both of which have been rather neglected in recent faunal and check
lists. One of these is a mountain, the other an island form. I am of the opinion
that both should be recognized as subspecies.

It should be understood that any black ring, or pair of black rings,
completely or partially split with red, is considered a triad, provided it is
bounded on either side by a white ring. The beautiful red coloration of these
snakes fades badly in preservative, so it is sometimes difficult to determine
which rings in life were white, and which red. In triads wherein the red rings
completely split the black, it will be found that the black borders are usually
more irregular or serrated on the side where they abut the red than where they
contact the white. This will often aid in counting triads in old specimens
wherein the red has faded to cream or buff. And it must be remembered that
many snakes have at least some triads composed of a single black ring, unmarked

76 San Disco Society oF NaturaL History

by any red, or in which the red is represented only by a light lateral spot. This
kind of complete, or almost complete, suppression of red is not the result of
preservative, but is characteristic of the snakes of some areas. Such a single
black ring is counted as a triad, because it is bordered on each side by a white
ring.

There will be some borderline triads with respect to which it will be
difficult to decide whether they should be considered split or not. In these the
dorsal black joining the two side black rings comprises irregular black lines
following scale edges. These have been judged not to break the continuity of
the red ring, unless a black line is at least one scale wide.

I should summarize the Coral King Snakes of the Pacific Coast as follows:
Lampropeltis multicincta multicincta (Yarrow), 1882.

Sierra Coral King Snake.
Range: The Sierra Nevada of California from Kern County north to
Shasta County. Intergrades with L. m. multifasciata via the Siskiyou
and Cascade ranges to Klickitat County, Washington; and in the San
Gabriel, San Bernardino, and San Jacinto mountains of southern
California.

Lampropeltis multicincta multifasciata (Bocourt), 1886.
Coast-Range Coral King Snake.
Range: The Coast Range of California from Del Norte County south
to northern Lower California, where it intergrades with L. m. agalma
in the Sierra Juarez.

Lampropeltis multicincta agalma Van Denburgh and Slevin, 1923.

San Pedro Martir Coral King Snake.
Range: The San Pedro Martir Mountains, Lower California, Mexico.

Lampropeltis multicincta herrerae Van Denburgh and Slevin, 1923.

Todos Santos Coral King Snake.
Range: South Todos Santos Island, off Ensenada, Lower California,
Mexico.

The question whether the specific name of the coral king snakes of the
Pacific Coast should be multicincta or zonata has been the subject of considerable
argument, hinging upon the uncertainty of identification of the type specimen
of Coluber (Zacholus) zonatus Blainville, 1835, and the alternative applicability
of Bellophis zonatus Lockington, 1877. I shall not review this question, which
will be found fully discussed in the papers by C. E. Burt (1936), and J. L.
Peters (1938). Personally, I have adopted the view that the identity of the
Blainville specimen is indeterminate! and that Lockington’s name cannot be
used for reasons advanced by Peters. Those who disagree may assign the name
L. zonata zonata to the coastal form, L. zonata multicincta to the Sierran, and
L. zonata agalma to that of the San Pedro Martirs.

As to the applicability of the names multicincta and multifasciata to the
Sierran and coastal forms, respectively, it may be noted that, while the type
locality of multicincta is given as Fresno, the specimen probably came from the

~ See Bull. S. D. Zool. Soc., No. 18, p. 47, 1943, for a discussion of the tail length
of Blainville’s type as a possible clue to identification.

KLAUBER—CORAL KING SNAKES 77

mountains to the east, since it is well known that in California this species is a
mountain or, rarely, a foothill dweller. The description leaves no doubt that
the type was collected in the Sierras, rather than in the Coast Range, since only
one or two of the triads are completely split by red.

Similarly, although the type locality of multifasciata is even more uncertain,
being given only as “Californie”, the fact that a majority of the triads are
completely split with red, and the wide separation of the bordering black rings
of the anterior triads, as can be seen in Bocourt’s figures, make it quite certain
that this specimen represents the coastal subspecies.

With regard to the adequacy of the differentiation between multicincta and
multifasciata, I should say that they seem to be territorially quite consistent,
except where the two mountain ranges which they inhabit conjoin. I have
examined 30 specimens from the Sierras and of these 28 have less than half the
triads split dorsally. Three have none split, while four others have only one or
two split, it being noted that the first ring on the neck, and the first anterior
to the vent, are likely to be divided even though the others are not. Of the
two which fail to follow the mode, MVZ 19324, from Generals Highway, near
Sequoia National Park, Tulare County, has 20 out of 27 split; while MVZ
24395 from Wolfboro, Calaveras County, has 18 out of 33 split, thus barely
failing to pass the test.

As to the Coast Range specimens, I have examined 19, of which 18 have
more than half the triads split; 6 have every one split, while 5 others have only
one or two in which the black rings merge. The single specimen which fails
under this rule for segregation is MVZ 18178 from Covelo, Mendocino County;
this has 13 triads divided out of 29, and thus barely fails of proper allocation.
This is one of the most northerly of the available Coast Range specimens and
may indicate the beginning of the area of intergradation, which is continued into
the Siskiyous.

The following table will serve to summarize the almost complete segregation
of the Sierra and Coast Range specimens in California with respect to the

nature of the triads:
Per cent of triads

with uninterrupted Coast Sierra
red rings + Range Nevada
Oto 99 8
10to 19.9 9
20 to; 29:9 5
30 to 39.9 4
40 to 49.9 1 2
50 to 59.9 1 1
60 to 69.9 1
70 to 79.9 2 1
80 to 89.9 3
90 to 100.0 1

TOTAL 19 30

78 San Dreco Socrety oF Naturat History

It should be understood that these data on triads have reference only to
those on the body. The tail rings are less conclusive, as they are often solid
black even in the Coast Range specimens.

Multicincta reaches its extreme differentiation from multifasciata in the
region of Yosemite National Park where specimens with the red completely
suppressed are occasionally found (Klauber, 1932). In these snakes lateral
spots, usually cream in life, rather than red, may appear on one or two of the
black rings; these represent the final vestiges of red rings.

In multifasciata the most extreme specimens are found in the Santa Cruz
Mountains. Here every triad is likely to have the red dorsally complete, and
most of the red rings are much wider than both the white and black rings taken
together, so that the snakes are predominantly red. They are very brilliant and
handsome in life. In these wide-banded snakes the total body triads are reduced
in number; however, there is considerable variation in both forms, so that
multifasciata cannot be segregated from multicincta by the number of rings.

There seem to be no important differences between the two subspecies in
scutellation, at least none that can be used as a key character. Multicincta has
a somewhat higher average ventral scale count than multifasciata and has 21
scale rows (in place of the more normal 23) less frequently than the coastal
form.

At the northern end of the range the number of specimens available is still
too few to define the ranges of the two subspecies. A single specimen from Del
Norte County is multifasciata, while one from northeastern Shasta County is
multicincta; both run true to expectation, as they are at the north ends of the
Coast Range and the Sierra Nevada, respectively. Between lies Siskyou County,
from which there are three specimens at hand; of these, one resembles the
coastal, and two the Sierran form. This will probably prove an area of inter-
gradation.

I have seen only four specimens from Oregon and Washington, where the
species has lately come to light as reported by Gordon (1935), Fitch (1936),
Johnson (1939), and Lewis (1942). Three of these are multicincta, although
one is from Curry County, Oregon, not far from the coast (LMK 32309, 3 mi.
north of Illahe). The other Oregon specimen is from Jackson County, and
is also multicincta. Of the two Washington specimens, both from Klickitat
County, one falls in each subspecies, as determined by the triad criterion.
Additional specimens must be had before we can decide whether these extreme
northern snakes are predominately of the coastal or Sierran form.

At the south ends of the Coast Range and the Sierra Nevada, where they
merge into the Santa Monica, San Gabriel, San Bernardino, and San Jacinto
ranges, the snakes are also mixed, yet in two of the ranges there are indications
of a predominance of one subspecies or the other, as shown in the following

table:

Range Multifasciata Multicincta
Santa Monica 8 2
San Gabriel 9 16
San Bernardino 1 11
San Jacinto 53 14

KLAUBER—CoRAL KING SNAKES 79

Specimens from foothill points have been allocated to the nearest mountain
range. It will be seen that there is a predominance of the coastal multifasciata
in the range nearest the coast (Santa Monica), and of multicincta in the ranges
farthest inland (San Bernardino and San Jacinto). A good many of these
southern mountain snakes fall in the 40 to 60 per cent range of split triads.

Still farther south, in San Diego County, 5 out of 24 specimens have less
than half the triads split so that the population is to be considered multifasciata.
Although most of the triads are divided, the red is not usually as wide as in
the snakes of the central Coast Range. A single specimen from Orange County
also belongs to this subspecies; this was found DOR on the coastal plain 2 mi.
south of Irvine, in a dry-farming area of several thousand acres given over
to the cultivation of lima beans, a most unusual habitat for this snake, which,
in arid southern California, is seldom found below an altitude of 4000 feet.

From the Sierra Juarez of northern Lower California a single specimen is
available, MVZ 10493 from Laguna Hansen. Linsdale (1932) has discussed
this snake; he concluded that because it, and several other specimens from
California, have some pink or red on the snout, agalma is invalid, for a pink
snout was the differential character upon which agalma was originally described,
a jet-black snout being the mode in the Californian snakes. It will be recalled
that pyromelana of Arizona has a white snout.

It is true that occasional specimens of multifasciata, particularly from the
Santa Cruz Mountains, have some red spots or blotches on the top of the head.
However, the coverage is not so great as in the three available specimens of
agalma from the San Pedro Martir Mountains. In the latter the prefrontals,
especially, are almost entirely pink. The supralabials of agalma are less maculate
than the other multicincta subspecies. A galma also differs from the Coast Range
form in having a higher number of triads on the body and relatively more black
than red per triad, as compared with typical multifasciata. I am therefore of the
opinion that agalma should be recognized as a valid subspecies, the Laguna
Hansen specimen being considered a multifasciata-agalma intergrade.

Herrerae of South Todos Santos Island is :aperficially much like the
occasional black and white specimens of multicincta from Yosemite National
Park; for in herrerae the pink, if present at all, is represented only by a few
anterior lateral buff spots. Were it not for the queer Yosemite pattern phase,
the Todos Santos snakes would be sharply differentiated from all mainland
specimens; as it is, we must search for some other difference besides the absence
of red. Probably the best key character wherewith to segregate them is the
condition of the last supralabial, which in herrerae is touched with black
posteriorly, but is clear in the few bicolor specimens from Yosemite. Herrerae
is also peculiar in that in five out of the nine available specimens the supraoculars
are fused to the parietals, a condition which I have not observed in any
multicincta. This is an important deviation from the king snake mode, even
though it is not consistent enough to be used as a key character. Occasionally
an upper postocular is fused to a parietal.

Of course, it is to be remembered that one or more split triads are found
in almost all mainland snakes; therefore the segregation of herrerae from any of

80 SAN Disco Society oF NATURAL History

the other subspecies of multicincta is usually quite simple. It is surprising to find
this relative of a species which is characteristic of the Transition Life Zone, on
this small coastal island, with a maximum altitude of 313 feet and only a mile
and a quarter long, where it has been able to maintain itself during the long
period required to differentiate so markedly from the mainland form. It is
relatively large for a coral king snake; all 9 originally collected by Mr. J. R.
Slevin are adults; they range in length from 755 to 862 mm. Being an island
form I should have considered it a full species were it not for the occasional
Yosemite phenotypes and the obvious close relationship with the mainland
form.

The brilliance of the red coloration in these Californian King Snakes is
remarkable. For purposes of comparison it may be noted that a freshly-shed
individual from the mountains of San Diego County had Scarlet rings anteriorly,
and Brazil Red toward the tail. The capitalized colors refer to Ridgway’s
Standards, 1912.

KEY TO THE CorRAL KING SNAKES OF THE PACIFIC COAST

A. — More than half of the triad rings of the body include

transverse red” bands which cross the dorsum.

B. Top of snout back to the frontal black; or if
predominantly red or pink, total body triads

lessthat 40. fer Ween L. multicincta multifasciata

BB. Top of snout back to the frontal predominantly
red or pink, and with body triads exceeding
Bio hiroka: 2) 4 ei Meee eee SN aE ee en L. multicincta agalma

AA. More than half of the triad rings of the body entirely

black, or with lateral red* areas which are not con-

fluent dorsally.

C. Usually some red areas laterally or dorsally;
last supralabial untouched by black posteriorly.
ee Be EN an CON OLED Wea A L. multicincta multicincta

CC. No red or pink laterally or dorsally, except a
few lateral spots; last supralabial touched with
black posteriorly; supraoculars often fused to
Patte cals. Chae ice ure eee L. multicincta herrerae

2May be yellow, cream or buff in preserved specimens, but must not be confused with
the really white rings between triads. A black ring even though unsplit by red, is considered
a triad if bordered on either side by a truly white ring.

KLAUBER—CoRAL KING SNAKES 81

ACKNOWLEDGMENTS

In conclusion I wish to thank Dr. Alden H. Miller and Mr.
Thomas Rodgers of the Museum of Vertebrate Zoology, University of
California; Dr. R. B. Cowles of the University of California at Los
Angeles; Miss Margaret Storey of the Natural History Museum,
Stanford University; Dr. Howard R. Hill, Los Angeles Museum; Mr.
Joseph R. Slevin of the California Academy of Sciences; and Mr.
James R. Slater of the College of Puget Sound, for the privilege of

examining the coral king snake specimens in these collections.

BIBLIOGRAPHY

BLAINVILLE, H. D. DE
1835. Description de Quelques Espéces de Reptiles de la Californie. Nouv.
Ann. Mus. d’Hist. Nat., Vol. 4, pp. 233-296.
BLANCHARD, FRANK N.
1921. A Revision of the King Snakes: Genus Lampropeltis. U. S. Nat.
Mus., Bull. 114, pp. vi + 260.
Bocourt, F. (witH DumeriL, A. and Mocquarp, F.)
1870-1909. Mission Scientifique au Mexique. Les Reptiles, pp. 1-1012.
Burt, CHArLEs E.
1936. The Nomenclature of Western Coral King Snakes, Lampropeltis
zonata Versus L. multicincta. Copeia, No. 2, pp. 94-98.
FitcH, Henry S.
1936. Amphibians and Reptiles of the Rogue River Basin, Oregon. Amer.
Midl. Nat., Vol. 17, No. 3, pp. 634-652.
GorDON, KENNETH
1935. Boyle’s and Coral King Snakes in Oregon. Copeia, No. 1, p. 46.
1939. The Amphibia and Reptilia of Oregon. Oregon State College,
Studies in Zool., No. 1, pp. 1-82.
JoHNson, Murray L.
1939. Lampropeltis zonata (Blainville) in Washington State. Occ.
Papers Dept. of Biol., College of Puget Sound, No. 1, pp. 2-3.
KiLAuBer, L. M.
1932. A Coral King Snake of Peculiar Coloration. Yosemite Nature
Notes, Vol. 11, No. 9, p. 1.
1943. Tail-Length Differences in Snakes, with Notes on Sexual Dimor-
phism and the Coefficient of Divergence. Zool. Soc. of San Diego,
Bull. No. 18, pp. 1-64.
Lewis, THomas H.
1942. Additional Records for Washington Snakes. Copeia, No. 2, p. 129.
LINSDALE, JEAN M.
1932. Amphibians and Reptiles from Lower California. Univ. Calif.
Pubs. in Zool., Vol. 38, No. 6, pp. 345-386.

82 SAN D1EGO SOCIETY OF Natura History

LockincTon, W. N.
1876. Description of a New Genus and Species of Colubrine Snake.
Proc. ‘Gal: Acad, Set, Ser: 1, Vol..7, pp. 92-93.
PETERS, JAMES L.
1938. The Name of the Western Coral King Snake. Copeia, No. 2, p. 93.
STEJNEGER, LEONHARD
1902. The Reptiles of the Huachuca Mountains, Arizona. Proc. U. S.
Nat. Mus., Vol. 25, pp. 149-158.
VAN DENBURGH, JOHN
1922. The Reptiles of Western North America. Occ. Papers Cal. Acad.
Sci., N&. 10, 2 vols., pp. 1-1028.
VAN DENBURGH, JOHN, and SLEVIN, JOSEPH R.
1923. Preliminary Diagnoses of Four New Snakes from Lower California,
Mexico. Proc. Cal. Acad. Sci., Ser. 4, Vol. 13, No. 1, pp. 1-2.
Yarrow, H. C.
1882. Descriptions of New Species of Reptiles and Amphibians in the
United States National Museum. Proc. U. S. Nat. Mus., Vol. 5,
pp. 438-443.

TRANSACTIONS

; OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY
VoL. X, No. 7, pp. 83-90 DECEMBER 30, 1943
Y < <a at Com Pers Py,
76,498 aren
) THE SUBSPECIES GAN 14 1944
OF THE RUBBER SNAKE, CHARINA ee
BY

LAURENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

I have lately examined somewhat over 150 specimens of Charina
to determine whether, with additional material at hand, subspecies
hitherto proposed, or new forms, might be validated. It is my conclusion
that the coastal and Great Basin snakes should be subspecifically
differentiated, as proposed by Van Denburgh, and that a new subspecies
from the mountains of southern California should be recognized. Before
discussing the territorial trends and relationships, I shall first describe
the new form as

Charina bottae umbratica subsp. nov.
SOUTHERN CALIFORNIA RUBBER SNAKE

1929. Charina bottae (part) Klauber, Copeia, No. 170, p. 19.

Type—No. 12,101 in the collection of the San Diego Society of Natural
History. Collected in Fern Valley, near Idyllwild, Riverside County, California,
by Clyde Searl, July 1, 1929. Fern Valley is in the San Jacinto Mountains at
elevation 5800 ft. The snake was found under a slab of fallen pine bark.

Diagnosis—A subspecies distinguished from C. bottae bottae by its reduced
number of scale rows; and from both bottae bottae and 6. utahensis by a low
ventral scale count, and the configuration of the frontal, supraocular, and
parietal plates.

Description of the Type—A male, probably immature; length over-all 264
mm., length of tail 32 mm. The tail is complete. There are 41 scale rows at
mid-body, the lowest lateral row being conspicuously larger than the others.
The second row is slightly enlarged and contains scales of varying sizes. All
dorsal scales are smooth and are rounded posteriorly. The ventrals number 184;
anal entire; subcaudals 31. Small anal spurs are in evidence. The supralabials

84 San Dtgeco Society oF NaturAL History

are 9-9, the fourth and fifth entering the eye; the infralabials are 11-11. The
nasals are separated above but not below; there is a single loreal on each side.
The ocular rings number 6-6; there is a single preocular on each side, which
touches the frontal, a pentagonal supraocular, and two small postoculars. The
scales on top of the head comprise a large, overhanging rostral, which is wider
than high; a pair of internasals, not quite separated by the rostral; two pairs of
prefrontals, the posterior being slightly larger; a frontal which is much wider
than long and is only a trifle convex posteriorly; and a single parietal, slightly
wider than the frontal, but about the same in length. The scales on top of the
head are unusually regular and symmetrical for Charina. On the under jaw
there is a small triangular mental followed by a pair of first infralabials in
contact on the median line. The first two pairs of gulars are slightly enlarged.
The color (as preserved in alcohol) is fawn above and buff below, without
spots or lines.

Paratype——A single paratype is available. This is No. 39 in the collection
of Pasadena Junior College. It was taken in Switzer Canyon, near Lake
Arrowhead, San Bernardino County, California, by H. C. James, May, 1925.
It is a male 295 mm. over-all, with a tail length of 41 mm. The scale rows
are 39, ventrals 190, anal entire, subcaudals 33. The supralabials are 9-9 (none
touches the eye) ; infralabials 10-11, ocular ring 7-7. In this specimen the frontal
is longer than the parietal, and is slightly convex posteriorly. In color this
individual (formalin preserved) is somewhat darker than the type.

Range.—While at present known only from the localities where the type
and paratype were collected, this subspecies may well be expected from moist
areas throughout the San Jacinto and San Bernardino mountains, and possibly
in the San Gabriels and other southern California ranges.

Discussion—So far C. bottae has not been collected in the Coast Range
south of Carmel, Monterey County, nor in the Sierra Nevada south of Sequoia
National Park, Tulare County. The latter point is about 180 miles from
Switzer Canyon, the nearest of the umbratica localities. But as the characters
separating this form from the other subspecies are differences of degree rather
than innovation, and as intervening mountains make an almost-continuous range
possible, I deem it best not to consider it a full species.!

Umbratica has fewer scale rows than bottae bottae, being similar in this
character to 6. utahensis. The ventral counts are lower than in either of the
other subspecies, although there will be overlapping when more specimens are
available. At present the lowest ventral count known from bottae is 192 in
CAS 13741, from Carmel, Monterey County. The lowest utahensis also has
192; this is LA 800 from Clear Creek, near Delta, Shasta County.

While the head scales of Charina are notably variable, the two specimens
of the new subspecies have plates of a shape not seen in any other rubber snake
T have examined. Most Charina have a frontal somewhat wider than long, but,
if unsutured, always conspicuously angular or semicircular posteriorly, where it

\Charina is known to occur on Mt. Pinos, northern Ventura County, but as no
specimens from there are presently available, it is not known to which subspecies this area
should be assigned.

KLAUBER—RUBBER SNAKES 85

contacts the parietal. The parietal is a narrow crescent-shaped scale, usually
divided in b. bottae, and entire in utahensis. In the two umbratica specimens
the frontal is almost straight across posteriorly, and the parietal is likewise almost
straight. In 6. bottae and utahensis the supraocular (or the posterior, if there
are two) deeply indents, with a sharp point, the suture between the frontal and
parietal, whereas in umbratica this point of contact is quite blunt. This supra-
ocular point may be used as a criterion with those specimens of bottae or
utahensis wherein, as occasionally occurs, either the frontal or the parietal is so
split up that its identity is lost.

THE VALIDITY OF THE GREAT BASIN SUBSPECIES
In 1920 Van Denburgh described the Utah Rubber Snake, Charina bottae

utahensis, based on the reduced scale rows of several Utah specimens as
compared with those from California. Subsequently, Ruthven (1926) and
V. M. Tanner (1933) pointed out that the high variability of the scale rows
in this genus causes a considerable overlap, especially between specimens from
California and Utah, and therefore they considered utahensis invalid. Stejneger
and Barbour have not recognized utahensis in any of the four editions of their
check list appearing subsequent to its description, nor did Stull (1935) in her
check list of the Boidae.

I believe the Great Basin subspecies should be recognized. These snakes
can be segregated from those of the Pacific Slope, with a satisfactory degree of
consistency, using the dorsal scale rows as the key character, provided state
lines are not adhered to; for the specimens in some California areas should be
allocated to the Great Basin form, while at least one Nevada county is within
the range of the western subspecies. Southern California specimens are allotted
to umbratica. The snakes most difficult to assign under this scheme are those
from the northwest, which, in part, must be considered an area of intergradation.
I regard the Pacific subspecies as being composed of individuals having 45 or
more scale rows, allocating to the Great Basin group those with 44 or less.

These two subspecies may then be summarized thus:

Charina bottae bottae (Blainville), 1835
PaciFIC RUBBER SNAKE

1835. Tortrix bottae Blainville, Nouv. Ann. d’Hist. Nat., Vol. 4, p. 289.
Type locality: California. Type specimen in Mus. d’Hist. Nat.,
Paris.

1849. Charina bottae Gray, Cat. Spec. Snakes Brit. Mus., p. 113.

1852. Wenona isabella Baird and Girard, Proc. Acad. Nat. Sci. Phila., Vol.
6, p. 176. Type locality: Puget Sound, Washington. Type speci-
men: USNM 7299.

1852. Wenona plumbea Baird and Girard, Proc. Acad. Nat. Sci. Phila., Vol.
6, p. 176. Type locality: Puget Sound, Washington. Type speci-
men: USNM 4492.

1888. Charina brachyops Cope, Proc. U. S. Nat. Mus., Vol. 11, p. 88. Type

86 San Disco Society oF NATURAL History

locality: Point Reyes, Marin County, California. Type specimen:
USNM 15524.

1920. Charina bottae bottae Van Denburgh, Proc. Cal. Acad. Sci., Ser. 4, Vol.
10, No. 3, p. 31.

Range.—The Coast Range of California north from Monterey County into
Oregon; also the Sierra Nevada north from Tulare County to Placer County
(including Douglas County, Nevada). The Placer County-Plumas County
area is one of intergradation between bottae and utahensis, as are also the
Siskiyous and Cascades to the Puget Sound region. Preliminary indications
(somewhat uncertain because of inadequate material) suggest that bottae
predominates in these northern sections, particularly toward the coast. The
snakes of Vancouver Island may belong to this subspecies. It is important, from
the standpoint of nomenclature, to note that the type specimens of both isabella
and plumbea have 45 scale rows, although specimens from around Puget Sound
often have 43 rows.

Charina bottae utahensis Van Denburgh, 1920
GrEAT BAasIN RUBBER SNAKE

1878. Charina plumbea Yarrow and Henshaw, Ann. Rept. Chief of Engineers,
App. NN, p. 1639.

1892. Charina bottae (part) Cope, Proc. U. S. N. M., Vol. 14, p. 592.

1920. Charina bottae utahensis Van Denburgh, Proc. Cal. Acad. Sci., Ser. 4,
Vol. 10, No. 3, p. 31. Type locality: Little Cottonwood Canyon,
Wasatch Mountains, Wasatch County, Utah. Type specimen: CAS
38421.

Range.—Suitable mountain and forested (non-desert) areas in north-
central Utah, western Wyoming, Idaho, western Montana, central Nevada,
western British Columbia, eastern Washington, eastern Oregon, and _north-
eastern California (eastern Siskiyou, Modoc, eastern Shasta and Lassen
counties). Intergrades with C. bottae bottae along the Cascades from Puget
Sound down to Siskiyou County, California, and along the Sierra Nevada to
Placer County, south of which area the population is bottae bottae. Parts of
the range of utahensis comprise a series of montane islands.

While this division into subspecies involves some territorial ambiguities,
wherein some individuals have 43 and some 45 scale rows, nevertheless in much
of the area inhabited by this snake there will be no misplaced or doubtful
specimens. There can be no question as to the existence of a real difference
between the subspecies, in spite of the local variations to which attention has
been called by Ruthven and Tanner. Thus, if we place all the snakes of the
Cascades in the bottae category (notwithstanding some of them have only 43
rows of scales and should be considered intergrades) we have the following
statistics of the two subspecies: Mean of 111 specimens of bottae 46.43 + 0.20;
mean of 39 specimens of utahensis 41.77 + 0.19; coefficient of divergence 10.6
per cent; P=0.0001—. Thus, in spite of weakening the difference by assigning
the intergrades to bottae, there is no question as to the significance of the

KLAUBER—RUBBER SNAKES 87

difference between the two. Within the greater part of the two territories, as
I have outlined them, the scale-row criterion will properly classify almost every
specimen.

These calculations are based on the maximum number of rows that can be
found near the middle of the body. The dorsal scale arrangements of these
snakes are much more irregular than is usual in the colubrids and crotalids, and
it is by no means easy to count the rows with accuracy or to be sure that the
maximum series has been found. The absence of pattern and the smoothness
of the scales accentuate the difficulty, especially in young specimens. Counting
can be facilitated by brushing the snake lightly, while moist, with an indelible
pencil. The color tends to follow scale edges, thus outlining them in purple,
which disappears when the snake is returned to alcohol.

While most rubber snakes are unicolor dorsally, specimens from the
southern Sierras are often rather conspicuously spotted on the lower lateral
tows or the edges of the ventrals. Young specimens from most areas are pink
or tan in life, while the adults are olive or gteenish-brown. They change so
considerably in preservation that a widespread collection of live specimens will
be necessary to determine whether there are any consistent territorial color
phases. Unless umbratica is a much stunted form, we may expect the same color
range in it, the fawn type and paratype being adolescents.

This genus is remarkable for the great variability of its head scales.
Stejneger (1890) and Tanner (1933) have pointed out that the subspecies
hitherto proposed, which have been premised on various head-scale contacts and
arrangements, that is isabella and plumbea of Baird and Girard, 1852, and
brachyops Cope, 1888, break down with the availability of adequate material.
Often the scales on one side of the head suggest one subspecies as defined,
while those on the other favor a second. As an example of this extreme
variability, I find among six specimens collected on the trash racks of a water
intake at Prospect, Jackson County, Oregon, two individuals with head plates
cut up into small scales to such an extent that according to the usual keys
they would come out Lichanura rather than Charina. Yet the other four are
quite normal. This instability is less frequent in the parietal, which is usually
divided in the Pacific subspecies and entire in the Great Basin. Whether entire
or sutured, it is angular or crescent-shaped in both of these but nearly straight
in umbratica. The parietal may, therefore, be used in some cases to reinforce
the scale rows as a differential criterion between bottae and utahensis; however,
the snakes of the northwest generally have undivided parietals, which would
place them in the Great Basin category, were this character used as a key.
Thus we have another case in which two characters differentiating a pair of
subspecies are not modified simultaneously.

The applicability of the name bottae to the coastal form is somewhat
doubtful. The type locality of Blainville’s specimen is unknown, being stated
merely as “Californie.” The scale rows of the type, as recorded by Blainville,
are too low for the Pacific subspecies; they were originally given as 39, but
Cope (1900, p. 729) stated that he inspected the type in Paris and found it
had 43. Even this would be questionable for a specimen from coastal California,

88 SAN Deco Society oF NAtuRAL History

where Botta is known to have collected. Both Blainville’s original figure and a
later one by Jan and Sordelli show that the type had a divided parietal. The
ventrals (202) are too high for umbratica and the frontal-parietal suture is
curved. Thus, while there is a questionable point here, there is nothing of
sufficient certainty to warrant a change in the application of a long-established
name.

The following may be stated with regard to territorial trends in scale rows:
In general, the snakes with the highest numbers of rows are found in the
southern Sierra Nevada and around San Francisco Bay. There is a gradual
decrease from both points northward along the mountain chains, through the
Puget Sound area into British Columbia. Eastward there is a decline to a
minimum in southeastern Idaho and extreme northern Utah. The southern
California snakes (umbratica) are also low.

There seems little or no sexual dimorphism in the ventrals of Charina.
It is easy to sex these little boas, the presence of anal spurs indicating the males,
as was first pointed out, I think, by Dr. Frank N. Blanchard. Grouping the
sexes, 87 specimens of bottae bottae have a mean of 205.02+0.59 ventrals;
extreme range 192-220; coefficient of variation 2.66 per cent. In utahensis 30
specimens have a mean of 204.80=-0.89; extreme range 192-214; coefficient of
variation 2.37 per cent. These variabilities compare favorably with those found
in other species of snakes. The figures show the importance of the low counts
in umbratica, and that bottae and utahensis do not differ conspicuously from
each other. Within bottae, specimens from the southern Sierra Nevada have
the highest numbers of ventrals, while the minima are found in the central
Coast Range. Sufficient specimens are not available to determine trends in
utahensis.

The hemipenes of bottae, as noted in two specimens from Glen Ellen,
El Dorado County, California, may be described as follows: The organs are
single-lobed. The sulcus is single; there are no spines, only a few ridges. The
basal section is smooth, except for a ridge along one side of the sulcus. From
this ridge (which may be tubular) other less-prominent ridges branch and
rebranch, in a regular pattern, forming almost plane surfaces, bounded by ridges.
At the top the sulcus terminates in a rosette surrounded by a transverse set of
ridges.

ACKNOWLEDGMENTS

I wish to acknowledge the courtesy of Mr. Joseph R. Slevin,
California Academy of Sciences; Miss Margaret Storey, of Stanford
University; Dr. Alden H. Miller and Mr. Thomas L. Rodgers, Museum
of Vertebrate Zoology, University of California; Dr. R. B. Cowles,
University of California at Los Angeles; Dr. Howard R. Hill, Los
Angeles Museum; and Mr. Robert P. Hays, Pasadena Junior College,
in permitting the examination of the specimens of Charina in these
collections. To Mr. Charles M. Bogert I owe the locating of the

KLAUBER—RUBBER SNAKES 89

paratype of umbratica in Pasadena. Mr. C. B. Perkins and Mr. C. G.

Abbott have made useful suggestions with respect to the manuscript.

KEY TO THE SUBSPECIES OF CHARINA BOTTAE

A. — Scale rows 45 or more; parietal usually divided........................ bottae
AA. Scale rows 44 or less; parietal usually entire
B. Ventrals more than 191; posterior edge of
frontal sharply angular or semicircular; sharp
point of supraocular penetrating to some depth
between) trontaly and. parietal 21a utahensis
BB. Ventrals less than 192; posterior edge of
frontal only slightly convex; supraocular with
blunt end, penetrating little between frontal
aniclnpratieta etre meet e ee ase Bhs MAE umbratica

SUMMARY

The rubber snakes may be divided into three subspecies, Charina
bottae bottae of the Pacific Slope, C. b. utahensis of the Great Basin,
and C. b. umbratica in southern California. The numbers of scale rows
and ventrals are the best key characters for segregating these subspecies.

BIBLIOGRAPHY

Barrp, S. F., and Grrarp, CHARLES
1852. Descriptions of New Species of Reptiles, Collected by the U. S.
Exploring Expedition under the Command of Capt. Charles Wilkes,
U.S.N. Proc. Acad. Nat. Sci., Phila., Vol. 6, pp. 174-176.
BLAINVILLE, H. D. DE
1835. Description de Quelques Especes de Reptiles de la Californie.
Nouv. Ann. Mus. d’Hist. Nat., Vol. 4, pp. 233-296.
Cope, E. D.
1888. On the New Species of Charina from California. Proc. U. S. Nat.
Mus., Vol. 11, pp. 86-87.
1900. The Crocodilians, Lizards, and Snakes of North America. Rept.
U. S. Nat. Mus. for 1898, pp. 153-1270.
GIRARD, CHARLES
1858. United States Exploring Expedition during the years 1838, 1839,
1840, 1841, 1842 under the Command of Charles Wilkes, U.S.N.
Herpetology, pp. xvii + 496 + Atlas of Plates. Philadelphia.
JAN, G, and Sorbet, F.
1860-1881. Iconographie Générale des Ophidiens. Pp. 100 + 3 vol.
Atlas. Milan and Paris.

90 San Dreco Society oF NAaturAL History

RUTHVEN, A. G.
1926. Notes on Utah Reptiles. Occ. Papers Mus. Zool., Univ. Mich.,
No. 179, pp. 1-4.

STEJNEGER, LEONHARD
1890. On the Snakes of the Genus Charina. Proc. U. S. Nat. Mus.,
Vol. 13, pp. 177-182.
STEJNEGER, LEONHARD, and BARBOUR, THOMAS
1923. A Check List of North American Amphibians and Reptiles.
Second Edition, pp. x + 171. Cambridge.
1933. Same. Third Edition, pp x1v + 185. Cambridge.
1939. Same. Fourth Edition, pp. x1v + 207. Cambridge.
1943. Same. Fifth Edition, pp. xix + 260. Cambridge (Bull. M. C. Z.
Vol! 93, INom1):
STULL, OLIVE G.
1935. A Check List of the Family Boidae. Proc. Boston Soc. Nat. Hist.,
Vol. 40, No. 8, pp. 387-408.
TANNER, V. M.
1933. A Study of the Variation of the Dorsal Scale Rows of Charina
bottae (Blainville). Copeia, pp. 81-82.
1939. Notes on Charina bottae in Utah: Reproduction. The Great Basin
Naturalist, Vol. 1, No. 1, pp. 27-30.
VAN DENBURGH, JOHN
1920. Description of a New Subspecies of Boa (Charina bottae utahensis)
from Utah. Proc. Cal. Acad. Sci., Ser. 4, Vol. 10, No. 3, pp.
31-32.
1922. The Reptiles of Western North America. Occ. Papers Cal. Acad.
Sci., No. 10, 2 vols., pp. 1-1028.

TRANSACTIONS

OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY

Vo.uME X, No. 8, pp. 91-126, plates 6-7, fig. 1, map

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THE SIDEWINDER, CROTALUS CERASTES,
WITH DESCRIPTION OF A NEW SUBSPECIES

LAURENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

SAN DIEGO, CALIFORNIA
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MAP SHOWING
RANGES OF THE SUBSPECIES OF

CROTALUS CE RASTES
C.c.cerastes
C.c. laterorepens

THE SIDEWINDER, CROTALUS CERASTES,
WITH DESCRIPTION OF A NEW SUBSPECIES

LauRENCE M. KLauBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

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INTRODUCTION "i

The sidewinder, or horned rattlesnake, is one of the most characteristic
animals of our southwestern deserts. While by no means restricted to sandy
areas, its peculiar method of progression, a sort of flowing sideways, is
particularly adapted to traversing such a yielding medium as sand.

Color differences between sidewinders from diverse areas have been often
noted—there seems no doubt that they tend to resemble the hue of the sand
or soil on which they live, varying through shades of cream, tan, pink,
light-brown, and gray. In addition to these color modifications, certain
differences in lepidosis and pattern, between the sidewinders of the Mojave
and Colorado deserts, have been observed. In fact, there is no difficulty in
distinguishing adult specimens, and most juveniles, from the two particular
areas of these deserts which contain the most characteristic variants, but the
allocation of specimens from other areas is not so clear-cut. I am of the
opinion, however, that the recognition of a Colorado Desert subspecies is
warranted.

The type locality of C. cerastes is given by Hallowell (1854) in this
rather indefinite way: “Borders of the Mohave river, and in the desert of the
Mohave .. . The river of the Mohave spreads itself out in the desert and
there loses itself, and upon the floating sand hills near it these animals are
found.” Whether the type specimen is still in existence is not known, although
there is some possibility that it may be Acad. Nat. Sci. Phila. No. 7098. In
fact, there is, in this case, no certainty that the type specimen was collected at
the stated type locality, the situation being analogous to that so frequently
existing between Say’s type specimens and the observations relating to the
position of Long’s Expedition when similar specimens were said to have been
seen. At any rate, the description of the type leaves little doubt that it was
a juvenile female and that it came from the Mojave Desert, and quite likely
from near the river, but just where along the river is not known.

94 SAN DtEGo SocretTy oF NATURAL HIstTory

A New SUBSPECIES FROM THE COLORADO DESERT

In order to illustrate the differences between the snakes of the two deserts,
I propose to compare two series, one from the region of the Mojave River
at the middle of its course—for adequate specimens are unavailable from
around the sink—and the other from the immediate vicinity of the type
locality of the new subspecies. With these as bases, I shall then indicate the
variations and allocations of the inhabitants of other districts. Since the
paratype series of the new subspecies is of greater interest than a single
specimen, I shall condense the description of the holotype. The new
subspecies may be known as

Crotalus cerastes laterorepens subsp. nov.

COLORADO DESERT SIDEWINDER
Plate; fig. 2.

Type.—No. 34074 in the collection of LMK, collected at The Narrows,
San Diego County, California, at 7:45 p.m., June 6, 1941, by C. B. Perkins
and Charles E. Shaw. Preserved June 13, 1941.

Diagnosis —A subspecies of Crotalus cerastes differing from the typical
form of the Mojave Desert in having 23 rather than 21 scale rows, a higher
number of ventrals, and a black instead of a brown proximal lobe of the
rattle-matrix. Other differences of morphology and pattern are discussed
hereafter.

Description of the Type-—An adult male. Length over-all (before setting
in preservative) 604 mm.; length of tail 51 mm. The head length is 27 mm.
The rattle string comprises 8 segments and is incomplete. The width of the
proximal rattle is 7.8 mm. The scale rows are 25-23-18; all rows except the
lowest on either side are keeled; the mid-dorsal row exhibits conspicuous
tubercles or bosses, which are also present, but somewhat reduced, on the
next row on each side; they are most evident at mid-body. There are 12
scale rows around the middle of the tail. The ventrals number 148 and
the subcaudals 23, most of them being entire. The anal is undivided. There
are 13-13 supralabials, and 14-14 infralabials. The rostral is wider than high;
it is contacted by six scales: a first supralabial and prenasal on each side, and
a pair of internasals. There are three canthals on each side between the
internasals and supraoculars. The other scales on top of the head are highly
irregular, and are rugose; there are about 24 scales anterior to the supraoculars.
The minimum intersupraoculars are 5+5. The nasals are divided below, but
not above, the posterior section being the larger. The loreals are 1-1; the
preoculars 2-2, the upper being the larger; the post- and suboculars number
6-6. There are three rows of scales on each side between the supralabials and
the orbit. The supraoculars are extended into prominent horn-like but flexible
processes, jutting above and beyond the eyes. The head scales in the parietal

KLAUBER—NEw SIDEWINDER 95

region are tubercular, becoming keeled and forming regular rows toward the
neck.

The head is buff above, but is given a grayish cast by the presence of
many punctations, which are especially concentrated on the snout. There is
a postocular dark stripe of punctated gray-brown on either side, which
terminates above the last supralabial. There is a prominent light cross-dash
on each supraocular, narrowing outwardly and edged with a fine black line.
In the parietal region there are 4 large and several smaller spots of gray-brown
edged with darker. Below, the mental and first three infralabials are punctated;
otherwise the lower surface of the head is unmarked.

The body is ivory or buff dorsally but is given a grayish cast by the
presence of many dark punctations. On the body there is a longitudinal series
of 38 major dorsal blotches of yellow-brown, emphasized by many darker
punctations. These are about 21/ scales long by 5 scale rows wide, and are
quite irregular in shape. They are partly edged with black. The centers of
these blotches were yellow in life. The dorsal punctations are somewhat less
evident, both within and between blotches, on the central row. Laterally there
are smaller secondary blotches, irregular both in size and shape; there are
4 series of these, the lowest of which engages the outer edges of the ventral
scutes, which otherwise are immaculate. All of these side blotches consist of
concentrations of punctations, often restricted to a single scale. On the
tail there are 5 or 6 irregular cross-marks, the first brown, the others black.
These black tail marks are conspicuously the darkest maculations on the
snake. There are some irregular black marks on the underside of the tail.
The proximal rattle-matrix lobe is black, as seen through the rattle, while the
second lobe is black anteriorly.

Summary of Paratypes——As paratypes I have used 95 specimens of
laterorepens from the Borego area of San Diego County. Most of these
were collected along the road from the foot of Sentenac Canyon to the San
Diego-Imperial County line, 3 miles east of Benson’s Dry Lake, a distance of
2014 miles. This road passes through The Narrows. A few are from the
Borego Valley, north of The Narrows, while some are from the now-abandoned
San Felipe townsite to the southeast; but all are from within a radius of 13
miles of The Narrows, the type locality of this subspecies. These specimens
are ILMIKe 1762) -1858,- 2209-10; “4571-2, -4645-7,. 4827, 4875, 4931, 4958,
5052-5, 5173-5, 9507, 21098, 21426, 22275, 22357, 23009, 23233, 23640,
23858, 23860, 23917, 23954-6, 23998, 24006, 24019-20, 25423, 25445, 26729-31,
26823, 26847-8, 26857, 26865-6, 26937-8, 27240, 28113, 28228, 28682-3,
28728, 28750, 29085, 29118-9, 29271, 29277, 29898, 30719, 31930, 31999,
32307, 32977-8, 33044, 33058-9, 33123, 33333-5, 33342, 34035, 34176-9,
54391234570, 39179. 35187-8, 35305, 35997-9, 35997, 39634-.

The statistics of the principal countable characters in the paratype series
(including the holotype itself) are set forth in Table 1. These will be discussed
later in pointing out salient differences between the Borego laterorepens and
Mojave cerastes.

96 SAN Disco Society oF NATURAL HIsTorY

As to the patterns in this series of paratypes, they have almost universally
a grayish or clay-colored cast. The ground color is in reality buff or ivory,
but the dorsums are much speckled with brown or dark-gray. The youngest
specimens are often the grayest.

The dorsal blotches are usually evident, but are obsolescent in scme
specimens, although not to the extent characteristic of certain other areas.
The dorsal blotches may be dark-gray, or light-brown, but retain little or none
of the yellow characteristic of these specimens in life. The same is true of
the postocular dark streak. The mid-dorsum is usually less punctated than
other areas of the back. The dorsal blotches are often irregularly outlined in
black. The lateral secondary blotches are fairly distinct in some specimens,
but are highly irregular and indefinite in others. When most distinct they
comprise single scales, heavily punctated with dark-brown or even black. The
edges of the ventrals are usually punctated. Head markings (except the
postocular dark streaks) are highly irregular, and often absent; however, light
supraocular cross-marks are generally present. Except in the juveniles, black tail
marks are conspicuously evident; these are rather irregular in arrangement,
seldom being in the form of even rings.

In a series of live specimens from the Borego area the following colovs
were found, using the designations of Ridgway, 1912. The ground color is
Light Buff to Pearl Gray, heavily punctated with Brownish Olive dots. The
postocular streaks are Honey Yellow to Isabella Color, as are also the centers
of the dorsal blotches. The ventrum is White to Marguerite Yellow. The
posterior tail rings are Black. It was noted, when these specimens were
preserved in alcohol, that the yellow almost disappeared. Also, there seems
little doubt that the colors of live sidewinders are somewhat affected by
temperature in the manner so much more evident in lizards.

Comparison of Subspecies——For purposes of comparison with the type
seties of the new subspecies, I have taken a series of 107 specimens of cerastes
cerastes from the desert plains surrounding the middle Mojave River. These
comprise the best available series from anywhere in the vicinity of the indefinite
type locality, given as the “Borders of the Mohave river and in the desert of
the Mohave.” I have used them as a sort of informal type series. Most of
the specimens comprising this lot were collected along highway U. S. 395 in
the 30 miles between Adelanto and Kramer Junction, San Bernardino County,
California. There are also scattered representatives from along U. S. 66 as
far east as Daggett, Minneola, and Newberry; from westward at Lovejoy, Piute,
and Pinnacle buttes in northeastern Los Angeles County; and from Muroc |
Dry Lake in southeastern Kern County. All of the area thus included
comprises but a small part of the Mojave Desert—a part contiguous to the
middle stretch of the Mojave River.

The major countable characters of these two type series are compared
in Table 1. Out of 12 characters (9 independent) 8 are found to be
significantly different in the two areas, employing the usual level of P=0.05.
But significance—that is, the real existence of a difference not chargeable to

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the exigencies of sampling—is not to be accepted as the sole criterion of relative
importance. We must also take into consideration the extent of differences,
as measured by the coefficient of divergence, and the nature of the characters
involved. I consider the subcaudals, and more especially the tail rings, as of
minor importance since neither is determinable with high accuracy—the
subcaudals because of the feathering at the rattle, and the tail rings because
of the irregularity of these marks. The latter, however, while not important
numerically, are of importance from the standpoint of color.

If we take the characters which are both significant and involve a
coefficient of divergence exceeding 5 per cent, we have scale rows, ventral
scutes, scales between the supraoculars, tail rings of the females, and the
width of the button. These are important and definite differences, although
not equally valuable as key characters. Tables 2 and 3 give the arrays of
the first two. Although overlapping is evident, the important extent of the
differences will be clearly seen.

As estimates of the overlapping in the ventral scale counts of these series of
cerastes and laterorepens. we may use the criterion of the standard range, as
suggested by Simpson (1941) and Simpson and Roe (1942). This is the
estimated range that a sample of 1000 specimens would have, and is determined
by multiplying the optimum estimate of the standard deviation of the population
by 3.2414 and adding the result to, and subtracting it from, the mean. The
standard ranges of the ventrals thus computed are as follows:

MALES FEMALES
Gerdasiesm a el at: See tee 128.90 to 144.88 132.75 to 148.63
Laterorepens plete ndneie oes. ee 137.94 to 152.00 139.69 to 157.11

The percentage of overlapping of the populations may be determined by a
formula suggested in a previous paper (Klauber, 1943, p. 56). The results
are 4.0 per cent for the males and 6.7 per cent for the females. The actual
overlaps, in my relatively small samples, are 4.4 per cent in the males and
9.0 per cent in the females.

The divergence in the size of the button (Table 1) is an indication of
a body-size difference between the two subspecies, for the button constitutes
a residual record closely correlated with body size just after birth. This
difference is further validated by the maximum sizes found in the two type series,
as follows:

MALES FEMALES
Cerastesne se eee 555 mm. 583 mm.
Waterorepens a oe ea 660 mm. 767 mm.

Cerastes is the only rattlesnake known in which the females consistently
exceed the males in size.

Scale

Rows

Ventral
Scutes
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154

19
20
21
22
23
24,
25

Total

cerastes

KLAUBER—NEw SIDEWINDER
TABLE 2
Distribution of Scale Rows in the Type Series
Mojave Borego
cerastes laterorepens
1
90 14
6 >
10 67
5
5
107 96
TABLE 3
Distribution of Ventral Scutes in the Type Series
MALES FEMALES
laterorepens cerastes laterorepens

eR
NONMVONM VO WN

RBS MOM UD BWP He

me Mm UW OW DHA DM KN WNW LO

100 SAN Dreco Society oF NATURAL History

The following color notes were made on live specimens from the two areas:

Mojave Borego
cerastes laterore pens
Ground color Fawn to light-brown Ivory to tan
Punctations Smaller and less evident | Heavy and conspicuous
Dorsal blotches and Red-brown to dark- Yellow to brownish-
postocular dark streak brown, sharply olive, indefinitely
outlined outlined
Proximal matrix-lobe Red-brown to dark- Black
in adults brown

The last-named item comprises one of the most easily applied and
consistent key characters wherewith to distinguish the two subspecies; un-
fortunately, it is mot applicable to the juveniles, for the black color of
laterorepens does not become fully evident until the acquisition of the third
or fourth rattle at the earliest, and is sometimes not present until the seventh
or eighth rattle is attained. It should be understood that this criterion refers
to the rattle-matrix itself, as seen through the rattle, and not to the last dark
marks on the tail, although these also are usually red-brown or dark-brown in
cerastes and black in laterorepens. The lateral areas of the matrix are more
important than the top and bottom, which are often blotched with light, even
in adult laterorepens.

SEGREGATION OF SUBSPECIES

Although the proper classification of individuals from the two type
areas involves little or no difficulty, this is not true of some other areas,
partly because the material at hand is somewhat inadequate, but mostly since
the subspecific differences are not so consistent or sharply drawn. As is
usually the case, the characters are not modified simultaneously; in the present
instance the two easiest-applied key characters—number of scale rows and
color of proximal rattle-lobe—are not always transformed coincidently.

The material available comprises some six hundred specimens distributed
as shown in Table 4.

The areas most poorly represented are the eastern portions of Riverside and
San Bernardino counties, in California; some parts of Arizona (particularly
Mohave and northern Yuma counties); and Sonora. This situation is the
result of a lack of collecting activities rather than the rarity of the snakes,
which are, in fact, quite common in some districts from which few or no
specimens are available in study collections.

KLAUBER—NEw SIDEWINDER 101

TRABLES4

Localities of Specimens of Crotalus cerastes

Available for Study

Number of
State County Specimens

California San Diego 125
Imperial 53

Riverside 8

San Bernardino 99

Los Angeles 48

Kern 5)

Inyo 26

Mono 1

Nevada Nye 9
Clark 20

Utah Washington 17
Arizona Yuma 34
Mohave 1

Maricopa 27,

Pinal 12

Pima 7.

Sonora Z.
Lower California 11
Uncertain 34
629

Using the number of scale rows and ventrals, and the color of the proximal!
rattle-matrix as guides, a considerable part of the range of the sidewinder
may be allocated to one subspecies or the other on the combined trends of all
three characters. The areas thus assigned are as follows:

To cerastes To laterorepens
Western San Bernardino County Lower California
Los Angeles County San Diego County
Kern County Imperial County
Inyo County Yuma County
Mono County West-central Riverside County
Nye County

Clark County

It should be understood that the listing of a county in this tabulation
dces not indicate that sidewinders are found throughout the county; the
tabulation refers only to the section in which they occur. For example, cerastes
inhabits only the extreme northeastern corner of Los Angeles County.

102 SAN Disco Society oF NATURAL History

The segregations thus made account for about two-thirds of the sidewindet’s
range. The decisions with respect to the rest of the area involve determinations
based on inadequate material, or upon giving pre-eminence to one differential
character as compared to the others. With regard to the latter, I have
decided to make the color of the basal lobe of the rattle-matrix the primary
criterion, not because I deem it more important than scale rows or ventrals,
but because, by its use, there is less subspecific overlapping, and the identification
of single specimens is made more simple and direct. However, it must be
re-emphasized that it is applicable only to fully grown adults. Further,
occasional deviations are to be expected in any territory. For example, a
specimen of c. cerastes from near Furnace Creek Inn, Death Valley, has a
black matrix, as have several snakes from the islands in Lake Mead.

Upon this basis we can make the following additional allocations:

Washington County, Utah: Assigned to cerastes based on 21 scale rows
and matrix color; the ventrals are intermediate, being somewhat higher than
in typical cerastes.

Eastern San Bernardino County, California, and Mohave County, Arizona:
Assigned to cerastes. All characters justify this allocation, but the material
is inadequate for a final determination.

Eastern Riverside County, California: Assigned to laterorepens, although
specimens with 21 scale rows slightly outnumber those with 23. This makes
the Riverside-San Bernardino line the approximate boundary in California
between the two subspecies. Along this line intergradation is to be expected,
as is suggested by specimens from Twentynine Palms and Blythe Junction

(= Rice).

Arizona, east of Yuma County: Maricopa County seems quite certainly as-
signable to laterorepens. Pinal and Pima counties are likewise, on the criterion of
matrix color, although both scale rows and ventrals are low for the southerly
subspecies. As nearly as can be determined from two specimens, Sonora
should also be considered laterorepens territory.

RANGES AND LOCALITY RECORDS

The ranges so determined may be summarized thus:

Cerastes: The desert regions of eastern (but not extreme southeastern)
California, southern Nevada, southwestern Utah, and northwestern Arizona,
including the following: extreme southern Mono, Inyo, eastern Kern, north-
eastern Los Angeles, and San Bernardino counties in California; southern Nye,
extreme southern Lincoln, and Clark counties in Nevada; southwestern
Washington County, Utah; and extreme northwestern, and west-central
Mohave County, Arizona.

Laterorepens: The desert areas of central and eastern Riverside, north-
eastern San Diego, and Imperial counties in California; northeastern Lower

KLAUBER—NEw SIDEWINDER 103

California, Mexico, from San Francisquito Bay north; northwestern Sonora,
Mexico; and Yuma, Maricopa, Pinal, and Pima counties, Arizona.

The detailed locality records available are given in the lists below. These
omit a number of doubtful localities, especially some given in the popular
literature. The term “sidewinder” is rather widely used throughout the
southwest as a designation for any small rattlesnake, and therefore it has
been frequently reported from areas where its occurrence is quite impossible.
For example, some accounts of the Hopi Snake Dance mention sidewinders
as being used, whereas the snakes referred to are really Crotalus viridis nuntius.

CERASTES

CALIFORNIA: Mono County—Chalfant; Inyo County—Laws, Bishop,
Lone Pine, Alico Siding, Keeler (also 4 and 5 mi. se.), base Inyo Mountains
(1 mi. s. of Keeler), 8 mi. ne. of Olancha, Cowan Station, Dunmovin, Linnie,
Panamint Valley, Ballarat, Panamint Mountains (vic. Goler Wash), 4 mi. w.
of Townes Pass, Mesquite Valley, also Mesquite Spring (n. end Death
Valley), Stovepipe Wells (also 4 mi. sw. and 1 and 6 mi. e.), 7 mi. sw. of
Boundary of Death Valley National Monument in Boundary Canyon, 1 mi.
sw. of Hole-in-the-Rock Spring, Beatty Junction (12 mi. n. Furnace Creek
Inn), Furnace Creek (also 6 mi. nw.), Furnace Creek Ranch, 3 mi. nw. of
Borax Mines, Funeral Mountains (4 mi. s. of mouth of Echo Canyon), Echo
Canyon, 1 mi. w. of Ryan, Bennetts Well, 10 mi. s. of Shoshone (on Cal.
127), 12 mi. n. of Trona (S.B.Co.), Borax Flats Water Station; Kern County—
Brown (also 6 mi. e.), Leliter, China Lake, Inyokern (also 8 mi. s. and 8 mi.
se.), 1 mi. nw. of Freeman Junction, Terese, Randsburg (also 2 mi. n.), Red
Rock Canyon, Cinco, Neuralia, Mojave (also 5 mi. n. and 5 mi. e.), Boron
(= Amargo) (also 3 and 4 mi. w.), Muroc, Muroc Dry Lake (= Rogers
Dry Lake) (also n. end of lake); Los Angeles County—15 mi. e. of Lancaster
(also 22, 24, and 28 mi. se.), Piute Butte, Pinnacle Butte, Lovejoy Buttes,
Lovejoy Springs, Peck’s Butte (also 1 and 2 mi. s.), Wilsonia, 8 and 9 mi. e.
of Llano; San Bernardino County—1l4 mi. e. of Inyokern, Borax Flat, Atolia
(also 2 and 3 mi. s.), Kramer, Kramer Hills, Kramer Junction (also 3, 5,
6, and 7 mi. n.; 1 and 2 mi. e.; 1 mi. w.), Adelanto (sidewinders have been
collected in every mile of the 30 miles between Kramer Junction and Adelanto),
2 and 4 mi. s. of Adelanto, 15 mi. e. of Wilsonia (L.A.Co.), Phelan, L. A.
Co. line w. of Victorville, Victor Valley near Victorville, 6 and 12 mi. n. of
Miller’s Corner, Jimgrey (also 2 and 3 mi. e.), Hawes (also 2 mi. w.), Eads,
Hinkley, Lenwood, Hodge, Wild, Helendale, Bryman, Oro Grande, Victor-
ville (also 2 and 3 mi. e.), Hesperia, Lucerne Valley, Sidewinder Well,
Stoddard Well, Ord Mountains, Coolgardie, Leach Spring, Bicycle Lake,
Camp Irwin, Calico Mountains, Yermo, Cronise, Baker (also 32 mi. n.),
Barstow (also 3 and 5 mi. ne., and 8 mi. s.), Nebo, Daggett (also 5 mi. e.),
Gale, Minneola, Newberry Spring, Rancho Larga Vega, Troy (also 4 mi.
e.), Hector, 3 mi. s. of Lavic, Ludlow, Siberia, 4 mi. e. of Bagdad, Rock
Corral, 6 mi. e. of Lone Star, Twentynine Palms (also 8, 9, and 16 mi. w.),

104 SAN Disco Society oF NATURAL HIstTory

Blythe Junction (= Rice), 8 mi. w. of Clark Mountain, 3 mi. sw. of Kelso,
Fenner, Goffs, Klinefelter, Needles.

Nevapa: Nye County—Sarcobatus Flat, North Amargosa, Amargosa
Desert, 3 mi. w. and 18 mi. se. of Beatty, Death Valley National Monument
Boundary at Highway 58, 15 mi. n. of Ash Meadows, Pahrump Valley;
Lincoln County—Quartz Spring; Clark County—Indian Springs Valley, Indian
Springs (also 2 mi. w.), Pahrump Valley, Vegas Valley, Las Vegas (also 16
and 22 mi. nw., and 6 mi. sw.), Erie, Dry Lake (RR Sta.), Glendale (also
1 and 3 mi. w., 3 mi. sw., 14 mi. e., 1 mi. n.), Lovell, near St. Joe (= bet.
Glendale and Logandale), Mormon Mesa near Virgin River, Mesquite, Lower
Muddy Valley, Atlatl Rock (Valley of Fire), 1 mi. s. of St. Thomas, Virgin
Valley, Hemenway Wash, Islands in Lake Mead, 3 mi. above Boulder Dam,
Boulder City, 114 mi. nw. of Fort Mojave.

UraH: Washington County—Beaver Dam, Beaver Dam Mountains, near
Arizona State Line (on U.S. 91), St. George (also 5 mi. n.), Sugar Loaf
(near St. George), Red Hill (n. of St. George), 14 mi. s. of Bloomington,
2 mi. w. of Indian Reservation Farm, Hurricane. (Alpine, Utah County,
reported in the literature, is now-considered inaccurate. )

Arizona: Mohave County—U. S. 91 just s. of Utah line, near Kingman,
1 mi. s. of Yucca, 6 mi. n. of Topoc.

LATEROREPENS

CALIFORNIA: Riverside County—2 and 5 mi. e. of Cabazon, Snow Creek,
Snow Creek Hatchery, Whitewater (also 3 mi. se.), Palm Springs RR.
Station~(also 1 and 3 mi. se., and 1 mi. nw.), Palm Springs (also 5 and 6
mi. nw., 4, 9, and 12 mi. se.), Cathedral City (also 1 and 2 mi. nw., 3 mi. s.,
and 3 mi. se.), 3 mi. s. junction of Cal. 74 and Cal. 111 (also 3 mi. w.), Indian
Wells (also 4 and 6 mi. w.), La Quinta (also 1 mi. e.), Indio, Sand Hills
(e. of Indio), Coachella (also 6 mi. s.), Torres, Oasis, Flying Sphinx Ranch
(near Oasis), Mecca, Box Canyon (near Mecca), Myoma, Garnet (also 3
and 4 mi. n., and 5 mi. ne.), Edom, Little Morongo Canyon (Little San
Bernardino Mountains), Wide Canyon (and 7 mi. nw.) (Little San
Bernardino Mountains), Pushawalla Canyon (Little San Bernardino
Mountains), Fargo Canyon (Little San Bernardino Mountains), Shaver
Summit, Chuckwalla Mountains, Dos Palmas Valley (w. side), North Shore
Road (Salton Sea), Salton, Hayfield, Desert Center (also 5 and 8 mi. e.,
and 5 and 17 mi. w.), Chuckwalla Valley, Hopkins Well (also 5 and 6 mi.
w.), Blythe (also 17 mi. w. and 2 mi. e.), 2 mi. s. of Blythe Junction (San
Bernardino County), near Neighbors; San Diego County—Beatty’s Ranch
(Borego Valley), De Anza Ranch (Borego Valley), Borego Springs, 6 mi.
e. of Borego P.O. (also 2 and 5 mi. s.), 8 mi. ne. of The Narrows, Yaqui
Well, The Narrows (type locality of laterorepens), Benson’s Dry Lake,
Imperial County line 3 mi. e. of Benson’s Dry Lake, (sidewinders have been
collected in every mile of the 20 miles from the foot of Sentenac Canyon via

KLAUBER—NEw SIDEWINDER 105

Yaqui Well, The Narrows, and Benson’s Dry Lake to the Imperial County
line), San Felipe (abandoned townsite), La Puerta (= Mason Valley),
Carrizo Spring (also 5 mi. nw.), Dos Cabezas; Imperial County—Fish
Springs, Truckhaven, Arroyo Salada (at U.S. 99), Tule Wash (at U.S. 99),
Arroyo Grande (at U.S. 99), Kane Spring (also 5, 10 and 12 mi. nw., and
4 and 5 mi. se. on U.S. 99), Kane Spring Junction (U.S. 99 and Cal. 78)
(also 7 mi. w.), Trifolium, half way between Westmorland and Kane Spring,
5 and 8 mi. w. of Westmorland, Vendels, Echo Island (Salton Sea), Sandy
Beach (Salton Sea), New River at Salton Sea, Myers Creek Bridge (U.S.
80), Ocotillo (= Millers), Painted Gorge (also 6 mi. sw.), 5 mi. s. of
Coyote Mountain, Coyote Wells (also 2 and 5 mi. e. on U.S. 80, and 2 mi.
w.), Yuha Plain (10 mi. s. of Coyote Wells), Petrified Forest (sw. cor. Yuha
Plain), Plaster City, Dixieland (also 1 mi. w.), Seeley (also 16 mi. w.),
Laguna Station (near present town of Seeley), 11 mi. w. of Imperial, Calexico,
4 mi. e. of Bond’s Corner, Holtville (also 4, 6, 8, 10, 12, mi. e. on U.S. 80),
Date City (also 10 mi. e.), Midway Well (Junction U.S. 80 and Cal. 98; also
2 mi. e. and 3, 4, and 6 mi. w.), Gray’s Well (also 2, 4, and 6 mi. e. on
U.S. 80), 10, 11, 14, 17, 18, 20, and 26 mi. w. of Yuma on U.S. 80, Ogilby
Junction, Springers, 1 mi. w. of Little Valley Wells Maint. Station, Pilot
Knob (also 6 mi. n.), 3 mi. n. of Bard, 3 mi. n. of Potholes, Araz, Andrade,
Coyote Valley (se. corner of county), Black Hills.

Lower Carirornia: Nw. of Laguna Salada, Cocopah Mountains, e.
base of Cocopah Mountains, Pattie Basin, San Felipe, San Felipe Bay, San

Francisquito.

Sonora: Punta Pefiasco, 9 mi. ne. of Punta Penasco en route to
Sonoyta. (Dr. Seth Benson states that he has seen sidewinder tracks at
Kino Bay.)

ARIZONA: Yuma County—Kofa Mountains, Yuma (also 1 and 3 mi. s.,
and 5, 8, 10, 11, 15, and 25 mi. e.), Yuma Mesa, 5 mi. ne. of Somerton,
VY, mi. n. of San Luis (Sonora), Monument 200 (15 mi. from Colorado
River), Araby, Dublin, Ligurta, Welton (also 5 mi. w., and 17 and 19 mi.
e.), Welton Mesa, Tacna, Pembroke, Kim, Mohawk (also 5 and 9 mi. e.).
Mohawk Mountains (also e. base), Chrystoval (= Stoval), 1 mi. e. of
Aztec, 25 mi. e. of Tule Tank; Maricopa County—Vulture, 1 mi. s. of
Morristown, Coldwater (= 18 mi. w. of Phoenix), Agua Caliente, Gila Bend
(also 10 and 28 mi. e., and 10 and 15 mi. s.), Maricopa Mountains (19 mi.
e. of Gila Bend), Phoenix, Tempe, Salado Valley (near Tempe), Mesa (also
9, 10, and 12 mi. e., and 14 mi. ene.), Desert Wells, Stewart Mountain Dam;
Pinal County—Foot of Superstition Mountain (23 mi. e. of Mesa), Sacaton,
Casa Grande (also 5 mi. w.), Casa Grande National Monument, Coolidge
(also 5 mi. e.), 3 mi. se. of Picacho, 8 mi. e. of Red Rock; Pima County—
Growler Valley, Bates Well, Growler Pass (11 mi. sw. of Ajo), Ajo (also 6
mi. ne. and 20 mi. n.), 5 mi. ne. of Sells, Marana, Santa Cruz River (near
Marana), Cortaro, 12 mi. n. of Tucson. (The following records are to be
deemed doubtful without further confirmation: Fort Buchanan and Tubac,
Santa Cruz County; and Tombstone, Cochise County.)

106 SAN Disco Socrety oF NATURAL HIstTory

INTRASUBSPECIFIC COLOR VARIATIONS °

Having discussed the variations of the two subspecies, in determining to
which the snakes of some areas should be allocated, I shall now comment
upon less-important differences of color and pattern. Most of the color
varieties of the sidewinder are localized, as shown by differences found within
rather restricted areas.

One of the most clearly defined color phases occurs in the sand hills
some 18 miles west of Yuma, and from these hills westward across the desert
to the edge of the irrigated area of the Imperial Valley. These specimens of
laterorepens are generally pink, with dorsal blotches almost obsolete; where
present, the spots are burnt-orange in color. Dorsally these snakes are heavily
punctated with brown dots, but superficially they appear almost unicolor, except
for the conspicuous black tail rings. The blotches become obsolete only in
the adults. Similar almost-patternless sidewinders are found to the west of
the Imperial Valley, especially in the vicinity of Coyote Wells and Painted
Gorge, where the same type of desert, characterized by sandy mesquite
hummocks, occurs. Strangely enough, specimens of c. cerastes from the sand
hills on the floor of Death Valley, between Stovepipe Wells and Furnace Creek,
have the same characteristic loss of blotches. These Death Valley snakes are
stunted, as indicated by the size of the proximal rattles when they have
reached parallelism.

In southwestern Utah, and in nearby corners of Arizona and Nevada,
most of the specimens are decidedly pink or reddish. The dorsal blotches
are distinct. The postocular dark stripe is often extended into a hook around
the commissure, or there may be a break, with an extra spot there. Thomas
Rodgers informs me that a specimen from Atlatl Rock, Valley of Fire, Clark
County, Nevada, which was found on sloping sandstone, was a beautiful pink
in life.

The snakes of the Coachella Valley in Riverside County, are much like
those from Borego Valley, in San Diego County—ash gray and much punctated.
The black tail rings are broad and conspicuous, even more so than in the
laterorepens patatypic series.

The most definitely marked snakes, with wide dorsal blotches clearly
and evenly outlined, are from Nye County, Nevada, especially from the
country lying northeast of Death Valley. Snakes from around Boulder Dam,
as evidenced by those collected on the islands as the water rose in the reservoir,
ate also clearly marked.

Some specimens from around Yuma have obsolete blotches, similar to
those from parts of Imperial County. Further southeast into Pima County
they are tan or gray, with blotches moderately evident.

Of the two specimens from Sonora one is pink, the other gray. Miss
Margaret Storey tells me that the specimen from’ Punta Pefiasco had brilliant
orange blotches in life.

KLAUBER—NEw SIDEWINDER 107

The most southerly specimen from Lower California, that from San
Francisquito, was probably tan in life, with fairly distinct spots. Its most
notable character is the large size of the black tail blotches.

MorPHOLOGY

The largest specimens of c. cerastes that I have seen are: male, 555 mm.;
female, 587 mm. A male measuring 628 mm. from intergrading territory
has been noted. The smallest specimen (not including the broods mentioned
elsewhere) was 171 mm. In c. laterorepens the largest male measured 665
mm., the largest female, 767 mm. The smallest measured 190 mm. Both
minimum juveniles were collected in the fall.

The sidewinder is a stout-bodied snake, as compared to other rattlers.
The weight-length curve (based largely on the subspecies laterorepens) was
found to be approximately W = 930 L-4*, W being expressed in grams and L
in meters (Klauber, 1937, p. 44). As the exponent is greater than 3, side-
winders get proportionately stouter as they age.

It is also a large-headed and broad-headed snake (Klauber, 1938, pp. 8,
38, and 48). Using the methods hitherto discussed, I have redetermined the
head (7) to length over-all (L) regression lines separately for the subspecies
cerastes and laterorepens, with the following results: for the Mojave series of
cerastes, H = 5.8 + 0.0368L; and for the paratypic series of laterorepens
H = 60 + 0.0378L. Both H and L are expressed in millimeters. There
is little difference between these lines; they give the following head lengths
at an assumed standard body length of 550 mm.; cerastes 26.0 mm.; laterore pens
26.8 mm. The coefficient of divergence at this body length is therefore 3.03
per cent, not an important difference. There is noted a tendency of some of
the largest adult females of both subspecies to have larger heads than indicated
by these equations. However, the positive value of the constant term in both
equations indicates that juveniles have proportionately larger heads than adults,
as is the case with most vertebrates.

The fangs of c. cerastes are proportionately shorter than those of c.
laterorepens, as shown by the following average figures for 5 adults of each
subspecies :

Tbe [Je A/F
Mojave cerastes 120 5.69
Borego laterorepens 98 4.83

Cerastes is found to have a somewhat shorter fang than would be expected
from consideration of the usual relationship existing between subspecies having
different adult sizes (Klauber, 1939a).

The palatine teeth in laterorepens are 3-3, the first being slightly smaller
than the other two. The pterygoid teeth are 8-8 and are slightly shorter and

108 SAN Disco Society oF NATURAL History
heavier posteriorly. The dentary teeth are 8-8 or 9-9, also growing smaller
posteriorly (scaphiodont).

The tail-length regression equations (see Klauber, 1943, p. 51), derived
from the two type series, are as follows (both T and L being expressed in mm.) :

Cerastes males tf —— 6.0 ONOZE
Laterorepens males E36 st O090E
Cerastes females == —— 20) Dea 30. 06pi15
Laterorepens females fe—-— 0) sie 0 0631e

It will be observed that there is a considerable ontogenetic change in the
males, whose tails grow proportionately longer as they age; in the case of the
females the constant term in the regression equations is too small to be im-
portant. The coefficients of divergence between the two subspecies at an
assumed standard body length of 550 mm. are: males 14.1 per cent; females
4.0 per cent, cerastes having the longer tail. The male difference is undoubtedly
significant. The respective coefficients of sexual dimorphism are: cerastes
39.1 per cent; laterorepens 29.4 per cent. This is a higher sexual difference
than is found in most rattlesnakes.

ECOLOGICAL PREFERENCES

Sidewinders prefer the sandy areas of the desert, yet they are by no means
restricted to sand, being often found in places where the surface is baked hard.
and where stones are strewn about. But they do occur in the most extreme
arenaceous areas and they are rare or absent on rocky hillsides where no
soft ground is nearby. If the territory is diverse, sidewinders are more likely
to be found in the flats, particularly about sand hummocks, or along sandy
washes. For example, they are plentiful along the sandy wash of San Felipe
Creek, from Yaqui Well to The Narrows and beyond; but up the rocky side-
canyon of the Sentenac they have never been taken, although this territory
has been hunted assiduously. Nor are they present in In-Ko-Pah Gorge,
above the Myers Creek crossing on U.S. 80. But where they are able to
follow washes and thus reach isolated valleys, they will persist there, as is
the case in La Puerta Valley of eastern San Diego County. In flats of this
kind they are by no means restricted to places where the soil is loose or
sandy. I should say that the ideal territory for the sidewinder comprises
desert flats with scattered brush, and where sand hummocks, crowned with
mesquite, or sandy washes are prevalent. And they can exist in terrain
which is entirely sand, as shown by their presence in the sand hills some
20 miles west of Yuma in Imperial County, where the strip of dunes is fully
5 miles wide.

The sidewinder seems to be driven out by irrigation, being absent in the
cultivated sections of the Imperial Valley, where it was plentiful before 1900.
The same is true of the area southwest of Yuma. I have found one laterorepens
DOR at Date City, near Holtville, where both sides of the road were cultivated,

KLAUBER—NEw SIDEWINDER 109

but this was near a neglected field which had reverted to desert. The inability of
the sidewinder to survive such an ecological change is in contrast with some
other snakes, such as the desert gopher snake, Pituophis catenifer deserticola,
which is now much commoner in the cultivated than in the primitive areas
of the desert.!

Laterorepens is a lower-altitude subspecies than cerastes; I have no records
much above 2000 feet. Without doubt it is found slightly higher at La
Puerta, in San Diego County, and near Ajo and to the northwest of Tucson,
in Pima County, Arizona. From these points it ranges down to below sea
level around the Salton Sea. Cerastes, on the other hand, ranges from below
sea level in Death Valley to at least 4500 feet in Sarcobatus Flat, Nye County,
Nevada, and 4200 feet at Chalput, Mono County, and near Bishop, Inyo
County, California. Much of the range of cerastes, particularly the western
Mojave near the probable type locality, is at an altitude of about 2500 feet.
Eventually the division between the two subspecies may be found to follow
the contours of the eastern Mojave.

Where the sidewinder is thoroughly at home it is likely to be quite
plentiful. In the western Mojave Desert in the Barstow-Kramer-Adelanto
triangle, I judge it to be the commonest nocturnal snake, with the possible
exception of Arizona elegans occidentalis and Crotalus scutulatus. In San
Diego County, along the Yaqui Well-Narrows-Benson’s Dry Lake road, it is
the fourth commonest snake, being exceeded by Phyllorhynchus decurtatus
perkinsi, Sonora occipitalis annulata, and Arizona elegans occidentalis.

CONDITIONS GOVERNING ACTIVITY;
EXAMPLE FreLp NOTES

Sidewinders are primarily nocturnal animals. As a proof one may cite
the fact that in many thousands of miles of travel on the desert only one was
found active on the highway in the daytime, while 129 were collected along
the road at night. Yet, while their activities are essentially nocturnal, they
are found abroad, but usually resting, in daylight, especially in the temperate
sun of the early spring and the late fall, or early in the morning in the summer
before the heat has become unbearable.

Some idea of the hours when sidewinders have been found on the road
may be gained from the data of Table 5. While necessarily this table
represents the hours of activity of the collector as well as his quarry, the
gradual increase in numbers after twilight proves that cerastes is not primarily
crepuscular. As to the later evening hours, while it is true that such hours
are not as often used by us for hunting as the earlier evening, yet there have
been many occasions when collecting obviously diminished as the ground
cooled below the optimum temperature. So we may judge that when the most

1 For frequency statistics see Klauber, 1939b, Table 12, p. 52.
2 For detailed statistics see Klauber, 1939b, Table 18, p. 60.

110 SAN Disco Soctgty oF NATURAL History

favorable temperatures are experienced in the early evening, the snakes do go
below ground later. Unfortunately, there having been little collecting in the
hours just before dawn in the summer, we are unable to determine whether
this is a time of activity, as might be expected from the fact that: temperatures
then would be most to the liking of the snakes.

TABLE)

Time of Evening when Sidewinders have
been found Active on Roads

Time Subspecies
Cerastes Laterorepens
6:30- 6:59 1
700-7229 2 4
7:30- 7:59 9 10
8:00- 8:29 6 7
8:30- 8:59 9 10
9:00- 9:29 5 6
9:30- 9:59 9 16
10:00-10:29 2 3
10:30-10:59 3 4
10051129 1 3
MB OST 59 1
47 66

Table 6 shows the air temperatures at times when sidewinders have been
found on the road at night. It indicates that these snakes, particularly of
the subspecies cerastes, are not seriously affected by moderately low temperatures.
The data of this table might be misunderstood in one particular. I do not
think that cerastes deliberately seeks evenings of lower temperature than
laterorepens; rather, the western Mojave, being at a higher altitude than the
western Colorado desert, and less protected by adjacent mountains from the
strong west winds of the spring evenings, cerastes finds it necessary, because of
the requirements of food and mating, to be abroad under less favorable cir-
cumstances. At any rate, one is repeatedly surprised at the conditions which
these snakes withstand; I have found them when the cold wind was so strong
as literally to sweep them across the smooth surface of the highway. I judge
their spring season of activity to be as early as that of any snake found on
the desert, which is surprising since some of the other forms range into more
temperate areas, and should thus be more accustomed to low temperatures,
while the sidewinder is restricted to the desert. Of course, it is to be remembered
that ground temperatures are even more important than air temperatures;
there is evidence that under some conditions the snakes seek the highways
because they have remained warmer than the adjacent sand.

KLAUBER—NEw SIDEWINDER 111

TABLE 6

Air Temperatures at Times when Sidewinders
have been found Active on Roads

Air Temperature Subspecies
Degrees Fahr. Cerastes Laterore pens

SI
So
Re re WR TW DW WwW A

28 47

The season of greatest activity is from the last week in April to mid-June.
Thus May is the most productive single month. The season probably lags
from one to two weeks in the higher altitudes.’

Certain features of sidewinder activities may be best described by citing a
few typical incidents.

On April 25, 1932, near Glendale, Nevada, two c. cerastes were caught
on the road at 7:45 and 8:25 p.m., with a strong, cold wind blowing; and
another snake, probably of the same species, escaped. This occurred before
temperatures were recorded, but I suspect it was well below 60° F.

May 30, 1932: In the afternoon a small specimen was uncovered just
below the surface of the sand dunes 18 miles west of Yuma. The head may
have been above ground, but at least it was not seen by my son, who found it.

April 19, 1936: At about 9 o'clock in the morning, I found a large
sidewinder by following its track. This was at the eastern edge of the sand hills.
14 miles west of Yuma. The snake was stretched out, quite inactive, in the

3 For statistics on a snakes-per-mile-of-travel basis see Klauber, 1939b, Table 7, p. 46.

i SAN DtgEGo Soctety oF NATURAL History

shade of a creosote bush. A year later (May 2, 1937) a large female (length
730 mm.) was found in the same locality; this was at 8:30 am. This snake
was at the foot of a mesquite, about 18 inches from the entrance to a hole,
and with head and neck resting in a hollow in the mesquite trunk. The snake
was entirely in the shade. It was warm, with little wind.

In the evening of the same day, I had a sidewinder come directly at me
in his best fighting pose. As I stepped out of the way his purpose was clear—
he was seeking refuge under the car. This is an example of how stories of
rattlers attacking people may start. Parenthetically, it may be said that among
people living in the desert, sidewinders are reputed to be both pugnacious and
very poisonous. As to the first, I have not found them conspicuously different
from other rattlers; as to the second, Githens and George (1931), and Githens
(1935) place cerastes in an intermediate position in the scale of rattlesnake
venoms from the standpoint of virulence. Since the sidewinder is a relatively
small snake, and the venom is not exceptional, it is less dangerous than its
larger congeners. With regard to venom yield, I can report that 169 sidewinders,
about half of which were adults, produced an average of 22.8 milligrams of
dried venom per snake. Several small groups of adults averaged from 40 to 50
milligrams, and two sets averaged 65.8 and 68.4 respectively. It is evident
that in exceptional cases secretions of at least 80 milligrams must be reached.
This compares with yields of from 100 to 300 milligrams in the larger species
of rattlesnakes, and even exceeding 1000 in exceptional cases (Klauber, 1936a,
p: 209)".

September 4, 1937: A small sidewinder bit itself while being captured.
It was dead next day—the only one of a number of snakes caught at this time
which did not survive the trip.

September 25, 1938: James Deuel found at the foot of Borego Mountain,
a Lampropeltis getulus californiae (ringed phase), which was eating a
laterorepens head first. When released the sidewinder seemed uninjured.

June 1, 1940: This evening at 8:30 a laterorepens was found in Pushawalla
Wash, Little San Bernardino Mountains, Riverside County, coiled in the sand
beside the road. The temperature was 80°F. The snake had scooped out a
little hollow so that its back was even with the surface of the ground. This is a
quite characteristic method used by the sidewinders, for they are frequently
found in the daytime, flush with the ground surface (see Gloyd, 1937, p. 124,

for illustration), if the temperature is moderate.

On April 24, 1943, James Deuel found a sidewinder 3 miles northeast
of Barstow; it was neatly coiled in the sand, exposed to the sun, at 9:10 a.m.,
and was reluctant to move, until disturbed with a stick.

COLLECTING

There are several methods of collecting sidewinders, such as driving on
desert paved roads at night, hunting around sand dunes with a flashlight, or
tracking snakes in the early morning. The latter method was preferred by

KLAUBER—NEw SIDEWINDER 113

several professional collectors whom I interviewed. It was their practice to
start out at daylight, either among mesquite hummocks or along the bank
of a sandy wash, particularly where there were sand slides. Of course, they
were searching for any snakes, but particularly the rattlers C. cinereous.
C. scutulatus, and C. cerastes, sidewinders being often found in association
with one or both of these. An early start is necessary for two reasons: to find
the tracks before they have been obliterated by the morning wind, and to
come upon the snakes before they have sought refuge in holes. It is the
custom of the snakes to lie about the holes or on the edges of the brush for a
part of the morning, the time depending on the season and the advent of an
uncomfortable degree of heat. In fact, in the early spring and late fall cerastes
may spend much time above ground, either coiled in the sand, in a hollow
scooped out for the purpose, or stretched out under the protection of a
bush. I have been told by experienced collectors that nearly every track
results in a capture by this method, and that a territory continuously worked
will soon become depleted of snakes.

In the early days of my own collecting, I hunted at night in sand dunes
with a flashlight. This is not a particularly fruitful method, as the snakes’
cryptic coloration makes them difficult to see among the twigs and desert
debris. This scheme was therefore soon superseded by the method of driving
along paved desert roads at night, picking up the snakes found crossing the
road in the glare of the headlights of the car. Because of the large area
covered, and the fact that, under certain temperature conditions, the snakes
stay on the road to profit from the warmth retained in the pavement, this
is a much more prolific scheme than the older one. Full details of this method
and its results will be found elsewhere (Klauber, 1939b).

LOCOMOTION

The peculiar method of locomotion of the sidewinder, which gives the
snake its name, is a side-flowing or looping motion whereby only vertical forces
(rather than transverse) are applied to the supporting surface. The track
comprises a series of short and separated lines, set at an angle of about 30 deg.
with the direction of progression, each line approximating the length of the
snake. If the track be in fine sand, and undisturbed by wind, the impressions
of the ventral scutes can be clearly seen, for there is no transverse or sliding
motion. Each section of the track has a fairly evident J-shaped mark (made
by the head and neck) at one end, and a T-shaped terminus (made by the tail)
at the other. The direction of progression, if one desires to track the snake, is
that toward which the hook of the J is pointing; however, the T-shaped mark
left by the tail is nearer to the destination than the J made by the head. -

All snakes tend to sidewind, if placed on a smooth or very yielding
surface, but, of course, they are not as adept as a sidewinder, which uses the
same motion even on a firm surface, although he can use a more normal
method if he wishes. It is said that the viper Cerastes of the Sahara Desert has
developed a motion quite similar to that of this rattlesnake.

114 SAN DtsGo Society oF NATURAL HIstory

The sidewinding motion, although it appears clumsy, not only facilitates
movement across a soft medium, but is relatively speedy. I think a sidewinder

can travel as rapidly, if not more so, than any other rattlesnake.

To describe the motion of the sidewinder analytically, it is necessary to
segregate several elements of motion which the snake executes simultaneously.
Assume a snake outstretched on the ground, with the body lying at an angle
of about 30 deg. with the projected line of travel, but with the tail, rather than
the head, pointing obliquely toward the objective. Using the central part of
the bedy as an anchor, the head is projected forward in such a way that the

OF ~TRAVEL aN

~ DIRECTION

Text Fic. 1. Showing consecutive position of the snake’s body in relation to the
tracks. The solid track-outlines have already been made; the dotted outlines are yet to be
made. Only the solid-black sections of the snake’s body are in contact with the ground; the
rest of the body is raised sufficiently to clear the ground. (After Mosauer, with

modifications. )

anterior third of the snake makes an angle of about 60 deg. with the rest of
the body; in so doing the head and neck form an angle of 30 deg. with the
line of progression, but on the opposite side of the line from the angle made
by the body. In advancing the head and neck they do not touch the ground
until they reach a new anchorage point, although they are raised so slightly
that one must have the eye close to the ground to note the clearance. Now
the head and an inch or so of neck are brought into contact with the ground
and serve as a new anchor, and from this anchor the snake lays out his body
forward toward the objective—not directly toward it, but at an angle of 30 deg.
As the body is being laid out, with the tail toward the objective, there is a
sharp crook at the neck so that the stationary head faces in the direction of
progression, although the body is being advanced further in that direction than
the head itself. Then again the head reaches forward for a new anchorage

KLAUBER—NEw SIDEWINDER 115

and again the body is laid out in advance of it. But always in transferring
from one anchorage to the next, whatever part of the body is in transition is
raised slightly above the ground, so that the surface offers no resistance to this
part of the movement, and no track is made; tracks are left only where the
body is laid down obliquely in advance of the head. By this means virtually
no transverse forces are applied to the supporting medium; only vertical forces
are exerted, and the track is not’a sinuous line, but a series of separate short
straight lines, advancing in echelon toward the destination. Each section of
track has a length equal to that of the snake; and, as only vertical forces are
applied, with no sliding, the imprints of each ventral scale are to be seen in

the sand (Plate 6).

Primarily, sidewinding represents the most efficient use of a loose supporting
medium, such as sand, which ‘can offer little resistance, and therefore little
reaction, to transverse forces directed across the surface (as do the sticks,
stones, and irregularities of ordinary firm ground by which the normal snake
aids his sinuous progression), but can exert a considerable resistance to forces

applied vertically.

A person watching a sidewinder will see no resemblance between the
motion as I have described it—throwing out the head, laying down the body,
throwing out the head again, etc.,—and what the snake seems to be doing; for,
as I have said, the snake telescopes these operations by executing several
simultaneously. Long before the tail has been placed, at the end of the laying
down sequence, the head has already reached out for a new anchorage, so
that the moving snake is never, even for an instant, fully outstretched along
any one of his tracks. As a matter of fact, he is always touching at least two
tracks at once, with that portion of his body between tracks slightly clearing
the ground surface; and at the time the head is first touching its next
anchorage, the tail-tip is just leaving the track two steps behind, so that for a
moment the snake may actually contact three tracks at once.

From the standpoint of step-by-step analysis, this is an accurate statement
of what the snake does, but from a pictorial standpoint, it could hardly be
worse. A sidewinding snake may best be described as one with a loose S-curve
in his body; as the curves undulate the body appears to flow smoothly (and
with a most unexpected rapidity) sidewise across the sand. Why the motion
is said to be sideways is this: if a line be drawn from the snake’s head to his
tail and this be considered the snake’s axis (although he is not outstretched, but
in an S-curve), the line of motion will be seen to be perpendicular to this
axis. In ordinary sinuous snake-motion it will be remembered that the pro-
gression is in the direction of an imaginary axis from tail to head. But a
sidewinder goes sideways as compared to the direction that a normal snake
having the same S-curve would take—hence “sidewinding.”

For an excellent discussion of the details of sidewinding, including a
description of the muscles employed, attention is directed to the papers by
the late Dr. Walter Mosauer (especially 1930, 1932b, and 1935a).

116 San Dteco Socrety oF NATURAL History

The sidewinder is rounded rather than sharply angled at the line of the
abdomen. Some species which normally inhabit sandy areas—Sonora occipitalis
and Chilomeniscus cinctus, for example—have a definite angle along the edge
of the ventrals, which serves as a boundary of the abdominal crawling surface,
but this the sidewinder lacks.

Foop

Sidewinders feed primarily on mammals (Dipodomys and Perognathus
particularly) and lizards (especially Cnemidophorus and Uma). There is
evidence that lizards are preferred by the young snakes, and mammals by the
adults. This is shown by the following table:

Cerastes Laterorepens
Mammals Lizards Mammals Lizards
Specimens containing
food 11 5 13 10
Mean size of snakes,
mm. 467 403 457 341
Limiting sizes, mm. 345-587 299-503 324-606 260-543

The preference of the younger snakes for lizards probably results from
the greater prevalence of small lizards which the little sidewinders can
successfully engulf, as compared with young mammals. The fact that the
two lizards most often found in sidewinder stomachs—whiptails and fringe-
footed sand lizards—are primarily diurnal, suggests that the snakes must often
find their prey in holes. However, H. T. Woodall found a sidewinder eating
a whiptail at mid-day above ground. This was at Twentynine Palms, in June.
One Cnemidophorus removed from a sidewinder stomach showed a fang mark
on its head. Uta stansburiana stejnegeri is another lizard which is eaten, and
Van Denburgh (1922) reports one having swallowed a horned toad
(Phrynosoma platyrhinos platyrhinos). I have been surprised to find no
Coleonyx variegatus in sidewinder stomachs, although this gecko is common
almost everywhere the rattlesnake occurs and is nocturnal. Possibly the soft-
bodied geckos would only remain recognizable for a short time.

One sidewinder (a DOR, length 530 mm.) was found to contain a bird
a California Yellow Warbler (Dendroica aestiva brewsteri). Another had
swallowed a caterpillar, but as it also contained a Uma, it is possible it may
have secured the lizard while the latter was eating the caterpillar. C. B. Perkins,
at the San Diego Zoo, feeds adult sidewinders a full grown mouse, at intervals

of a week or 10 days.

REPRODUCTION

Sidewinders usually mate in the spring. However, fall matings do occur,
as reported by Lowe (1942). A fall mating in captivity was also observed
at the San Diego Zoo, Oct. 21, 1938.

KLAUBER—NEw SIDEWINDER 117

J. R. Slevin found a pair of c. cerastes mating under a bush near Barstow
at about 5 o'clock in the afternoon, during the last week in April. The male
measured 425 mm., the female 530.

I have found two mating pairs of laterorepens. The first was on April
21, 1935, in mesquite sand hills one mile east of La Quinta, Riverside County.
This was at 9 am. on a clear warm day. They were found by tracking. The
snakes were lying quietly on the top of a dune in the shade of a bush. The
tracks showed that there had been considerable joint activity nearby. The
snakes were caught in a net with as little disturbance as possible and were
placed in a wooden box with a hinged cover. Thereafter during the day
(which was spent in alternate traveling and hunting) they were examined at
approximately hourly intervals. They were beginning to separate, although
still in copulation, at 4:25 p.m., and were fully separated sometime before
6 p.m. The left hemipenis of the male was used. The male measured 596
mm.; the female was 663 mm. in length and very stout-bodied.

The second pair was found at 8:50 p.m., May 6, 1939, on a branch
road up Fargo Canyon, Little San Bernardino Mountains, Riverside County.
There was a strong cold wind blowing up the canyon; evidently they had
sought the warmth of the pavement. Although carefully placed in a box
they were separated when we reached Indio at 9:20. The male measured
543 and the female 578 mm.

A c. cerastes from Piute Butte, Los Angeles County, gave birth to 10
young Oct. 15, 1939. The mother measured 542 mm.; the young 161 to 169
mm., with a mean of 166 mm. A laterorepens from Borego Valley, San Diego
County, was brought to the San Diego Zoo May 15, 1937, and gave birth
to 6 young Nov. 4, 1937. The young shed on Nov. 11. They varied in length
from 176 to 187 mm.; mean 182 mm. A brood of 7 born to a mother said
to have been taken in Riverside County, measured from 165 to 183 mm.; mean
176 mm. Another brood of 5 from a mother of uncertain origin measured
172 to 189 mm.; mean 179 mm.; date of birth Nov. 28, 1942. A sixth born
dead, was only 152 mm. long. It is believed that natural births in the wild
occur in the early fall; about Sept. 10 to Oct. 1, judging from the first appear-
ance of young in the field.

Gravid females of cerastes have been noted to measure from 469 mm. to
560 mm., and of laterorepens from 434 mm. to 663 mm. The following
numbers of eggs have been noted: cerastes 7, 8, 10, 10, 12, 13; laterorepens 6, 7,
88 11, 16.

As is usually the case with the rattlers (Klauber, 1936b, p. 8) the males
seem to range abroad more than the females, particularly in the spring, so
that they generally exceed the females in collections, although the actual
populations are probably little, if any, unbalanced. For example, on an evening
drive April 21, 1940, along the Adelanto-Kramer Junction road, 6 sidewinders

were collected, all males, 5 adults and one adolescent.

118 SAN Disco Society oF NaturAL History

The hemipenes are bilobed, with divided sulcus. The lobes are spinous
at the base and frilled distally, terminating in calyculate apices. There is a
sharp division between the spinous and frilled areas. The spines are rather
short and heavy, reaching their largest size on the outer shoulders, where they
form a conspicuous patch. They decrease in size by degrees around the edges of
the patch, to tiny points, which renders an accurate count impossible. However,
there are few small spines in the centers of the patches, differing in this respect
from viridis. On the side opposite the sulcus the spines on each lobe extend
to a second patch in the crotch. The spines per lobe vary in number from
about 45 to 75, depending somewhat on how many of the little points are
deemed large enough to be considered true spines. About 60 per cent are
on the shoulders, the balance in the crotch or in the line between. Occasionally
this mesial patch is almost completely separated from that on the shoulder.
Compared with other rattlesnakes the spines are of medium length and rather

heavy.

The frills or fringes are reticulate, rather than laminate, and have papillose
edges. They number from 15 to 21 on each lobe.

The lobes have a length about 11/4 times their thickness.

The male organs of c. cerastes closely resemble those of laterorepens.
The lobes of the former appear to be proportionately somewhat slimmer,
with a few more fringes and a few less spines.

In the proportionate size of the mesial patch of spines, cerastes is most
like pricet and willardi. Adamanteus and enyo have more spines in this area;
all other rattlers fewer. In the viridis group they are usually absent entirely. In
having reticulated, rather than laminated fringes, cerastes is like durissus,
molossus, adamanteus, and tigris. In the ratio of lobe length to thickness,
cerastes (especially the subspecies laterorepens), is like molossus and enyo, most
other rattlers having proportionately slimmer lobes. On the whole, except for the
reticulated fringes, cerastes appears to resemble enyo most closely.

RELATIONSHIPS

The species Crotalus cerastes is rather sharply differentiated from all
other rattlesnakes, not only because of the horn-like supraocular, but in its
method of progression, and in the superiority of the females in size. In
hemipenial characters it seems closest to enyo, and this is the case also in the
prominent tubercles of the mid-dorsal scales. Thus we might be disposed to
consider it as allied to the durissus group; however, it differs from the members
of that group in having a proportionately large, and especially a broad, head,
a short tail, and in the growth-equation of the rattle. In these characters,
and likewise in the occasional scales between rostral and prenasal, it resembles
mitchellii; yet cerastes has a relatively small rattle, proportionate to body size,
while mitchellii is at the other extreme. Thus, while it seems to be nearest to
enyo and the mitchellii group, it differs conspicuously from both.

KLAUBER—NEw SIDEWINDER 119

Coues (1875) considered the sidewinder sufficiently different from other
rattlesnakes, by reason of the horn and the roughness of the head scales, to
warrant placing it in a separate subgenus Aechmophrys. This distinction is
not recognized by modern herpetologists.

KEY TO THE SUBSPECIES OF Crotalus cerastes

Proximal lobe of rattle-matrix brown in adults; scale rows usually 21; ventrals
in males 141 or less, and in females 144 or less C. c. cerastes

Proximal lobe of rattle-matrix black in adults; scale rows usually 23; ventrals
exceed 141 in males, and 144 in females C. c. laterorepens

ACKNOWLEDGMENTS

I am greatly indebted to the following individuals and institutions for
the loan of specimens: Mr. Joseph R. Slevin, California Academy of Sciences;
Dr. Doris M. Cochran, United States National Museum; Miss Margaret
Storey, Stanford University; Mr. Thomas Rodgers, Museum of Vertebrate
Zoology, University of California; Dr. Raymond B. Cowles, University of
California at Los Angeles; Dr. Howard R. Hill, Los Angeles Museum; Mr.
Charles M. Bogert, American Museum of Natural History; Mr. M. Graham
Netting, Carnegie Museum, Pittsburgh; Dr. Vasco M. Tanner, Brigham
Young University; Dr. Ross Hardy, Dixie Junior~ College; Mr. Robert H.
Rose, Boulder Dam National Recreational Area; Mr. O. N. Arrington; and
Mr. Earl Sanders.

I am grateful to the following for assistance in making scale counts
and measurements: Messrs. Charles E. Shaw, Lawrence H. Cook, James F.

Deuel, Philip M. Klauber, and Robert J. Menzies.

I am indebted to the following for field notes, life notes, and similar data:
Mr. C. B. Perkins, Mr. Joseph R. Slevin, Miss Margaret Storey, Dr. Raymond
B. Cowles, Mr. Charles M. Bogert, Mr. Charles E. Shaw, Mr. James F. Deuel,
Ens. Paul Breese, Mr. Robert J. Menzies, and the late Lieut. Harold T.
Woodall.

I have profited from the editorial suggestions of Mr. C. B. Perkins and
Mr. C. G. Abbott, who kindly read the paper in manuscript. The illustrations
were prepared by Mr. L. C. Kobler, except the photograph of the sidewinder
track which was furnished by Dr. Raymond B. Cowles from his collection.

SUMMARY

A new subspecies of the sidewinder or horned rattlesnake, Crotalus cerastes
Hallowell, 1854, is described as C. c. laterorepens. This occurs in the desert
regions of southeastern California (Riverside County and southward), north-
eastern Lower California, northeastern Sonora, and southwestern Arizona. The

120 SAN Dteco Society oF NATURAL History

typical subspecies, C. c. cerastes, is found in the Mojave Desert of California
from southern San Bernardino County north to southern Mono County; and
in southern Nevada, southwestern Utah, and northwestern Arizona. The
differences between the subspecies entail scale counts, pattern, and color.
Intrasubspecific differences are pointed out. Life history notes are presented.

BIBLIOGRAPHY

BocERT, CHARLES M.
1941. Sensory Cues Used by Rattlesnakes in Their Recognition of
Ophidian Enemies. Annals N. Y. Acad. Sci., Vol. 41, art. 5,
pp. 329-344.

Camp, CHARLES L.
1916. Notes on the Local Distribution and Habits of the Amphibians
and Reptiles of Southeastern California in the Vicinity of the Turtle
Mountains. Univ. Calif. Publ. Zool., Vol. 12, no. 17, pp. 503-544.

Cope, Epwarp D.
1900. The Crocodilians, Lizards, and Snakes of North America. Rept.
U. S. Nat. Mus. for 1898, pp. 153-1294.

CouEs, ELLIOTT
1875. Synopsis of the Reptiles and Batrachians of Arizona; with Critical
and Field Notes, and an Extensive Synonymy. Rept. on Explora-
tions and Surveys West of the 100th Merid., Vol. 5, Zool.,
pp. 585-633.

Cow es, RAyMonp B.

1920. A List and Some Notes on the Lizards and Snakes Represented
in the Pomona College Museum. Jour. Ent. and Zool. (Pomona
Coll.), Vol. 12, no. 3, pp. 63-66.

1938. Unusual Defense Postures Assumed by Rattlesnakes. Copeia, No.
I pp? 13516:

1941. Observations on the Winter Activities of Desert Reptiles. Ecology,
Vol.'22, no. 2, pp. 125-140.

GITHENS, THomas S.
1935. Studies on the Venoms of North American Pit Vipers. Jour. of
Immunology, Vol. 29, no. 2, pp. 165-173.

GITHENS, THomas S., and GeorcgE, I. D.
1931. Comparative Studies on the Venoms of Certain Rattlesnakes. Bull.
Antivenin Inst. Amer., Vol. 5, no. 2, pp. 31-34.

GLoyp, Howarp K.
1937. A Herpetological Consideration of Faunal Areas in Southern
Arizona. Bull. Chicago Acad. Sci., Vol. 5, no. 5, pp. 79-136.
1940. The Rattlesnakes, Genera Sistrurus and Crotalus. A Study in
Zoogeography and Evolution. Chicago Acad. Sci., Spec. Pub. No.
4, pp. VII + 266.

KLAUBER—NEw SIDEWINDER 121

HALLOWELL, EDwARD
1854. Descriptions of New Reptiles from California. Proc. Acad. Nat.
Sau Piilas Vole 7, pp. 91-97.

KLAuBER, L. M.

1931. A Statistical Survey of the Snakes of the Southern Border of
California. Bull. Zool. Soc. San Diego, No. 8, pp. 1-93.
1936a. A Key to the Rattlesnakes, with Summary of Characteristics.

Trans. San Diego Soc. Nat. Hist., Vol. 8, no. 20, pp. 185-276.
1936b. A Statistical Study of the Rattlesnakes. I Introduction. II Sex
Ratio in Rattlesnake Populations. III Birth Rate. Occ. Papers
San Diego Soc. Nat. Hist., No. 1, pp. 1-24.
1937. A Statistical Study of the Rattlesnakes. IV The Growth of the
Rattlesnake. Occ. Papers San Diego Soc. Nat. Hist., No. 3,
pp. 1-56.
1938. A Statistical Study of the Rattlesnakes. WV Head Dimensions.
Occ. Papers San Diego Soc. Nat. Hist., No. 4, pp. 1-53.
1939a. A Statistical Study of the Rattlesnakes. VI Fangs. Occ. Papers
San Diego Soc. Nat. Hist., No. 5, pp. 1-61.

1939b. Studies of Reptile Life in the Arid Southwest. Bull. Zool. Soc.
San Diego, No. 14, pp. 1-100.

1940. A Statistical Study of the Rattlesnakes. VII The Rattle, Part 1.
Occ. Papers San Diego Soc. Nat. Hist., No. 6, pp. 1-62.

1943. Tail-length Differences in Snakes, with Notes on Sexual Di-
morphism and the Coefhcient of Divergence. Bull. Zool. Soc. San
Diego, No. 18, pp. 1-76.

Lowe, CHARLES H., Jr.
1942. Notes on the Mating of Desert Rattlesnakes. Copeia, No. 4,
pp. 261-262.

Merk, SETH E.
1905. An Annotated List of a Collection of Reptiles from Southern
California and Northern Lower California. Field Columbian Mus.,
Zool. Ser., Vol. 7, no. 1, pub. 104, pp. 1-19.

Merriam, C. Harr
1893. (Field notes in Stejneger, 1893)

MosAvuEr, WALTER

1930. A Note on the Sidewinding Locomotion of Snakes. Amer. Nat.,
Vol. 64, pp. 179-183.

1932a. Adaptive Convergence in the Sand Reptiles of the Sahara and
of California: A Study in Structure and Behavior. Copeia, No.
appe 72278:

1932b.On the Locomotion of Snakes. Science, Vol. 76, no. 1982.
pp. 383-585.

1932c. Ueber die Ortsbewegung der Schlangen. Zool. Jahrb., Zool. u.
Phys., Vol. 52, pp. 191-215.

122 SAN DtEGo SocreEty oF NATURAL HIstory

1933. Locomotion and Diurnal Range of Sonora occipitalis, Crotalus
cerastes, and Crotalus atrox, as Seen from Their Tracks. Copeia,
No. 1, pp. 14-16.

1935a. The Reptiles of a Sand Dune Area, and Its Surroundings in the
Colorado Desert, California: A Study in Habitat Preference.
Ecology, Vol. 16, no. 1, pp. 13-27.

1935b. How Fast Can a Snake Travel? Copeia, No. 1, pp. 6-9.

1936. The Toleration of Solar Heat in Desert Reptiles. Ecology, Vol.
17, no. 1, pp. 56-66.

1937. Sense and Nonsense about the Sidewinder. Westways, Vol. 27,
HOw SPs 22).

Mosauer, WALTER, and Lazrger, EpGar L.
1933. Death from Insolation in Desert Snakes. Copeia, No. 3, p. 149.

PERKINS, C. B.
1938. The Snakes of San Diego County with Descriptions and Key.
Bull. Zool. Soc. San Diego, No. 13, pp. 1-66.

RipGway, ROBERT
1912. Color Standards and Color Nomenclature, pp. 1-43, 53 plates.

Srmmpson, G. G.
1941. Range as a Zoological Character. Amer. Jour. of Sci., Vol. 239,
pp. 785-804.

Simpson, G. G., and Roz, ANNE
1942. A Standard Frequency Distribution Methed. Amer. Mus. Novi-
tates, No. 1190, pp. 1-19.

STEBBINS, R. C.
1943. Diurnal Activity of Crotalus cerastes. Copeia, No. 2, pp. 128-129.

STEJNEGER, LEONHARD
1893. Annotated List of the Reptiles and Batrachians Collected by the
Death Valley Expedition in 1891, with Descriptions of New
Species. North American Fauna, No. 7, pp. 159-228.
1895. The Poiscnous Snakes of North America. Rept. of U. S. Nat.
Mus. for 1893, pp. 337-487.

Van DENBURGH, J.
1922. The Reptiles of Western North America. Occas. Papers Calif.
Acad. Sci., No. 10, vol. 1, Lizards; vol. 2, Snakes and Turtles,
pp. 1-1028.

Van DENBURGH, J., and SLEVIN, J. R.
1913. A List of the Amphibians and Reptiles of Arizona, with Notes
on the Species in the Collection of the Academy. Proc. Calif.
Acad. Sci., Ser. 4, vol. 3, pp. 391-454.

KLAUBER—NEw SIDEWINDER PLATE 5

Fic. 1. Crotalus cerastes cerastes.
Adult male from 2 miles south of Kramer Junction, San Bernardino County, California.

—
a
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Fic. 2. Crotalus cerastes laterorepens.
Adult male from Borego Valley, San Diego County, California.

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TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY
Vot. X, No. 9, pp. 127-130, fig. 1 AuGusT 18, 1944

NEW OCCURRENCES OF FOSSIL TAPIR
IN SOUTHERN CALIFORNIA'

Jae 4 Comp, Pr

FA 2
BY A Zacleny

7 ee So (“sue 281944

CHESTER STOCK
California Institute of Technology

Discovery of fossil tapirs in California occurs with such infrequency
that new finds are worthy of special notice. This is all the more desir-
able since known occurrences within the state have made available only
fragmentary remains. Unfortunately, the two specimens herein de-
scribed are likewise incompletely preserved.

A single lower cheek-tooth (figure 1a) was found in April, 1943,
by Howard Duncan while making an excavation for a cesspool on his
property located at 1844 Palm avenue, National City, San Diego
County. It was encountered at a depth 12 feet below the surface in
decomposed granitic debris below a layer of hardpan. The specimen
was deposited in the paleontological collections of the California Insti-
tute of Technology through the courtesy of Clinton G. Abbott, Director,
San Diego Natural History Museum.

The geological formations in the area where the tooth was found
include a terrace cover, Pleistocene in age, and the underlying San
Diego deposits that are regarded as not older than middle Pliocene.
The San Diego formation is both marine and continental in origin,
and its upper strata, where they are weathered, can not always be
readily distinguished from the superimposed Pleistocene beds. The
latter comprise marine and littoral boulders, sands and silts. Thus the
age of the tapir tooth, in the light of information available, may be
Pleistocene. It is definitely not older than middle Pliocene.

The tooth, No. 3340 C.I.T. Coll., figure 1a, belongs to the left

1 Contribution No. 370, Balch Graduate School of the Geological Sciences, California
Institute of Technology, Pasadena, California.

128 San DtEGo Society oF NATURAL History

side of the lower jaw, and is apparently a second molar. It resembles in
size and in shape a tapir tooth (M/2?) described by J. C. Merriam
(1913, pp. 170-173, figs. la-lc) as Tapirus haysii californicus from
the late Cenozoic auriferous gravels of the Sierra Nevada, California.
It is smaller than lower teeth in the type specimen of Tapirus merriami,
described by Frick (1921, pp. 311-314, figs. 26-28) from the Bautista
Pleistocene of Riverside County. Although the dimensions of the San
Diegan tooth are slightly greater than those of Merriam’s specimen,
the ratio of crown width to crown length (19.2mm. : 28mm.) is more
like that in the latter than in Tapirus haysit.

The transverse axes of the two principal crests on the occlusal
surface diverge toward the inner border of the tooth. The crown
shows little or no wear from occlusion with other teeth, but it has
been damaged in the course of preservation. A tiny tubercle or
rudimentary ledge is located near the inner posterior base of the
metaconid. The paralophid is not completely preserved at the inner
end. It forms a straight trending ridge, situated low on the crown.
The cingulum in front of it is more feebly developed than in the
tooth from the auriferous gravels. The posterior cingulum resembles
that in the latter. Roots are not preserved in the San Diegan specimen.

A second occurrence of tapir was brought to the attention of the
Los Angeles County Museum by Pat Escobar of West Los Angeles.
An upper tooth was found in 1943 while digging a trench for a sewer
for a housing project located between Lomita and San Pedro, Los
Angeles County. The excavation is approximately 600 feet west of
the intersection of Anaheim boulevard and Vermont avenue, and 150
feet south of Anaheim boulevard. With the tapir tooth and presum-
ably found at the same site is a third upper molar of Equus occidentalis.
According to Dr. W. P. Woodring, who examined the locality, the
deposits whence the teeth are said to have come are probably upper
San Pedro (Pleistocene) in age. The equine molar is similar to teeth
of E. occidentalis found elsewhere in upper San Pedro beds. This
species occurs in the asphalt of Rancho La Brea, and is one of the
characteristic late Pleistocene mammals of the region.

The tapir tooth (left P4/?), figure 16, is relatively narrow in
anteroposterior diameter, and is rectangular in shape. The crown shows
moderate wear. The enamel surface of the inner wall of the protocone
is broken away and this defect tends to emphasize the rectangularity
of the crown. In reality, however, the transverse diameter along the

Strock—FossiL TApIR IN CALIFORNIA 129

protoloph is greater than that along the metaloph. The tooth does
not exhibit the anteroposterior extent of crown seen in molar teeth
of tapirs. In the presence of a relatively narrow V-shaped notch on
the outer wall between protocone and metacone the tooth resembles
more a fourth than a third pre-molar. The parastyle is a strong and
distinct cusp, but the cingulum along the anterior border of the tooth
is only weakly developed. At one point the thick enamel, forming the
front wall of the protoloph, practically eliminates this ledge where it
projects toward the anterior border of the crown. No ledge is present
at the inner opening of the valley between protoloph and metaloph.
Only the stubs of roots remain. The two inner roots are fused to a
distance of at least one centimeter below the crown.

The upper dentition of Tapirus, near haysii californicus, described
by J. C. Merriam (1913, pp. 172-175, fig. 2) from Cape Blanco,
Oregon, includes, unfortunately, only the three molars. It represents
a larger individual than that to which the upper San Pedro tooth be-
longed. In the front molars of the Cape Blanco specimen the anterior
cingulum seems likewise to be a thin ledge. Measurements (in milli-
meters) of the tooth from the upper San Pedro deposits are: antero-
posterior diameter, 21.1; greatest transverse diameter (approximately),
28.

Fig. 1. a, Tapirus haysu californicus Merriam. M/2?, crown of tooth is
stained but not worn; Pleistocene?, National City, San Diego County, Calif.
b, Tapirus, cf. haysu californicus Merriam. P4/?, upper San Pedro Pleistocene,
Los Angeles County, Calif. Occlusal views, natural size.

130 SAN DrecGo Society oF NATURAL HIstTory

BIBLIOGRAPHY
Frick, CHILDS
1921. Extinct vertebrate faunas of the badlands of Bautista Creek and
San Timoteo cafion, southern California. Univ. Calif. Publ., Bull.
Dept. Geol., vol. 12, no. 5, pp. 277-424.

Hay, OLIver P.
1927. The Pleistocene of the western region of North America and its
vertebrated animals. Carnegie Instn. Wash. Publ. No. 322B, p.
76, map 11 on p. 334.

MerrIAM, JOHN C.
1913. Tapir remains from late Cenozoic beds of the Pacific coast region.
Univ. Calif. Publ., Bull. Dept. Geol., vol. 7, no. 9, pp. 169-175.
StirTon, R. A., and H. W. WeEDDLE
1929. The California tapir Tapirus haysii californicus Merriam from Santa
Barbara County, California. Univ. Calif. Publ., Bull. Dept. Geol.
Sci., vol. 18, no. 7, pp. 225-226.

TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

Vor7X,)No: 10) pp213i-132 Marcu 9, 1945

A NEW RACE OF KANGAROO RAT FROM THE

ARGUS MOUNTAINS, CALIFORNIA ee

ae A

25998

Curator of Birds and Mammals, San Diego Society of Natural History

A collection of mammals made for the San Diego Society of
Natural History during August, 1931, by S. G. Harter and Vernon
Safford, in the region about Junction Ranch, Argus Mountains, Inyo
County, central-eastern California, has, with the exception of the pocket
gophers, hitherto not received detailed study. Contained in the lot is
a good number of kangaroo rats, including a series of Dipodomys
mohavensis. It is now found that these differ consistently in several
characters from topotype specimens and from specimens taken elsewhere
within the range of this species. The Argus Mountains form therefore
seems worthy of a new name and may be known as

Dipodomys mohavensis argusensis subsp. nov.

Arcus MOouNTAINS KANGAROO RAT

Type.—From Junction Ranch, 5725 feet altitude, Argus Mountains, Inyo
County, California; no. 9552, collection of the San Diego Society of Natural
History; adult male; collected by Samuel G. Harter, August 13, 1931.

Characters——As compared with topotype specimens of Dipodomys
mohavensis from Warren Station, Kern County, California, and with specimens
from Freeman Canyon, on the desert slope of the foothills of the southern
Sierra Nevada, in Kern County, Dipodomys mohavensis argusensis is slightly
larger in size, darker colored dorsally, has a darker, almost black arietiform
and a well-marked, black nose; it has also a larger ear. Cranially, D. m.
argusensis has slightly more inflated bullae, with heavier rostrum, in fully
adult specimens.

LAURENCE M. Hury LiBRAR~

132 SAN Disco Society oF NATURAL HIstory

Measurements—T ype: Total length, 297; tail, 172; hind foot, 44; ear 13.
Skull (type): Greatest length, 40.2; width across bullae, 24.5; spread of max-
illary arches, 24.0; greatest length of nasals, 15.6; greatest width of rostrum
near end, 4.5; width of maxillary arch at middle, 5.5.

Range.—So far as known, the region about Junction Ranch, Argus
Mountains, Inyo County, California.

Remarks.—Grinnell, on page 61 of “A Geographical Study of the Kan-
garoo Rats of California” (Univ. Calif. Publ. Zool., 24, 1, 1922), calls
attention to the larger mastoid bullae of a small series of 7 specimens of
Dipodomys mohavensis from the region about the Providence Mountains, as
compared with those from the type locality. The series from the Argus
Mountains studied by the writer not only shows this character, but the specimens
are also slightly larger in size and darker in coloration than topotypes.

The eastern section of the Mohave Desert is of extreme interest to the
naturalist and zoogeographer. Here are to be found several short, forest-
bearing mountain ranges, whose summits are like verdant islands. They are,
in fact, biological islands, and are the home of birds and mammals that are
racially different from those of either the plains below or the more distant,
forest-clad mountain ranges to the north.

These specialized “island” inhabitants might be divided into two classes:
they may be the last vestiges of relict populations, or they may represent racial
divergences of advancing populations. Certainly the chipmunks (Eutamias)

found on the higher brushy and wooded slopes of the Providence Mountains
would come under the former classification, while the races of pocket gophers
(Thomomys) and kangaroo rats (Dipodomys), such as the one herewith
described, would seem to reflect the results of advancing populations.

It is interesting to note that four novelties (including the present
one)—two mammals and two birds—all described within the last decade, have
been found living in the Argus Mountains. They are: Thomomys bottae
argusensis, Argus Mountains Pocket Gopher; Dipodomys mohavensis argus-
ensis, Argus Mountains Kangaroo Rat; Pipilo fuscus eremophilus, Argus
Mountains Brown Towhee; and Oreortyx picta eremophila, Desert Mountain
Quail. All, in the writer’s estimation, fall in the second of the above
classifications.

Specimens examined.—Dipodomys mohavensis mohavensis: 8 from
Warren Station, 3275 feet altitude, Kern County, California (type locality) ;
1 from Walker Pass, 4500 feet altitude, Kern County, California; 35 from Free-
man Canyon, east slope of Walker Pass, Kern County, California; 1 from Fair-
mont, Antelope Valley, Los Angeles County, California; 1 from Rock Creek,
Los Angeles County, California; 1 from 15 miles south southeast of Lancaster,
Los Angeles County, California; 5 from Hesperia, San Bernardino County,
California; 1 from Mojave River near Hesperia, San Bernardino County,
California. Total, 53.

Dipodomys mohavensis argusensis: 15 from Junction Ranch, 5725 feet

altitude, Inyo County, California (type locality).

TRANSACTIONS

OF THE
SAN DIEGO: SOCIETY, OF NATURAL HISTORY

VoLuME X, No. 11, pp. 133-216, 2 maps

THE GECKOS OF THE GENUS COLEONYX
WITH DESCRIPTIONS OF NEW SUBSPECIES

LauRENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY

Marcu 9, 1945

Zoowagy
> e
MAR 28 1949
LigRaRr*
TABLE OF CONTENTS

Page

ANELOGUCH OM oe Fe 615.200 te en ee Sue NR en een ene a ed De eee 1B
Felistoricall 5° ef. one bee ae er aIG, We HN. BAS ee nee ee 135
Discussion: of, Chatacters: 2.01 222 / -k re I Se, ele eee ee 137
Genus Coleonyx Gray. ooeecceeece oes A Nel IT a calles SNe a 138
Coleonyxtvantécatus vanegatus (Baird) ie 2 ee 138
Coleonyx:variepatus abbotts subspsnov.:.9 2 ee ee A 154
Coleonyx variegatus peninsularis subsp. nov. .......-.-..-2----0----0-0--e- ee 160
Calenr: aegis sonar erin alga nar eee A 162
Gcleonyxsvanepatus idevini subsp: MOV, 24:2 Sen ee eee 167
Coleonyx variegatus utahensts subsp: ‘Nov. 27.22 2 ee 171
Coleonyx vaniepatus sbogertt, subsp» mov.qe = ee ee ee 176
Coleonyx. fasciatus. (Boulenget). 1) Ses 2) ee ee ee eee 182
Coleonyx brews Stejnegetyc- 52 nk ae OSs spa a de ee 184
Coleonyaselegans elegans: Gray. te 2) ice oe ent eine eee 191
Coleonyx-elegans: nemoralisssubsp nov...) ase es 195
Coleonyxsmitratus, (eters). 22 2s eee et a eee ee 199
incerpenetic Relationships. .ce ase eR erent a Oe Mere aU eee 203
Generig Differentiation a2 cies te ee Ne SBE ls Sor ee ee eee 204
Keyato) the, Species ands subspeciesyot Coleone 22.2 ne eee 205
Acknowledgments) \s:20 5 eed ene ober pe RI eter Ae eee 206
Bibliography: ies 0e. Cece Wh Se Ee ae ae he Ae Sa ee ee Se ae eae 206
Sinmiimatyc eA tee A AR eae tele See 28 Doe ceane ee ee 213

Wap siAets ince oe. eesti. Ne ac cece tine As Meee eee eee 214

THE GECKOS OF THE GENUS COLEONYX

WITH DESCRIPTIONS OF NEW SUBSPECIES
BY

LauRENCE M. KLAUBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

INTRODUCTION

The geckos of the genus Coleonyx are found only in North
America. These lizards occur from the arid areas of the southwestern
United States southward to the jungles of Panama. While they have
sometimes been placed in a family known as the Eublepharidae, erected
by Boulenger in 1883 to comprise three genera characterized by the
possession of single parietal bones, more recently they have been re-
included in the far more generalized and widespread family Geckkon-
idae, partly as a result of the work of Noble (1921, p. 1), who deter-
mined that the genera assigned to the family Eublepharidae were
probably of polyphyletic origin. For this reason the latter family is
no longer generally recognized as valid, and the same may be said
of the subfamily Eublepharinae suggested by Gadow (1901, p. 512).
This decision has, however, been questioned by Walls (1942, p. 623),
premised on eye structure.

HISTORICAL

Coleonyx is a genus of geckos moderate to small in size, with bodies largely
covered with granular scales. The genus was proposed by Gray (1845, p. 162)
for the species elegans, type locality Belize, British Honduras.

In 1851 A. Duméril (in Duméril and Dumeéril, p. 45) described Gymno-
dactylus scapularis, type locality Petén, Guatemala; in 1858 he renamed the same
species Gymnodactylus coleonyx (p. 483). Both names are now considered
synonyms of C. elegans, Gray.

In 1858, Baird (p. 254) described Stenodactylus variegatus, type locality
the Colorado Desert. This was placed in the genus Coleonyx by Cope (1866b,
p. 310). It was distinguished from elegans by its lack of scattered tubercular
scales among the granules on the dorsum.

Peters (1863, p. 41) described Brachydactylus mitratus from Costa Rica.
While this also has been sometimes synonymized with Gray’s C. elegans, it has
more recently been considered a valid species (Schmidt, 1928).

In 1885 Boulenger placed Baird’s variegatus in the genus Eublepharis
(originally set up for certain Asian forms), and described two new species:
E. dovit (p. 233) from Panama, and E. fasciatus from Ventanas, western
Durango, Mexico (p. 234). Dovii is sometimes considered valid (e.g. Werner,

136 San Disco Society oF NaturAL History

1912, p. 8), sometimes a synonym of elegans Gray, 1845 (e.g. Wettstein, 1934,
p. 19), and sometimes a synonym of mitratus Peters, 1863 (Schmidt, 1928,
p- 194). Most authors, including Stejneger (1893, p. 163) and Van Denburgh
(1922, p. 58), have placed fasciatus in the synonymy of variegatus Baird, 1858;
but more recently Taylor (1935, p. 203), having available an pd dicomel
specimen, which he himself collected in southern Sinaloa, has demonstrated
the validity of Boulenger’s fasciatus.

Stejneger (1893, p. 162) felt that Boulenger’s generic arrangement, which
was based’on the relative sizes of claw sheaths, was artificial in that it split the
American forms among two genera, one of which, Eublepharis, was largely
southwest Asian. He suggested a rearrangement on a basis of the presence or
absence of enlarged chin shields. This had the very practical effect of placing
all the American species in the genus Coleonyx, while the African and Asian
species were assigned to Hemitheconyx and Eublepharis, respectively. This is
the generic arrangement currently followed by most authors, (e.g. Werner, 1912,

p. 7; Parker, 1930, p. 603; M. A. Smith, 1935, p. 125).
While Stejneger did not recognize C. fasciatus of Boulenger, he described

the Texas specimens hitherto assigned to C. variegatus as a new species, C.
brevis, based on a shorter snout, smaller anterior nasals, and more numerous
labials. Recently H. M. Smith (1933, p. 301) has pointed out additional and
more consistent differences between brevis and variegatus. He likewise extended
the known range of the Texan species.

Thus to date eight species of Coleonyx have been described, all North

American, of which five are usually recognized as valid, as follows:

C. elegans Gray, 1845; southern Mexico and northwestern Central
America.
Synonyms: Gymnodactylus scapularis A. Dumeéril, 1851.
Gymnodactylus coleonyx A. Dumeéril, 1858.

C. variegatus (Baird), 1858; eastern and southern California,

southern Nevada, southwestern Utah, western and southern Arizona;
and Lower California and northwestern Sonora, Mexico.

C. mitratus (Peters) , 1863; Salvador to Panama.
Synonym: Eublepharis dovu Boulenger, 1885.

C. fasciatus (Boulenger), 1885; western Durango and Sinaloa,
Mexico.

C. brevis Stejneger, 1893; southern New Mexico, southwestern
Texas; and Coahuila, Nuevo Leén, and eastern Durango, Mexico.

These species may be readily divided into two groups: variegatus, fasciatus,
and brevis in the north, characterized by uniformly granular dorsal scales; and
elegans and mitratus in the south, with mixed tubercular and granular scales
on the dorsum. There is a considerable territorial gap in central Mexico between
the two groups, as far as their ranges are known at this time.

KLAUBER—GENUS COLEONYX 137

DISCUSSION OF CHARACTERS

This investigation started in an endeavor to determine whether the banded
individuals of Coleonyx variegatus from coastal southern California might be
recognized as a valid subspecies. Gradually other territorial differences within
the species became evident, so that, as is frequently the case, it became desirable
to widen the scope of the research.

On the species level in this genus, it has long been known that the most
useful differential characters are the presence or absence of enlarged dorsal
scales, the shape of the snout, the nature of the scales on the toes (especially
the claw sheath), and the number and arrangement of the preanal pores. I have
found, on the subspecific level, that the number of gulars touching the mental,
the shape of the mental, the extent of separation of the prenasals and also of
the supranasals, the number and configuration of body bands, and the head
markings are of some value, although by no means all criteria show differences
between every pair of subspecies.

Countable characters, or those which are present or absent, are much more
useful in keys than those of the ratio or proportionality type (e.g. ratio of
width of orbit to length of snout), and are therefore employed wherever possible.
Some characters often used in classifying lizards are rather inefficient, or suffer
from particular limitations, in this genus. For example, the preanal pores are
clearly evident only in the adult males of some species, thus reducing the useful
specimens as far as this criterion is concerned. In the young males of variegatus
the pores are represented by uncolored indentations, and in the females by
enlarged, but flat scales, which, however, sometimes are centrally depressed.
Even in the adult males, although they are usually yellow or brown, the pores
sometimes lack color (probably the result of seasonal inactivity), and occasion-
ally one on the edge of the series, even when colored, will be seen to be
rudimentary, represented by a tiny brown dot. As by no means all enlarged
scales in the adult males have pores, even though they may seem a part of the
pore series, it is evident that counts in females, based only on enlargement or
faint indentations, are of doubtful value. This is less true in the tubercular
species wherein the females’ scales of the pore series are more definitely indented.

The enlarged supralabials can be counted with a fair degree of accuracy
by ending the count where the plates are no longer definitely enlarged, that is
where they approximate in size, the granular scales comprising the adjacent
head covering. This is usually somewhere below the middle of the orbit. The
infralabials, on the other hand, curve inward, out of view when the mouth is
closed. in such a way that the posterior members of the enlarged series are
concealed. They are difficult to limit and it is probable that observers would
often disagree as to their number. Thus, while there are average subspecific
differences in labials, they have little practical diagnostic value.

It is unfortunate that most scale series in Coleonyx are either so small,
so irregular in arrangement, or so indefinite as to point of termination, that it is
impractical to count them; for I have little doubt some would prove useful in
diagnosis, particularly the scales on the top of the head between the eyes or

138 SAN Dreco Soctety oF NAtTuRAL History

along the snout, the lamellae on some finger or toe, or the scales bordering the
eyelids. In the large species elegans, some of these series are of sufficient size and
definiteness to be counted with accuracy, but in variegatus, where they are most
needed, they are too minute and irregular to be of much practical value.

Of the variable but definitely countable scales, I have found those touching
the mental (or the mental together with the first infralabials) to be most
useful. These can be counted quite readily, even in the smallest specimens,
and there is seldom a questionable decision to make respecting any single scale.

The shape of the mental is of some importance but suffers from the usual
limitations of characters involving proportions or shape.

Notwithstanding the notable ontogenetic pattern change which takes place
in C. variegatus, certain localized modes are clearly evident, so that some rather
consistent territorial pattern types may be defined. However, these characters
of pattern, whether of head, body, or both, are generally useful only with
respect to adults, for the juveniles from all areas are much alike.

I have always found it desirable, before utilizing characters in diagnosis,
to ascertain their variability within homogeneous series, for obviously their
importance in distinguishing between forms must be dependent on their intrasub-
specific variation, or rather its opposite, consistency. In the present case the
best series of Coleonyx available to me is one from desert San Diego County.
Hence I shall use this series to define and describe the variability of Coleonyx
_ variegatus variegatus, and shall start the discussion of the genus with this
subspecies, rather than with elegans, which was first described.

Genus COLEONYX Gray
Coleonyx Gray, 1845, Ann. and Mag. Nat. Hist., Vol. 16, p. 162. Type
species elegans.

Brachydactylus Peters, 1863, Mon. Berl. Acad., p. 41. Type species mitratus.

There is a single parietal bone. The skin is soft and unattached to the
bones of the skull. The males have preanal pores in an angular series. There
is an ear opening. The eyes are large; the pupils vertically elliptical; the eyelids
are functional and are internally pigmented with black. The tail (when
original) is subcircular in cross section and regularly tapering, without transverse
constrictions. The lower surfaces of the digits are sheathed with a uniform row
of strongly imbricate transverse lamellae. They terminate in pointed claws
which are partly or entirely hidden by two shell-like lateral scales capped by a
pointed terminal. The dorsal scales are finely granular with or without tubercles
scattered among them. The ventral scales are flat and imbricate. The gulars
contacting the mental are similar to those which follow, although they may
be somewhat enlarged; there are seldom as few as 3 contacting the mental.

Coleonyx variegatus variegatus (Baird)
Desert BANDED GECKO
1858. Stenodactylus variegatus Baird, Proc. Acad. Nat. Sci. Phila., Vol. 10,
p. 254. Type specimen: USNM 3217. Type locality: Colorado Desert.

KLAUBER—GENUS COLEONYX 139

1866. Coleonyx variegatus Cope, Proc. Acad. Nat. Sci. Phila., Vol. 18, pp.
125, 310.

1885. Eublepharis variegatus (part) Boulenger, Cat. Lizards Brit. Mus., Vol.
eps 233.

Type.—The type specimen, USNM 3217, has apparently been lost. It is
figured in Baird, 1859, plate 23, figs. 9-18. It is a specimen of the speckled
desert type and therefore an adult, probably a female.

The type locality is sometimes cited as the “Rio Grande and Gila
Valleys” but this is merely the range as given in Baird’s original description.
The place of collection of the type specimen was the “Colorado Desert,” which,
from the itinerary of the collecting party, may be taken as somewhere along the
southern border of what is now Imperial County, California.

Diagnosis —On the specific level variegatus may be distinguished from
the southern Mexican and Central American members of the genus by the
absence of tubercles among the scales on the dorsum. From brevis it may be
segregated by the arrangement of the preanal pores. which, in brevis, are divided
medianly into two groups by the interposition of small scales without pores,
while in variegatus the scales carrying the pores are almost invariably confluent
at the apex of the series. From fasciatus, variegatus may be distinguished by
its blunter snout and wider head, more slender digits, and differences in pattern
and color. The body bands of fasciatus are darker than any variegatus, and
there are only 3 between limb insertions, instead of 4 as in most variegatus.

On the subspecific level variegatus variegatus may be segregated from
the subspecies subsequently to be described as follows: from slevini by the
higher number of gulars in contact with the mental; from bogerti by having
fewer preanal pores, on the average, and a deeper mental; from abbotti by the
loss of the nuchal light line and the presence of spots on the head inthe adults.
from utahensis, sonoriensis, and peninsularis by the narrow transverse adult
body bands, which are usually little or no wider than the interspaces (and are
often considerably narrower), whereas, in the other three the bands are much
wider than the intervals between. Further, the variegatus variegatus bands,
having light centers, have a double-barred effect not evident in the others. All
of these pattern differences have reference to adults, and to the majority of the
specimens found in the center of dispersal of each subspecies. There are
occasional deviates, even in these centers, and intergrades are naturally to be
expected in areas of contact between subspecies.

Range.—The subspecies variegatus variegatus is found in the deserts of
southeastern California from northern Inyo County southward; southern Nye
and southwestern Clark counties, Nevada; Arizona from the central mountain
area west and south, but not including the section east of the line Casa Grande-
Covered Wells; extreme northwestern Sonora, and extreme northeastern Lower
California, Mexico.

Example Series —As its name suggests, Coleonyx variegatus is an extremely

variable species, especially in markings and, to a lesser extent, in size and form.
An examination of adequate series, however, soon discloses that the variations

140 San Disco Society oF NaturaL History

are not haphazard; on the contrary, they fall into fairly consistent geographical
patterns, although somewhat complicated by ontogenetic changes in markings
and color. Several of these warrant subspecific recognition.

For purposes of orientation I shall first discuss the typical subspecies; _
fortunately in this case the original name was applied to the population which
comprises the central core of the complex. Following this I shall describe the
subspecies which occupy various sections of the peripheral territory.

However, variegatus variegatus itself ranges over so large an area that it is
subject to some intrasubspecific variability, although without sufficient consis-
tency—at least as far as can be determined from the available material—to
warrant further nomenclatorial subdivisions. I shall therefore first describe a
still more restricted series and discuss its internal variations in order to deter-
mine the extent of the dispersion existing in a homogeneous population.
Following this I propose to outline the differences apparent in adjacent territories
within the subspecies, and finally shall proceed to the descriptions of the new
subspecies.

The best available homogeneous series is from the Borego area of north-
eastern San Diego County. Here these geckos are quite common and may
readily be collected by driving along the paved highway at night, picking them
up in the glare of the headlights. This area is not far distant from the type
locality, which is somewhere along the Mexican border, about 50 miles across the
desert to the southeast. I have available from the Borego area 143 specimens,
mostly in my own collection. In addition, there are at hand 183 specimens of
the subspecies from other areas. The following description is based on the
Borego series only.

Morphology.—A lizard of average habitus, not particularly depressed,
either in head or body, as are many geckos. The limbs are relatively short and
weak; however, when adpressed they overlap. The head is wedge-shaped, both
horizontally and vertically; the snout is rather blunt. The nostrils are large
and circular, and lie below the canthal ridge. The ear openings are quite
prominent, but vary considerably in relative size. They are usually elliptical in
shape with the axis oblique, sloping forward and downward. The eyelids are
functional and are lined with black pigment within. The pigment shows
through the skin.

Specimens undeformed in preservation have a mid-dorsal groove, with a
low rounded ridge on either side. Sometimes there is a pair of secondary
ridges. A short mid-ventral umbilical line is present.

The largest specimens in this series (but not the maxima for the subspecies)
are two females each measuring 74 mm., snout to vent. Six other females range
from 70 to 72 mm., so this length is not to be considered exceptional. The
longest male is 69 mm.; there are several measuring 67 mm. Approximately this
size difference in favor of the females is found in all subspecies. The smallest
individual measures 43 mm.; this, however, must not be taken as a minimum
juvenile size, but results from the method of collecting, which is not so prolific
of young as that of prying off rocks or overturning debris.

KLAUBER—GENUS COLEONYX 141

The tails, when complete and original, are about equal to the body in
length, the difference seldom exceeding 10 per cent in either direction.

Table 1 gives several measurements of three example adults of each sex.

TABLE 1

Dimensions of Example Adult variegatus variegatus
Borego Series (in mm.)

Males ; Females

Snout to vent 63.0 62.0 65.0 69:05" 71.0 771.0
Length of head 1D Sali 16> 7A 16:8" 17-4
Width of head eo Oy 0 VES Ge 1251
Depth of head bh ROSIE WILD, Siler 47-2. 83
Snout to center of orbit 80° “7.2 - 7-6 S3r (7:9) > 82
Snout to ear 14.5 13.2 14.6 147) OR 5:2
Snout to arm insertion 26.0 26.0 25.0 27.0) ZION 26:0
Between limb insertions 34.5 34.0 36.5 38.5 39.0 41.0
Arm to end of fourth finger 19S PIS 5 S200 20.0 21.0 - 19.0
Leg to end of fourth toe 23:0" 26:9. 275 29:0. 927). 26.0
Length of cloacal bones 1:9 P16 eG = * **
Distance between points :

of cloacal bones S22) 78s 77, F-02655 276

* Too small to measure accurately.

There is no difficulty in sexing adults; the males may be easily recognized
by the presence of preanal pores, a postanal swelling, and lateral postanal bony
spurs, usually termed cloacal bones.

The variation in the preanal pores in the males is as follows (the first figure
indicates the number of pores while the second—in parenthesis—gives the
number of individuals with that tally): 4 (1), 5 (10), 6 (36), 7 (15), 8 (10),
9 (2); mean 6.39; coefficient of variation 15.9 per cent. Only the adult males
are included, since one cannot be sure that all uncolored depressions exhibited
by the juveniles will develop into active pores.

The scales containing the pores are in an obtuse angular series with the
point forward. The two wings of the angle meet in the center.

A considerable variation in the development of the preanal pores will be
noted, even though only adult males be considered. This is believed to be a
seasonal effect. At the peak of development (or activity) the pore occupies a
considerable part of the scale in which it is centered, although not reaching the
comparative size noted in elegans and some other species. At the time of
maximum activity the pores of preserved specimens will be found to contain
a clear yellow core, which may readily be lifted out as a solid cylinder.

Occasionally, in fully developed males, imperfect pores, smaller than the
others, are to be noted; these are usually at the posterior end of a series. While

142 SAN Disco Society oF NATURAL History

pores generally center in the scales which carry them, asymmetrical settings
have been noted. Rarely, there are two pores in a scale.

The cloacal bones in variegatus variegatus adult males are moderately wide.
They curve outward and forward. They are ridged and sloping on the upper
edge, with the points forward. They are somewhat striated and the ridge is
often notched. These processes are flexible and have a surface of skin, which is
shed like that on the rest of the body. The use of these spurs has been dis-
cussed by Greenberg, 1943. The females have rudimentary spurs which are
mere tiny points.

Coleonyx variegatus lays two eggs, which, at the time of deposition, are
quite large. The egg carried on the right side is considerably anterior to that
on the left. A specimen 74 mm. long from Yaqui Well, San Diego County,
contained eggs measuring 92 by 18/2 mm. on May 25. Another from 2!
miles northeast of Vincent, Los Angeles County, with a head and body length
of 70 mm., contained eggs 92 by 202 mm. on June 21. It is presumed that
the eggs would normally be laid at about this time.

The hemipenes are single, with single sulcus. They are distally enlarged and
terminate in fine reticulations.

Both males and females have paired post-anal sacs, with slightly oblique
openings.

Scalation—The head is covered above and below with non-imbricate
circular granules, except for the scutes which border the mouth, nostrils, and
eyes. The granules are somewhat enlarged on the snout and where they contact
the labials. The rostral is the largest of the head scales except the mental; it is
pentagonal in shape, with shorter sides engaging the first supralabials, and longer
concave edges contacting a pair of prenasals. The rostral is wider than high and is
pointed at the top. The enlarged supralabials, which are not always easy to
assign a definite count, since they gradually decrease in size posteriorly until
they become granules, number as follows: 6 (1), 7 (73), 8 (139), 9 (48),
10 (2); mean 7.91 + 0.04; coefficient of variation 8.92 per cent. The last
enlarged scales are generally anterior to a point below the center of the orbit.
The first supralabials are the largest of the series, being both higher and longer
than those which follow. There is a pair of slim, triangular prenasals which
broadly engage the rostral and barely touch the first supralabials at their lower
ends; this contact is occasionally prevented by the interposition of a granular
scale. The two prenasals usually touch medianly, but such contact is prevented
in 42 specimens out of 138 (30.4 per cent) by the presence of one or more
granules between. There is usually a very small subnasal, and two postnasals,
the upper being the larger. There is a circular supranasal on each side, larger
than the postnasals. The supranasals do not contact each other, as there are
one or more granules interposed. Counting the minimum number of scales
bridging this gap we have the following dispersion: 1 (4), 2 (28), 3 (83),
4 (18), 5 (5); mean 2.94; coefficient of variation 26.13 per cent.

The eyelids are edged with enlarged and imbricate scales forming a serrated
line. There are about 17 scales bordering the upper lid and 19 or 20 the lower.

KLAUBER—GENUS COLEONYX 143

The scales at the ends of each series are quite small and therefore difficult to
count. Posteriorly they are pointed.

The mental is the largest single scale on the head. Along the edge of the
mouth it equals or is slightly wider than the rostral. It is somewhat wider than
deep, the ratio being about 0.9. The sides are slightly convergent posteriorly.
The bottom edge is an arc of rather short radius.

The infralabials usually number 8 to 10, this being the count with the
mouth closed. Since these scales curve over the edge of the mouth, the posterior
members of the series are not visible unless the mouth be open. The visible
enlarged members of this series usually terminate somewhat posterior to the
enlarged supralabials, that is, back of the center of the orbit. The first infra-
labial is considerably deeper than the rest of the series, although not so wide as
some of those which follow. There is more reduction in size in the infralabials
from front to back than in the supralabials.

The gular scales contacting the mental can be counted definitely and
constitute a character differentiating some of the subspecies. In this series the
dispersion is as follows: 4 (4), 5 (32), 6 (48), 7 (34), 8 (15), 9 (4),
10 (0), 11 (1); mean 6.30 + 0.10; coefficient of variation 18.7 per cent. The
total gulars touching the mental, together with the first infralabial on either side,
are: 8 (1), 9 (25), 10 (36), 11 (36), 12 (31), 13 (6), 14 (2), 15 (0),
16 (1); mean 10.75 + 0.11; coefficient of variation 12.1 per cent.

The dorsum is covered with granules of similar size to those on the head.
There are no enlarged tubercles. In some specimens the granules become slightly
larger posteriorly. On the sides the scales are somewhat enlarged, flatter, and
imbricate; at the points of insertion of the limbs, however, they remain small.
Ventrally the scales are much larger than the dorsal granules; they are imbricate
with rounded ends. There is a tendency toward an increase in size both
posteriorly and medianly. Many of these scales have tiny indentations (single
or in pairs) on the posterior edges; these may be analogues of the apical scale
pits of snakes.

The regularity of the posterior abdominal scales is broken by an umbilical
line, which is bordered by irregular scales. The largest abdominal scales are in
a preanal patch, which, in the males, includes the angular series containing the
preanal pores.

The tail is covered with annular rows of subrectangular and imbricate
scales cf larger size than those on the body. These are largest ventrally. Where
the tail is thickest there are about 36 to 42 scales in a ring. In regenerated tails
(which are thicker than originals) the scales are somewhat enlarged, and the
rings rather irregular.

The scales on the arms are mostly imbricate, except on the lower or inner
side of the upper arm. The scales are largest on the upper side of the wrist.
The palmar surface is covered by tubercular scales; the upper sides of the hands
and fingers are covered by strongly imbricate scales. Below on the digits there
are series of rectangular lamellae. These are so thick and strongly imbricate
as to form a series of transverse ridges, thus comprising a longitudinal row of

144 SAN Dreco SocteTry oF NATURAL HIstory

corrugations. The claws are sharp and delicate; they are formed by lateral
compression of scale-like members, and are, therefore, hollow. They are held
between a pair of lateral shell-like scales, the infero-laterals of Parker, 1926. On
the top, the crevice between these (out of which the claw issues) is capped by a |
single, pointed scale (the terminal); while on the bottom the crevice is covered
by a broader, shorter, and blunter, scale. There are about 17 lamellae on the
fourth finger.

The thighs are covered with enlarged imbricate scales on the anterior surface
and small granular scales behind. On the lower section of the legs the relation-
ship is reversed; the larger scales are behind, the smaller forward. The coverings
of the feet and toes, and the housing of the claws, are similar to those on the
fingers. There are about 22 lamellar corrugations on the fourth toe.

Pattern and Color.—In the juvenile stage variegatus variegatus is not
materially different in pattern from the other subspecies, although the head is
somewhat lighter in color, and the dorsal cross-bars are both lighter and narrower
(compared with the interspaces) than the others.

The juvenile pattern may be described thus: The head above is light-brown,
somewhat darker on the snout and laterally. The canthal ridge is light; below
this there is a dark bar. The labials, particularly the upper, are mottled with
brown; sometimes the color is so disposed that individual scales are either
entirely light or dark. Below each eye there is a narrow light line which passes
backward above the ear; these lines join to form a loop on the anterior part of
the neck. This postparietal or nuchal light loop is highly characteristic of the
juveniles of all variegatus subspecies, but it is less perfect—that is, less definitely
and evenly outlined—in variegatus variegatus juveniles than in the other sub-
species.

On the neck there is a broad, brown cross-band with pointed extensions
carried forward laterally toward the ears. These comprise the posterior borders
of the light nuchal loop.

On the body there is a series of medium-brown cross-bands, usually 4
between limb insertions; they are approximately as wide as the cream-colored
interspaces or somewhat narrower. They often curve forward laterally. The
ventral surface is clear white. The tail (if not regenerated) is also barred;
the rings may be slightly darker than those on the body.

The change from the juvenile to adult pattern is quite marked in this
subspecies, and there is a considerable variability in the manner of its occurrence.
The head first becomes mottled and finally spotted with brown on a cream
background. The mottling begins to appear at a body length of from 35 to
45 mm. and constitutes the earliest change from the pattern of abbotti. In its
final phase the identity of the light nuchal loop is often completely lost, and in
any case it becomes ill-defined and irregular. The light canthal lines are
generally retained, and the darkest marks on the head are the dashes which
comprise their lower borders; these dark lines run from the anterior point of
the eye to the nostril.

The rostral is generally light in the center and dark laterally. The prenasals
are often conspicuously dark. The serrated scales bordering the upper eyelids

KLAUBER—GENUS COLEONYX 145

are faintly tinged with gray. The upper labials are mottled; usually 2 or 3 are
entirely punctated. The mental is generally clear, and the infralabials less
maculate than those above.

Conspicuous changes also take place in the body marks. The more or less
regular brown cross-bars of the juveniles become increasingly irregular. Usually
they become narrower (along the body) and a series of brown spots or
blotches appears in the widened interspaces, first laterally, then entirely across.
The bars themselves become lighter in the middle, resulting in a double-barred
effect. Much spotting (usually in the form of smaller dots than the spots in
the interspaces) appears along the sides between limb insertions. The lower
surface remains immaculate cream-color. Sometimes the dorsal bands break
up into spots to such an extent that uniform spotting results, all vestiges of
bands being lost. Such seems to have been the case in the type specimen.

The tail rings also become lighter, particularly in the centers so that they
are double-barred. Spots occasionally appear in the interspaces, but not so
generally as on the body. One or two of the terminal rings may be complete
ventrally. Regenerated tails are irregularly spotted.

Mid-dorsal light lines, such as characterize sonoriensis, are rare but do
occur. I might mention that the mid-dorsal groove and its bounding ridges
sometimes give a false impression of a light vertebral line.

The cross-bars on the body in this Borego series usually number 4, but
other numbers are occasionally encountered. Out of 128 specimens in which
the bands are still evident, one has 3 on one side of the body and 4 on the
other, eight have 4 on one side and 5 on the other, and seven have 5 straight
across; the other 112 have 4 bands. The tail rings vary from 10 to 14, 11 being
the mode.

With this Borego series spread out before me, there is hardly a half-dozen
adults which, in the relative widths of body bands and interspaces, and in their
nature, might be confused with either sonoriensis or utahensis.

The peritoneal lining in Coleonyx is not dark as in many desert lizards
(Klauber, 1939, p. 75). However, the eye socket is lined with black as can be
seen, not only on the edges of the lids, but through the skin. In fact the skin
is so transparent that most specimens have a pinkish cast in life. However,
even in the desert specimens there is sometimes a yellowish pigment between
the dorsal brown bars.

The following color notes, using Ridgway’s standards, were made on a live
young adult from Cornville, Yavapai County, Arizona: The spots on the head
are Natal Brown upon a background of Dull Lavender. The dorsal body bars
are Raisin Black, with Deep Olive Buff interspaces. The legs are Vinaceous
Lilac above and Deep Lavender below. The ventrum is White.

A pair of adults from The Narrows, San Diego County, exhibited the
following colors: Head spots, Prussian Red to Haematite Red; ground color
between head spots, Light Grayish Vinaceous to Vinaceous-Buff; body spots,
Hay’s Brown, Sorghum Brown, Vandyke Brown; between spots but within
blotches, Brownish Vinaceous; interspaces, Olive Buff; tail bands, Natal Brown
to Bone Brown; interspaces on tail, Avellaneous; legs above, Vinaceous Buff;

146 San Deco Society oF NaAtTuRAL History

underside of legs, Light Vinaceous Fawn; underside of head, body, and tail,
White.

Miss Atsatt (1939, p. 244) has discussed the effects of changes in
temperature and light on the color of Coleonyx. It tends to become darker
at lower temperatures, and, probably, in stronger illumination.

Intrasubspecific Trends.—Having described variegatus variegatus, as exem-
plified by the Borego series, I shall now survey briefly the trends in other areas
inhabited by the subspecies. Also, the dispersions of the countable characters
will be given for all available material. The following additional specimens have
been surveyed, beyond the Borego series of 143.

Northeastern Lower California, Mexico 6
Imperial County, California 30
San Diego County, California 6
Riverside County, California aA
San Bernardino County, California 26
Los Angeles County, California
Kern County, California

Inyo County, California

Nye County, Nevada

Clark County, Nevada

Mohave County, Arizona
Yavapai County, Arizona
Maricopa County, Arizona
Pima County, Arizona

Yuma County, Arizona
Northwestern Sonora, Mexico

Total 183

The list, of course, excludes all specimens allocated to other subspecies.
Also, pattern notes were made on some 68 additional specimens of variegatus
yvariegatus which are not included above.

With the material at hand I have been unable to find any important terri-
torial trends in morphology within the subspecies. Specimens approaching the
area of intergradation with bogerti usually have an increased number of preanal
pores.

As I have stated elsewhere, variegatus variegatus is the largest of the sub-
species. Females exceeding 70 mm., snout to vent, are by no means uncommon
on the floor of the desert. The largest specimen I have measured is one from
Alameda Junction, 3 miles west of Garnet, Riverside County, which is 77 mm.
in body length. Individuals exceeding 70 mm. have been collected in San
Diego, Imperial, Riverside, San Bernardino, and Los Angeles counties in
California, and Yuma County, Arizona. The largest male measured 69 mm.
The smallest available individual of this subspecies measures 33 mm. Mountain
specimens do not attain the extreme length of those of the desert, and the same
is true of some of the fringe territory approaching the ranges of other subspecies.

The complete distribution of the preanal pores in the males, for the
subspecies as a whole, is as follows: 3 (1), 4 (1), 5 (16), 6 (80), 7 (36),

—"

i)
NAN OOOONNM Oe Ff

KLAUBER—GENUS COLEONYX 147

8 (26), 9 (3); total 163; mean 6.45. It will be observed that 6 is strongly the

mode in this subspecies, about half the specimens having that number.

There are apparent no important territorial trends in scalation in variegatus
variegatus as a whole, when compared with the Borego series.

The dispersion of the gulars contacting the mental in the subspecies as
a whole is as follows: 4 (13), 5 (70), 6 (116), 7 (73), 8 (37), 9 (8),
10 (1), 11 (1); total 319; mean 6.26. The gulars contacting the mental and the
first infralabials taken together are: 8 (9), 9 (60), 10 (95), 11 (76), 12 (55),
13 (15), 14 (5), 15 (3), 16 (1); total 319; mean 10.61. The gulars contacting
the mental tend to be fewer in the geckos from the northern Mojave Desert;
they are most numerous in specimens from around Yuma.

The prenasals fail to make contact in 76 cases out of 324, or 23.5 per cent.
Contact most frequently fails in the area between Yuma County, Arizona, and
the eastern slope of the coastal range in Imperial and San Diego counties,
California.

For the subspecies variegatus variegatus as a whole, the dispersion of the
granules bridging the gap between the supranasals is as follows: 1 (13), 2 (85),
3 (180), 4 (33), 5 (9), 6 (1); total specimens 321; mean 2.82. The number
of these scales tends to be low in the upper Mojave Desert and toward Death
Valley, and reaches a maximum in the Yuma area and across the line in
northwestern Sonora.

With regard to pattern variations in the subspecies as a whole, we find
the following; In Imperial County, especially along the southern border, at
some point of which the type specimen was probably collected, the patterns are
similar to those of the Borego series. If any difference is present, it lies in an
increased tendency toward a complete loss of the bands in the fully-grown
adults—to have them break up into spots, as was the case in the type. The same
is true in the desert area of northeastern Lower California, as far south as San
Felipe, the southern known limit of this subspecies in the peninsula.

In San Bernardino County—the Mojave Desert—there is a tendency to
retain the dorsal bands. They narrow and become double-barred by the light-
ening of the interiors, and the interspaces become filled with spots; but the
bars do not break up as completely as is often the case with the specimens from
the Colorado Desert further south. The Los Angeles County desert specimens
are similar to those of the Mojave.

Further north in Inyo County there is more variability, resulting, no
doubt, from the complex terrain. In general, the body blotches tend to be
darker and are often wider. They are conspicuously double-barred. The head
spots are clear.

In southern Nye County and in southwestern Clark County, Nevada,
tendencies toward utahensis are found, the bars becoming wider, more irregular,
and with the light centers less evident. Those from Las Vegas and Boulder
City northeastward are to be considered utahensis, though not as extreme as
typical Utah specimens; some, in fact, from this area of intergradation, more
nearly resemble variegatus variegatus.

148 SAN Dreco Society oF NaturaL History

In Mohave and Yavapai counties, Arizona, the markings are quite dark,
especially when from mountain areas. The bars in the adults are sometimes
wider than the spaces between. Further south in Maricopa County, the blotches
are still dark, the double-barred effect is quite clear, and the head spots
prominent. In Yuma County the patterns are much like those of Imperial
County, except that the dorsal pattern, in breaking up in the adults, tends to
split into relatively larger spots. Where the bars are retained, they are double
and narrower than the interspaces. There is little trend toward sonoriensis,
except in an occasional vertebral light line. The specimens in extreme north-
western Sonora, while still variegatus variegatus, do show occasional sonoriensis
tendencies, both in the widening of the dorsal bands and the greater frequency
of a light mid-dorsal line.

The number of dorsal body bars and the tail rings do not show any terri-
torial tendencies in variegatus variegatus. Four bars on the body are strongly
the mode everywhere. The following tabulation gives the distribution for the
subspecies as a whole. The figure 1/2 indicates a specimen with a higher number
on one side and a lower on the other—as is the case where one of the bars
hasia! Y-shape:1521(l)5 44( 239) 4720 (19), > (23).

Relationships with Other Subspecies —V ariegatus variegatus comprises the
largest and most centrally located subspecific population. It intergrades with
the new subspecies abbotti, utahensis, bogerti, and sonoriensis, all of which are
peripheral variants. Areas of intergradation will be discussed under each of
these. Variegatus variegatus may be related to peninsularis through abbotti,
or directly via the east coast of Lower California, or through some presently
unknown subspecies; this cannot be decided until much more material shall
have become available from central Lower California.

Field Notes——The method of collecting desert reptiles by driving on
paved roads at night (Klauber, 1939) has greatly improved both our herpe-
tological collections and our knowledge of reptile habits. This has been the
case with Coleonyx variegatus variegatus, which was found to be much more
common at night in the Colorado Desert than any other lizard. or, in fact, than
all other snakes and lizards taken together. As many as 30 have been en-
countered on the road in a single night. Often we have collected only a few of
those seen and have even failed to keep a full record of them. As they
were formerly secured only by the hard work of prying off rock flakes,
and overturning debris, night driving has converted a rare into a common
species. From the coastal side of the mountains, where we are still restricted
to the older methods, comparatively few specimens are even yet available.

Variegatus variegatus has been met with on the road in a great variety of
situations. It is found in every kind of desert habitat to an elevation of about
4000 ft.; in rocks, brush, cacti, and on sandy flats and washes, although
probably most plentiful in the latter. But it is not equally common in all parts
of the desert; for example, it is less frequently encountered on the run from
Adelanto to Kramer Junction, in San Bernardino County, than between The
Narrows and Bensons Dry Lake, in San Diego County, although the ecologi-

cal conditions are not greatly different in the two areas. It occurs among stony

KLAUBER—GENUS COLEONYX 149

outcrops and also in our most extreme arenaceous territory—the sand dunes
some 17 miles west of Yuma. It seems to have been driven out of the irrigated
sections of the Imperial and Coachella valleys, although found where the land
has reverted to desert. The vertical distribution is from below sea level in the
Salton Basin and Death Valley to about 4000 ft. (Wheaton Spring, 4025 ft.,
Big Pine, 4002 ft.); probably it reaches 4500 or 5000 ft. in the mountains
of Inyo and northeastern San Bernardino counties. La Rivers (1942, p. 56) has
recorded a specimen from Quartz Spring, Lincoln County, Nevada at 4500 ft.
This should probably be assigned to the subspecies utahensis.

If one is traveling slowly the geckos are rather easily seen on the road, as
they are almost white laterally. Usually they cross slowly and appear like bits
of paper blown along the highway. They will often remain quiet if the car
is stopped quickly enough to hold the headlights on them; if they are passed,
so they are again in the dark, they will occasionally escape. No eye shine has
been observed. Where paved roads are not available these geckos may be
secured by hunting afoot with a flashlight; however, this scheme is not nearly
so fruitful as collecting by auto.

Although their skins are so thin as to be partially translucent, and they
have the appearance of being delicate and fragile, they must actually be quite
hardy. For while they prefer warm spring nights, they have been found active
on nights when the temperature was 60° F. or less, and with so violent a gale
blowing that it was difficult to understand how they could cling to the highway
surface. I think I have seen them out under more extreme weather conditions
than any other desert reptile except possibly Crotalus cerastes. On occasional
spring trips when the weather conditions were particularly adverse, Coleonyx
was the only reptile seen. However, I have never encountered one in full day-
light except on one occasion when no place of concealment was available. This
was on a tiny island in Lake Mead (June 16, 1936) which was about to be
inundated for the first time by the rising water of the dam. There were two
geckos in plain sight; one was shedding in a peculiar fashion—possibly burned
by the sun.

The following records of the number of variegatus variegatus found on
the road each month will give an indication of seasonal activity:

March 1
April 45
May 340
June 130
July 28
August 24
September 8
October 12

Total 588

150 SAN Drieco Society oF NATURAL History

This result is partly an index of our own activity rather than that of the
geckos; I therefore present another table covering only the Borego area in San
Diego County, and reduced to a mileage basis:

Specimens
Month Specimens Miles per 100 miles

March 0 42 0.0
April 34 330 10.3
May 274 1413 19.4
June 102 731 13.9
July 13 Dale 6.0
August 19 170 112
September i, 385 1.8
October 10 127 7.9

Total 459 3413 13.4

Even these records are not to be deemed highly accurate since there is
certain to be some carelessness in recording so common a creature, but without
doubt May is the month of greatest activity.

The best trips disclose specimens at the rate of about one specimen every
two miles (24 specimens in 49 miles, May 16, 1941, Borego area).

As to the time of night when they are most active, I have records of about

* 400 specimens of which the time of collection was noted. These are as follows:

Live
Time Specimens

3:30) tor 3:99 1
6:00 to 6:29 1
6:30 to 6:59 1
7:00 to 7:29 21
7330\to2 7259 42
8:00 to 8:29 58
8:30 to 8:59 P24
9:00 to 9:29 54
9:30 to 9:59 47
10:00 to 10:29 34
10:30 to 10:59 34
11-00 to 11:29 18
U30 to; tl-59 10
12:00 to 12:29 1
12-30) ito 12359

1:00 to 1:29 2
1-3 0Fto elo. 1
2:00%to. 2:29 0
2-30 ito. 2 2299 0
3:00 to 3:29 1
3:30 to 3:59 1

Total 3

KLAUBER—GENUS COLEONYX 151

It should be pointed out that this table does not give a true indication of
the relative time of activity, since much more collecting was done early in the
evening than later. The lizard may be just as active in mid-summer after
midnight as before; we have data on only a few trips during those hours. But
in the spring they certainly are more active early in the evening before the
desert has cooled off. The earliest specimen was recorded at 5:55 PM, October
30, 1939; the latest at 3:37 AM, August 16, 1936 (the latter by Miss Elizabeth
Sprague). It has been my experience that the first individuals may be out before
darkness is complete, but they are not most active until later. This is correctly
indicated by the table, since, while the later hours (after 11:00 PM) are not
fairly represented, the early hours are; for these trips were begun at dusk, and
if the geckos had been out they would have been recorded. We conclude that
maximum activity is reached about two hours after sunset. In the summer
this may continue all night, but in the spring most of the lizards retire as the
desert cools off.

The air temperatures at times when Coleonyx has been found may be
summarized as follows:

Temperature
Deg. F. Specimens
60-64 12
65-69 12
70-74 32
75-79 51
80-84 115
85-89 47
90-94 16
95-99 1
Total 286

It will be noted that 40 per cent were taken at temperatures of 80° to 84°
F., inclusive. As the average temperature, when all our desert night drives are
taken into consideration, was distinctly below this range, there is no question
but that a definite preference is here indicated, and that the optimum temper-
ature for this species is slightly above 80° F. The highest air temperature
noted when a specimen was collected was 96° F., and the lowest 60° F. I am
quite certain I have collected Coleonyx at temperatures below the latter figure,
but unfortunately no temperature records were then made. It is not at all
unusual to find them out in the heavy winds which often sweep across the
desert on spring nights.

I have collected this subspecies in the daytime under a variety of objects,
such as rock flakes, fallen yucca stems, boards, advertising signs, and other
items of debris. In parts of the desert where such kinds of cover are not avail-
able, they no doubt take refuge in the many mammal holes, as do most of the
other reptiles. One was found in a puddle at night after a brief summer shower.

152 SAN Disco SociETy oF NATURAL History

A captive Coleonyx was observed digging in the sand, alternately with
front and hind feet. A shedding specimen has been observed to detach and
eat patches of the loosened skin; or other captives would remove the patches
from their fellow.

They feed readily in captivity (Derbonne, 1934). Greenberg (1943) has

reported extensively on various characteristics of behavior in captivity.

Coleonyx variegatus variegatus, as seen in the field at night, often runs
with the tail curved over the back. Sometimes in the glare of headlight or
flashlight the tail will be observed to wave from side to side. The tail is easily
broken and will be dropped if the lizard be seized by it. These geckos some-
times emit faint squeaks when caught.

The food comprises insects and other arthropods including beetles, grass-
hoppers, and sowbugs.

Coleonyx variegatus is an important food element on the menu of the
nocturnal desert snakes, particularly Phyllorhynchus, which, when not large
enough to engulf a full-grown lizard consumes its tail at least. Coleonyx eggs
are also eaten by Phyllorhynchus in the spring and early summer. A red racer,
Masticophis flagellum piceus, when caught, was observed to disgorge a Coleonyx
(May 18, 1930).

Throughout the Southwest there is a widespread fear of Coleonyx among
the Mexicans; it seems often thought to be a young Gila Monster. Yet the
same fear exists where the Gila Monster does not occur. The fishermen on
Cedros Island, when shown specimens of abbotti, stated that they were “muy
malo.” Vorhies (1917, p. 367) and Strecker (1928, p. 10) comment on the
general fear of this little harmless and delicate lizard. Sanderson (1941,
p. 156) mentions a similar reaction toward Coleonyx elegans in British Hon-
duras. But fear of geckos is by no means restricted to this genus; Boulenger
(1890, p. 108), Saunders (1912, p. 1341), and M. A. Smith (1935, p. 129)
report the same apprehension with regard to the related genus Eublepharis, in
India.

Locality Records—Lower CattForNniA, Mexico: San Felipe, Colorado
River Delta, Colorado Desert. CatiForNIA: San Diego County—Beattys
Ranch, Tubbs Canyon, Borego Valley, Borego Springs, San Felipe Valley,
Banner, Scissors Crossing (also 2 mi e.), Sentenac Canyon (also foot of
canyon), Yaqui Well, The Narrows, Bensons Dry Lake (variegatus has been
collected in every one of the 19 miles of Highway Cal. 78 from Yaqui Well
via The Narrows and Bensons Dry Lake to the Imperial County line 3 mi. e.
of Bensons Dry Lake), San Felipe Wash, San Felipe (abandoned townsite) ,
La Puerta, Agua Caliente Spring, Carrizo; Imperial County—Fish Springs, Sea
View (also 2 mi. s.), Salton Sea (nw. of Sea View), Truckhaven (also 3 and
4 mi. n.), Winona, Tule Wash (at U.S. 99), San Felipe Wash (at U.S. 99),
6, 8, 9, and 12 mi. e. of Bensons Dry Lake (San Diego County), Kane
Springs Junction (U.S. 99 and Cal. 78) (also 4, 5, 8, and 9 mi. w.), Kane
Springs (also 5 mi. n., 3 mi. e., and 3 mi. se.), Harpers Well, Coyote Wells
(also 2 and 3 mi. e.), Plaster City, Seeley (also 2 mi. s.), El Centro, Calexico
(also 15 mi. e., 3 mi. w.), 6 mi. e. of Bonds Corner, 9 mi. se. of Date City,

KLAUBER—GENUS COLEONYX 153

Midway Well (intersection U.S. 80 and Cal. 98), (also 5 mi. w.), the sand
hills 14, 15, 16, and 17 mi. w. of Yuma, 4 mi. n. of Bard, Palo Verde, Winter-
haven (= Fort Yuma = Camp Yuma), Colorado Desert (type locality) ;
Riverside County—Banning, Cabazon, Snow Creek, Alameda Junction (= 29
Palms Junction) (also 1 mi. n.), Palm Springs (also 1 mi. n. and 3 mi. se.),
Indian Wells, 4 and 6 mi. e. of Indio, Coachella, Coachella Valley, Mecca
(also 4 mi. e., and 1 and 3 mi. se.), Box Canyon (near Mecca), Caleb Siding,
Hidden Springs, Shavers Well, Edom, Thousand Palms, 2% mi. sw. of
Berdoo Camp, the following canyons on the sw. slope of the Little San Bernar-
dino Mountains—Lone, Wide, Thousand Palms, Fan Hill, Pushawalla, Berdoo,
Fargo, (also on Aqueduct Road between mouths of Berdoo and Fargo
canyons)—, south base of Coxcomb Mountains, Hopkins Well, 5 mi. s. of
Vidal (San Bernardino County), east end Riverside Mountains, Blythe (also
7 mi. w. and 5 mi. n.); San Bernardino County—Colton, Slover Mountain
(near Colton), Reche Canyon (near Colton), 18 mi. w. of Victorville, Dead-
mans Point, Lucerne, Adelanto (also 2 and 4 mi. s., and 5, 10, 11, 14, and 15
mi. n.), Kramer Hills, Kramer Junction (U.S. 395 and U.S. 466) (also 2, 4,
6, 9, and 10 mi. s. and 7 mi. n.), Yucca Valley (e. of Morongo), Twentynine
Palms (also 2 and 7 mi. w.), 40 mi. nw. of Barstow, 3 mi. s. of Two Springs,
7 mi. n. of Baker, Pisgah, Cima, Wheaton Spring, Needles, Beal, Vidal;
Los Angeles County—Lovejoy Springs, halfway between Victorville (S.B.Co.)
and Pearblossom, Littlerock, 2/2 mi. ne. of Vincent; Kern County—Mojave,
Inyokern; Inyo County—Little Lake, Big Pine, Owens Valley, Emigrant
Ranger Station (Death Valley), east side of Death Valley (opposite Bennett
Wells), Mesquite Spring (Death Valley), Goler Canyon, 2 mi. n. of Sour-
dough Spring (Goler Canyon, Panamint Mountains), Bruce Canyon (Argus
Mountains), 10 mi. s. of Shoshone. Nevapa: Nye County—Amargosa River
(32 mi. ne. of Beatty), Beatty; Clark County—Colorado River 5 mi. above
California border, Jean, 6 mi. se. of Indian Springs Ranch. Arizona: Mohave
County—4 mi. se. of Hoover Mine, 26 mi. n. of Chloride, Kingman (also 12
mi. n., 3, 4, and 7 mi. nw.), Fort Mohave, Toroweap Valley (south of Tuweep,
Grand Canyon National Monument); Coconino County—Bright Angel Can-
yon, Grand Canyon National Park; Yavapai County—Cornville, McCloud
Mountains, near Yarnell, Congress Junction (also 3 mi. n., 4 mi. ne., and 3
mi. s.), 3 mi. w. of Octave, 5 mi. n. of Wickenburg (Maricopa County) ;
Maricopa County—2 mi. n. and 2 mi. nw. of Wickenburg, Wittman, Phoenix
(also 20 mi. sw.), near Mesa, Big Horn, Agua Caliente, Sentinel, Gila Bend
(also 16 mi. e.), Maricopa Mountains (30 mi. e. of Gila Bend), Gila River
(below Gillespie Dam), Bella Vista (on Gila Bend-Casa Grande Road);
Pima County—Bates Well, 2 mi. e. of Dowlings Well, Gunsight, Covered
Wells; Yuma County—Bouse, 2 mi. n. of Stoval, 5 and 10 mi. e. of Mohawk,
Pembroke, Tacna, Wellton, Wellton Mesa, 3 mi. w. of Dublin, Dome Rock
Mountains, Yuma (also 2 mi. s.), Gila River (near Yuma), Somerton, | mi. n.
of San Luis (Sonora). Sonora, Mexico: Paso MacDougall (n. of Sierra
del Pinacate), Sierra Blanca (s. of Sierra del Pinacate), Salina del Pinacate
(Bahia de Adair), between Salina del Pinacate and Salina Grande (Bahia de
Adair), Punta Penasco, and 10 mi. sw. of Sonoyta.

154 SAN Disco SociETY oF NATURAL HIsrory

Coleonyx variegatus abbotti subsp. nov.
SAN DiEGAN BANDED GECKO

1897. Coleonyx variegatus (part) Wan Denburgh, Occ. Papers Calif. Acad.
Sci., No. 5, p. 40.

Type-—No. 34,790 in the collection of L. M. Klauber. Collected in
Proctor Valley, San Diego County, California, February 28, 1942, by William
Moore. Proctor Valley runs between Jamul and Upper Otay Reservoir.

Diagnosis——A_ subspecies of Coleonyx variegatus characterized by the
retention of the banded pattern of the juveniles into the adult stage. From the
typical subspecies it may be distinguished by the presence of a narrow and
evenly-outlined nuchal light loop in the adults. It differs from utahensis,
peninsularis, and sonoriensis in having dark dorsal bands in the adults which
are approximately equal to the interspaces, instead of being much wider as in the
other three forms. It has fewer gulars in contact with the mental than slevini.
From bogerti it differs in having fewer preanal pores, in possessing a deeper
mental, and in the lack of spotting on the heads of the adults.

Description of Type—A gecko with a stubby snout and relatively short
and delicate limbs, which overlap. The tail is round. The ear openings are
elliptical and oblique. There is a distinct vertebral groove, bounded on either

side by a rounded ridge. The type is an adult male.

Except for the scales bordering the mouth, nostrils, and eyes, the head
is covered above and below with non-imbricate circular granules, uniform in
size posteriorly, but considerably enlarged on the snout and bordering the
labials. The rostral is pentagonal in shape, pointed at the top, and wider than
high. The two edges touching the nasals are longer than those engaging the
first supralabials, and are concave. The supralabials number 6-6, gradually
decreasing in size posteriorly. They are replaced by granules just forward of
the center of orbit. The infralabials visible when the mouth is closed number
8-7. The anterior are much larger than those which follow. There is a pair
of long thin prenasals, which narrowly touch the first supralabials, broadly
contact the rostral, and are slightly in contact with each other on the median
line. There are two small postnasals on either side, the upper being somewhat
larger. The lower might be considered a subnasal. There is a pair of triangular
supranasals, medianly separated from each other by 2 granules. Both upper
and lower eyelids are edged with enlarged scales which are conspicuously
serrated, especially the posterior. The upper scales number about 16; the lower
17. The last are quite small, with points directed outward. The mental is
conspicuously the largest head scale. It is slightly wider than the rostral, and
is somewhat wider than deep, the dimensions being 2.3 mm. wide by 2.1 mm.
deep. The sides are divergent anteriorly; the lower or posterior edge is semi-
circular in form. This edge is contacted by 5 gulars. The mental and the first
infralabial on each side, when taken together, are contacted by 9 gulars.

* Named for Mr. Clinton G. Abbott, Director of the San Diego Society of Natural

History, a friend, editorial guide, and scientific associate for many years.

KLAUBER—GENUS COLEONYX 155

The dorsum is covered by granules of similar size to those in the head,
there being no enlarged tubercles. On the sides the scales become somewhat
enlarged and imbricate, posteriorly first, since the granules are smallest under
the arms. On the underside the scales are imbricate and considerably larger
than those on the dorsum.

There is a group of especially enlarged scales in the preanal region; of
these, six, which are conspicuously larger than the rest, contain preanal pores.
These are not colored brown, presumably because of the date of collection; the
scales containing the pores are obtusely angular in arrangement with the point
forward; the two scales forming the apex of the angle are in contact.

The scales on the arms are mostly imbricate, except on the inner side of
the upper arm. They are largest back of the wrist. The palmar surface is
tubercular. On the inner surfaces of the digits there are rows of rectangular
transverse lamellae, of which there are 13 on the fourth finger. The claws are
retracted and barely visible; they are held between a pair of shell-like lateral
scales, with a narrow and pointed scale closing the gap above, while below it
is closed by the last of the lamellae.

The legs are covered with granular scales on the upper and inner surfaces;
they are imbricate below and are largest on the outer areas of the thighs. There
are 18 lamellae on the fourth toe.

The cloacal spurs are rather narrow and there is little if any slope to the
ridge. They are about 1 mm. in length.

The tail is covered by annular rows of subrectangular and imbricate scales
of larger size than any on the body. They are largest ventrally. At the thickest
part of the original tail (part is regenerated) there are about 29 scales in a ring.

The principal dimensions of the type, in mm., are as follows: Body length,
snout to vent 53; head length 14.5; head width 8.7; rostral to mid-orbit 6.4;
rostral to mid-ear 11.5; width of orbit 3.2; arm fully extended, measured to
end of fourth finger 18; leg fully extended, measured to end of 4th toe 24;
length of tail (regenerated) 45; distance between points of cloacal bones 6.

The head is medium brown above, the color being carried by the tips of
the granules. There is a thin, cream-colored line of horse-shoe shape extending
from below and behind each eye and above each ear to a loop on the neck.
Theze is also a short light line on the canthus rostralis on either side. The
upper eyelid edges are strongly marked with gray. The inner surface of each
eyelid is black, except for the edge scales. The rostral is brown, although
lighter in the center. The upper labials are punctated, the first, third, and
sixth being especially dark. The mental is lightly speckled, as are the lower
labials. the fifth being particularly dark. There are scattered punctations on
the gulars adjacent to the infralabials.

There is a single wide, dark ring on the neck, which laterally curves for-
watd toward the ears, thus comprising the posterior dark border of the light
postparietal or nuchal loop previously mentioned. On the dorsal surface of
the body, between limb insertions, there are 4 brown crossbars; they are slightly
wider than the interspaces, especially mid-dorsally. They fade out on the sides,

156 SAN Disco Society oF NaturaL History

the ventral surface being almost immaculate white; however, tiny punctations
are here and there discernible. The light areas between dorsal bands are also
punctated. The upper surfaces of the limbs, out to the ends of the fingers and
toes, are brown; the inferior surfaces, even of the digits, are lightly speckled.

On the tail there are only 2 rings, for the rest has been regenerated and
is spotted.

Paratypic Material—The following is a list of the paratypes which have
been available. All are from the coastal side of the mountains, or on the
divide.

Los Angeles County
LMK 2011 San Francisquito Hydroelectric Plant 2

Riverside County
LMK 2725 Moreno

San Diego County
LMK 30 Cottonwood
843 Foster
21,249 El Capitan
24,050 San Pasqual
25,303 Jacumba
27,770 Foot Agua Tibia Mountain
32,797 Pala
32,817 Mission Gorge
32,821-2 Black Mt., near La Mesa
34,666 Jacumba
34,786 Mission Gorge
SDSNH_ 16,702 Rincon
16,988 Mission Gorge
16,989 Jamul
17,012 De Luz
CAS 13,200 Poway

Northern Lower California
LMK_ 2,593 Ensenada
6,553 65 mi. se. of Tecate
24,390 San José (lat. 31°)
Cedros Island
LMK 5265-6, 27,726, 30,295; SDSNH 15,970-1; CAS 59,625, 79,875-9;
MCZ 45,721.
In addition I tentatively assign the following to this subspecies:
MVZ 12,447 Kern River Canyon, 3 mi. above the mouth, Kern County,
California
USNM 62,247 Calmalli, central Lower California, Mexico.

Stanford Nos. 15-18, San Jacinto, Riverside County, California, although
their condition is such as not to permit a final decision, probably should be
considered abbotti. Specimens from the vicinity of Colton, San Bernardino

KLAUBER—GENUS COLEONYX 157,

County, although from the coastal side of the mountains, seem nearer
variegatus variegatus.

Range.—Coastal and cismontane southern California and northern Lower
California from the San Gabriel Mountains, Los Angeles County, south to the
west slope of the San Pedro Martir Mountains of Lower California, Mexico.
Also Cedros Island off the Pacific Coast of Lower California.

Morphology.—This is a smaller and seemingly slimmer gecko than those
typical of the desert areas to the east. The longest specimen is 68 mm.; this
is a particularly light female from De Luz, San Diego County; the next
longest is a female 61 mm. in length from the foot of Agua Tibia Mountain.
The longest male measures 56 mm. I should estimate that the average adult
body length of abbotti is about 15 mm. below that of variegatus variegatus.

The tails, when complete and original, which is seldom the case, are
slightly longer than the bodies. The excess tail length is more pronounced in
the young specimens.

The preanal pores in the males vary from 5 to 7, the counts being as
follows: 5 (2), 6 (6), 7 (6); average 6.28.

The cloacal bones are rather narrow and sharp. In specimens from Cedros
Island they are wider.

Scalation—The rostral is wider than high. The prenasals are long and
slim; they touch the first supralabials, and are in contact on the median line in
all except one specimen from Cedros Island. The supranasals are separated by
from one to three granules, the counts being 1 (1), 2 (19), 3 (17); mean 2.43.
The supranasals are larger than the postnasals. Of the latter, there are two on
each side, the upper being the larger. The supralabials number from 6 to 9;
they average lower in the Cedros Island specimens than those from the main-

land.

In the mainland specimens the scales bordering the upper eyelids are large
and serrated; the scales of the lower lids are not so conspicuous. The scales
bordering the eyelids in the Cedros Island specimens are smaller and less
pointed.

As is the case in the other subspecies, the circular granules on the snout
are larger than elsewhere on the head, this being especially true of those border-
ing the supralabials.

Although the mental is sometimes only as wide as the rostral, it is usually
conspicuously wider. It is almost as deep as wide, the depth averaging about
9 of the width. The lower edge is a circular arc of moderate radius. The
gulars contacting the mental number as follows: 5 (11), 6 (17), 7 (6), 8 (2),
9 (1); mean 5.81. Those contacting the mental, together with the first infra-
labial on either side, number as follows: 8 (1), 9 (13), 10 (14), 11 (5),
12); 1342) s:meanr9.97.

Pattern and Color—This subspecies is characterized by the slight onto-
genetic change in pattern which it undergoes and by an abundance of pigment
evidenced by punctations scattered over the body.

158 San Dreco Society oF NATURAL History

The head continues brown throughout life, with almost no spotting or
mottling. The nuchal light loop remains clear and evenly outlined, as do also
the light bars on the canthus. The body bands, exclusive of the dark band on
the neck, number 4 in all mainland specimens available to me; they are approxi-
mately equal to the interspaces in width. Spots are occasionally present in the
interspaces when the adult stage is reached, this being especially the case in
the Cedros Island individuals. A close examination of the interspaces discloses
the presence of many fine dots on the scale tips, thus reducing the contrast
between the brown and cream areas; however the bars remain even-edged in
the adults. The upper surfaces of the limbs are also speckled, so that they appear
grayish. Some of this speckling is evident on the belly, on the ventral surfaces
of the limbs, and on the digits; and even the claws themselves are frequently
brown in color.

The rostral is always dark, although the center may be lighter than the
edges. The supralabials are heavily punctated. The mental may be clear or
somewhat mottled. The infralabials are usually darkened and the adjacent gulars
are irregularly speckled.

Where the tail has not been broken, the rings number from 9 to 12.
Usually only the posterior are ventrally complete. In the mainland specimens
the tail rings are little if any darker than the body bars. Regenerated tails are
speckled, spotted, or irregularly barred.

The Cedros Island specimens differ somewhat from the mainland in
pattern. The body bars tend to be wider than the interspaces. One specimen
has 5 bars on one side and 4 on the other, while two others have 5 dorsally,
but 4 laterally. There is a greater tendency for spots to develop between bars
in the adults. The tail rings are darker than the body bars and are more often
complete ventrally. There is less marking on the limbs and digits, both on
the upper and lower surfaces, and the claws are light. One specimen has a
pattern reminiscent of peninsularis; and altogether these island individuals are
not so sharply differentiated from either peninsularis or variegatus variegatus
as are those from the mainland.

In a live adult from Cedros Island the dorsal -bands were Saccardo’s
Umber and the interspaces Ecru Olive, by Ridgway’s Color Standards.

Relationships with Other Subspecies—Abbotti undoubtedly intergrades
with variegatus variegatus in some of the mountain passes, there being a number
low enough for these geckos to range through. Intergradation probably takes
place toward the coastal side, as specimens from the eastern slopes of the
mountains seem to be variegatus variegatus; in fact, in some areas the desert
form has encroached on the cismontane terrain, as is the case in the vicinity of
Colton, San Bernardino County, where the population is nearer variegatus
variegatus than abbotti.

At the southerly end of its range abbotti may eventually be found to
intergrade with peninsularis, or another desert subspecies occupying the Vizcaino
Desert may be interposed.

I have tentatively placed in the abbotti category a gecko from Calmalli,
central Lower California (USNM 62,247). This specimen has been damaged

KLAUBER—GENUS COLEONYX 159

so that the scale arrangements on the head cannot be determined. The body
bars equal the interspaces in width, thus favoring abbotti rather than peninsularis
in this character.

Another specimen of doubtful allocation is MVZ 12,447 from the Kern
River Canyon, 3 miles above its mouth, a locality probably not over 10 miles
eastward from Bakersfield. This is the only specimen known to me from the
San Joaquin Valley side of the Sierra Nevada. The markings are those of
abbotti, but the gulars are much enlarged, there being only 4 in contact with
the mental. Were future specimens from this area to show similarly low counts,
a subspecific segregation might be warranted.

Field Notes——The dark color of this gecko renders it diffcult to see at
night, which probably accounts for its being so rare in collections, compared to
variegatus variegatus, which is so easily collected by driving on the desert roads
at night. I think that abbotti, although having a considerable range, is not so
common anywhere as is the type subspecies in the desert.

Most of the specimens in collections were found under stones, rock flakes,
or cap rocks. The latter have been the source of a number of San Diego County
individuals. Some were found while searching for Xantusia henshawi, although
they are much rarer than the granite night lizard, at least in these retreats.
For example, on March 20, 1926, a single gecko was found under a granite
slab at Jamul. This was a cold day when lizards were almost dormant. Fifteen
Xantusia henshawi were collected under this and other slabs and flakes.

On April 20, 1927, a Coleonyx y. abbotti was found under a cap rock on
a granite boulder at San Pasqual, and three days later another was collected
under the same conditions near Foster. Another was found under a cap rock
at Moreno, Riverside County, May 25, 1930. The preference for cap rocks
seems to indicate that the geckos cannot stand the temperature changes which
Xantusia henshawi must sustain under the thin flakes below which it is so often
found. Of course, both lizards may be common within the deeper crevices to
which the collector cannot gain access.

While not entirely restricted to areas where granite is available, abbotti
seems to prefer such territories. L. H. Cook, collecting on the south end of
Cedros Island, found four specimens by turning over rocks in the daytime.

Locality Records ——Catirornia: Los Angeles County—San Francisquito
Hydroelectric Plant No. 2, Owensmouth; Riverside County—Moreno, San
Jacinto; San Diego County—De Luz, foot of Agua Tibia Mt., Pala, Rincon,
Escondido, Lake Hodges, San Pasqual, Poway, Foster, El Capitan, Harbison
Canyon, Mission Valley, Alvarado Canyon, Mission Gorge, Black Mt. (near
La Mesa), Jamul, Proctor Valley (type locality), Dulzura, Cottonwood (=
Barrett P.O.), Jacumba. Lower CatirorniA, Mexico: Rodriguez Dam,
Ensenada, 65 mi. se. of Tecate, San José (lat. 31°), Cedros Island.

Additional records tentatively assigned to this subspecies: Kern River
Canyon, 3 mi. above the mouth, Kern County, California; Calmalli, central
Lower California, Mexico.

160 SAN Dreco Society oF NAturRAL History

Coleonyx variegatus peninsularis subsp. nov.
San Lucan BANDED GECKO

Type.—No. 37,210 in the collection of the Museum of Comparative
Zoology, Harvard University. Collected at La Paz, Lower California, Mexico,
by Miguel L. Cornejo, Jr., 1933.

Diagnosis—A subspecies of Coleonyx variegatus distinguished from all
other subspecies, except slevini, utahensis, and sonoriensis, by the wide transverse
dark bars on the body as compared to the narrow light interspaces. From slevini
it may be segregated by its having a higher number of scales in contact with
the mental. From sonoriensis it may be distinguished by the following pattern
differences: peninsularis has less superimposed spotting on the head and body
blotches, the anterior ends of the nuchal light loops are higher, and it has a
pair of prominent canthal light lines, with an unspotted brown area between,
not possessed by sonoriensis. The Cape form lacks the mid-dorsal light line
characteristic of most Sonoran specimens. From utahensis it differs in having
even, rather than highly irregular, edges of the dorsal bands, and in having
straighter light canthal lines.

Description of Type——An adult male. Except for the scales bordering the
mouth, nostrils, and eyes, the head is covered above and below with non-
imbricate, circular granules, approaching uniformity in size. They are slightly
enlarged on the snout and bordering the supralabials. The rostral is pentagonal
in shape and is wider than high. The two edges touching the nasals, which
are somewhat concave, are longer than those engaging the first supralabials.
The supralabials number 7-7 and decrease in size posteriorly. The infralabials
also number 7-7. There is a pair of prenasals, lunar in shape, which contact
the first supralabials at a point, broadly contact the rostral, and are in contact
with each other on the median line. There are two postnasals, of which the
upper is the larger. There is a pair of supranasals considerably larger than
the postnasals; they are separated from each other by 3 granular scales. Both
upper and lower eyelids are edged with enlarged scales, which are somewhat
serrated. The mental is the largest head scale. It is wider than the rostral and
is slightly wider than deep, the dimensions being about 1.9 by 1.7 mm. The
sides are nearly parallel; the lower or posterior edge has a rather flat curve.
This edge is contacted by 6 gulars. The mental and the first infralabial on
each side combined are contacted by 10 of the small circular gulars.

The dorsum is covered by granules of a size similar to those in the head,
there being no enlarged tubercles. On the sides the scales become somewhat
enlarged and imbricate. On the ventrum the scales are imbricate and much
larger than those on the dorsal surface.

The scales in the preanal region are further enlarged and include a row
carrying 8 brown preanal pores, obtusely angular in arrangement with the point
forward. The scales containing the two middle pores are contiguous.

The scales on the arms are mostly imbricate. On the lower surfaces of the
digits there are series of rectangular transverse lamellae.

KLAUBER—GENUS COLEONYX 161

The legs are covered with granular scales on the upper surfaces but are
imbricate below. They are largest on the thighs. There are transverse lamellae
on the toes. The claws on both fingers and toes are fine and delicate. They
are retractile between a pair of laterals and a terminal.

There is a pair of cloacal bones which are broad and ridgedy one being
conspicuously notched. They are about 1 mm. in length.

The tail is covered by annular rows of subrectangular and imbricate scales,
which are largest ventrally. At the widest part of the tail there are about 31
scales per ring.

The principal dimensions, in mm., are as follows: Body length, snout to
vent 42; head length 11.5; head width 7.3; rostral to mid-orbit 5; rostral to
mid-ear 9.5; width of orbit 2.5. The tail is incomplete.

The head is brown above and cream-colored below. There is a thin light
line of horse-shoe shape extending from behind and slightly below each eye
and above the ears to a loop on the neck. There is also a well-defined light
line along the canthus rostralis on either side. The inner surface of each eyelid
is black, except for the edge scales. The rostral is light in the center but is
brown along the upper border. The mental is unmarked. The labials, both
upper and lower, are lightly punctated with gray at various points. Back of
the enlarged labial scales the granules which border the mouth are punctated
with gray-brown.

There is a single wide, dark ring on the neck, which laterally curves forward
across the ear openings, thus comprising the posterior dark border of the light
nuchal loop. On the dorsal surface of the bedy. between limb insertions there
are 4 transverse brown bars, each of which is darker posteriorly. The last is
broken at one side by an irregular light mark. The middle bars have a width of
about 4 times the interspaces. The ventral surface is immaculate. The upper
surfaces of the limbs, out to the ends of the fingers and toes, are stippled with
brown.

Summary of Paratypes—The definition of this subspecies is handicapped
by lack of adequate material. There are only two paratypes: USNM 67.382.
a faded juvenile, probably a male, from between Loreto and Comonds; and
MVZ 26 989. an adult female from San José del Cabo. USNM 62.247 from
Calalli is definitely not of this form. One cannot tell from his brief description
whether the two specimens reported by Mocquard (1899, p. 300) from Santa
Rosalia and Mulegé should be assigned to peninsularis or to one of the two
subspecies inhabiting the northern half of the peninsula of Lower California.
Therefore, the known range of peninsularis may be described as the eastern side
of the peninsula of Lower California from Lat. 26° N. southward.

Peninsularis is probably a small subspecies. The type, although only 42
mm. from snout to vent, has well developed preanal pores and cloacal bones.
MVZ 26.989 measures 49 mm. and contains eggs. The third specimen is a
juvenile only 29 mm. in body length. None of the three has a complete tail.

The three specimens differ in no important itern of scalation. The enlarged
labials seem to number 7 in all cases. There are 6 gulars in contact with the

162 San Dreco Soctety oF NATURAL HIstory

mental, and 10 in contact with mental and first supralabials. The prenasals
meet in the median line in all three. In two specimens 2 granules separate the
supranasals, while the type has 3 interposed.

In pattern they are also much alike. The dorsal blotches are wide, separated
by comparatively narrow light lines. The blotches are darkest posteriorly and
are somewhat pointed toward the rear. There is some evidence of a mid-dorsal

light line in MVZ 26,989.
On the head the light nuchal loop is clearly defined, ending just below the

eye on each side. The canthal light lines are clear and even.

Relationships with Other Subspecies—Peninsularis most nearly resembles
sonoriensis in all characters, the principal divergence between them being in
the adult pattern. It is also close to slevini, from which it differs in having a
higher number of gulars in contact with the mental.

Locality Records——Known only from La Paz, San José del Cabo, and the
trail between Comondiu and Loreto, southern Lower California, Mexico. The
species occurs at Mulegé and Santa Rosalia but it is not certain whether this
subspecies is the one found there.

Coleonyx variegatus sonoriensis subsp. nov.
SONORAN BANDED GECKO

1897. Coleonyx variegatus Van Denburgh, Proc. Acad. Nat. Sci. Phila., Vol
49, p. 460.
Type—No. 72,140 in the collection of the Museum of Zoology, Universi-
ty of Michigan. Collected by Morrow J. Allen, June 25-29, 1932, 5 miles

southeast of Hermosillo, Sonora, Mexico.

Diagnosis ——A subspecies of Coleonyx variegatus characterized by wide
dorsal bands, much wider than the interspaces, and by the frequent presence
of a vertebral light stripe splitting the bands into paired rectangles. It differs
from slevini in having a greater number of scales in contact with the mental,
and from peninsularis in the frequent presence of the mid-dorsal line and in
the head-spotting of the adults. From variegatus variegatus it is distinguished
by its adult retention of the postparietal light loop and the virtually unbroken
body bands, except for the vertebral stripe. From utahensis, abbotti and bogerti
it may be segregated by the relative widths of body bars and interspaces.

Description of Type—An adult male. The limbs are rather short and
delicate; they overlap when adpressed. The tail is round; the terminal portion
is regenerated. The eyelids are functional; the ear openings are oblique.

Except for the scales bordering the mouth, nostrils, and eyes, the head is
covered above and below with non-imbricate, circular granules, rather uniform
in size. They are slightly enlarged on the snout and somewhat more so where
they contact the supralabial scutes. The rostral is pentagonal in shape and is
considerably wider than high. The two edges touching the nasals are longer
than those engaging the first supralabials; these edges are somewhat concave.
The supralabials number 8-7 and decrease in size posteriorly; this count is

KLAUBER—GENUS COLEONYX 163

discontinued where they are no longer conspicuously larger than the adjacent
granules, which is just anterior to the middle of the eye. The infralabials
(using the same limitations) are 8-8. The infralabials also decrease in size
posteriorly, those touching the mental being much the largest of the series.
There is a pair of prenasals, lunar in shape, which touch the first supralabials,
and broadly contact the rostral; they are separated from each other by a pair
of granules in tandem. There are two small postnasals on each side, and a pair
of subtriangular supranasals, separated from each other by 3 granules. Both
upper and lower eyelids are edged with enlarged scales which are very slightly
pointed at the outer edges. The upper scales number about 19, the lower 21.
The mental is conspicuously the largest head scale. It is slightly wider than
the rostral and is somewhat wider than deep, the dimensions being 2.4 mm. by
2.0 mm. The sides converge posteriorly and the lower edge is a circular arc.
This edge is contacted by 6 granules. Twelve granules touch the mental and
the first infralabial on either side, when taken together.

The dorsum is covered by granules of a size similar to those in the head,
there being no enlarged tubercles. On the lower sides the scales become some-
what enlarged and imbricate, posteriorly first, since the granules are smallest
under the arms. On the belly the scales are imbricate and definitely larger
than those on the dorsum; they increase in size toward the mid-ventral line,
and to a small extent posteriorly.

The scales in the preanal region are further enlarged and include a row
with conspicuous, brown preanal pores. The latter total 6; the scales containing
them are obtusely angular in arrangement with the point forward. Posteriorly
there is a rapid decrease in scale size toward the vent.

The scales on the arms are mostly imbricate, except on the lower side of
the upper arm. They are largest back of the wrist. The palmar surface is
tubercular. On the lower surfaces of the digits there are series of rectangular,
transverse and strongly imbricate lamellae. The claws are extremely delicate.

The legs are covered with granular scales on the upper surfaces and
imbricate below. They are largest on the anterior upper areas of the thighs.
The scales on the soles of the feet are tubercular. There are imbricate lamellae
on the underside of the toes. The claws are similar to those on the fingers.
They are sheathed between a pair of shell-like laterals, capped by a somewhat
shorter terminal. Normally, in preserved specimens, they protrude well beyond
the sheath.

The cloacal bones are about 1.1 mm. in length. They are slender and
pointed.

The tail is covered by annular rows of subrectangular and imbricate scales
of larger size than any on the body. They are largest ventrally. At the widest
part of the tail there are about 38 scales in a ring.

The principal dimensions in mm. are as follows: Body length, snout to
vent 53; head length 15.5; head width 9; rostral to mid-orbit 6.4; rostral to
mid-ear 11.7; width of orbit 3; arm fully extended, measured to end of 4th

164 San Disco Society oF NaturaAL History

finger 16; leg fully extended, measured to end of 4th toe 22; length of tail
(partly regenerated) 45; distance between points of cloacal bones 6.8.

The head is light-brown above, conspicuously spotted with darker. It is
cream-colored below. There is a thin cream-colored line of horse-shoe shape
extending from the supralabials to a loop on the neck. There is a light V-shaped
mark in front of the eyes, with the point between them. The upper eyelids are
faintly punctated posteriorly. The inner surface of each eyelid is black, except
for the edge scales. The rostral is light in the center but brown on either edge.
The mental is unmarked. There are dark spots on the first, fourth, and last
supralabials; and on the third, and last infralabials.

On the neck there is a pair of brown rhomboidal blotches bounded by
narrow light lines. On the dorsal surface of the body, between limb insertions.
there are 4 transverse brown bands, all of which are split centrally by a light
mid-dorsal line which runs from the postparietal loop to the base of the tail.
Laterally, the bands terminate in a spotted area; they are somewhat darker on
the edges, and show faint spots within. The ventral surface is immaculate cream-
colored. The upper surfaces of the limbs, out to the ends of the fingers and
toes, are spotted and punctated with brown.

On the tail there are 4 rings, followed by a spotted regenerated section.

Paratypic Material —There is available a series of 11 topotypes (MZUM
72,141-51), and 6 other specimens (MZUM 72,152a-f) from 15 miles south-
“east of Hermosillo. Dr. E. H. Taylor has kindly loaned me the following
EHT-HMS material: Nos. 10,538-41, 10,561-65, from 5 miles southwest of
Hermosillo, and Nos. 10,542-60 from La Posa, 10 miles north of Guaymas.
Miscellaneous specimens include CAS 53,410 from Tepoca Bay and MVZ
20,839 from Ensenada del Perros, Tiburén Island, Gulf of California. A
specimen (USNM 16,808) recorded from Nogales, Arizona, should probably
be allocated to this subspecies; yet knowing how frequently early specimens were
attributed to the point of shipment rather than that of collection, I hesitate
to include this subspecies in the fauna of the United States until more material
from around the U. S. boundary shall become available. Other specimens
available from between Nogales and Tucson are not sonoriensis. Specimens
from northwestern Sonora, from Punta Pefasco and beyond. while showing
some trends toward sonoriensis, are clearly nearer variegatus variegatus.

Range-—West-central Sonora, Mexico, from Tepoca Bay to Guaymas
and inland to Hermosillo; also Tiburén Island in the Gulf.

Morphology.—Sonoriensis is rather small in size compared with variegatus
variegatus. The largest specimen out of about 40 adults, a female, measures
58 mm., rostral to vent. The tail is incomplete. The largest male measures
>5 mm. Few specimens have complete, original tails. Where they are complete,
the tails approximately equal the body in length. Aside from this matter of
ultimate size, I find no consistent difference between sonoriensis and variegatus
variegatus in head or body shape. There seems to be a considerable individual
variation in the shape of the snout and the ear opening. The preanal pores in

KLAUBER—GENUS COLEONYX 165

the males are distributed as follows, the figure in parentheses indicating the
number of specimens: 4 (1), 5 (2), 6 (7), 7 (3), 8 (3), 9 (1); mean 6.61.
The cloacal bones are somewhat narrow.

Scalation—The pentagonal rostral is wider than high. The prenasals are
long and lunar in shape. They usually touch the first supralabials but occasion-
ally a granule is interposed. The postnasals are small; sometimes they are two
in number, sometimes three; the upper is the largest. The supranasals are
larger than the postnasals. The prenasals are not in contact in 2 out of 48
specimens, or 4.17 per cent. The separation of the supranasals is effected by
the following numbers of granules (counting the minimum number bridging
the gap): 1 (2), 2 (12), 3 (31), 4 (3); mean 2.73. The visible enlarged
supralabials usually number 7 or 8, but sometimes 6 or 9, or even 10. The
scales on the edges of the eyelids are somewhat less pointed than in variegatus
variegatus. The granules on the top of the snout are larger than those on top
of the head back of the eyes; the largest, however, are on the sides immediately
above the supralabials. The mental is usually somewhat wider than the rostral
and is slightly wider than deep. The sides are posteriorly convergent and the
bottom is a circular arc. The number of gulars contacting the mental are as
follows:=) = (9), 16-.(12) } 7. (14),,.8 (10); 9 (2). 10) (1); ‘mean’6.73: “The
numbers of gulars contacting the mental and the first infralabials, when taken
together, are as follows: 9 (8), 10 (11), 11 (14), 12 (10), 13 (2), 14 (1),
15 (0), 16 (1); mean 10.83. The gulars contacting the infralabials are often

larger than those touching the mental.

Pattern and Color—The outstanding pattern characteristics of this sub-
species are the wide transverse body bands (as compared to the narrow inter-
spaces) and the mid-dorsal light line, which is present, at least to a slight
degree, in nearly every adult. In this species an ontogenetic pattern change is
evident, as it is in all subspecies; in this case it involves a lightening of the
dorsal color, and the superposition of brown spots on the light-brown of the
head and body blotches. But the main blotches, instead of breaking up into
paired, transverse bands, or into irregular blotches or spots, as in variegatus
variegatus or bogerti, are retained almost without change in relative size. The
longitudinal borders are usually curved, but are not serrated, or highly irregular,
as in utahensis. The nuchal light horse-shoe is retained without much loss of
definition, which is quite different from variegatus variegatus. Only rarely are
spots developed in the light interspaces, in the manner so characteristic of
variegatus variegatus, bogerti and utahensis. The greatest ontogenetic change is
the development of the mid-dorsal light line. The transverse body bars in this
subspecies are seldom other than 4. Out of 48 specimens one has 4 on one
side and 3 on the other; another has 4 in the middle on the back, but 5 on
either side; the other 46 have 4 bars.

The rostral is always mottled and there are conspicuous collections of
punctations on several of the supralabials; these are often restricted to single
scales. The infralabials are less marked. The mental is often spotted, usually
on the upper edge or laterally. In this it differs from slevini. The rings on

166 SAN Dreco Society oF NATURAL HIstTory

original tails are approximately equal to the interspaces. They number about
11 to 14. The last few are complete ventrally. Regenerated tails are usually
spotted.

Of the three available specimens which are not from either the vicinity of
Guaymas or Hermosillo, MVZ 20,839 from Ensenada del Perros, south end
of Tiburon Island, seems to be without material difference from the others.
The vertebral stripe is quite clear. CAS 53,410 from Tepoca Bay has definite
variegatus variegatus tendencies, the blotches bee more broken up than in most
sonoriensis; thus, although I sessile: it in the latter category, it approaches
intergradation. MVZ 10,165 from Sierra Alamo, 30 miles west of Caborca,

also is an intergrade, but has been placed in the sonoriensis category.

Relationships with Other Subspecies——Sonoriensis most closely resembles
peninsularis, from which it has only moderate pattern differences. Nevertheless,
there cannot now be intergradation between the two, as the Gulf of California
divides their ranges. Although good series of sonoriensis are available from two
localities, we lack specimens from both the north and northwest. Intergradation
with variegatus variegatus in northwestern Sonora is strongly indicated; contact
with bogerti in the north is much less certain. Vertebral light lines in some
specimens of bogerti may suggest a close relationship.

In the nature of the superimposed spotting on head and body, and in the
shape of the cloacal bones, an affinity with brevis is suggested. However, there
is a wide gap between the ranges of the two forms, as far as at present known.
Although sonoriensis is to be expected to the south of Guaymas, there is no
indication of any trend toward fasciatus, for the latter is different in many
important characters, as Taylor (1935, p. 203) has pointed out.

Field Notes—‘‘Taken only at night on the open desert with the exception
of two or three specimens found under stones on a hillside.” Morrow J. Allen
(1933. p. 4).

“Thirty-eight specimens of Coleonyx variegatus (Baird) were collected in
Sonora, and with one exception, in which the specimen was found hidden under
a rock, they were discovered at night running about over gravelly soil near the
beach and in the mountains . . . The specimens of this species usually run with
the tail lifted, often curved over the back. The bright light from my lantern
tended to bewilder them, and they were caught at night with little difficulty.”
E. H. Taylor (1936, p: 479).

Locality Records—This subspecies has been collected at the following
points, all in Sonora, Mexico: In the vicinity of Hermosillo (5 miles south-
west, 5 miles southeast [type locality], and 15 miles southeast); at and near
La Posa (10 miles northwest of Guaymas); Tepoca Bay; Sierra Alamo (30
miles west of Caborca); and Ensenada del Perros at the south end of Tiburon
Island. It has also been reported from San Miguel de Horcasitas, which is
some 70 miles slightly east of north from Hermosillo (Wan Denburgh, 1897,
p. 460). A specimen of this subspecies said to be from Nogales, Arizona, is
to be considered of doubtful origin until verified by additional material.

KLAUBER—GENUS COLEONYX 167

Coleonyx variegatus slevini’ subsp. nov.
SANTA INEZ ISLAND BANDED GECKO

1922. Coleonyx variegatus (part) Wan Denburgh, Occ. Papers Calif. Acad.
Sci., No. 10, vol. 1, p. 58.

Type—No. 51,697 in the collection of the California Academy of Sciences.
Collected on South Santa Inez Island (Lat. 27° N.) on the Gulf of California
coast of Lower California, Mexico, on May 13, 1921, by Joseph R. Slevin.

Diagnosis—A_ subspecies of Coleonyx variegatus characterized by the
enlargement of the gular scales bordering the mental, so that this subspecies
averages fewer scales making this contact than any other. It also usually has
5 instead of 4 body bars as in the other subspecies.

Description of Type—An adult male. This lizard has relatively short
legs, although when adpressed they overlap. The tail is round. The snout is
somewhat blunt. The eyelids are functional, and the ear openings evident.

Except for the scales bordering the mouth, nostrils, and eyes, the head
is covered above and below with non-imbricate, hemispherical granules, ex-
ceedingly small but rather uniform in size. They are slightly enlarged on the
snout and are conspicuously so where they border the labials. The rostral is
pentagonal in shape and is wider than high; the two edges touching the nasals
are longer than those engaging the first supralabials. The enlarged supralabials
number 7-7 and decrease in size posteriorly; this count is discontinued where
the labials become no larger than the adjacent granules, which is just forward
of the center of the orbit, and refers to those which can be seen with the mouth
closed. The infralabials also decrease in size posteriorly, and to a more marked
extent than the upper labials, those touching the mental being much the largest
of the series. Those exteriorly visible number 7-8. The nostril is below the
line of the canthus rostralis. There is a pair of prenasals, triangular in shape,
which contact the first supralabials, the rostral, and are broadly in contact with
each other on the median line. There are two small postnasals on either side,
and a pair of subcircular supranasals, separated from each other by 3 granules.
The post- and supranasals are smaller than the prenasals. Both upper and lower
eyelids are edged with serrated scales. The upper scales number about 17; the
lower 15, but there seems to be an abnormal gap at the end of the series. The
mental is conspicuously the largest head scale. It is somewhat narrower than
the rostral but is deeper; its dimensions are 2.1 mm. wide by 2.0 mm. deep.
The sides are nearly parallel; the lower or posterior edge is semicircular in
form. This edge is contacted by 4 enlarged round gulars. The mental and the
first infralabial on each side are contacted by a total of 8 scales. The scales
contacting all infralabials are conspicuously ee compared to those covering
the central gular area.

The dorsum is covered by granules of similar size to those on top of the
head. There are no enlarged tubercles. On the sides the scales become somewhat

* Named for my good friend Mr. Joseph R. Slevin, Curator of Reptiles, California
Academy of Sciences, San Francisco, to whom I have been greatly indebted for many
favors and courtesies during the past twenty years.

168 SAN Dieco Society oF NAatTuRAL History

enlarged and imbricate, posteriorly first, since the granules are smallest under
the arms. On the ventrum the scales are imbricate and definitely larger than
those on the dorsum. They are subcircular in shape and increase in size
posteriorly.

The scales in the preanal region are further enlarged and include a row
with conspicuous, brown preanal pores. The latter total 6; the scales containing
them are obtusely angular in arrangement with the point forward. The two
middle scales are very closely in contact so that the pores touch.

The scales on the arms are mostly imbricate, except on the outer part of
the forearm and the lower side of the upper arm. They are largest on the outer
forearm. The lower surfaces of the digits are covered by rectangular transverse
lamellae, which form corrugated ridges. The claws are extremely delicate, and
are held between a pair of shell-like lateral scales, capped by a third.

The legs are covered with granules on the inner surfaces, but with imbricate
scales outwardly and below. The claws on the toes are fine and delicate, similar
to those on the fingers. There are 21 lamellae on the fourth toe.

There is a spurlike cloacal bone protruding laterally and upward on either
side just posterior to the vent. These bones are about 1.2 mm. in length. At
the top they are ridged rather than conical.

Most of the tail has been lost, and the regenerated portion is quite short.

The original part is covered by annular rows of subrectangular and imbricate
_scales. They are largest ventrally. At the thickest part of the tail there are about
40 scales in a ring.

The principal dimensions in mm. are as follows: Body length, snout to
vent, 56; head length 15; head width 10; rostral to mid-orbit 7.5; rostral to
mid-ear 12; width of orbit 3.6; arm fully extended, measured to end of 4th
finger 17.5; leg fully extended, measured to end of 4th toe 23; distance between
points of cloacal bones 9.

The head is mottled light-brown above and cream-colored below. There
is a thin and rather irregular cream-colored line of horse-shoe shape extending
from below each eye and above each ear to a loop on the neck. It is interrupted
on each side of the neck in a manner characteristic of this subspecies. There
is also a short light mark on the canthus rostralis on either side. The bordering
scales of the upper eyelids are marked with gray, particularly posteriorly. The
inner surface of each eyelid is black, except for the edge scales. The rostral
is light in the center but has a dark spot on either side. The mental is unmarked.
The upper labials are punctated with brown, but not uniformly, some being
dark, others almost immaculate. The posterior infralabials are slightly stippled.

There is a single wide, dark transverse band on the neck, which laterally
curves forward under the ears, thus comprising the posterior dark border of
the light postparietal or nuchal loop previously mentioned. On the dorsal
surface of the body, between limb insertions, there are the mottled and irregular
remnants of 5 transverse brown rings, which are much wider than the inter-
spaces. There are rows of brown dots in the interspaces, thus increasing the
mottled effect. The ventral surface is cream-colored and immaculate. The

KLAUBER—GENUS COLEONYX 169

upper surfaces of the limbs, out to the ends of the fingers and toes, are punc-
tated with brown.

Summary of Paratypes.—In addition to the type, 19 other specimens, CAS
51,698-51,716, are available from South Santa Inez Island. In the following
summary, where statistics are given, the type is included, thus making a total
of 20.

In the paratypic series there are 10 adults exceeding 50 mm., snout to vent,
and 4 young adults in the 40-50 mm. range. Only one specimen out of 20
has a complete tail. This is a higher ratio of loss than is customary, even in this
species with its notably fragile tail, so it is presumed that this subspecies has
some active enemy. Some of the juveniles probably lost their tails in the course
of capture. The specimen with a complete tail (CAS 51.712) measures 45 mm.
in body length, the tail being 41 mm. The longest individual, a female, meas-
ures 60 mm.; the shortest is 33 mm. The largest male measures 57 mm.

Of the males sufficiently adult to have active preanal pores, 1 has 5 pores, 5
have 6, and 2 have 7; average 6.13. The mental varies in width from slightly
less than the rostral to considerably wider. The mental is usually about as long
as wide; in some the sides are parallel, others narrow posteriorly. The posterior
edge is curved in a relatively flat arc. The gulars in contact with the mental
vary from 3 to 5; average 3.95. This is lower than in any other subspecies. The
gulars in contact with the mental and first infralabials (taken together) vary
from 7 to 9; average 7.90. The supranasals are separated bv either two or
three granules; average 2.4. No specimen has the prenasals separated; the
median contact is broader than in most other subspecies. The visible, enlarged
supralabials number from 6 to 8 and the infralabials the same.

In pattern this subspecies is characterized by the irregularity of the trans-
verse body bands, even in the juveniles, for they are often broken or branched
so that there are more on one side than the cther. Out of 20 specimens only
3 have 4 bands; 7 others have 4 on one side and 5 on the other, while 10 have
5 bands, a much higher proportion with this number than any other subspecies.
The only specimen with a complete tail has 13 dark bars thereon, of which the
last 5 are complete ventrally.

In the young the body bars are slightly wider than the interspaces dorsally,
and each bar is darker posteriorly. They are not even-edged. They narrow
laterally, but at their termini on the sides they tend to spread into thin lateral
lines, usually with a forward projection. The body marks are much lighter
than the tail bars.

As the lizards age, the dorsal marks become lighter and widen at the
expense of the interspaces. The cross bands become somewhat mottled, and a
series of spots appears in the interspaces, so that in the largest specimens the
cross bands almost lose their identity, and the general effect is of a mottled
surface. A vertebral light stripe is sometimes in evidence anteriorly.

The top of the head is brown and unicolor in the young and somewhat
mottled or spotted in the adults; the spots, when present, are superimposed
on the normal brown. The light postparietal loop, so characteristic of the
species, remains narrow and well-defined, even in the largest of the adults;

170 San Disco Society oF Natura History

however, it has a peculiar wavy irregularity and often has a short gap or two
in the vicinity of the ear. Sometimes the light line, which comprises the posterior
edge of the dark neck band, curves forward and almost joins the postparietal
loop.

The rostral is usually dark, particularly at the sides. The upper labials are
alternately light and dark, although the spotting does not rigidly follow scale
edges, nor is the pattern regular. The mental is clear. The lower labials
are spotted, although less so than the supralabials. These spots often extend
onto the adjacent gulars.

The cloacal bones are rather broad, with the points at the anterior end of
the ridge, and are usually without notches.

To the subspecies slevini I also tentatively assign two specimens (CAS
51,463-4) from the larger nearby island of San Marcos. Both of these speci-
mens are young males. One has 6 preanal pores, the other 7. The prenasals
are in contact and the supranasals separated by 3 granules. One has 3 gulars
in contact with the mental, the other 5. The body rings are irregular but are
wider and posteriorly more pointed than in typical slevini, thus resembling
peninsularis. The body bands are 4 in one specimen and 4-5 in the other.

Relationship with Other Subspecies——Slevini is most nearly related to
peninsularis and sonoriensis, as would be expected geographically. It is readily
distinguished from all other subspecies by the low number of gulars in contact
with the mental. Taking only the 20 specimens from South Santa Inez Island,
we have the following distribution: 4 have 3 scales; 13 have 4 scales; and 3
have 5 scales; average 3.95. Of the two specimens from San Marcos Island,
one has 3 and the other 5 in contact with the mental. With respect to all other
subspecies out of 621 specimens examined, not one has only 3 scales in contact.
Those with 4 gulars touching the mental are occasional in several other sub-
species, but they are comparatively infrequent; for example, in variegatus
variegatus only 13 out of 319, or 4.1 per cent number 4 (none less); whereas in
slevini those with 4 or less number 18 out of 22, or 82 per cent. This difference
is highly significant.

The high number of specimens having 5 body bands (not counting the
one on the neck) is also characteristic of this subspecies. In the adult retention
of the narrow postparietal light loop slevini resembles peninsularis and abbotti,
and to a less extent sonoriensis, utahensis, and bogerti, although the wavy
nature and frequent breaks in the line are solely characteristic of slevini. The
spotting on the upper labials is more conspicuous in slevini than in the other
subspecies. In the wide character of the bands in the adults the resemblance
is nearest utahensis and sonoriensis, although without the vertebral light line
which is often—but not always—present in the latter. There is also, in the
nature of these bands, a resemblance to peninsularis, especially in the specimens
from San Marcos Island, which in this character, must be considered inter-
grades. These, in fact, in their intermediate nature, justify the subspecific
status assigned to the Santa Inez Island individuals, which I should otherwise
have considered a full species.

KLAUBER—GENUS COLEONYX 171

Field Notes——Mr. Slevin tells me he found these geckos on Santa Inez
in a rocky ledge about three feet high. They were captured by pulling off

the crumbling pieces of rock. Those on San Marcos Island were under stones.

Locality Records——Reported only from Santa Inez and San Marcos
Islands, gulf coast of Lower California, Mexico.

Coleonyx variegatus utahensis subsp. nov.
UraH BANDED GECKO

1920. Coleonyx variegatus Pack, Copeia, No. 88, p. 101.

Type—No. 35,792 in the collection of L. M. Klauber. Collected at
Watercress Spring, Washington County, Utah, by Dr. Ross Hardy, April
16, 1941. Watercress Spring is about 1 mile northwest of Saint George.

Diagnosis—A subspecies of Coleonyx variegatus characterized, in the
adult stage, by the possession of wide, irregular dorsal bands, the irregularity
being increased by the merging of the bands with intercalated spots. The bands
in utahensis are relatively wider than those of the other subspecies except slevini,
sonoriensis, and peninsularis. From the first named, utahensis differs in having
a greater number of gulars in contact with the mental; from sonoriensis in having
bands not so wide compared with the interspaces, with less regularity of the
interspaces, and usually without a mid-dorsal light line. From peninsularis
it differs in having more irregular dorsal blotches, which are not so wide com-
pared with the interspaces, and in having more wavy light canthal lines. It is
noted that where utahensis intergrades with variegatus variegatus there is first a
narrowing (along the body) of the dorsal bands, which coincidentally begin
to exhibit lighter centers (not present in utahensis) thus producing the double-
barred effect characteristic of many variegatus variegatus adults.

Description of Type-—An adult male. Except for the scales bordering
the mouth, nostrils, and eyes, the head is covered above and below with non-
imbricate, circular granules, rather uniform in size, except that they are enlarged
on the snout and bordering the labials. The rostral is pentagonal in shape and
is higher than wide; it is noticeably pointed at the top. The two edges touching
the prenasals are longer than those engaging the first supralabials. The supra-
labials number 7-7 and decrease in size posteriorly, the first being considerably
longer than the rest. The infralabials number 8-8, and also decrease in size
posteriorly. Such enlarged members of both labial series—to the extent that
they can be seen with the mouth closed—terminate under the middle of the eye,
the infralabials somewhat posterior to those of the upper series. There is a pair
of prenasals, long and narrow; their contact with the first supralabials is prevented
by the presence of a tiny scale on either side. They broadly contact the rostral but
are prevented, by a single small scale, from meeting each other on the median
line. There are three postnasals on each side, the upper being the largest. There
is a pair of triangular supranasals, separated ftom each other by 2 granules.
Both upper and lower eyelids are edged with enlarged scales which are slightly
serrated, particularly posteriorly. The upper scales number about 19; the lower
15. The mental is conspicuously the largest head scale. It is slightly narrower

172 SAN Dreco Socrety oF Natura History

than the rostral and is somewhat wider than deep, the dimensions being about
2.7 mm. wide by 2.2 mm. deep. The sides converge posteriorly, the lower edge
being a circular arc of small radius. This edge of the mental is contacted by 5
granules. The mental and the first infralabial on each side, taken together, are
contacted by 10 granules.

The dorsum is covered by granules similar in size to those on the head.
There are no enlarged tubercles. On the sides the scales become somewhat
enlarged and imbricate, first at the middle of the body, since the granules are
smallest near the arms and legs. On the underside the scales are imbricate
and definitely larger than those on the dorsum; they increase in size posteriorly.
There is a midventral umbilical line bordered by about 14 scales on either side.
The scales in the preanal region are enlarged and include a row of 7 with
preanal pores. These are not brown, possibly because of the use of formalin
preservative. The scales containing the pores are obtusely angular in arrange-
ment with the point forward.

The scales on the arms are mostly imbricate, except on the lower side of
the upper arm. They are largest on the inner edge of the wrist. The palmar
surface is tubercular. On the lower surfaces of the digits there are series of
rectangular transverse lamellae which are so imbricate as to form sharp corru-
gations. The claws are extremely delicate; they are brownish in color.

The legs are covered with granular scales outwardly and imbricate anter-
iorly. These scales are largest on the anterior lower areas of the thighs. The
scales on the soles of the feet are granular. The transverse lamellae on the 4th
toe number 21. The claws are similar to those on the fingers, fine and delicate,
and colored brown. The sheaths, beyond which the claws protrude, comprise
a pair of laterals, and a somewhat shorter, pointed terminal.

The cloacal bones are rather sharp, without conspicuous ridges; they are
about 1.3 mm. in length.

The tail is covered by annular rows of subrectangular and imbricate
scales of larger size than any on the body. They are largest ventrally. At the
widest part of the tail there are about 35 scales in a ring.

The principal dimensions in mm. are as follows: Body length, snout to
vent 63; head length 14.4; head width 10.1; rostral to mid-orbit 7.2; rostral
to mid-ear 13.5; width of orbit 3.5; arm fully extended, measured to end of
4th finger 19; leg fully extended, measured to end of 4th toe 24; length of
tail 67; distance between points of cloacal bones 8.

The head is spotted brown above and cream-colored below. There is a
thin cream-colored line of horse-shoe shape extending from behind each eye and
above each ear to a loop on the neck. It is rather straight across the neck and
has a characteristic dip at the ear. There is also a light wavy line on the canthus
rostralis on either side. The upper eyelid edges are marked with gray, particu-
larly posteriorly. The innersurface of each eyelid is black, except for the edge
scales. The rostral is light in the center but punctated on either side. The
mental is speckled anteriorly. The prenasals are dark. The labials, both upper
and lower, are punctated with brown, but not uniformly, some being dark,
others almost immaculate. Back of the enlarged labial scales the granules which

KLAUBER—GENUS COLEONYX 173

border the mouth are punctated with gray-brown. A dark mid-gular line is
faintly visible through the skin; it is not on the surface. There is aj single
wide, dark ring on the neck, which laterally curves forward to the ears, thus
comprising the posterior dark border of the light postparietal or nuchal loop
previously mentioned.

On the dorsal surface of the body, between limb insertions, there are 4
transverse, highly irregular, light-brown bands, slightly wider laterally, and
dorsally much wider than the irregular interspaces. These bars terminate on
the sides, the ventral surface being immaculate cream-colored. The upper sur-
faces of the limbs, out to the ends of the fingers and toes, are spotted with
brown.

On the tail there are 13 irregular rings, none of which completely circles
the tail.

Paratypic Material—There have been available 29 specimens from the
collection of Dixie Junior College, Saint George, Utah, and 17 from the private
collection of Dr. Ross Hardy, all from Washington County, Utah. I have also
examined CAS 55,219-20 and 65,116 from Saint George, Utah. In my own
collection there are four specimens, Nos. 36021-4, from the type locality,
Watercress Spring.

Eighteen specimens from Clark County, Nevada, have been at hand. Those
from the Las Vegas-Bunkerville area and the vicinity of Lake Mead are to be
considered utahensis, although not typical in all characteristics; indeed some
are nearer to variegatus variegatus than to utahensis, showing this to be an area
of intergradation. Other specimens from around Indian Springs are also nearer
variegatus; however, one from Beatty, Nye County, has a pattern somewhat like
utahensis. Nine specimens from Mohave County, Arizona, and nine from Inyo
County, California, have been examined. None of these is typical utahensis,
although one from Littlefield, Arizona, is nearer utahensis than variegatus
variegatus.

~ Range.—Washington County, Utah, extreme northwestern Mohave Coun-
ty, Arizona, and northeastern Clark County, Nevada. Intergrades with
yvariegatus variegatus on the southern and western periphery of this range.

Morphology.—The description which follows is based on the material
from Utah, omitting the Nevada and Arizona specimens which may be
variegatus variegatus intergrades in some characters. This is a moderately large
subspecies, although it does not reach the size of the central desert form. The
largest female has a snout to vent length of 68 mm.; the largest male 64 mm.
The smallest individual is 33 mm. The tail, when complete and original, is
about as long as the body.

The preanal pores in the males vary from 5 to 8; the counts being as
follows:> >) (l’)"6"(17)5: 7 1(6)2-8 (2); mean 6.35. The cloacal bones are
rather sharp and narrow, without conspicuous ridges.

Scalation—The rostral is wider than high; it is rather pointed above.
The prenasals are long and slim; the contact with the first supralabials is very
narrow, and in some cases is prevented by the interposition of a granule. The

174 SAN Disco Society oF Natura History

prenasals are in contact on the median line in 32 cases out of 52, a lower pro-
portion than in any other subspecies. The supranasals are separated by from
two to four granules, the dispersion of the separating scales being as follows:
2 (23), 3 (28), 4 (1); mean 2.58. The supranasals are larger than the post-
nasals; of the latter there may be 2 or 3, the upper largest; the lower might be
considered a subnasal.

The enlarged supralabials number 6 to 9, 7 or 8 being the more usual.
The visible infralabials number 7 to 10, 8 being the most frequent number.
The enlarged infralabials extend further posteriorly than the supralabials. The
scales of the eyelid edges are only moderately serrated. The granules on the
snout are considerably enlarged, compared to those on top of the head; those
touching the labials are particularly large.

The sides of the mental are not parallel but converge posteriorly; some-
times the posterior boundary of the entire scale approaches a circular arc. The
gulars contacting the mental number as follows: 5 (5), 6 (16), 7 (22), 8 (6),
9 (3); mean 6.74. Those contacting the mental and the first infralabial on
either side total as follows: 9 (3), 10 (13), 11 (20), 12 (11), 13 (5); mean
11.04. These contacting scales average higher in number than in any other
subspecies. .

Pattern and Color—This subspecies is characterized by a considerable
ontogenetic change, involving a net increase in pigment. In the young the usual
4 transverse brown body bands are in evidence. These are even-edged and are
about as wide as the interspaces, or the dark bars may be slightly narrower.
There are no lateral marks. The heads are somewhat lighter brown than the
body marks, and are without conspicuous spots. The nuchal light loop is
clearly outlined. The tail rings are of the same color as those on the body.
The limbs are speckled, particularly on the upper side.

As the lizards age the dark body bands widen conspicuously at the expense
of the interspaces, although not reaching the proportional width evident in
sonoriensis and peninsularis. At the same time, as is the case in most of the
subspecies, spots appear in the interspaces; however, in utahensis these usually
become confluent with the main transverse bars, which not only widens the
latter but causes their borders to become highly irregular. The bars seldom
lighten in the center, thus not attaining the double-barred effect so common in
variegatus variegatus.

The head becomes conspicuously spotted with dark-brown on a light-
brown background; and the band on the neck, also, is mottled, although not so
conspicuously. The postparietal or nuchal light loop remains evident, although
it becomes somewhat wavy, square on the neck and with a dip downward at
the ear. Forward it terminates just below the center of the eye. Light canthal
lines are often in evidence, but they also are wavy, becoming lyre-shaped.

The rostral is usually light in the center with dark lateral rings. The pre-
nasals are characteristically dark, much more so than in variegatus variegatus,
wherein they are usually light at the top, where they approach each other. The
upper eyelids are generally tipped with gray, especially posteriorly. The labials

KLAUBER—GENUS COLEONYX 175

are mottled, the upper more than the lower. The mental may be clear but is
usually lightly punctated near the edge of the mouth. Some of the gulars may
be spotted. The limbs are darkly suffused with brown above or may be spotted.
this being particularly true of the hind legs. They are sometimes punctated
below. Some have the palmar surfaces darkened. The claws are often brown.
A few specimens have the ventrum lightly stippled posteriorly. Nearly all
specimens have a dark line below the surface in the mid-ventral position under
the head. This may be evident only in this subspecies because of the use of
formalin preservative in the series available to me.

The following color notes, using Ridgway’s standards, were made on live
adult specimens kindly sent me by Dr. Ross Hardy from the type locality: The
dorsal body bars are Mummy Brown with interspaces of Naples Yellow. The
spots on the head are Dark Vinaceous-Drab upon a background of Light
Vinaceous-Drab. The upper leg surfaces are Light Vinaceous-Drab with
Lavender below. The ventrum is White with a pinkish tinge of blood showing
through.

Relationships with Other Subspecies—This subspecies is obviously an
offshoot of variegatus variegatus, with which it probably intergrades over a
considerable area, especially in central Clark County, Nevada. In addition
to these areas of contact, specimens of variegatus variegatus from the mountains
scattered throughout its desert range are likely to show utahensis tendencies in
an increase in the width of the dorsal blotches. But in most of their respective
ranges they are quite distinct. In addition to the differences in pattern,
utahensis has a more pointed rostral. The first supralabials in variegatus
variegatus are higher in relation to the position of the nostril than in utahensis.
The eyelid scales are more pointed in the typical desert subspecies. In utahensis
the mental more often exceeds the rostral in width than in any other subspecies.
Superficially, utahensis is most like sonoriensis and peninsularis, yet is separated
from the ranges of these two subspecies by the wide desert area inhabited by
variegatus variegatus. This seems more likely a case of the retention of original
characters than one of parallel development.

Field Notes.—Dr. Ross Hardy advises me that most of the specimens col-
lected at Watercress Spring (the type locality) were found beneath red-
sandstone slabs.

I have collected specimens in the evening in the Boulder City, Nevada.
area by driving on the roads at night. I found two in the daytime on an island
in Lake Mead when they were forced to the peak by the rising water.

Locality Records —UtaH: Washington County—Saint George, Red Hill
(1 mi. n. of Saint George), Watercress Spring (type locality), Diamond
Valley, Washington Fields (3 mi. s. of Washington), Ivins, Zion National
Park, Beaver Dam Slope, Gunlock, summit 25 mi. w. of Saint George, Castle
Cliffs. Arizona: Mohave County—Littlefield, Lake Mead (20 mi. above
Boulder Dam). Nevapa: Clark County—Glendale, near mouth of Virgin
river, mouth of Boulder Wash, Boulder City, 8 mi. s. of Railroad Pass, Mes-
quite Dry Lake, mouth of Kyle Canyon (Charleston Mts.), and various islands
in Lake Mead. The Las Vegas—Boulder Dam area is one of intergradation

176 San Dreco Society oF NAturRAL History

between utahensis and variegatus variegatus, and many specimens from the
above mentioned localities in Clark County show pattern mixtures of both
subspecies. La Rivers (1942, p. 56) records two specimens from Lincoln
County, Nevada, (2 mi. s. of Carp on Muddy River at 3100 ft. and Quartz
Spring at 4500 ft.) which may tentatively be assigned to this subspecies. A
specimen occasionally mentioned in the literature from Farmington, Davis
County, Utah, is no longer available. This is far beyond the known range of
the species, and unless verified by additional material is to be considered an
inaccurate record.

Coleonyx variegatus bogerti subsp. nov.
Tucson BANDED GECKO

1893. Coleonyx variegatus (part) Stejneger, North American Fauna, No.
75 p: 162:

Type.—No. 32,486 in the collection of L. M. Klauber. Collected by Lee
W. Arnold, at Xavier, Pima County, Arizona, July 17, 1939. Xavier is a
railroad siding across the Santa Cruz River from San Xavier Mission, and about
10 miles south of Tucson.

Diagnosis—A subspecies which differs from the others in having a higher
number of preanal pores in the males, bogerti usually having 8 or more, while
the others generally have 7 or less. In addition, it differs from all except
variegatus variegatus in the adult pattern. In bogerti the transverse bars on
the dorsum are usually equal to, or narrower than, the interspaces and have
edges darker than the centers; in utahensis, sonoriensis, peninsularis, and slevini
the bars exceed the interspaces in width and lack the double-barred effect
resulting from the lighter interiors characteristic of the Tucson subspecies.
Bogerti adults have heads which are conspicuously spotted or mottled, which is
not true in abbotti. The mental of bogerti is wider, in proportion to its depth,
than is the case in variegatus variegatus.

Description of Type—An adult male. The head is covered above and
below with non-imbricate, circular granules, rather uniform in size, except that
they are enlarged on the snout and where they border the labials. The rostral
is pentagonal in shape and is wider than high. The two concave edges touching
the prenasals are longer than those engaging the first supralabials; the latter
are convex. The supralabials number 7-7 and decrease in size posteriorly; the
enlarged series terminates just anterior to the center of the orbit. The infra-
labials visible with the mouth closed number 8-8. There is a pair of prenasals,
lunar in shape but wider above, which contact the first supralabials, the rostral
and are broadly in contact with each other on the median line. There are two
circular postnasals, and a tiny subnasal on either side. There is a pair of
supranasals, larger than the postnasals and separated from each other by 3
granules. Both upper and lower eyelids are edged with a series of enlarged

* Named for Mr. Charles M. Bogert, Curator of Reptiles, American Museum of
Natural History, whose interest in herpetology I have watched expand from a boyhood
hobby to important and admirable research.

KLAUBER—GENUS COLEONYX 177

scales, which are serrated at the outer edges. The upper row numbers about
17; the lower 14. The mental is conspicuously the largest head scale. It is
equal in width to the rostral and is considerably wider than deep, the dimensions
being about 2.7 mm. wide by 2.2 deep. The sides are curved; the posterior
edge comprises a rather flat circular arc. This edge is contacted by 7 gulars.
The mental plus the first infralabial on each side are contacted by 11 gulars.

The dorsum is covered by granules of similar size to those in the head,
there being no enlarged tubercles. On the sides the scales become somewhat
enlarged and imbricate; the lateral granules are smallest at the limb insertions.
On the underside the scales are imbricate and definitely larger than those on
the dorsum. They are longest posteriorly and medianly.

The scales in the preanal region are further enlarged and include a row
with conspicuous, brown preanal pores. The latter total 10; the scales contain-
ing them are obtusely angular in arrangement with the point forward. The two
central scales overlap, one being somewhat forward of the other.

The scales on the arms are mostly imbricate, except on the outer edge.
The palmar surface is tubercular. On the lower surfaces of the digits there
are series of rectangular transverse lamellae, which form a serrated ridge. The
claws are extremely delicate. They protrude from a sheath comprising a pair
of large laterals, capped by a shorter, pointed terminal.

The legs are covered with granular scales on the upper surfaces and
imbricate below. They are largest on the lower areas of the thighs. The
lamellae on the toes and the claws are similar to those on the fingers.

There is a spurlike cloacal bone protruding laterally and upward on either
side just posterior to the vent. These cloacal bones are about 1.3 mm. in length.
At the top they are ridged rather than conical, the point of the ridge being
forward. There is no notch in the ridge.

The tail is covered by annular rows of subrectangular and imbricate scales
of larger size than any on the body. They are largest ventrally. At the widest
point there are about 41 scales in a ring.

The principal dimensions in mm. are as follows: Body length, snout to
vent 64, head length 17.2, head width 9.8, rostral to mid-orbit 7.4, rostral to
mid-ear 13.4, width of orbit 3.2, arm fully extended, measured to end of 4th
finger 20, leg fully extended, measured to end of 4th toe 25, length of tail
(regenerated) 48, distance between points of cloacal bones 7.1.

The head is light-brown above, spotted with dark-brown. There is a wide
cream-colored line of horse-shoe shape extending from below each eye and just
above each ear to a loop on the neck. There is also a short, wavy light line on
the canthus rostralis on either side. The scales edging the upper eyelids are
posteriorly marked with gray. The inner surface of each eyelid is black, except
for the edge scales. The rostral is light in the center but punctated with brown
on either side. The mental is also punctated anteriorly. The labials, both upper
and lower, are speckled with brown, but not uniformly, some being dark, others
immaculate. Back of the enlarged labial scales the granules which border the
mouth are punctated with gray-brown. The gular surface is cream-colored and
without spots.

178 San Disco Society OF NATURAL History

There is a single wide, dark ring on the neck, which laterally curves forward
under the eyes, thus forming the posterior dark edge of the light postparietal
loop previously mentioned. However, an irregular light line passes backward
from the loop, almost separating the dark nuchal band into two parts. On the
dorsal surface of the body, between limb insertions, there are 4 transverse brown
bars, the centers of which are somewhat lighter than the borders. These bands
are narrower than the interspaces. There are several dark spots in the inter-
spaces. The sides are spotted but the ventral surface is immaculate cream-
colored. The upper surfaces of the limbs, out to the ends of the fingers and
toes, are punctated or spotted with brown.

On the tail there are only 2 rings, since most of it has been regenerated.
The new portion is speckled.

Paratypic Material —I have had available for study 83 specimens from
Pima County, most of them from Tucson or from points within 15 miles of
that city. In addition there are 47 specimens from Pinal County, 11 from
Gila, 3 from Graham, one from Greenlee, and 4 from Santa Cruz.

Range.—Southeastern Arizona from the vicinity of Casa Grande, Pinal
County, south to the Mexican border, and northeast to the Roosevelt Reservoir,
thence southeastward to the New Mexican line at Duncan, Greenlee County.
It undoubtedly occurs in extreme southwestern New Mexico, westward of
the Continental Divide, but I have seen no specimens from that state. The line
of separation from variegatus variegatus is taken as the line Casa Grande-
Covered Wells, but additional specimens will be required to define the area of
intergradation with greater assurance.

Morphology.—The description which follows is based only on the speci-
mens from the Tucson area. Bogerti, while probably not reaching the extreme
size of variegatus variegatus, is nevertheless as large or larger than any of the
other subspecies. The largest specimen is a female measuring 69 mm., snout to
vent. The largest male is 64 mm. The smallest individual from the Tucson
area is 30 mm.; a specimen 28 mm. in length is available from the Sierra Ancha
Mountains, Gila County.

In body proportions and the shape of the snout this subspecies seems to
be similar to variegatus variegatus. There is considerable variation in the ear
opening, both as to size and shape; it may be either round or elliptical.

The preanal pores in the males are distributed as follows in the individuals
fromabout Tucson: 8 (12), 9 (15), 10 (7); 11- (2), 12 Cl); mean9:05.
While this is a considerably higher average than that of any other subspecies, it
is obvious that there is some overlapping and therefore these pores do not com-
prise an invariable key character. In bogerti there is a tendency of the median
scales in the series carrying pores to overlap.

Scalation.—The rostral, which is pentagonal in shape, is always wider than
high. The crescent-shaped prenasals taper almost to a point, where they contact
the first supralabials. There are usually two postnasals, of which the upper is
the larger; a small infranasal is sometimes present. The supranasals are larger
than the postnasals, and are triangular or circular in shape. The prenasals are

KLAUBER—GENUS COLEONYX 179

usually broadly in contact on the median line; this contact is prevented by the
interposition of granules in only 3 out of 81 specimens. The supranasals are
separated by the following numbers of granules (counting the minimum series
iveachcase)/c) 1=1(3,)),. 2 (16),°3 (61) 54" (1); mean’ 2.74." The. enlarged
supralabials usually number 7 or 8; the infralabials 7 to 9. The scales bordering
the eyelids form a serrated series. The scales on the snout are conspicuously
enlarged, compared with those on top of the head; however, the largest head
scales are those contacting the mental or labials. The mental is usually slightly
wider than the rostral; in some specimens it is conspicuously wider, but in others
equal or even a trifle narrower. The mental is more semicircular in shape and
is shorter in proportion to its width than is the case in variegatus variegatus.
The average ratio of depth to width in bogerti is about 0.8. The number of
gulars contacting the mental varies as follows: 4 (5), 5 (13), 6 (34), 7 (20),
8 (5), 9 (3); mean 6.20. The gulars in contact with the mental and first
infralabials on either side when taken as a unit are 7 (1), 8 (4), 9 (10),
10527), 11 (25), 12 (8) 13" (4) 3-mean 1041.

Pattern and Color—The young of bogerti are much like the other sub-
species in color and pattern. The cross bands on the body are even-edged and
somewhat narrower than the interspaces; occasionally they are much narrower.
The tail rings may be of the same shade as the body bars or slightly darker.
The head is unicolor light-brown on top—lighter than the body bands. The
nuchal light loop is narrow and moderately well-defined. Canthal light lines
are present.

The first ontogenetic change is the appearance of spots on the head. These
may be in evidence when the body length reaches 35 to 38 mm. This is fol-
lowed by the appearance of spots between body bars and punctations along
the sides.

The fully adult gecko has very conspicuous dark-brown spots on top of
the head. The postparietal or nuchal light loop is widened and without even
borders; in some specimens it almost disappears. The light canthal dashes are
retained; with a light cross mark in front of the eyes, they form a triangle.
The rostral is light in the center but dark at the sides. The apex is usually
light, but may be punctated. The prenasals are dark, usually to their points
of contact. The supranasals are light. The labials are irregularly spotted. The

mental is usually marked anteriorly, but may be clear.

The ontogenetic change in the body bars generally involves the interposition
of spots between the dorsal bars. At the same time the bars become rather
irregular. The anterior and posterior borders become darker than the centers,
thus giving a double-barred effect. The width of the bars remains about equal
to the interspaces. There is also considerable lateral spotting. The ventral
surface of the body is immaculate. If the tails remain complete, spots appear
between the original bands. Regenerated tails may be spotted or speckled. The
arms are usually suffused above, while the legs are spotted.

Of the specimens from the Tucson area, 68 out of 75 have 4 body bars,
counting only those between the limb insertions. One has 3 on one side of the
body and 4 on the other; five have 4 bars on one side of the body and 5 on the

180 San Disco Society oF NaturaAL History

other; and a single specimen has 5 complete bars. The tail bands vary from
9 to 14, 11 or 12 being the most prevalent. Somewhat less than half of the
posterior bands may completely encircle the tail, but they are rarely as distinct
ventrally as above. Mid-dorsal light lines are evident in only two specimens
out of 75.

A live specimen from Tucson, sent me through the courtesy of Dr. Charles
T. Vorhies, had the following coloration, by Ridgway’s Standards: The head
spots were dark Purple-Drab on a background of Light Vinaceous-Drab. The
nuchal ring was Pale Grayish Vinaceous. The dorsal body bars were Dusky
Brown on the edges and Vinaceous-Drab in the centers. The interspaces were
Colonial Buff. The venter was White. The anterior tail rings were Warm
Blackish-Brown with Colonial Buff interspaces. The legs were Light Purple-Drab

above and Light Brownish-Vinaceous below.

Intrasubspecific Trends——Specimens from the vicinity of Florence, Pinal
County, are much like those from near Tucson, with no great reduction evident
in the number of preanal pores in the males; 12 specimens average 8.83, com-
pared with a mean of 9.05 in the Tucson specimens. The pattern is unchanged
—double-barred primary blotches with conspicuous spots between, especially
laterally. There is a somewhat greater tendency toward a mid-dorsal light line,
characteristic of sonoriensis, particularly in specimens from about Picacho.
Only two specimens out of 47 have other than 4 cross bars, one with 5 on one
side, one with 5 straight across.

From further west in Pinal County, from such points as Covered Wells,
Gunsight, Dowlings Well, and Bates Well, there are too few specimens to
make sure whether they should be assigned to bogerti or variegatus variegatus.
There are 7 preanal pores in two males, so that tentatively I have placed them
in the latter category.

The specimens from Santa Cruz County are also too limited in number
to make a definite allocation, particularly as only one is a male and two are
juveniles. The one adult suggests bogerti rather than sonoriensis. One specimen
(USNM 16,808) recorded from Nogales. Arizona, is certainly sonoriensis,
but the locality records of these early specimens, owing to the method of
cataloguing, are often of doubtful accuracy.

From mountain areas in Gila County, there are eleven specimens available.
Their assignment to this subspecies is only tentative. Most of the available
specimens are juveniles; the few adults indicate that the inhabitants of this
area are probably stunted. The juvenile head marks are retained with little
spotting in the adults. The body bars are somewhat wider than in the Tucson
specimens, and some irregularity is in evidence. One specimen has 3 bars on
one side, 4 on the other; another has 4 and 5, and still another 5 and 6. The
number of preanal pores suggests that they be considered bogerti.

Two males from Graham County have 8 preanal pores. The body bars
are somewhat darker and wider than in the Tucson specimens. These speci-
mens show no brevis tendencies; undoubtedly the geckos of this area should
be considered bogerti.

KLAUBER—GENUS COLEONYX 181

The only specimen at hand from Greenlee County (Cornell 777) is the
most easterly available representative of the species variegatus. It is fortunately
a male and, having 9 preanal pores, shows no tendency toward brevis; quite
the contrary, in fact, for the pores are very close together at the apex, whereas
in brevis they are not only few in number (generally 4) but likewise are separ-
ated at the anterior point of the series. The dorsal bars are rather wider than
in most bogerti; unfortunately we have no way of knowing whether the full

growth has been reached, although the pores are fully developed.

Relationships with Other Subspecies—Bogerti is closest to variegatus
variegatus, with which it probably intergrades over a wide area in eastern
Maricopa County, western Pinal County, and western Pima County. It may
contact sonoriensis in north central ‘Sonora, but material from south of Nogales
and Douglas will be required to determine this. The relationships with the
other subspecies are through variegatus variegatus.

There is no evidence of a direct connection between bogerti and brevis,
notwithstanding their close territorial Proximity; they may in fact, overlap.
Brevis more nearly resembles sonoriensis than bogerti, and if it should ever be
demonstrated that brevis and variegatus are conspecific, the connection is likely
to be found in Mexico.

Ecological and Field Notes——Bogerti, when collected in the daytime, has
been found under stones, fallen advertising signs, debris in a city dump, and
the dried carcass of a steer. I have caught them by driving on the road at night
in the vicinity of San Xavier Mission and Wrightstown (east of Tucson),
but they seemed not nearly so plentiful as is variegatus variegatus in the Borego
area. Two were found together at 11:20 P. M. with the air temperature 79°
and another at 11 P. M. with the temperature 91°. Dr. A. I. Ortenburger
unearthed a specimen while digging for a snake; it was found about 11% feet
below the surface and 8 feet from the entrance of the mammal burrow in which
it had taken refuge (1926, p. 103). Charles E. Shaw reports specimens having

eaten beetles, grasshoppers, and sow bugs.

Locality Records—ArizoNna: Pima County—Tucson (also 6 and 8 mi. w.
and 6 mi. n.), Tucson Mountains, “A” Mountain (near Tucson), Fort
Lowell, Wrightstown, Sabino Canyon (also 2 and 3 mi. w.), Steam Pump
Ranch (13 mi. n. of Tucson), San Xavier Mission (also 3 mi. n.), Xavier
(type locality), Tanque Verde Ranch (also 3/2 mi. e.), Foothills Station
(Santa Catalina Mountains), Romero Canyon (Santa Catalina Mountains),
Saguaro National Monument, base of Santa Rita Mountains, 11 and 27 mi. w.
of Tucson (on the road to Sells), Silverbell; Santa Cruz County—Tubac,
Cayetano Mountains (near Calabasas) ; Cochise County—Tombstone, Turner,
Huachuca Mountains; Greenlee County—Duncan; Graham County—9¥2 mi.
sw. of Safford, 7 mi. se. of Solomonsville; Gila County—Sierra Ancha Moun-
tains, Roosevelt Reservoir, Miami (also 16 mi. w.), Rice; Pinal County—Cool-
idge Dam (may be Gila County), Southwestern Arboretum, Florence Junction
(also 1, 2, 3, 4, 5, 6, 7, 8, and 11 mi. s.), Florence (also 2, 3, 5, 6, 8, and 9 mi. n.,
30 mi. e., 1 mi. s., and 3 mi. w.), Coolidge, Canyon del Oro, 3 mi. se. of
Picacho, Sacaton (probably variegatus variegatus intergrade), Casa Grande

182 San Disco Society oF NaturAL History

(also 5, 6, and 12 mi. w., this being an area of intergradation with variegatus
variegatus). The area of intergradation between bogerti and variegatus varie-
gatus in central Pima County has not yet been determined with accuracy.
Probably there is a gradual change. I have tentatively assigned to variegatus
variegatus specimens from Gunsight, Covered Wells, Bates Well, and 2 mi. e.
of Dowlings Well.

Bogerti undoubtedly occurs in Hidalgo County, New Mexico, but no
definite records are available to me.

Coleonyx fasciatus (Boulenger)
BLiack BANDED GECKO

1885. Eublepharis fasciatus Boulenger, Cat. Liz. Brit. Mus., Vol. 1, p. 234.
Type specimen in the collection of the British Museum. Type locality
Ventanas, Durango, Mexico.

1893. Coleonyx variegatus (part) Stejneger, N. Am. Fauna, No. 7, p. 162.
1935. Coleonyx fasciatus Taylor, Univ. Kan. Sci. Bull., Vol. 22, no. 9, p. 203.

Type.—The type specimen in the British Museum has not been seen. In
addition to the original description, an expanded description of this specimen,
with a figure, will be found in Gunther, 1893 (p. 84, plate 31, fig. A).

Diagnosis——A species without dorsal tubercles, differing in this respect
-from elegans and mitratus. From brevis and all subspecies of variegatus it differs
in being without enlarged postnasals and in having more robust limbs and
digits. It has three, instead of four or more, dark dorsal bands between limb
insertions, and the bands are much darker—being black or almost black—
than in any brevis or variegatus.

Description—Only two specimens of this gecko are known, the type and
a specimen EHT-HMS 10,537 (field number 556) collected about 10 miles
south of Presidio, Sinaloa, Mexico, by Dr. E. H. Taylor, June 19, 1934. Dr.
Taylor has kindly permitted me to examine his specimen, upon which the
following description is based. I have used my own terminology rather than
copying Dr. Taylor’s own excellent description (Taylor, 1935) in order to
parallel the descriptions of the other species and subspecies.

The specimen is an adult male. The head is covered above and below
with non-imbricate, closely-set, circular granules, rather uniform in size, except
that they are much enlarged on the snout and where they border the labials.
The rostral is pentagonal in shape and is wider than high. The sides engaging
the first supralabials are shorter than the two concave edges touching the
prenasals. The supralabials number 6-7 and decrease in size posteriorly; the
enlarged series terminates just behind the center of the orbit. The infralabials
visible with the mouth closed number 7-7. There is a pair of prenasals, lunar
in shape, but wider above, which contact the first supralabials, the rostral and
touch each other on the median line. (These scales are separated by a granule
in the type.) The nostril is circular and moderate in size; it is below the canthal
ridge. The postnasals are not enlarged; about six granules contact each nostril

KLAUBER—GENUS COLEONYX 183

posteriorly. There is a pair of supranasals, separated from each other by 3
granules. Both upper and lower eyelids are edged with a series of enlarged
scales, of which the posterior are pointed. The upper row numbers about 18;
the lower 23. The mental is conspicuously the largest head scale. It is wider
than the rostral and is considerably wider than deep, the dimensions being
about 3.0 mm. wide by 2.1 deep. The sides are posteriorly convergent; the
posterior edge comprises a circular arc. This edge is contacted by 6 gulars.
The mental, together with the first infralabial on each side, is contacted by
9 gulars. The side granules touching the labials are larger than those contacting
the mental.

The dorsum is covered by granules of larger size than those in the head;
there are no enlarged tubercles. On the sides the scales become somewhat
enlarged and imbricate; the lateral granules are smallest behind the arms. On
the underside the scales are imbricate and definitely larger than those on the
dorsum. They are largest posteriorly and medianly. The after edges are
circular. The scales which contact the umbilical line are irregular.

The scales in the preanal region are further enlarged and include a row
of rectangular scales with conspicuous, brown pores. The latter total 11; the
scales containing them are obtusely angular in arrangement with the point
forward. The median scale of the series is triangular and contains a pore.

The scales on the arms are mostly imbricate. They are largest on the
inner edge of the forearm, and smallest on the lower side of the upper arm.
The palmar surfaces are tubercular. The digits are sheathed with imbricate
scales; on the underside of each there is a series of lamellae, with semi-circular
outer edges, which are so strongly imbricate as to constitute a corrugated ridge.
These are heavier than in variegatus. The fingers terminate in sharp claws,
which are housed by, and are probably retractile between, a pair of lateral
shell-like scales, larger than the corresponding members in variegatus. The upper
crevice between the shells is capped by a single sharply pointed scale; below, the
crevice is closed by the terminal lamella. The claws protrude less than in
variegatus, thus showing a tendency toward elegans. There are 13 lamellae
on the fourth finger.

Of the scales on the legs the largest are on the anterior edge of the thigh,
and the smallest on the posterior. The toes are sheathed similarly to the fingers.
There are 18 lamellae on the fourth toe.

There is a spurlike cloacal bone protruding laterally and upward on either
side just posterior to the vent. These spurs are shorter than those in variegatus
specimens of similar size. There is a postanal swelling and paired postanal sacs.

The tail is covered by annular rows of subrectangular and imbricate scales
of larger size than any on the body. They are largest ventrally. At the widest
point there are about 25 scales in a ring.

The principal dimensions in mm. are as follows: Body length, snout to
vent 58/2, head length 14%, head width 9.7, rostral to mid-orbit 7.4, rostral
to mid-ear 12.8, width of orbit 3.5, arm fully extended, measured to end of 4th
finger 17, leg fully extended, measured to end of 4th toe 23, length of tail
(regenerated) 53, distance between points of cloacal bones 7.2.

184 SAN Drisco SociETy OF NATURAL HIstory

The head is very dark-brown above, and almost black posteriorly and on
the outer edges. There is a wide tan-colored line of horse-shoe shape extending
from below each eye and just above each ear to a loop in the parietal area.
The posterior part of this loop is wider and further forward than in abbotti.
There is also a short, straight, tan line on the canthus rostralis on either side.
The scales edging the eyelids are light, except that those above are posteriorly
marked with gray. The inner surface of each eyelid is black, except for the
edge scales. The rostral, mental, and labials are unspotted. The gular surface
is clear, but is tinged with greenish on either side below the infralabials.

There is a single wide, dark band on the neck, which laterally curves
forward across the ears, thus forming the posterior dark edge of the light
parietal loop previously mentioned. An extension of this band is carried back-
wards, just above each arm insertion, to join the first body band. On the
dorsal surface of the body, between limb insertions, there are 3 transverse,
even-edged, dark bands, the centers of which are dark-brown and the edges
black. These bands are more than twice as wide as the interspaces. The bands
terminate on the sides. The interspaces are tan. The ventral surface is
immaculate yellowish-tan. The upper surfaces of the limbs, out to the ends
of the fingers and toes, are grayish.

On the tail there is one wide, black ring and the beginning of a second.
Posterior thereto the tail has been regenerated and comprises a gray ground
_ color with darker spots. Although the tail of the type specimen is also regen-
erated, it can be seen from the figure that the rings in this species are fewer
and wider than in variegatus or brevis, and probably they tend to be more
complete ventrally.

Relationships with Other Species—This is a well-differentiated species.
While it has many homologous items of morphology, scalation, and pattern with
both variegatus and brevis, it is so well separated that the territorially nearest
specimens of variegatus (sonoriensis) or brevis, except for a darkening in color,
seem to show no particular fasciatus tendencies.

Field Note—“The specimen was obtained late in the afternoon ensconced
beneath a pile of small logs in the forest, June 19, 1934. Here the trees (really
only overgrown shrubs, usually about 15 to 20 feet high) were thick, and be-
ginning to leaf out, since the rains had begun just a short time previously.”
Taylor, 1935, p. 204.

Locality Records —At present this species is known only from two points:
Ventanas, Durango (type locality), and 10 miles south of Presidio, Sinaloa,

Mexico.

Coleonyx brevis Stejneger
Texas BANDED GECKO
1858. Stenodactylus variegatus (part) Baird, Proc. Acad. Nat. Sci. Phila.,
Vol. 10, p. 254.
1880. Coleonyx variegatus Cope, Bull. U. S. Nat. Mus., No. 17, p. 13.

KLAUBER—GENUS COLEONYX 185

1885. Eublepharis variegatus (part) Boulenger, Cat. Lizards Brit. Mus., Vol.
cpt 233.

1893. Coleonyx brevis Stejneger, North American Fauna, No. 7, p. 163.
Type specimen: USNM 13,627. Type locality: Helotes, Bexar Co.,

Texas.

Type.—Stejneger originally distinguished brevis from variegatus premised
on its having a shorter snout, less developed anterior nasals, and more numerous
labials. Although he named a type specimen, No. 13,627, which is still in the
U. S. National Museum, he did not describe it. Strecker (1933, p. 77) has
given an interesting picture of the type locality, Marnock’s Ranch on Helotes
Creek, some 22 miles northwest of San Antonio, Bexar County, Texas.

Diagnosis —Brevis may be distinguished from the southerly members of
the genus by the absence of enlarged tubercular scales among the granular
scales of the dorsum. Although the characters first used by Stejneger to dis-
tinguish brevis from variegatus, proved, with the availability of more material,
to lack consistency, the species are indeed different, other divergences having
been discovered. H. M. Smith, in 1933, pointed out that the preanal pores
are fewer in brevis (seldom other than 4), and are separated into two lateral
series by unpitted scales at the apex, while in variegatus the two series contact
at the apex; he also discussed the pattern differences, and differences in the
cloacal spurs. In brevis the supranasals are somewhat smaller and more widely
separated than in variegatus. From fasciatus, brevis may be distinguished by
the lower number of anal pores and by the spotting of the adults.

Range.—Brevis occurs from central New Mexico south throughout the
trans-Pecos region of Texas, and in the Rio Grande area southwest of a line
from Crockett County to Kleberg County, with a northeast extension into
Medina and Bexar counties. In Mexico it has been collected in northern
Tamaulipas, northern and central Nuevo Leén, southern Coahuila, and western
Durango. No doubt this range will some day be considerably enlarged, par-
ticularly into northern Coahuila and Chihuahua.

Material—The following specimens have been available for study:

New Mexico 2
Trans-Pecos Texas 61
Middle Southwestern Texas 42
Extreme Southern Texas 39

Coahuila, Mexico iE
Nuevo Leon, Merico 5
Tamaulipas, Mexico 1
Durango, Mexico 5

Morphology.—Brevis is a lizard of moderate body proportions, not
particularly flattened either in head or body. The legs are relatively short and

186 SAN Dreco Society OF NATURAL HIsToORY

weak; however, when adpressed they overlap. The head is wedge-shaped and
the snout not especially pointed. It seems slightly blunter than in variegatus.
The nostrils are circular. The ear openings are conspicuous; they differ con-
siderably in relative size and may be either circular or elliptical. The eyelids
are functional and are interiorly pigmented with black. There is a mid-dorsal
groove bounded on either side by a rounded ridge. There is a conspicuous
umbilical line. The peritoneum is unpigmented.

Out of the 162 specimens which I have studied, the largest is a female,
measuring 59 mm., snout to vent. This is from Langtry, Texas. Other speci-
mens closely approach this length, which therefore is not exceptional. The
longest male measures 56 mm. Thusybrevis is considerably smaller than varie-
gatus variegatus, wherein females exceeding 70 mm. are not unusual. The
smallest brevis measures 22 mm.

The tails are seldom original and complete. When complete they approxi-
mate the body in length, tending to be slightly shorter, proportionally, in the
adults.

The lateral cloacal spurs are relatively wider than in variegatus, with a
ridge paralleling the side of the tail. The spur bends slightly forward so there

is an anterior point. Notches are occasionally present.

The dispersion of the preanal pores in the adult males is as follows:
3 (1), + (62), 5 (5), 6 (2); mean 4.11. The propensity to 4 is strongly
_evident. There are usually several median scales separating the two side
series of enlarged scales carrying the pores; these separating scales number as
follows: 0 (3), 1 (12), 2 (30), 3 (15), 4 (11); mean 2.27. While enlarged
scales are present in the females, and often have depressions corresponding to
the true pores in the males, they have not been tabulated, as there are some of
intermediate size, of which one cannot be certain. But in general, like the males,
the females possess a pair of enlarged scales on either side, separated by from
1 to 4 smaller median scales. This almost universal separation of the scales
carrying the pores is a character which readily distinguishes brevis from varie-
gatus.

Scalation—The head is covered above and below with hemispherical
granules, except for the scutes which border the mouth, nostrils, and eyes. The
granules are somewhat enlarged on the snout, particularly where they abut the
labials; in fact, the enlargement adjacent to the labials is generally greater than
In variegatus.

The rostral is the largest of the head scales except the mental. It is
pentagonal, with concave sides contacting the prenasals. The prenasals are
lunar in shape and wider above. They usually touch the first supralabials.
Often the prenasals meet on the median line, but in 53 cases out of 155 (34.2
per cent) this contact is preyented by the interposition of a granule. The
supranasals are quite small, and are widely separated from each other by the
small scales of the snout. The minimum scales bridging the gap between the
supranasals are as follows: 3 (30), 4 (92), 5 (29), 6 (1); mean 4.01. This

tabulation omits a single peculiar specimen from southern Coahuila having only

KLAUBER—GENUS COLEONYX 187

one scale interposed; this will be discussed later. There are 2 to 4 postnasals
(the lower might be considered a subnasal), the distribution being as follows:
2x(148)).3. (128)42(20); mean:2.57.

The enlarged supralabials decrease in size posteriorly, gradually merging
into granules. The first is the largest of the series, being the highest and
usually the longest as well. The enlarged scales terminate somewhat anterior
to a point below the center of the eye. The counts of the supralabials are as
follows: 5 (1), 6 (32), 7 (124), 8 (104), 9 (13), 10 (3); mean 7.38.

The eyelids are edged with pointed scales numbering about 18 to 20 above
and 20 to 23 below.

The mental is the largest scute. It is usually considerably wider than
the rostral. Posteriorly it is semi-circular in shape, seldom having parallel
anterior sides, as in some variegatus. The enlarged infralabials, visible with
the mouth closed, generally number 6 to 8.

The gulars contacting the mental are usually smaller than those touching
the anterior infralabials. Those touching the mental are distributed thus: 4 (1),
5 (15), 6 (54), 7 (56), 8 (21), 9 (7); mean 6.66. The total gulars touching
the mental, together with the first infralabial on either side are: 8 (3), 9 (12),
10 (55), 11 (48), 12 (24), 13 (9), 14 (1), 15 (1); mean 10.75.

The dorsal surface of the body is covered by granules somewhat like those
on the head. They are usually larger posteriorly and medianly, although this
is not always the case. The smallest body scales are at the limb insertions.
Ventrally the scales become imbricate and are considerably larger than those
on the dorsum. These ventral scales are usually largest posteriorly and medianly.

The tail is covered with annular rows of imbricate scales, usually larger than
any on the body.

The arms and legs are covered with scales which are usually imbricate,
although granular posteriorly, especially at points of insertion.

The palmar surfaces and scales are covered with tubercular scales. The
fingers and toes are sheathed with overlapping scales. The lower surfaces are
covered with strongly imbricate scales, thus presenting a corrugated surface.
There are about 14 corrugations on the fourth finger and 16 on the fourth toe.
The claws are delicate and hollowed out below. They are housed and are re-
tractible between two large lateral shell-shaped scales, capped above by a long
thin pointed scale (the terminal).

Pattern and Color—In the juvenile stage brevis is not greatly different
from the various subspecies of variegatus. There are four brown even-edged
cross bands on the body between limb insertions, and another on the neck
forming the posterior border of a light nuchal loop. The body bars are about
equal in width to the interspaces. The bars terminate on the sides, the ventrum
being immaculate. The head, anterior to the nuchal loop, is lighter than the
body bars. It is somewhat mottled and with light canthal lines. The nuchal
loop is wider than in variegatus. The tail rings are usually narrower than the
interspaces. They number 6 to 11, averaging about 9. Sometimes the tail
bands are darker than the body bands. The limbs are punctated above.

188 SAN Disco Society oF NATURAL History

The change to the adult pattern is as extensive as in variegatus variegatus,
but somewhat different in character. The earliest changes take place on the
head, which begins to show spots when a length of about 30 mm. is reached.
Then spotting begins on the body, first laterally and between blotches, but
gradually all over. The essential difference from the variegatus subspecies lies
in the even distribution of the body spots in the final adult dorsal pattern; for
in brevis the dorsal spots are evenly superimposed over almost the entire sur-
face, producing a leopard-like effect, whereas in variegatus they are accentuated
in, or restricted to, the areas between the dark cross bars. Simultaneously, the
original body bands grow at the expense of the interspaces, which are often
reduced to thin, highly irregular light lines. The nuchal loop also becomes
thin and wavy. The spots are dark-brown, while the bands, now highly irregu-
lar and often confluent, present a somewhat lighter ground color. A few
specimens become so completely spotted that all traces of the original bands
are obliterated. On the other hand, some specimens retain the dorsal bands
into the adult stage, the spotting being most in evidence laterally. Original
tails retain the rings of the juvenile state but add spots between. Sometimes the
bands become lighter in the centers. Regenerated tails are heavily spotted.

The rostral in the adults is usually light in the center, with dark wings.
The snout is mottled, the even canthal lines of the young being lost. The
labials are mottled or punctated, and the entire under-jaw is usually spotted
or blotched with brown; or there may be areas in which the usual white pigment
is lost, thus giving the appearance of a dark blotch.

The ventral surface of the body is often marked with scattered, fine black
dots, which may only be evident with magnification.

The limbs of the adults are spotted; the fingers and toes are punctated
with black or brown, both above and below.

Intraspecific Trends—The series of brevis which has been available to me
inadequately represents both the extreme northern section of the range (New
Mexico), and the southern, (lower Coahuila and eastern Durango). There
may therefore be territorial trends that I am unable to determine, which may
ultimately involve subspecific segregations.

With regard to the Texas specimens, the geckos of the Bexar-Medina area
seem to be somewhat different, in some characters, from the others. The median
scales separating the two series containing the preanal pores average somewhat
lower here than elsewhere, and there is a slightly greater tendency toward
a separation of the prenasals. At the same time there is a somewhat reduced
number of scales between the supranasals. It is also noted that in this area
the ontogenetic pattern change seems to take place later; that is, larger speci-
mens are found still retaining the juvenile barred pattern, with spots only
evident laterally.

Specimens from trans-Pecos Texas are more prominently spotted in the
gular region than those from southern Texas or Mexico.

USNM 113,063 collected by Dr. Hobart M. Smith near Saltillo, Coahuila,

Mexico, is a peculiar specimen in several particulars. The anterior nasals are

KLAUBER—GENUS COLEONYX 189

broadly in contact, and the supranasals are quite large and are separated by only
one scale. This is the only individual that has less than 3 scales between the
supranasals, out of 153 which have been checked for this character. The pattern
is also peculiar; the three posterior body bands have coalesced to form a central
longitudinal dark line, bordered with light, and then again by dark. Three other
specimens, which Dr. Smith tells me were collected only a mile or so away, and
in a similar environment, show none of these peculiarities. Therefore this
specimen must be considered a freak, until such time as others showing similar
abnormalities might come to light.

Relationship with Other Species——There are no present indications of
intergradation between brevis and variegatus, although they seem more closely
related to each other than either is to fasciatus, the only other non-tubercular
species. It appears significant that bogerti, the subspecies of variegatus territor-
ially nearest to brevis in the United States, is the most divergent from it in the
important character of the preanal pores. These forms may, in fact, overlap
in southwestern New Mexico. Dr. Smith advises me that Mr. H. W. Parker
has reported a typical variegatus from El Paso in the British Museum collection.
However, knowing how frequently in the past our southwestern reptiles have
been recorded in museums as coming from the place from which they were
sent. rather than the actual point of collection, I deem it best not to consider
overlapping proved until verified by other specimens. Of the variegatus
subspecies, brevis seems nearest related to sonoriensis and intergradation be.
tween these two, while improbable, is not impossible. If it occurs it will be
across southern Chihuahua, although the mountains here are high enough to
render any present contact quite doubtful.

Field Notes.—“1 found it rather abundantly in the rocky hills of the first
plateau northwest of San Antonio, but did not observe it in that region north
of that point either on the Guadalupe or the Llano. It is found in holes under
stones, towards evening, and generally in pairs, a peculiarity I have not observed
in any other lizard. Its manners are also peculiar. It carries its thick tail coiled
vertically on one side of its back like a spitz dog. Its movements are quick
but feeble, and its short legs forbid the speed of other lizards. Coleonyx is one
of the few genera of Gecconidae which have eyelids, and as these are thick,
and their movement in winking is slower than in other lizards, the physiognomy
is quite peculiar. When handled, this species chirrups and squeals feebly like
a singing mouse. One specimen which I took was about to shed its skin, so I
placed it in a jar to observe the process. This took place in the night, for next
morning it was so clear and its color so bright, that it looked as though gctten
up for some special occasion. As no trace of the skin could be found, I suppose
that it ate it, after the manner of the Batrachia. In life, the colors are very
elegant; the pale cross-bands are citron-yellow, and the brown ones bright
chestnut. The inferior surfaces and all parts of the limbs are flesh or rose

color.” E. D. Cope, 1880, p. 13.

“T collected a single example of this little gecko in the Chisos foot-hills.
It was running around among rocks on a hillside, just before dusk and on

190 SAN Dreco SoctEty oF NATURAL History

account of its rather feeble movements was easily captured. In the living

specimen the bands were brown and sulphur-yellow.” J. K. Strecker, 1909, p. 13.

“This little gecko is abundant in the vicinity of Helotes and has been col-
lected within two or three miles of San Antonio. It is found in rocky places and
the specimens represent two types—one banded, the other spotted.” J. K.
Strecker, 19225 p17.

“Fifteen specimens of Coleonyx brevis were found in hiding under small
flat rocks. In no instance was more than one found under the same rock, and
no two specimens were found near one another. The peculiar mouse-like squeak
of this species is quite characteristic of small geckos.” J. K. Strecker, 1933, p.
78, describing collecting at the type locality of C. brevis, Marnock’s Ranch on
Helotes Creek, 22 mi. nw. of San Antonio, Bexar County, Texas.

“They are scarce and found in rocky sections only. I have taken but four
of them during the past fifteen years. They seem to prefer certain special
areas. I found two under rocks on top of a certain hill; then two years later
found one at the same locality (near Somerset, Texas).” A. J. Kirn, by letter,
Sept. 10, 1943.

Locality Records—New Mexico: Santa Fe County—Waldo; Otero
County—Alamogordo. Texas: El Paso County—El Paso; Hudspeth County—
near El Paso; Culberson County—Apache Canyon (40 mi. n. of Van Horn) ;
Reeves County—Pecos, Vitro Draw; Jeff Davis County—Cherry Canyon,
Musquiz Creek (612 mi. s. of Fort Davis), Kingston Ranch (Madera Canyon
on n. side of Davis Mts.) ; Pecos County—Sheffield, Iraan; Crockett County—
17 mi. w. of Ozona; Brewster County—Y2 mi. w. of Banta, 14 mi. n. of Ter-
lingua, Tornillo Creek (10 mi. above San Vicente); and the following in the
Chisos Mountains area—3 mi. s. of Chisos Mts., Glenn Spring, Juniper Canyon,
The Basin (at 5100 and 5200 ft.), foothills e. and 6 mi. ne. of The Basin,
flats ne. of Chisos Mts., desert slope n. of Nugent Mt. (at 3250 ft.), Green
Gulch (3 or 4 mi. above Burnham Ranch), Wade (Pine) Canyon, Govern-
ment Spring (Burnham Ranch), Oak Creek, foothills e. and ne. of Chisos
Mts., Cottonwood Creek (9 mi. ne. of Chisos Mts.), 4 mi. w. of Chisos Mts.
CCC Camp, Hot Springs; Terrell County—Sanderson (also 15 mi. e.); Val
Verde County—¥2 mi. ne. of Shumla, Del Rio, Pecos River Canyon near
bridge and near mouth, Devil’s River (near mouth and 3 mi. above big bridge) ,
Devil’s River Crossing; Kinney County—Brackettville; Medina County—Diver-
sion Lake (Rio Medina), 12 and 18 mi. n. of Castroville; Bexar County—San
Antonio (also rocky hill of first plateau nw.), Marnock’s Ranch (Helotes
Creek) (type locality), Helotes; Maverick County—Eagle Pass; Webb County
—15 mi. n. of Laredo; Zapata County—Zapata (also 15 mi. s.), 20 and 32 mi.
se. of Laredo, 2 mi. s. of San Ygnacio; Jim Hogg County—Hebbronville;
Kleberg County—Kingsville; Starr County—Rio Grande City (also 5 mi. e.,
and 3 and 5 mi. se.), Arroyo Los Olmos, 5 mi. w. of Roma; Hidalgo County—
7 and 10 mi. n. of La Joya, 25 mi. w. of Edinburg, Mission (also 10 to 12 mi.
nw. and 15 mi. wnw.). There is also a record from Live Oak Creek which might
be in Crockett, Zavalla, or Dimmit County. Mexico. CoanutLa: Monclova,
Saltillo (also 4 mi. w.). Nugevo LEON: 5 mi. s. of Sabinas Hidalgo, near

KLAUBER—GENUS COLEONYX 191

China, Ciénega (Ciénega de Flores), Mamulique Pass. TAMAULIPAS: Mier.
DurANGo: 6 mi. ne. of Pedricefa, 32 and 35 mi. wsw. of San Pedro (this
San Pedro is in Coahuila about 15 mi. ese. of Torredn).

Coleonyx elegans elegans Gray
YUCATAN BANDED GECKO

1845. Coleonyx elegans Gray, Ann. and Mag. of Nat. Hist., Vol. 16, p. 162.
Type specimen in the British Museum. Type locality: Belize, British

Honduras.

1851. Gymnodactylus scapularis A. Duméril, in Dumeril and Duméril, Cat.
Meth. Coll. Rept., p. 45. Type specimen in Muséum d’Histoire Naturelle
de Paris. Type locality: Petén (Department), Guatemala.

1858. Gymnodactylus coleonyx A. Dumeéril, Arch. Mus. Hist. Nat., Vol. 8,
p. 483. Same type specimen as G. scapularis above.

Type.—The type specimen in the British Museum, from Belize, British
Honduras, has not been examined. From the brief summary of characters
given by Gray in describing both the genus and the species, somewhat elaborated
by Boulenger (1885, p. 235), the type specimen seems quite representative of
the species as found in the lowlands of the Yucatan Peninsula.

Diagnosis—Elegans elegans differs from brevis, fasciatus, and variegatus
and its subspecies, in having tubercular scales scattered profusely among the
smaller granules of the dorsum. From mitratus it differs in having longer scales
sheathing the claws, and in the possession of a more triangular mental and first
infralabials. The postnasal depression is more evident in elegans than in mitratus.
Elegans elegans differs from e. nemoralis in having the upper prenasals more
completely in contact, and in the greater profusion of the dorsal and lateral
tubercles.

Range.—From central Veracruz eastward along the lowlands bordering the
Gulf of Mexico, and throughout the Yucatan Peninsula to northern Guatemala
and British Honduras.

Material —I have examined 15 specimens from Veracruz, 2 from northern
Oaxaca, 2 from Tabasco, 1 from Chiapas, 3 from Campeche, 28 from Yucatan,
15 from northern Guatemala, and 5 from British Honduras; total 71.

Morphology.—A lizard of moderate habitus, not particularly depressed.
The head is wedge-shaped, with a more evenly tapering snout (viewed verti-
cally) than variegatus. The neck is rather long and slender. The limbs are
relatively short and weak, yet they overlap; in fact, the tips of the toes reach
the elbows. The nostrils are large and circular; they lie well below the canthal
ridge. The ear openings are prominent; they are elliptical in shape, with the
long axis almost vertical, but advanced slightly forward at the lower end. The
eyes have vertically elliptical pupils. The eyelids are functional and are lined
with black pigment within. The peritoneum is uncolored.

Well preserved specimens show four median dorsal ridges, of which the
central two are somewhat more prominent than the outer. They are most

192 SAN Disco Society oF Natura History

evident posteriorly. There is a short mid-ventral umbilical line. The tail is
round in section.

E. elegans grows to a considerably larger size than the non-tubercular
species. The longest, a female from Potrero Viejo, Veracruz, is 97 mm. from
snout to vent. The longest male measures 92 mm. The shortest specimen is
36 mm.

The tails, when complete, are about equal to the body in length. The tails

of the juveniles are proportionately somewhat longer than in the adults.

The males may be recognized by their more prominent preanal pores;
however in this species the females have definite depressions corresponding to
the pores in the males. The lateral cloacal spurs are relatively much less in
evidence than in variegatus and the other non-tubercular species; in fact, they
are little more prominent than the nearby tubercles. They are short and flexible,
and flattened transversely into a sharp ridge. Postanal sacs are present; and
there are postanal swellings in the males.

The variation in the number of preanal pores is as follows: 7 (2), 8 (7),
9 (28), 10 (16), 11 (9), 12 (0), 13 (1); mean 9.43 + .15; coefficient of
variation 11.4 per cent. The females have been included in these statistics since
there is no sexual dimorphism in numbers of pores or depressions. The arrange-
ment of the scales carrying the pores differs somewhat from that in variegatus;
in elegans the two series meet in a sharper point mid-ventrally, for each side

, series curves inward slightly. Occasionally there are two pores in tandem at the
apex, and sometimes there are two pores within a single scale. At the season
of maximum activity the pores are relatively much larger in elegans than in
variegatus; in some specimens the core of the secretion, which may be lifted out
as a clear yellow cylinder, occupies almost the entire scale in which it centers.

Scalation—Except for larger scutes bordering the mouth and nostrils, the
head is covered above and below with small non-imbricate granules, inter-
spersed with enlarged tubercles. The rostral is the largest of the head scales
excepting only the mental; it is pentagonal in shape, with a wide base, two
shorter sides contacting the first supralabials, and longer, slightly concave sides
engaging the prenasals. The upper point is rather obtuse (about 135 deg.).
There is a series of enlarged supralabials which decrease in size posteriorly.
They generally number 6, 7, or 8, with an extreme range of 5 to 9; the mean is
7.09 + .06. These enlarged labials terminate approximately under the center
of the orbit, posterior to which the mouth is bordered by granules. There are
two prenasals on either side, the upper much the larger, and contacting its
fellow medianly. This contact is prevented by the interposition of an extra
scale in three specimens out of 69, and in one other the contact is prevented by
an elongated rostral. The lower prenasals are small and triangular. The supra-
nasals are so small that they do not stand out conspicuously, as compared with
the other scales on the snout, nor are the remaining scales touching the nasal
orifice especially enlarged. The scales posteriorly bordering the upper prenasals
from nostril to nostril, number as follows: 5 (1), 6 (7), 7 (22), 8 (20),
9 (13), 10 (3), 11 (1); mean 7.75 = .14. Posterior to the nostril, and extend-
ing as far back as the beginning of the third supralabial, there is a shallow

KLAUBER—GENUS COLEONYX 193

longitudinal depression in which the scales are conspicuously smaller than
those above and below. The eyelids are edged with serrated scales. The rest
of the head surface is covered with circular granules which are considerably
enlarged on the upper surface of the snout, and where they abut the supra-
labials. Those which contact the supralabials are often elliptical in shape.
Among the granules covering the head there are enlarged subcircular tubercles
scattered at fairly even intervals; these increase in size toward the neck. A few
tubercles are evident laterally below the commissure, but not on the under surface
of the head, which, except for the scutes bordering the mouth, is covered with
granules. The mental is large with the two sides converging posteriorly. The
posterior edge is slightly convex. The first infralabials are triangular in shape,
with points directed inward along the converging edges of the mental. The
remaining lower labials are subrectangular, decreasing in size posteriorly, until,
at a point somewhat back of the orbit, they are no longer conspicuously en-
larged. The visible enlarged scales number from 6 to 9, with 7 predominating; the
mean is 7.16 + .06. The granules contacting the mental and infralabials are con-
siderably enlarged, compared with those toward the center of the lower jaw. The
largest of these scales abut the posterior infralabials rather than the mental; they are
somewhat elongated in shape. The scales touching the mental number as follows:
3 (1), 4 (3), 5 (17), 6 (20), 7 (16), 8 (6). 9 (8); mean 6.37 + .17; coefficient
of variation 22 per cent. Those touching the first infralabials number 3 (13),
4 (84), 5 (42), 6 (2); mean 4.23 + .05; coefficient of variation 15 per cent.
It is to be understood that there is duplication in these tallies, in that one scale
in each instance touches both the mental and a first infralabial. Of the scales
touching the first infralabials, those nearest the second infralabials are usually
the largest.

The neck and dorsal surface of the body are covered with granules, among
which there are scattered tubercles that tend to become larger posteriorly, and
more closely spaced iaterally. The dorsal tubercles are often keeled and have
posterior peaks. Ventrally the granules and tubercles are replaced with uniform,
flat, imbricate scales, which increase in size posteriorly, and are especially en-
larged where they abut the series carrying the preanal pores.

On the tail the dorsal granules of the body are replaced with rings of
imbricate scales. These are enlarged ventrally. There are a number of rings
containing tubercles, which, however, are not evident ventrally; these rings
comprise about one-seventh of the total. Regenerated tails lack tubercles.

Tubercles are also present on the arms and legs, being most prevalent
on the posterior edge of the forearm and the upper sides of the legs. The
posterior edges of the upper arm and thigh are covered with granules.

The hands and feet have imbricate scales above and tubercular granules
below. The fingers and toes taper only slightly. They are sheathed with
imbricate scales dorsally, with a single row of lamellae below. The latter are
so imbricate as to form a corrugated ridge. There are about 17 lamellae on
the fourth finger and 20 on the fourth toe. The claws in preserved specimens
are completely hidden by the scales which sheath them; these comprise, on
each finger or toe, a pair of shell-like convex lateral scales, partly capped

194 San Disco Society oF NATURAL History

above by a long slim pointed scale (the terminal). In this subspecies these
sheaths are longer, relatively, than in any other subspecies. Only rarely is the
tip of a claw evident.

Pattern and Color.—In the juvenile stage elegans has a fairly simple pat-
tern, comprising three major body triads, each of which consists of a wide
central cross-band of fawn color, bordered in front and behind by a narrow
dark-brown edge. Between the triads are narrow interspaces of buff, approxi-
mately equal in width to the dark edges which bound the triads. All marks
terminate on the sides, the ventral surfaces being immaculate cream color. The
legs are punctated with brown. The tail is also marked by triads like those
on the body, except that posteriorly the bordering dark bars widen at the
expense of the light centers until the bands are uniform dark-brown or black.
Also posteriorly a few rings are complete ventrally. On the head there is usually
a light parietal loop with narrow dark borders, and a second light central
ellipse. There are dark vertical bars marking the upper labials.

In the adults a marked change takes place. The dark borders of the dorsal
triads widen at the expense of the light centers, so that they often become
confluent, especially dorso-laterally. Irregular dark spots may appear within
the triads. The light cross bars remain between triads; usually they are clearest
mid-dorsally where they are also widest. On the sides dark blotches appear,
conspicuously speckled by the tubercles, most of which have become almost
white. Thus, while the basic pattern of the three dorsal triads usually remains
“in evidence, it assumes a considerable irregularity.

The tail rings number 8 to 10 in the complete tails. The anterior rings
are triads, while the posterior are uniform dark-brown, almost black. The last
two or three may be complete ventrally. Regenerated tails are spotted or
speckled, rather than banded.

The head is either streaked with dark or may be marked by a series of
irregular concentric ellipses. The sides are mottled. There is generally a light
cross streak, between dark edges, on the snout. The rostral is light in the center,
but dark laterally. The legs are spotted. The lower surfaces of the head and
body are clear.

Intrasubspecific Trends—I have been unable to detect any important
intrasubspecific trends in elegans elegans. Specimens from Veracruz have a
slightly reduced number of scales in contact with the mental, as compared to
those from the Yucatan Peninsula. In the adult specimens from British Hon-
duras, or northern Guatemala, there is a somewhat greater tendency to have
the dark edges of the triads joined dorsally into hour-glass figures than is the
case further westward in Veracruz.

In certain areas of the Yucatan Peninsula some of the specimens have the
normal cross bars on the body replaced with longitudinal stripes. There is a
mid-dorsal light line, bordered on either side by a wider brown band. The sides
are mottled, the tubercles being light. Where the replacement of cross bars
by longitudinal stripes is only partial, it is effected anteriorly; in any case the
tail remains ringed. This is not an adult development, the aberrant pattern
being evident in some juveniles as well.

KLAUBER—GENUS COLEONYX 195

Relationships with Other Forms.—Elegans elegans is only distantly related
to the non-tubercular species, its nearest relative being fasciatus, to which it
shows some affinity (through nemoralis) in the robust digits, the long claw
sheaths, and pattern. While I consider it specifically distinct from mitratus,
because of the differences in the claw sheaths, and the shapes of the mental and
first infralabials, it should be noted that there is a trend within mitratus toward
elegans elegans, with respect both to the mental shape and pattern, just as would
be the case if it intergraded with elegans elegans. Thus, while intergradation
between the species elegans and mitratus is improbable, it cannot be deemed
impossible, since no specimens are available from the area (central and southern
Guatemala) where either intergradation or overlapping might occur.

Field Notes——Ruthven (1912, p. 311) states that two individuals were
seen under boards in the sheds at San Juan, Veracruz, Mexico.

Stuart (1935, p. 41) reports four specimens, collected at La Libertad,
Guatemala, in the bush or from the savanna country. He believes it possible
that the species is a wet-season form, which accounts for his not securing it
during the dry season.

In Yucatan they occur, among other places, in caves. “One, Puz Cave,
Oxkutzcab, July 20, on wall of an inner chamber 63 m. from the mouth, in
complete darkness, at a depth of about 20 m. below the surface, and beyond
two artificial walls with small doorways, food was insects; one, Gongora Cave,
July 17, 15 m. from mouth, food, two cave-crickets; one Ziz Cave, July 24,
12 m. from mouth. Another specimen, which was not saved, was seen in Xkyc
Cave, Calcehtok Hacienda, San Bernardo, August 7, 13 m. below the surface
and more than 35 m. from the mouth.” Helen T. Gaige, 1938, p. 297.

Sanderson, 1941, p. 156, reports it in cohune logs and stumps, in British
Honduras.

Locality Records.—Elegans elegans has been collected at the following
points: Mexico. VeERAcRUZ: Jalapa, Potrero Viejo (about 10 mi. e. of
Cordoba), El Potrero (or Potrero) (near Cordoba), near Orizaba, Tezonapa,
Presidio (Tierra Caliente), Cuatotolapam, San Juan. Oaxaca: San Cristdbal
(Rio Valle Nacional), Cosolapa. Tasasco: Tenosique, Teapa. CHIAPAS:
La Esperanza (near Acapetahua). CAMPECHE:Tuxpena Camp, Aposote, En-
carnacion. YUCATAN: Chichen Itza, Mayapan, Puz, Ziz, and Gongora Caves
(near Oxkutzcab), Xkyc Cave (near San Bernardo), Mujeres Island. BritisH
Honpuras: Belize (type locality), Benque Viejo, Skates Lagoon, Silk Grass,
Stann Creek Valley. GUATEMALA: Departamento de Petén; Piedras Negras,
Uaxactun, near La Libertad, Poza de la Jicoteo.

Coleonyx elegans nemoralis subsp. nov.
Cottma BANDED GECKO
1905. Coleonyx elegans Gadow, Proc. Zool. Soc. London, Vol. 2 of 1905,
p. 212.
Type—No. 10,509, in the E. H. Taylor-H. M. Smith collection.
Collected at Hacienda Paso del Rio, Colima, Mexico, July 8, 1935, by Dr.

196 San Disco Society oF NaturAL History

Hobart M. Smith. This location is shown on most maps as Paso del Rio, at
the great bend of the Rio Armeria.

Diagnosis —This subspecies is a form having tubercular scales scattered
on the dorsum, thus differing from variegatus, brevis, and fasciatus. From
mitratus, nemoralis differs in having elongated scales forming the claw sheaths,
which completely, or almost completely, hide the claws, whereas in the more
southerly form the sheaths are shorter and the claws prominently in evidence.
From elegans elegans, nemoralis varies in having the upper prenasals less often
in contact, with a shorter juncture when there is a contact, and in having fewer
tubercular scales, especially laterally. Also, the mental is more nearly triangular
in nemoralis.

Description of Type—An adult male. The head is covered above and
below with non-imbricate, circular granules, larger anteriorly and interspersed
with tubercles posteriorly. The rostral is pentagonal in shape and is wider than
high. There are two short sides touching the first supralabials and two longer
edges contacting the prenasals. The apex is rounded. The supralabials number
7-5 and decrease in size posteriorly; the enlarged scales end anterior to the
center of the eye. The infralabials visible with the mouth closed number 6-5.
There are two prenasals on each side, the upper being considerably the larger.
The upper prenasals are, however, smaller than in. elegans elegans; they are
triangular in shape and are prevented from meeting on the median line by a
granular scale. The supranasals are not enlarged compared with other con-
tiguous scales; they are small and circular. The scales from nostril to nostril along
the prenasals number 8. The nostrils are large and oval. There are shallow
horizontal depressions back of the nostrils, the scales within which are especially
small. The scales edging the eyelids are rounded rather than serrated. The ear
openings are long and narrow. The mental is the largest head scale; it is equal
in width to the rostral. It is triangular in shape, with the sides posteriorly
convergent. The posterior end comprises a short circular arc, which is contacted
by 5 gulars. The mental plus the first infralabial on each side are contacted
by 11 gulars. The first infralabials are triangular, with ends pointed inward
along the mental. They are much deeper (but not so long) as the succeeding
infralabials.

The dorsum is covered by granules of similar size to those on the head,
among which there are interspersed enlarged tubercles in irregular rows. There
are about 19 of these rows at mid-body; adjacent to the mid-dorsal line there
are about 23 tubercles between centers of limb insertions. The tubercles are
larger than those on the head and neck. Dorsally, they tend to be slightly
keeled; some are peaked posteriorly. Laterally and ventrally the granules and
tubercles are replaced with flat, imbricate scales which become larger posteriorly.
The lateral scales are smallest at the limb insertions. There is a short umbilical
line which interrupts the regularity of the adjacent scales.

The scales in the preanal region are further enlarged and include a row
with conspicuous preanal pores. The latter total 10; the scales containing
them are acutely angular in arrangement with the point forward. The two
median scales at the apex are in contact.

KLAUBER—GENUS COLEONYX 197

The scales on the arms are imbricate above; the enlarged scales, which are
analogous to the tubercles, are not sharply differentiated from the rest. The back
of the hand is covered with imbricate scales; the palmar surface is granular.
On the lower surfaces of the digits there are series of rectangular transverse
lamellae, which form corrugated ridges. There are 14 lamellae on the fourth
finger. The fingers terminate in a pair of large shell-like laterals, capped by a
long wedge-shaped terminal. The claws are completely hidden by the sheaths.

The legs are covered with granular scales on the upper surfaces, inter-
spersed with conspicuous tubercles. The toes are sheathed like the fingers. There
are 17 lamellae on the fourth toe. The claws are concealed by the large laterals.

The cloacal spurs are comparatively small. They are placed on each side
of, and slightly posterior to, the vent; they are hardly more conspicuous than the
dorsal tubercles. In this character there is a notable difference from the much
enlarged spurs of the non-tubercular forms.

The tail is covered by annular rows of subrectangular and imbricate scales
which are largest ventrally. There are tubercles dorsally in every sixth or
seventh ring; these, however, do not occur on the regenerated posterior section.

The principal dimensions in mm. are as follows: Length, snout to vent 88,
head length 23, head width 16, rostral to mid-orbit 12, rostral to mid-ear 21,
width of orbit 5%, arm fully extended, measured to end of fourth finger 25,
leg fully extended, measured to end of fourth toe 30, length of tail (part

regenerated) 63, distance between points of cloacal bones 92.

The head is light-brown above, streaked with dark-brown. There is a
light bar across the snout, and concentric light and dark semicircular stripes in
the parietal region. The labials are alternately light and dark. The gular sur-
face is light.

The dorsum is barred, alternately chocolate-brown and buff. These marks
make up three triads between limb insertions. Each triad consists of two dark
bars with a light center. These light centers are narrower dorsally than laterally
and are more irregular than the light interspaces between triads, which are
widest dorsally. The lateral areas are considerably mottled. The ventrum is
unmarked. The limbs are brownish above and lighter below. The original part
of the tail is alternately ringed with buff and dark-brown; the regenerated
portion is mottled.

Range.—The coastal area of Mexico from Colima to southeastern Oaxaca,
where there is intergradation with Coleonyx elegans elegans.

Summary of Paratypes——Four paratypes are available, one of which is a
topotype. Of the others, two are from Agua del Obispo, Guerrero, and the
last is from 4 to 5 miles north of Acapulco, Guerrero. All are in the Taylor—
Smith collection.

Sixteen additional specimens are available from southern Oaxaca, particu-
larly from the vicinity of Tehuantepec, but these are to be regarded as elegans-
nemoralis intergrades, and will be discussed later. Thus only the Colima and
Guerrero specimens are to be considered paratypes; and even those from

198 San Dieco Soctery oF Natura History

Guerrero may be less differentiated from elegans elegans than the ones from
the extreme west. The type is included in the numerical figures given below.

The longest specimens, a male and a female, both measure 88 mm. from
snout to vent. The smallest is 53 mm. long, with a 54 mm. tail. No other
specimen has a complete tail.

The preanal pores (or pits in the females) number 7, 7, 9, 9, 10. They are
arranged in a sharply angular series.

The upper prenasals are separated by a scale in 4 specimens out of the five
available. (In elegans elegans only 4 out of 69 have these plates separated.)
The prenasals narrow dorsally; in the single specimen in which contact is made
the median suture is quite narrow. The supranasals are so small as to be virtually
undifferentiated from the adjacent scales above. The scales from nostril to
nostril number 6, 7, 8, 9, and 10. There are shallow postnasal depressions
covered with small scales. The apex of the rostral is rounded. The enlarged
supralabials number 5 (1), 6 (1), 7 (5), 8 (3); the second is usually the
largest. The mental is triangular but has a rounded posterior apex. The en-
larged infralabials number 6 (2), 7 (2), 8 (6). The first infralabials are
considerably deeper than the rest of the series, with long points carried inward
along the mental. The gulars contacting the mental number 3 (1), 4 (1),
5 (2), 6 (1); those touching the first infralabials are 3 (1), 4 (7), 5 (2).
Nemoralis mentals are more pointed posteriorly and with fewer gulars in con-
tact than is the case in elegans elegans.

There are 19 to 21 rows of tubercles, counting across at mid-body; and
23 to 32 in a line mid-dorsally between points opposite the limb insertions.
The laterals are long, as in elegans; only rarely is the tip of a claw visible.

The head pattern consists of a series of alternating light and dark longi-
tudinal streaks joined by semicircles posteriorly. The labials are mixed light

and dark.

The body pattern comprises three dorsal triads, each consisting of a pair
of narrow dark-brown transverse bands, with buff or light-brown between.
The light interspaces separating the triads are cream and are widest mid-
dorsally. As the lizards age the dark bands widen within the triads, until
they sometimes coalesce mid-dorsally, forming hour-glass shaped figures.
There are lateral irregular blotches. Dorsally the tubercles tend to become
darker than the ground color of the dark blotches in which they are placed;
while laterally the tubercles tend to stay light, thus spotting the sides, although
less conspicuously than in elegans elegans. The only individual with a complete
tail has 10 rings.

Trends and Relationships.—Since Coleonyx elegans has been collected in
the coastal areas of both the Gulf and Pacific slopes, but not on the plateau
between, it seems probable that they do not occur there. If a connection
between the Pacific and Gulf coastal populations exists today, it is probably
across the Isthmus of Tehuantepec, where the terrain seems suitable for an
uninterrupted range. From southeastern Oaxaca (the southern end of this
isthmus) there are 16 specimens available. Five out of the 16 have separated

KLAUBER—GENUS COLEONYX 199

prenasals. I think, therefore, they should be considered elegans-nemoralis inter-
grades. Their other counts are as follows:
Preanal pores or pits 6 (1), 7 (1), 8 (6), 9 (6), 10 (0), 11 (1).
Supralabials 6; @il)) 7-16) S- (4):
Infralabials 6-12) 577, (7), 8G):
Scales, nostril to nostril 5 (1), 6 (6), 7 (6), 8 (2), 9 (1).
Gulars contacting

mental Dh) O12) i anon S46), 29 (O)em10) (1):
Gulars contacting first
infralabials Dea) 3s (GE4) (2):

The longest specimen, a male, measures 96 mm. snout to vent. Where
the tail rings are complete they vary from 8 to 10. I note no pattern differences
in this series, either from elegans elegans or typical e. nemoralis. As usual, the
posterior tail rings are solid, rather than triads, and are complete ventrally. Two
juveniles from this area show rather unusual patterns; they have wide, dark
bands which are only slightly lighter interiorly. These differ from the ordinary
juvenile pattern of elegans, in which the bands are medium-brown, only
narrowly edged with dark-brown fore-and-aft.

Field Notes——Gadow (1905, p. 212) states that this gecko is distinctly
a forest form. He found it a few miles from the coast of Guerrero in a moist
patch of thick lowland forest on the ground under stones and rotten stumps.

“All the local geckos are much feared for their supposedly venomous
qualities, but this genus especially is abhorred. One specimen was found 2 feet
underground in the crevices of adobe brick ruins.” Hartweg and Oliver, 1940,
p. 14, discussing the intergrades found near Tehuantepec.

Locality Records—Typical nemoralis has been collected at the following
points: CoLtima: Hacienda Paso del Rio (type locality). GUERRERO: Agua
del Obispo and 4-5 mi. n. of Acapulco. One reported by Gadow (1908,
p. 439) from Pacific Camp, near Copala, Guerrero, is no doubt assignable
to this subspecies. The following from southeastern Oaxaca are considered
locality records of nemoralis-elegans intergrades: Tehuantepec (also 2 km. ne.),
Mixtequilla (also mountains nearby) , Tapanatepec, Tres Cruces, Cerro Arenal,
Cerro de Chipehua, Cerro de Guengola, Ranchero Poso Rio, Palo Colorado,
Chimalapa (Santa Maria).

Coleonyx mitratus (Peters)
CENTRAL AMERICAN BANDED GECKO

1863. Brachydactylus mitratus Peters, Mon. Berl. Acad. Wiss., p. 41. Type
specimen in the Berlin Museum. Type locality: Costa Rica.

1875. Coleonyx elegans Cope, Journ. Acad. Nat. Sci. Phila., 2nd Ser., vol.
8, p. 118.

1885. Eublepharis dovii Boulenger, Cat. Liz. Brit. Mus., Vol. 1, p. 233. Type
specimen in the British Museum. Type locality: Panama.

1893. Coleonyx dovii Stejneger, North American Fauna No. 7, p. 163.

200 SAN Dreco Soctety oF NaturaAL History

1928. Coleonyx mitratus Schmidt, Field Mus. Nat. Hist., Zool. Ser., Vol. 12
no. 16:1; 194:

Type.—The type specimen, in the Berlin Museum, has not been seen. It
was collected in Costa Rica. Peters’ description is quite adequate.

Diagnosis—A Coleonyx with tubercular scales scattered among the
granules of the dorsum, thus differing from variegatus, brevis, and fasciatus.
From elegans elegans and e. nemoralis it differs in having shorter scales sheath-
ing the claws; these are exposed in mitratus but concealed (or with only the
tips showing) in elegans. Also, the first infralabials are quadrilateral in mitratus
and triangular in elegans. Further, there is a longitudinal area of small scales
back of the nostril in elegans not present in mitratus.

Range.—Honduras, El Salvador, Nicaragua, Costa Rica, and Panama.

Material—I have examined the following specimens: Honduras 17, El
Salvador 2, Nicaragua 6, Costa Rica 5, uncertain 1; total 31.

Morphology.—A lizard of moderate body proportions, not especially
flattened. The snout is somewhat blunt and the neck long. The limbs are
relatively short but overlap when adpressed. The nostrils are circular. The
ear openings are large and vertically elliptical. The pupils are vertically ellipti-
cal. The eyelids are functional and are internally lined with black pigment.
The peritoneum is unpigmented.

There are four median dorsal ridges, the central two being the more evi-
dent. There is a mid-ventral umbilical line, which interrupts the regularity of
the adjacent scales. The tail is circular in cross section.

This species, similar to elegans elegans in size, is considerably larger than
variegatus variegatus, the largest of the non-tubercular forms. The longest
specimen at hand is a male from Nicaragua, measuring 91 mm. from snout to
vent. The longest female is 88 mm. The shortest specimen is 41 mm. The
tails, when complete, approximate the body in length.

The males may be recognized by the presence of preanal pores and post-
anal swellings. The females have scale pits, analogous to the preanal pores in
the males. The lateral cloacal spurs in this species are less prominent than in
variegatus; they are little more evident than the adjacent tubercles. They are
longitudinally flattened and ridged.

The variation in the number of preanal pores or pits is as follows: 5 (2),
6 (5), 7 (11), 8 (10), 9 (3); mean 7.23 + .19; coefficient of variation 14.6
per cent. The females have been included in these statistics. The pore arrange-
ment is similar to that of elegans, the two series making an angle of about 90°.
Males captured at the season of maximum activity have large pores almost filling
the scales they occupy. In these there is solidified a yellow transparent core.

The hemipenes are undivided with single sulcus. They are distally en-
larged, the outer part being covered with reticulated fringes.

Scalation—The head is covered above and below with circular granules,
mixed with enlarged tubercles. There are large scutes bordering the mouth
and nostrils, of which the rostral is the largest. It has two short sides touching
the first supralabials, and two longer sides contacting the prenasals. The apex

KLAUBER—GENUS COLEONYX 201

is rather blunt. There is a series of enlarged supralabials which reduce in size
posteriorly; they generally number 6 or 7, but vary from 5 to 9, with a mean
of 6.87 + .10. The enlarged supralabials are supplanted by granules back of
the center of the eye. There are two prenasals on either side, the upper much
the larger. The lower may be considered the nasal itself, since there seems to
be no suture between it and the nostril. The upper prenasals are in contact
medianly in all specimens. The supranasals are small and undifferentiated from
the other scales on the snout, nor are the rest of the scales bordering the nasal
orifice enlarged. The granules posteriorly in contact with the upper prenasals,
from nostril to nostril, number as follows: 6 (9), 7 (15), 8 (5), 9 (1); mean
6.93 += .14. There is a small postnasal depression. The rest of the head surface
is covered with circular granules which are considerably enlarged on the upper
surface of the snout, and where they touch the supralabials. Those which
contact the supralabials are often horizontally elliptical in shape. Interspersed
with the granules covering the head there are enlarged tubercles; these increase
in size toward the neck. There are a few tubercles below the angle of the
mouth, but not on the under surface of the head. The mental is large, with
the two sides usually somewhat convergent posteriorly. The posterior edge is
slightly convex, or may be angular. The enlarged infralabials usually number
6 or 7; the range is 5 to 8, and the mean 6.68 + .08. The first infralabials
are generally quadrangular in shape, rather than triangular as in elegans, the
two sides contacting the mental and second infralabial being substantially
parallel, although the side touching the mental is considerably the longer of
the two. The subsequent infralabials are rectangular and decrease in size
posteriorly; the enlarged scales terminate somewhat behind the center of the
orbit. The granules contacting the mental and infralabials are considerably
enlarged, compared with those toward the center of the lower jaw. The scales
touching the mental number as follows: 4 (1), 5 (10), 6 (8), 7 (8), 8 (2),
9 (1); mean 6.10 + .21. Those touching the first infralabials number 2 (1),
3 (44), 4 (14); mean 3.22 + .06. There is duplication in these tallies, for
one scale in each instance touches both the mental and one first infralabial.

The neck and dorsal surface of the body are covered with granules inter-
spersed with tubercles, becoming larger posteriorly, and more closely spaced
laterally. There are about 21 to 23 irregular rows of tubercles across the
dorsum. Some tubercles are keeled and posteriorly pointed. The belly is
covered with flat, imbricate scales, which increase in size posteriorly; they are
especially enlarged in the area of the series carrying the preanal pores, beyond
which they decrease in size rapidly to the anal opening.

On the tail the dorsal granules of the body are replaced with rings of
imbricate scales, which are enlarged ventrally. Dorsally, a few of the rings
(about one in six) include dorsal tubercles. Regenerated tails lack tubercles.

Tubercles are evident on the posterior edges of the forearms and the upper
surfaces of the legs; they are especially profuse on the latter. The posterior
parts of the upper arm and thigh are covered with granules.

The hands and feet have imbricate scales above and tubercular granules
below. The fingers and toes taper slightly; they are covered with imbricate

202 SAN Disco SociETY OF NaturRAL History

scales dorsally, and a single row of lamellae below. The latter are wider than
long; they are imbricate and form a corrugated row. There are about 13 or
14 lamellae on the fourth finger and 16 to 18 on the fourth toe. The claws in
preserved specimens are clearly in evidence, thus differing from e. elegans and
e. nemoralis, in which they are hidden by the elongated sheaths. The sheaths
in mitratus comprise a pair of shell-like convex lateral scales, partly capped
above by a pointed terminal scale. They are relatively shorter than in elegans.

Pattern and Color—lIn the juvenile stage mitratus has a pattern com-
prising three body bars between limb insertions, separated by narrow cream or
buff cross bars. The bars may be either uniformly dark-brown or triads with
lighter centers. There is a parietal light loop ending at the eyes, and a light
stripe across the snout. The lateral areas are lighter, while the lower surfaces

are immaculate buff. The tail is banded with wide dark bars, of which the

posterior are complete ventrally. The legs are punctated.

In the adults a considerable pattern change occurs. The body bars become
indented laterally and the sides mottled; the same change takes place in the
tail. In specimens from some areas the dark bars maintain their identity; but
in others, especially those from Honduras, they become much spotted anter-
iorly. The head becomes spotted or mottled dorsally, and the parietal light
loop, while retained, is more irregular. The light cross bar on the snout is
almost lost in the general spotting. The sides of the head are irregularly
mottled. The legs are speckled or spotted. The rostral is light in the center
and dark laterally. The lower surfaces of the head and body are clear, except
that the infralabials are blotched.

While there are nearly always three body bands or triads, four have been
noted on one specimen from El Salvador. There are usually 9 or 10 bands
on the tail, if complete.

Intraspecific Trends and Relationships——The specimens from the vicinity of
San Pedro, northwestern Honduras, are somewhat different from the geckos
of Nicaragua and Costa Rica, and, were adequate material available from the
latter areas, a subspecific segregation might be warranted.

In the Honduran specimens the sides of the mental are more convergent.
The rostral is slightly more pointed and the proportionate median contact
between the upper prenasals somewhat reduced. They have, on the average,
fewer scales in contact with the upper prenasals, between the nostrils.

It is in pattern that the differences are most evident. In the juvenile
specimens, those from Honduras have body triads comprising light-brown
bands between dark edges, while in a juvenile from Costa Rica the blotches
are quite dark throughout.

In the adults the Honduran individuals are usually light, with the triads
much spotted interiorly; while the specimens from Nicaragua and Costa Rica
tend toward darker and more unicolor dorsal bands.

It may be noted that all these differences indicated a directional trend
within mitratus toward elegans. Assuming that these geckos are distributed
throughout the lowlands bordering the Caribbean Sea, it is not impossible that

KLAUBER—GENUS COLEONYX 203

intergrades between mitratus and elegans elegans may be found around the
head of Lago de Izabal in eastern Guatemala (from which area no specimens
are available) , although the divergence in the claw sheaths, first infralabials, and
postnasal depressions, renders this doubtful.

Boulenger, at the time he described dovi, placed mutratus Peters in the
synonymy of elegans (1885, p. 235). But Peters’ description of the claws
and the mental leaves no doubt that the type of mitratus belongs to the clawed
form which ranges from Honduras to Costa Rica, and this is further verified
by the type locality (Costa Rica). Thus, unless the geckos of Panama show
an unexpected difference from those of Costa Rica, dovii must be placed in
the synonymy of mitratus. Neither Boulenger’s description, nor Gunther’s
plate (1893, plate 31) indicates that there is such a difference, but the final
decision must await the availability of more material from Panama. So far as
I know, the type of dovi is the only specimen that has been collected there.

Field Notes—Cope (1879, p. 217) reports that Zeledon collected
mitratus in ant hills on the tableland near San José, Costa Rica.

Mr. Karl P. Schmidt has kindly permitted me to abstract the field notes
of his collecting trip to Honduras in the spring of 1923. Altogether, 17 speci-
mens of mitratus were taken, 13 at Hacienda Santa Ana west of San Pedro,
the rest at Lake Ticamaya. All were caught at night on the ground; those from
the former locality were mostly along a path following a hydroelectric penstock
down a hill from 1000 ft. to 500 ft. elevation. The vegetation was scrubby,
low forest interspersed with cohune palms. At Lake Ticamaya there were some
larger trees as well. Here the geckos were found in dry leaves on the forest floor.

One specimen had a miller in its mouth. On another evening two were
caught only a foot apart.

Locality Records —Honoburas: San Pedro, Hacienda Santa Ana (west of
San Pedro at 800 ft. altitude), Santa Ana (near San Pedro), Lake Ticamaya
(east of San Pedro, between Rio Chamelecon and Rio Ulua), Progresso, and
Atlantida (Dept.). Ex Satvapor: Divisidero (Dept. Morazan). NicaraGua.
Costa Rica (type locality): San José, Tableland near San José, Turrialba,
Bebedero, Chica (= Chira?) Island (Gulf of Nicoya). PANAMA.

INTERGENERIC RELATIONSHIPS

I have given some thought to splitting the present genus Coleonyx, into
two genera, one to include the tubercular members, the other, those covered
with uniform granules. The gap between these two is consistent, both in
characters and range. However, the directive approach between nemoralis and
fasciatus in some characters finally deterred me from this step, notwithstanding
I do not believe nemoralis to be the most primitive of the tubercular group or
subgenus.

Of the latter I consider mitratus the most primitive, since the toe sheaths,
and the scales on the snout are less highly specialized than in the others.
Elegans elegans I regard as a derivative of mitratus. The former in turn, di-
verges on the two coasts, separated by the central plateau of Mexico, with

204 San Dreco Society oF NaturAL History

e. nemoralis of the Pacific slope deviating sufficiently to warrant subspecific
recognition.

I believe variegatus and brevis originally spread northward from a primi-
tive forest type not greatly different from fasciatus. I visualize them as first
spreading through the southwest from some center in central Mexico. The
juvenile patterns of both would suggest that abbotti is probably the present
form most nearly retaining the ancestral pattern. There was first a division
between variegatus and brevis, as they came northward over divergent routes,
followed by a further subdivision within variegatus. Although variegatus varie-
gatus today clearly comprises the central core of the variegatus group, through
which the other subspecies are interrelated, I do not consider variegatus varie-
gatus as having retained the greatest number of primitive characters. I am
of the opinion that differentiation by isolation in the several fringe populations
was effected after the area was largely inhabited by a fairly uniform population.
As these fringe populations gradually diverged from the ancestral type, and
therefore from each other, the central population, variegatus variegatus also
changed, in some particulars more than did the fringes. We may assume that
this resulted from the greater dessication of the central area. This course of
development would explain why contiguous populations are not always most
alike. Thus, utahensis is somewhat like sonoriensis, and some bogerti resemble
abbotti in pattern. Brevis is closer to sonoriensis than to its nearest neighbor
bogerti. The enlarged chin scales of slevini are more probably a reversion than
the retention of an ancestral character.

GENERIC DIFFERENTIATION

On the generic level Coleonyx may be distinguished from the genera with
which it has some superficial resemblance, by means of the following key:

1 a Two enlarged laterals on either side of the claw sheath _—Lepidoblepharis
1b A single enlarged lateral on either side of the claw sheath 2
2 a_ Digits with mixed or irregular scale formations on the ventral surface 3
2b Digits with a regular series of transverse lamellae below 4
3 a Tail with transverse constrictions forming a

series of corrugations Hemitheconyx
3 Tail without transverse constrictions Aeluroscalabates
4 a Tail with transverse constrictions

forming a series of corrugations Eublepharis
4b Tail without transverse constrictions Coleonyx

In some specimens of Eublepharis much of the tail has been lost, with or
without regeneration; in such cases the transverse constrictions will not be evi-
dent, since they are not present in regenerated tails, and even in original tails
are not prominent anteriorly. The following additional differences between
Eublepharis and Coleonyx may be mentioned: Eublepharis has a relatively
larger, and particularly a wider, head; the ears are larger and are edged with
tubercles; the terminals are less sharp at the ends; the preanal pores form a

KLAUBER—GENUS COLEONYX 205

more obtuse angle (greater than 135°); there are usually 2 gulars posteriorly
in contact with the mental, while Coleonyx seldom has as few as 3.

These generic differences are by no means exhaustive; I have only super-

ficially examined a few specimens of the genera other than Coleonyx.

KEY TO THE SPECIES AND SUBSPECIES OF Coleonyx

Dorsum covered with uniform granular scales 2
Dorsum with enlarged tubercular scales scattered among the granules 10
Usually three black body bands between limb

insertions; digits robust fasciatus
More than three brown body bands between limb insertions,

or dorsum spotted; digits delicate 3
Scale series carrying the preanal pores in the males*

divided medianly by the interposition of one or more

small scales; seldom more than 4 pores brevis
Scale series carrying the preanal pores in the males*

continuous across the median apex; usually more

than 4 pores 4
Usually 4 or fewer gular scales in contact

with the mental variegatus slevini
Usually 5 or more gular scales in contact with the mental 5
Preanal pores in the males usually number

8 or more variegatus bogerti
Preanal pores in the males usually number 7 or less 6

Dark transverse body bars in the adults considerably

wider than the light interspaces 7
Dark transverse body bars in the adults about equal
to, or narrower than, the light interspaces; or bars
obsolete, replaced by uniform spotting 9
A mid-dorsal light, longitudinal line usually splitting
the dorsal body bars in the adults varlegatus sonoriensis

No light mid-dorsal line splitting the dorsal body
bars in the adults 8
Adults with the longitudinal edges of the dark body
bars highly irregular, often confluent with spots in
the interspaces variegatus utahensis
Adults with the longitudinal edges of the dark body
bars even, with narrow, uniform interspaces variegatus peninsularis
Dark body bands in the adults unicolor; top of head

unicolor; nuchal light loop narrow and clear variegatus abbott

* Although only the males have true preanal pores, the corresponding scales in the

females are usually enlarged and are sometimes pitted.

206 San Dreco Society oF NaturaL History

9b Dark body bands in the adults with lighter centers,
producing a double barred effect, or bars obsolete
and replaced by spotting; top of head spotted;

nuchal light loop irregular or obsolete variegatus variegatus
10 a Scales of the claw sheath shorter, with claws con-

spicuously in evidence mitratus
10 b Scales of the claw sheath longer, with the claws

hidden, or only the tips in evidence 11
11 a Prenasals usually in contact; tubercles in greater

profusion laterally elegans elegans
11 b Prenasals usually separated; tubercles fewer in num-

ber laterally elegans nemoralls

ACKNOWLEDGMENTS

I am grateful to the following individuals and institutions for the loan of
specimens, and for other assistance without which this study could not have
been made: Mr. Charles M. Bogert, American Museum of Natural History;
Mr. Joseph R. Slevin, California Academy of Sciences; Mr. M. Graham
Netting, Carnegie Museum; Dr. Howard K. Gloyd, Chicago Academy of
Sciences; Messrs. Karl P. Schmidt and Clifford H. Pope, Chicago Natural
History Museum; Dr. A. H. Wright and Mrs. K. Kapp, Cornell University;
Dr. Ross Hardy, Dixie Junior College; Mr. Albert J. Kirn, Somerset, Texas;
Dr. Howard R. Hill, Los Angeles Museum; Mr. Stanley Mulaik, Salt Lake
City, Utah; Mr. Arthur Loveridge, Museum of Comparative Zoology; Mr.
Thomas L. Rodgers, Museum of Vertebrate Zoology, University of California;
Dr. Hobart M. Smith, University of Rochester; Miss Margaret Storey, Natural
History Museum, Stanford University; Dr. Doris M. Cochran, United States
National Museum; Dr. Raymond B. Cowles, University of California at Los
Angeles; Dr. Edward H. Taylor, University of Kansas; Mrs. Helen T. Gaige,
University of Michigan; Dr. Charles T. Vorhies, University of Arizona.

I wish to thank Mr. Robert Menzies for his courtesy in making translations.

I am greatly indebted to Messrs. C. B. Perkins and C. G. Abbott for

criticisms and editorial assistance.

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KLAUBER—-GENUS COLEONYX 209

GaicE, HELEN T.
1936. Some Reptiles and Amphibians from Yucatan and Campeche,
Mexico. Carnegie Inst. Wash., Pub. No. 457, pp. 289-304.
1938. Some Reptilian Records from Caves of Yucatan. Carnegie Inst.
Wash., Pub. No. 491, pp. 297-298.

Gravee:
1845. Description of a New Genus of Night Lizards from Belize. Mag.
Nat. Hist., Vol. 16, pp. 162-3.

GREENBERG, B.
1943. Social Behavior of the Western Banded Gecko, Coleonyx variegatus
Baird. Physiol. Zool., Vol. 16, no. 1, pp. 110-122.

GRINNELL, J. and Camp, C. L.
1917. A Distributional List of the Amphibians and Reptiles of California.
Univ. of Calif. Pubs. in Zool., Vol. 17, no. 10, pp. 127-208.

GUNTHkR, A. L. C. G.
1885-1902. Biologia Centrali-Americana: Reptilia and Batrachia, pp. xx +
326. (Coleonyx section dated 1893).

Harpy, Ross
1939. Some Notes on Utah Reptiles. Proc. Utah Acad. Sci. Arts,
Letters, Vol. 16, p. 83.
1944. Some Habits of the Banded Gecko in Southwestern Utah. Proc.
Utah Acad. Sci., Arts, Letters, Vol. 21, pp. 71-73.

Hartwes, N. and OLiver, J. A.
1940. A Contribution to the Herpetology of the Isthmus of Tehuantepec.
IV. Misc. Pubs., Mus. Zool., Univ. Mich., No. 47, pp. 1-31.

KLaAuBer, L. M.

1932. Amphibians and Reptiles Observed Enroute to Hoover Dam.
Copeia, No. 3 of 1932, pp. 118-128.

1934. Annotated List of the Amphibians and Reptiles of the Southern
Border of California. Bull. Zool. Soc. San Diego, No. 11,
pp. 1-28.

1939. Studies of Reptile Life in the Arid Southwest. Bull. Zool. Soc.
San Diego, No. 14, pp. 1-100.

La Rivers, Ira
1942. Some New Amphibian and Reptile Records for Nevada. Jour.
Ent. and Zool. (Pomona College), Vol. 34, no. 3, pp. 53-68.

LinspALe, J. M.
1932. Amphibians and Reptiles from Lower California. Univ. Calif.
Pubs. in Zool., Vol. 38, no. 6, pp. 345-386.
1940. Amphibians and Reptiles in Nevada. Proc. Am. Acad. Arts and
Saz:,,Vol.:73,,no. 8) pp: 1975257.
bre EShs JR.
1940. Amphibians and Reptiles of the Roosevelt Reservoir Area, Arizona.
Copeia, No. 4 of 1940, pp. 260-265.

210 SAN DreGco Society oF NatTuraAL History

LOvVERIDGE, ARTHUR
1936. African Reptiles and Amphibians in Field Museum of Natural
History. Field Mus. Nat. Hist., Zool. Ser., Vol. 22, no. 1, (Pub.
360) “pp. dt”

McKez, E. D. and Bocert, C. M.
1934. The Amphibians and Reptiles of Grand Canyon National Park,
Copeia, No. 4 of 1934, pp. 178-180.

MEEK, S. E.
1905. An Annotated List of a Collection of Reptiles from Southern
California and Northern Lower California. Field Columbian
Mus., Zool. Ser., Vol. 7, no. 1, (Pub. 104), pp. 1-19.

Mocquarp, F.
1899. Contribution a la Faune Herpétologique de la Basse-Californie.
Nouv. Arch. Mus. Nat. Hist., Ser. 4, vol. 1, pp. 297-344.

Mutalik, STANLEY
1935. Tail Regeneration in Coleonyx brevis Stejneger. Copeia, No. 3 of
1935, p. 155.

Nose, G. K.
1921. The Bony Structure and Phyletic Relations of Sphaerodactylus and
Allied Lacertilian Genera, with the Description of a New Genus.
Am. Mus. Novit., No. 4, pp. 1-16.

OrTENBURGER, A. I. and ORTENBURGER, R. D.
1926. Field Observations on Some Amphibians and Reptiles of Pima
County, Arizona. Proc. Okla. Acad. Sci., Vol. 6, pp. 101-121.

Pack, Ele J:
1920. Coleonyx in Utah. Copeia, No. 88, pp. 101-2.

Parker, H. W.

1926. The Neotropical Lizards of the Genera Lepidoblepharis, Pseudo-
gonatodes, Lathrogecko, and Sphaerodactylus, with the Description
of a New Genus. Ann. and Mag. Nat. Hist., Ser. 9, vol. 17,
pp: 291-301.

1930. Three New Reptiles from Somaliland. Ann. and Mag. Nat. Hist.,
Ser. 10, vol. 6, pp. 603-606.

1932. Two Collections of Reptiles and Amphibians from British Somali-
land. Proc. Zool. Soc. London, pp. 335-367.

Peters, W.
1863. Ueber einen neuen Gecko, Brachydactylus mitratus aus Costa Rica.
Monatsber. Akad. Wiss. Berlin, for 1863, pp. 41-44.

Ripcway, Rost.
1912. Color Standards and Color Nomenclature, pp. iv + 43, plts. 1-53.

KLAUBER—GENUS COLEONYX Mit

RuTHVEN, A. G.
1907. A Collection of Reptiles and Amphibians from Southern New
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1912. The Amphibians and Reptiles Collected by the University of
Michigan—Walker Expedition in Southern Vera Cruz, Mexico.
Zool. Jahrb., 32, Abt. f. syst., pp. 295-332.

SANDERSON, I. T.
1941. Living Treasure, pp. 1-290.

SAUNDERS, H. F.
1912. “Poisonous” Lizards in India. Jour. Bombay Nat. Hist. Soc.,
Vol. 21, no. 4, pp. 1340-1.

ScHmiIpT, K. P.

1922. The Amphibians and Reptiles of Lower California and the Neigh-
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pp. 607-707.

1928. Reptiles Collected in Salvador for the California Institute of Tech-
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251), pp. 193-201.

1941. The Amphibians and Reptiles of British Honduras. Field Mus.
Nat. Hist., Zool. Ser., Vol. 22, no. 8, (Pub. 512), pp. 475-510.

ScHmipT., K. P. and Smitu, T. F.
1944. Amphibians and Reptiles of the Big Bend Region of Texas. Field
Mus. Nat. Hist., Zool. Ser., Vol. 29, no. 5, (Pub. 550), pp. 75-96.

Smitu, H. M.
1933. On the Relationships of the Lizards Coleonyx brevis and Coleonyx
variegatus. Trans. Kan. Acad. Sci., Vol. 36, pp. 301-314.

SmitH, M. A.
1933. Remarks on some Old World Geckoes. Records of the Indian
Museum, Vol. 35, part 1, pp. 9-19.
1935. The Fauna of British India including Ceylon and Burma. Reptilia
and Batrachia; Vol. 2, Sauria, pp. xiii + 440.

STEJNEGER, LEONHARD
1893. Annotated List of the Reptiles and Batrachians Collected by the
Death Valley Expedition in 1891, with Descriptions of New
Species. North Am. Fauna, No. 7, pp. 159-228, 394-398.

STEPHENS, FRANK
1918. Some Southern California Reptile Notes. Copeia, No. 54, pp.
3435.
1921. An Annotated List of the Amphibians and Reptiles of San Diego
County, California. Trans. San Diego Soc. Nat. Hist., Vol. 3, no.
4, pp. 7-09:

212 SAN Disco Society oF NATURAL History

STRECKER, J. K.

1909. Reptiles and Amphibians Collected in Brewster County, Texas.
Baylor Univ. Bull., Vol. 12, no. 1, pp. 11-15.

1915. Reptiles and Amphibians of Texas. Baylor Bulletin, Vol. 18, no.
4 Ppa LOZ.

1922. An Annotated Catalogue of the Amphibians and Reptiles of
Bexar County, Texas. Scientific Soc. San Antonio, Bull. No.
AS pPsaleoL-

1928. Common English and Folk Names for Texas Amphibians and
Reptiles. Contrib. Baylor Univ. Mus., No. 16, pp. 3-21.

1929. Random Notes on the Zoology of Texas. Contrib. Baylor Univ.
Mus., No. 18, pp. 3-12.

1933. Collecting at Helotes, Bexar County, Texas. Copeia, No. 2 of
1933% PPa/ alo:

1935. A List of Hitherto Unpublished Localities for Texas Amphibians
and Reptiles. Baylor Bull., Vol. 38, no. 3, pp. 35-38.

STUART, EaG:
1934. A Contribution to a Knowledge of the Herpetological Fauna of
El Petén, Guatemala. Occ. Papers Mus. Zool., Univ. Mich.,
No. 292, pp. 1-18.
1935. A Contribution to a Knowledge of the Herpetology of a Portion of
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SUMICHRAST, F.
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de l’Isthme de Tehuantepec. Bull. Soc. Zool. France, Vol. 15,
pp- 162-190.

TPAVrORMEs Et.
1935. Coleonyx fasciatus, a Neglected Species of Gecko. Univ. Kan.
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1936a. Notes on the Herpetological Fauna of the Mexican State of So-
nora. Univ. Kan. Sci. Bull., Vol. 24, no. 19, pp. 475-503.
1936b. Notes on the Herpetological Fauna of the Mexican State of
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VAN DENBURGH, J.

1897a. The Reptiles of the Pacific Coast and Great Basin. Occas. Papers
Calif. Acad. Sci., No. 5, pp. 1-236.

1897b. Reptiles from Sonora, Sinaloa and Jalisco, Mexico, with a Descrip-
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Vol. 49, pp. 460-464.

1922. The Reptiles of Western North America. Occas. Papers Calif.
Acad. Sci., No. 10, 2 vols., pp. 1-1028.

1924. Notes on the Herpetology of New Mexico, with a List of Species
Known from that State. Proc. Calif. Acad. Sci., Ser. 4, vol. 13,
no. 12, pp. 189-230.

KLAUBER—GENUS COLEONYX 213

VAN DENBURGH, J. and SLEvin, J. R.
1913. A List of the Amphibians and Reptiles of Arizona with Notes on
the Species in the Collection of the Academy. Proc. Calif. Acad.
Sci., Ser. 4, vol. 3, pp. 391-454.
VoruHIes, C. T.
1917. Poisonous Animals of the Desert. Agric. Exper. Station, Univ.
Ariz., Bull. 83, pp: 233-392:
WALLS, G. L.
1942. The Vertebrate Eye. Bloomfield Hills. Pp. xiv + 785.
WERNER, FRANZ
1896. Beitrage zur Kenntniss der Reptilien und Batrachier von Central
Amerika und Chile, sowie einiger seltener Schlangenarten. Verh.
Ges. Wien, Vol. 46, pp. 344-365.
1910. Ueber neue oder seltene Reptilien des Naturhistorischen Museums
in Hamburg. II Eidechsen. Hamburg Jahrb. wiss. Anst. 27,
pp. 1-46.
1912. Eublepharidae, Uroplatidae, Pygopodidae (in Das Tierreich, Lief.
33) (pp.1x ay 33%
WETTSTEIN, OTTO
1934. Ergebnisse der osterreichischen biologischen Costa Rica—Expedi-
tion 1930. Akad. Wiss. Wien, Vol. 143, pp. 1-39.
WELLBORN, V.
1933. Vergleichende osteologische Untersuchungen an Geckoniden, Euble-
phariden und Uroplatiden. Sitz. Gesel. Natur f. Fr. Berlin, Nos.
1-3, pp. 126-199.
Woopsury, A. M.
1931. A Descriptive Catalog of the Reptiles of Utah. Bull. Univ. Utah,
Vol. 21; no. 5; pp. x +7129:
Yarrow, H. C.
1883. Check List of North American Reptilia and Batrachia, with Cata-
logue of Specimens in U. S. National Museum. Bull. U. S. Nat.
Mus., No. 24, pp. 1-249.

SUMMARY

Coleonyx, a genus of geckos inhabiting the southwestern United States,
Mexico and Central America, has been surveyed. There are two groups, or
subgenera, one in which the bodies are covered with rather uniform granules,
the other having enlarged tubercular scales scattered over the dorsum. The
former occupies the southwestern United States and northern Mexico; the
latter southern Mexico and Central America. As far as is now known, the
two groups do not overlap territorially. Of the non-tubercular group there are
three species, variegatus, brevis, and fasciatus. Six new subspecies of variegatus
are described: slevini, peninsularis, abbotti, utahensis, bogerti, and sonoriensis.
There are two tubercular species: elegans and mitratus. A new subspecies of
elegans, nemoralis, is described. Ranges and field notes are set forth, and
relationships are discussed.

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TRANSACTIONS oP re080n
OF THE ME ike RA

SAN DIEGO SOCIETY OF NATURAL HISTORY

VOLUME X, No. 12, pp. 217-236, map

THE TRANSVERSE VOLCANIC BIOTIC PROVINCE
OF CENTRAL MEXICO AND ITS
RELATIONSHIP TO ADJACENT PROVINCES

RosBert T. Moore

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY
Aucust 31, 1945

110° 105° 100°

| SIX BIOTIC PROVINCES
| OF
MEXICO

CHIHUAHUA

PR : FAUNAL DISTRICTS
SINALOA OPATA

TARAHUMARE
TEPEHUANE

ALAMOS

TEBACA

SINALOA COASTAL
JALISCAN

OTOMI

TARASCAN

AZhEG
ORIZABA-ZEMPOALTEPEC

SIERRA MADRE OCCIDENTAL
CHIHUAHUA DESERT
IZ SIERRA MADRE ORIENTAL

MZ TRANSVERSE VOLCANIC

=OoeeProusuh —

MI SIERRA MADRE DEL SUR

105 100° 95°

THE TRANSVERSE VOLCANIC BIOTIC PROVINCE
OF CENTRAL MEXICO AND ITS
RELATIONSHIP TO ADJACENT PROVINCES*

Bid Ko <
2 g $Y g Ropert T. Moore ( sek

12 19409

Students of biotic relationships have overlooked the importance
of the great volcanic area, which stretches across the southern end of
the Central Plateau of Mexico, extending west and east to within fifty
miles of the Pacific and Atlantic Oceans respectively. Perhaps this is
due in part to a lapsus in our maps, which seldom have shown a name
for this range, but which is known and feared by the Indians as the
Sierra de Anahuac. The maps give one an impression of a series of
unimportant isolated peaks. The truth is quite different! Here vulcan-
ism has reached its zenith of power and destruction. It has created
hundreds of mighty cones, many extinct, massed and concentrated
transversely across an enormous volcanic belt, four hundred miles long
by sixty wide. Pumice, tuff and lava have been poured over the region
again and again in enormous quantities so that the valleys and pockets
of this range are conditioned by an entirely different set of environ-
ments from those of the two north-south sierras, and zoological life
within them has been profoundly and continuously modified by the
ever changing conditions for more than a million years. It is true that
there are scattered volcanoes in other areas of Mexico, but were
they lumped together, their total effect upon wild-life would not com-
pare with that of the mighty Anahuac.

According to Schuchert (1935, p. 129), “this mountainous area
of southern Mexico ..... extends from Cape Corrientes on the west
coast to Jalapa, Veracruz...... It has a wild rugged topography due
Shae ae to the vast quantities of volcanic material that have been ejected
from numerous vents and piled upon it: lavas, tuffs, and pumice, to-
gether with deposits of Pliocene and Pleistocene lakes.” Thayer (1916,
pp. 89-90) writes: “It appears that there have been two periods of
vulcanism in this province: the first in the Miocene and affecting

* Contribution from the California Institute of Technology, Pasadena, Calif.

220 SAN Disco Society oF NatTurAL History

all of Mexico; the second beginning in the Pliocene and continuing
to the present time and limited to this province.” Again Schuchert
(1935, p. 133) writes that in the late Pliocene and the Pleistocene “came
the very great epeirogenic elevation, which produced present Mexico,
elevating the land 3000-4000 feet in the north and 7000-8000 feet south
of Mexico City.” It was partly during this period that the great trans-
verse volcanic southern front was raised above the Central Plateau of
Mexico, which Thayer and Schuchert term the “Anahuac Plateau.”

The tremendous display of vulcanism, which persisted for a vast
period of time, not only continued to raise this front much higher, but
also produced the gigantic volcanic cones which exist today. The new
volcano, Paricutin, which arose from a cornfield in central Michoacan
and which was visited by the author when it was only a few weeks old
in March, 1942, and again six months later, although it has affected
an area over three hundred miles in diameter with a great deposition
of ash, can be classed, at least up to the present time, as only a
minor disturbance in the continuous transformation of the area for the
past two million years by much greater volcanoes of the region. High
as these are, reaching an altitude of 18,250 feet in the case of Mount
Orizaba, they are merely “stubs of former peaks, which have been re-
duced by erosion.” Thirteen of them range above 12,000 feet in alti-
tude and only three others higher than 12,000 feet are located outside
of this region; one of them, Mount Zempoaltepec in southeastern Oax-
aca, is deemed to be within the southern boundary of the province, from
the zoological viewpoint, making the fourteenth over 12,000 feet. An-
other, Mount Mohinora in extreme southwestern Chihauhua, which was
ascended and its birds collected in 1937 by the author, is believed by
him to be slightly lower than 12,000 feet from the performance of his
aneroid, although airplane pilots claim it is nearer 13,000.

The effect of the deposition of ash, not to speak of the enormous
quantity of igneous vapors constantly pouring into the atmosphere, on
the mammals, birds and other creatures of the area, can hardly be ex-
aggerated. On his visits to Paricutin, as well as on his ascents either
to the summits (or nearly to them) of three others of the great vol-
canoes of the region, the author personally has noted the disastrous
results of volcanic phenomena on bird-life and mammals. At Morelia,
nearly 80 miles as the crow flies from Paricutin, the trees and plants
appeared as if a black blizzard had struck them, every leaf being covered

with ash to a depth from one to two inches. At Patzcuaro, sixty miles

Moore—Brotic Provinces oF Mexico 221

away, the marginal waters of the lake had a consistency of heavy black
cream, a dense shroud of fine ash blotted out the sun and vision was
restricted as at twilight. Seed-eating birds hunted futilely over the
blanket of ash and insect-eaters found the leaves and trunks of trees
so covered that the dead remains of insects beneath could not be dis-
interred. As we approached the volcano, the blanket of ash increased
in depth until, at a distance of five miles, it averaged three feet and
everything had the appearance of death, with the exception of the black-
ened exterior needles on outer branches of pines. Such birds as sur-
vived were obviously weakened, while nests had been destroyed, so that
the effect upon them must have been profound. Yet a survival
characteristic seemed to be present in these sturdy residents. During
the first few months, when the ash plume was blown steadily toward
the east, residents, such as bluebirds, on the west side were observed
fearlessly carrying on their activities within a half mile of the belching
cone.

Fortunately the region receives summer rains and the ash is fertile.
The highly elevated region at the southern end of the plateau serves
as an enormous dam, four hundred miles in length, which has forced
its streams to flow towards the north and they in turn have developed
the great river system of the Lerma, the longest wholly contained within
the country. Here are the finest lakes of Mexico, one of them, Lago
de Chapala, being the largest of all, eighty miles long by thirty-five wide.
This fluvial system with its marshes is inhabited by interesting endemic
forms of bird-life, and, when its birds have been studied more thoroughly,
may be classed as a sub-faunal district.

The author deems this area, to which he has given the name,
“Transverse Volcanic,” one of the major biotic provinces of Mexico. In
fact, the preliminary statistical appraisal of our specimens from the area
would indicate that it may be the most important province of all, at
least in respect to the number of endemic forms occurring within it.
The author has made a survey, admittedly provisional, of all the bird
forms found within the boundaries of Mexico and has completed a
more careful study of those breeding within this province and the ad-
jacent provinces on each side. By a conservative estimate it would
seem that at least eighty forms are confined to the province, whereas
less than sixty are limited to the next largest biotic province, the Yuca-
tan Peninsula, and less than fifty to the third largest, the Veracruz.
Further study with more adequate material may either increase or reduce

222 SAN Disco Society OF NaATuRAL History

these figures, because new forms will be described which will be found
confined to the Transverse Volcanic and some, which now appear to
be confined to it, will prove to range into other provinces. The pro-
portions between the number of endemics probably will not be greatly

changed.

The author has ventured to make this preliminary and obviously in-
adequate study of the Transverse Volcanic Province and its ornithological
relationships to adjacent provinces, because of the truly great amount
of fresh material from it alone, now available to him in our collection.
This amounts to approximately fifteen thousand specimens of birds
alone, all collected during the past eleven years by the following Mexi-
can and American collectors: Mario del Toro Avilés, Pablo Roveglia,
and the American collectors Wilmot W. Brown and Chester C. Lamb.
Mr. Brown made special trips for the author in the middle 30’s to two
of the famous type localities of Swainson—Real del Monte, Hidalgo and
Temascaltepec, in the state of Mexico. Each collection consisted of more
than five hundred specimens, beautifully prepared, and the latter one
brought to light the diverse zonal characteristics of the Temascaltepec
region, part of it tropical and part temperate. By far the largest collection,
approximately ten thousand specimens, was amassed by Chester C.
Lamb, who, beginning with the year 1938, established a large num-
ber of collecting-sites over the entire province and for over six years
has made his home at Irapuato in the northern part of the province.
The only other large collection of birds obtained from the prov-
ince was that of Nelson and Goldman, who traversed it rather rapidly,
but displayed extreme acumen in the species they collected, which result-
ed in the description of many new forms. Theirs must be considered the
basic collection, although it does not nearly equal that of Mr. Lamb.

The most important analysis of the birds of any portion of the
province is that by Blake and Hanson (1942), who confined their work
to one mountain and its southern basal extension in Michoacan, that of
the Cerro de Tancitaro. This is an excellent paper and provides a great
deal of information, especially regarding the zones of Tancitaro and
the forms inhabiting them. That it is not completely adequate, is not
due to the fault of the members of the expedition nor to the authors,
but to the paucity of the series of birds collected, which amounted to
only 481. The high level of this paper is all the more remarkable when
one realizes that only an average of about three individuals, for each
one of the 144 forms treated, was taken by the collectors. This ac-

Moore—Brotic Provinces oF Mexico 223

counts in part for the uncertainty of the identification of some material.
Apparently the authors did not know that, by the time their first col-
lection was made, Mr. Lamb had already amassed several thousand
specimens from the state of Michoacan, and had collected every form
mentioned by Blake and Hanson (1942, p. 519) except some of their
“West Mexican Arid Tropical Fauna” forms, which I assign to the
Nayarit-Guerrero Province of the west coast, where Lamb has collected
them. In fact, it is only fair to Mr. Lamb, and not derogatory to Blake
and Hanson’s exceedingly fine article, to state that his collection con-
tains many forms not taken previously by anyone, such as Otus asio
sortilegus and Otus vinaceus seductus and others not mentioned herein,
as only ones requiring mention for the purposes of this paper are now
considered. More than twenty-five thousand specimens have been col-
lected by him in the Transverse Volcanic Province and in the other most
important of the adjacent provinces, specifically considered herein, such
as the Sinaloa, the Sierra Madre Occidental, the Chihuahua, the Sierra
Madre Oriental, the Tamaulipas and the Veracruz.

The present paper does not pretend to be a complete analysis of
this vast collection, but most series of critical forms have been examined
with reasonable care. The author regards it merely as a preliminary
effort to stimulate future research in the provinces and biotic districts
of Mexico. Undoubtedly, some of the conclusions and statements will
have to be modified in the light of more adequate material obtained
later and the author himself may amend some of them, when he has
examined more thoroughly the entire collection under his control.

Acknowledgment is herewith made of how deeply the author is
indebted to many museums and individuals for their courtesy in per-
mitting him to examine their material, these including the United States
National Museum, the Biological Survey, Museum of Comparative
Zoology and the American Museum of Natural History. Special
thanks should be given to Dr. Herbert Friedmann and Dr. Alexander
Wetmore of the United States National Museum; also to Major E. A.
Goldman and Mr. John W. Aldrich of the Fish and Wildlife Service
(formerly the Biological Survey), and Mr. George Willett of the Los
Angeles Museum. Mr. James L. Peters of the Museum of Comparative
Zoology was very generous in permitting the author to employ the
nomenclature adopted by him in his unpublished manuscript of the

Trochilidae for the Birds of the World.
Major Goldman’s assistance should be emphasized. It hap-

224 SAN Deco Society oF NATURAL History

pened that this article already was well advanced in manuscript
form when Major Goldman and the author decided to join in a sep-
arate paper on “The Biotic Provinces of Mexico,” which is now in the
hands of the publisher. When the author called attention to his proposal
to recognize the Sierra de Anahuac volcanic area as a separate “Trans-
verse Volcanic Province,” Major Goldman approved the decision as
well justified by the mammalian data and agreed to insert it as
one of the most important biotic provinces to be considered by our
joint paper. Since then Major Goldman has given generously of his
advice. Both of us have been astonished at the closeness of the bound-
ary lines of the provinces of Mexico, when determined independently
by the two authors from the standpoint of their ornithological, mamma-
lian and botanical life respectively. Major Goldman should be given
credit for names of provinces, mentioned in this report, except that of the
Transverse Volcanic. Throughout the tables only the reasonably cer-
tain breeding ranges of birds are considered and winter records are dis-
regarded. On the map, solid red lines enclose biotic provinces; broken
red lines mark off faunal districts within the provinces.

As to the affinities of adjacent provinces, the Sierra Madre del
Sur Province has very much in common with the Transverse Volcanic,
but the same is true only in a less degree of the two other adja-
cent high altitude provinces of Mexico, the Sierra Madre Occidental
and the Sierra Madre Oriental. As will be noted in Tables 3 and 4, of
the fifty-two forms selected because they occur in only two or three
provinces, fifteen are common to the Transverse Volcanic and the
Sierra Madre del Sur and to no others, seven are common to the Trans-
verse Volcanic and the Sierra Madre Oriental and to no others, and
eight are common to the Transverse Volcanic and the Sierra Madre
Occidental and to no others. Another adjacent province of fairly high
altitude, that of the Chihuahua of the Central Plateau, has seven forms
common to it and the Transverse Volcanic and to no others. The only
other high altitude province of Mexico is the far distant Chiapas High-
lands, which is isolated, and an analysis of its forms indicates that it
has less in common with the Transverse Volcanic. The same is even
more true of all the lowland provinces, the Nayarit-Guerrero Coast-
al Province and the Tehuantepec Province, they seeming to have no
form proved to be in common with the Transverse Volcanic, which is not
also common to at least one other province. Therefore, it would seem rea-
sonable to deduce from the above, that in spite of the fact that the Trans-

Moore—Brotic Provinces oF Mexico 225

verse Volcanic has within its borders Arid and Humid Tropical Zones,
this province may be characterized as having its chief affinities with
provinces ranging from 5000 feet in altitude up. The author does not
consider this study in any sense complete, so he will attempt very few
other deductions from the data supplied in the tables. One reason for
this decision is that the tables do not include the very large number of
high altitude forms of northern origin, which are found resident in more
than three provinces of Mexico, nor any of the migrant forms. Even
without this important unlisted northern element, it will be noted that
a great many of those listed as confined to the Transverse Volcanic in
the two tables, must have originated in the north.

Seemingly this paper constitutes the first attempt to delimit the
faunal districts of the province from an ornithological point of view,
if not from any other. In making this statement, the author is fully
acquainted with the pioneer work of students of other kinds of zoolog-
ical life, such as Hobart M. Smith (1940) in his “An Analysis of the
Biotic Provinces of Mexico, As Indicated by the Distribution of the
Lizards of the Genus Sceloporus,” and the study made by Nelson, as
quoted by Merriam (1895), showing the remarkable distribution of
the pocket gophers. It is possible to divide this province into seven
separate faunal districts and one of these, the Mount Zempoaltepec
region, herein believed to be only a sub-faunal district, may
eventually be given higher rank. Five forms are confined to this
last area, but for several reasons the author has deemed it best to in-
clude it tentatively with the birds of the Mount Orizaba area, to be
known as the Orizaba-Zempoaltepec Faunal District, and for the pres-
ent to consider the province as having only five faunal districts.
As this paper deals exclusively with birds, it seems better to employ the
term “faunal district” rather than “biotic district.”

Regarding the names given to the faunal districts, as well as to the
province itself, the author has followed a conviction which he has had for
some years, that names should be chosen with care and from the
following categories of available ones, in the order listed:

(1) Names of sedentary Indian tribes, whose boundaries correspond
fairly well with those of its zoological life. In making such a choice, it is well
to avoid the names of tribes which were or are of a nomadic nature, chang-
ing their locations frequently, since such names sound out of place in
studies of biotics, which deal more with sedentary than with transient
or migratory life. However, the employment of some aboriginal names, which
would meet the above requirements, might lead to confusion, as would be true

226 SAN Disco Society oF NaturaAt History

of the designation “Mexicano,” if one followed Lumholtz’ map of the Indian
tribes of Mexico, to which name he seems to refer the large group of Nahuatls,
including many component divisions, such as the Aztecs.

(2) Names of important topographical features of the country, such as
the four great sierras of Mexico. Considered under this category, the best name
for the province is “Sierra de Anahuac,” an ancient one given to the great
transverse volcanic range by its natives. Unfortunately, the name has been
employed by geologists for the main Central Plateau of Mexico and its use in
connection with the Transverse Volcanic Biotic would create endless confusion.

(3) Names descriptive of important physical characteristics of the whole
province. The name “Transverse Volcanic” was chosen on this basis.

(4) Names of political states are as a rule undesirable, but when their
boundaries coincide fairly well with that of the province to be named and no
name of the previous three categories is available or deemed desirable, such a
political name may be employed. When possible, it is well to avoid names that
are not of native origin, some designations of Mexican states, such as
“Durango,” being taken from political entities in Spain.

The names “Tarascan,” “Otom1” and “Aztec” for districts were
chosen under category (1) and seem particularly appropriate, since
the habitat of each tribe has almost the same limits as that of the zoolog-
ical life. The name “Mexicano” for the extreme western faunal district
of the province would have been confusing, not only with the name of
the country, but also with that of the state of Mexico, which lies in a
different part of this same province. In this case, the political name
“Jalisco” appears to be the least inappropriate.

The inner boundary lines of the faunal districts are tentative and
may be changed when a more complete analysis is possible of the avail-
able material. The greatest difficulty was encountered in determining
the eastern boundary of the Jaliscan district and the western boundary
of the Tarascan, since this line intersects a region, which is characterized
by intergrades in many plastic species.

We might conclude from Table 2 that the Orizaba-Zempoaltepec
District, with twenty-one forms confined to it and a total of forty-one
of the eighty-three endemic forms of the province occurring in it, is the
most important. Of these twenty-one, however, none represents a
genus and only two are full species. But, if this district should be di-
vided ultimately into two, the resulting new districts would each be
no stronger than the Aztec. The second most important district is
the Aztec with ten forms confined to it, consisting of one genus
and one full species, with a total representation of thirty-six of the

Moore—Brotic Provinces oF MExico 227

endemic forms of the province. The Jaliscan District is third in
importance with nine endemic forms. The Otomi District with only
three forms confined to it and only fifteen of the total endemic forms of
the province is unquestionably the weakest, and also probably has the
lowest average altitude with no very high mountains in it. Perhaps
its endemic weakness would be expected, because of the almost entire
absence of the higher zonal elements of the rest of the districts.

Some additional deductions may be made, provided one keeps
in mind the tentative nature of a report of this kind and the additional
information that may be obtained from more thorough collecting and
that is, that of the eighty-three forms confined to the province, four
appear in all five districts and not in any other provinces, and an
interesting total of eight appear in the three districts, the Tarascan,
Aztec and Orizaba-Zempoaltepec, which have the highest mountains
of the province. Mention has been made previously that the Trans-
verse Volcanic Biotic apparently has more affinities with the Sierra
Madre del Sur Province, than with any other province in Mexico and
in a lesser degree has important affinities with the Sierra Madre Orien-

tal, the Sierra Madre Occidental and the Chihuahua Provinces.

Table 3 gives statistics of one method of showing the relationships
between the Transverse Volcanic Province and the four other provinces
mentioned above. This Table disregards all forms which appear in
more than three provinces, in order to eliminate wide-ranging species
that are not greatly subject to the effects of environmental changes.
The author has not yet analyzed the component elements of these other
four provinces to the extent that he has the Transverse Volcanic, but this
much may be deduced on the evidence to date that, next to the Trans-
verse Volcanic, the Sierra Madre Occidental is the richest in endemic
bird-life, both in numbers and rank. Of the forty-five forms restricted to
the Sierra Madre Occidental, one is a genus Xenospiza; another genus,
Cyanocorax, occurs nowhere else north of Costa Rica; and of a third
genus, Otophanes, only a single specimen has been taken in any other
province and that probably not a breeding bird. Very little collecting
had been done in the Tepehuane Faunal District of the Sierra Madre
Occidental Province prior to the inception of our collecting program
in this massive mountain range. The veteran collectors, Nelson and
Goldman, to whom so much of our knowledge of Mexican zoological
life is due, crossed this range in one or two places, but were moving
rapidly and, although new forms were discovered, the total number of

228 SAN Dreco Society oF NAturRAL History

specimens taken was not large. That most expeditions, even that of
J. H. Batty in 1903, did not do extensive work in the higher portions
of the ranges, is indicated by the failure to obtain specimens of such
very remarkable endemic forms as Asio stygius lambi, Otophanes mcle-
odi and particularly such a conspicuous jay as Cyanocorax dickeyi. In
the spring of 1931, Alfred M. Bailey and H. B. Conover (1935), al-
though they were only a few days in the foothills of the Sierra Madre
Occidental west of Durango City, Durango, secured specimens which
provided the basis for the description of the new species and genus,
Xenospiza baileyi. The only intensive collecting that has ever been
done in the Tepehuane District is that of C. C. Lamb, who made his
first camp-site on May 20, 1934 at Santa Gertrudis, Sinaloa, at an
elevation of 6200 feet near the northern limits of the Tepehuane Dis-
trict. During the next four years, at least twenty collecting stations
were established along the higher parts of the range, covering the length
of it throughout Sinaloa and as far south as the vicinity of the city of
Tepic, Nayarit. Between four and five thousand specimens were taken in
this district by Mr. Lamb or my associates, but the great majority by
Lamb.

There is not nearly so firie a representation anywhere of the bird-
life of the Tarahumaran District to the north. Save for our own fair-
sized collections from the southern part of this district made by Mr.
Lamb at a few stations and by myself on a separate expedition to the
Barranca del Cobre, the only other major collections worth men-
tioning have been that of M. Abbott Frazar in 1887 and 1888,
the small collections made by John C. Cahoon about the same time, and
by R. R. McLeod from 1883 to 1885, as reported by van Rossem (1934).
Only Frazar and McLeod worked in the high sierras. In 1890, Carl
Lumholtz began his explorations, which carried him eventually the length
of the Sierra Madre Occidental and a small collection of birds was
preserved.

The Sierra Madre Occidental Province is clearly divisible orni-
thologically into three faunal districts, of which the two largest and
most important are the Tepehuane, occupying approximately the lower
half of the province, an area which is named for the Tepehuane Indians
who live in a large part of the region, and the almost equally important
Tarahumare Faunal District, occupying the major portion of the
northern part of the province. The higher portion of these two

MoorEeE—Biotic Provinces oF Mexico 229

districts is found for the most part in the Transition Zone, whereas the
lower portion of the northern district drops into the Upper Austral
Zone. The climate of the lower altitudes is rather dry, but the
Transition Zone on some mountains, such as the highest one, Mount
Mohinora, reaching an altitude of nearly 3500 feet above the general
level of the range and an altitude above sea level of somewhat less than
12,000 feet, is subjected to rather frequent rains in the summer months
and snow as late as the month of May. During this month of the year
1937, the ground about the author’s tent at the 10,500 foot level, was
covered with snow an inch and a half deep on two different days and
ice formed on the stream in front of his camp.

On the western side of the Tepehuane District, this province drops
rather abruptly, in some places precipitously, into the Upper Arid Trop-
ical Zone of Sinaloa, which is deemed by the author to have so little
in common with the Sierra Madre Occidental Province, that it is placed
by him in the Sinaloa Biotic Province. The author’s collection from the
Sinaloa Province is the largest and most adequate one which has been
taken in any single province of Mexico.

The author has gone far enough in his study of the birds of the
Sinaloa Province to convince him it should be divided into at least three
faunal districts. To the two southern ones he has given the names
“Sinaloa Coastal” and “Tebaca” Faunal Districts, the latter being
the name of the tribe which occupies a considerable portion of that
district. The northern district is called the Alamos Faunal District, in
view of the fact that it includes a considerable portion of a district in
southern Sonora and extreme northern Sinaloa, to which van Rossem
(1931) gave that name. How far to the south the Tebaca and Sinaloa
Coastal Faunal Districts extend is still uncertain, but further light will
be thrown upon this question when our large collections now available
from Nayarit are analyzed. Such material as the author has examined
would seem to indicate that these two districts and the Sinaloa Province
itself may extend as far as the Sierra de Vallejo in southern Nayarit,
where it comes close to the Pacific Ocean and acts as a barrier to prevent
coastal-loving, low-altitude forms from extending their habitats farther in
that direction. In this respect, the ornithological picture may not be the
same as the mammalogical or that obtained from the study of other cate-
gories of zoology. For these, the great Rio Grande de Santiago of central
Nayarit and its deep gorge may act as a barrier.

230 SAN Disco Society oF NATURAL History

Collecting in the Sierra Madre Oriental at high elevation has been
much less thorough and almost negligible, except for a few collecting
stations made by Mr. Lamb and by Sutton and his associates at medium
altitudes in the range. No attempt can be made here to define the faunal
districts of this province. However, the material that is available proves
that its bird-life is allied to that of the Transverse Volcanic Biotic.

I have already alluded to the importance of the Sierra Madre del
Sur Biotic as the one whose affinities are probably closer than any other
biotic to that of the Transverse Volcanic. Sufficient material! is avail-
able to make certain that there are at least three faunal districts involved,
but, since their outlines can be indicated only roughly, no attempt will
be made at this time to demark them, when it is obvious that the bound-
aries would have to be revised, as soon as the component elements of
the bird-life can be intensively studied.

BIBLIOGRAPHY

BarLey, ALFrep M. and Conover, H. B.: Notes from the State of Durango,
Mexico. Auk, vol. 52, pp. 421-424, 1935.

BLAKE, EmMMeET R. and Hanson, Harotp C.: Notes on a Collection of
Birds from Michoacan, Mexico. Zool. Series, Field Mus. Nat. Hist., vol.
22, no. 9, 1942.

Merriam, C. Hart: North American Fauna, no. 8, p. 30, 1895.

SCHUCHERT, CHARLES: Historical Geology of the Antillean-Carribbean
Region, 1935.

SmitH, Hosart M.: An Analysis of the Biotic Provinces of Mexico, As
Indicated by the Distribution of the Lizards of the Genus Sceloporus.
Anales de la Escuela National de Ciencias Biologicas {Mexico}, vol. 2,
no. 1, pp. 95-110, 1940. (In English and Spanish.)

THAYER, W.N.: The Physiography of Mexico. Jour. Geol., vol. 24, pp. 61-94,
1916.

VAN RosseM, A. J.: Report on a Collection of Land Birds from Sonora,
Mexico. Trans. San Diego Soc. Nat. Hist., vol. 6, no. 19, pp. 237-304,
1931.

vAN RossEM, A. J.: Critical Notes on Middle American Birds. Bull. Mus.
Comp. Zool., vol. 47, pp. 387-490, 1934.

OCONAVAWN EH

Moore—Brotic PRoviNcEsS OF MExico

TABLE 1

231

Showing Distribution of Forms of Bird-Life by Faunal Districts in the
Transverse Volcanic Biotic Province, Mexico.

x= Known from the district. X= Entirely confined to the district.

Jalis.
. Dendrortyx barbatus..........-.....-0---.00-- —
. Dendrortyx macroura macroura.........- a
. Dendrortyx macroura griseipectus...... —
. Dendrortyx macroura diversus.........-.- x
. Dendrortyx macroura oaxacae..........-- ==
. Lophortyx douglasii teres... x
. Colinus virginianus graysoni...........--- x
. Colinus virginianus nigripectus.........- =
. Colinus virginianus thayeri..........-..--.-- —
. Meleagris gallopavo gallopavo............ =
. Rallus longirostris tenutrostris............ =
. Coturnicops noveboracensis goldmani —
. Geococcyx velox velox... —
sMOEUSaSt0. SOTELEQUS coon soo ncn ccccstceomnke —
. Otus vinaceus sed uctus................--00--- —

Otus flammeus flammeus............---- x

. Otus guatemalae cassini..........-.-.---- a

Cynanthus latirostris propinquus........ —

. Thalurania furcata ridgwayi..........-..- ><

Lampornis amethystinus
GELS NUS ge oe ee eee ce —

. Xiphocolaptes promeropirhynchus

SCL ALC NE eee bo ee _

. Sittasomus griseicapillus

PAIS CONS Gare ote eo Ree Thad a xX

. Mitrephanes phaeocercus

PE OCCT GUS et tere retest =

. Empidonax difficilis immemoratus...... —
. Empidonax albigularis axillaris.......... —

Aechmolophus mexicanus «......2..2-+2---- —

. Megarynchus pitangua caniceps.......... xX
. Chionophilos alpestris chrysolaema.... x

@y tinal yea rains seco stseeact se evant ee

. Aphelocoma coerulescens sumichrasti —
. Aphelocoma sordida colimae.............. xX
. Aphelocoma sordida sieberit.............. —
. Aphelocoma unicolor unicolor...........- —

Cyanocitta stelleri coronata.............-.- —

. Cyanocitta stelleri: azteca...............-.-- —

Parus sclateri sclateri.............20-...00---- —

Tarase.

Otomi Aztec

-- x
— Xx
x x
-- x
— x
- Xx
— X
x x
xX —o
— x
— x
- x
— Xx
x x
-- x
x x
— x
— x

Oni-Zem

| * <*

| pdm |: Sd Stine. |

232

. Geothlypis nelsoni nelsoni
. Ergaticus ruber ruber...°.... 2.2... x
. Basileuterus belli belli =

y Car podacus mexicanus coccineus
: Car podacus mexicanus roseipectus —

SAN DtgGo Society oF Natura History

TaBLeE 1—Continued

Jalis.
. Certhia americana jaliscensis............-- xX
. Cistothorus platensis tinnulus............ —
. Heleodytes megalopterus

IDE Dello PL Cris Meret oes eS =

. Heleodytes megalopterus nelsoni...... ==
. Heleodytes jocosus gularis............ x
. Thryothorus felix grandis............... =:
. Thryomanes bewickii bairdi................ _
. Thryomanes bewickii_percnus............ x
. Troglodytes brunneicollis nitidus...... ==
. Henicorhina leucophrys mexicana...... =
. Salpinctes obsoletus notius...........0..---- x
. Catherpes mexicanus mexicanus........ x
| Moxostoma ocellatums. 4) a1. —_

. Toxostoma curvirostre deflexum —

. Toxostoma curvirostre vetula --

. Regulus satrapa aztecus........0.0..c.c00 —
. Ptilogonys cinereus cinereus......2....-.-- x

. Vireolanius melitophrys goldmani...... —
. Vireo nanus —

. Vireo gilvus amauronotus..........2.---- =
. Vireo gilvus eleanorae.... 2.2.0.0... —
. Neochloe brevipennis brevipennis...... —
. Dendroica aestiva dugesi........... x

. Geothlypis trichas melanops —

. Geothlypis chapalensis 0.0.0... —
. Geothlypis speciosa “=

Tarase.

Otomi

. Cassidix palustris —

. Agelaius gubernator grandis. A. —
. Piranga bidentata bidentata
. Carpodacus mexicanus centralis _

. Atlapetes torquatus virenticeps.......... x
. Pipilo torquatus alticola
. Pipilo torquatus nigrescens —
. Pipilo fuscus fuscus
. Pipilo fuscus toroi =

| *

Aztec

Nema lisse Te PACT pas pcaed cote ieisd— flies Ear OPM ooo tea ale |r ae |

Ori-Zem

esa tee ST eesti tee Pahoa he t|etreact [ates [Ey fat ev ials metsscit el had leat erepaee it~ eet fi

Moore—Brotic Provinces oF Mexico 233

TABLE 1—Continued
Jalis. Tarasc. Otomi Aztec Ori-Zem

80. Aimophila ruficauda acuminata ........ x x = x —

81. Aimophila rufescens pallida................ x x — — —

82. Melospiza melodia pectoralis.............. x —

83. Melospiza melodia adusta.................. —_ xX — — —
TABLE 2

The Number of Genera, Species, and Subspecies of Endemic Birds Recorded
From the Different Faunal Districts of the Transverse
Volcanic Biotic Province.

Addl _Add'l
Genera Species Subspec. Total
Continedstoy)aliscans ee _- 9 9

Gonfined tomMatascan=. 250 = 1 5 6
Continedito@tomin.. 4 ta ee — 1 2 3
@ontinedito Azteca. #8. fe ee citi: 1 1 8 10
Confined to Orizaba-Zempoaltepec..............-.....-- = 2 19 21

Common to Jaliscan and Tarascan, but not

imivOthers msec eee Re ee — — 4
Common to Jaliscan and Otomi, but not

it OCIetG Mee ee ew eet tes ee AMS — — 0
Common to Jaliscan and Aztec, but not

MV OLUICES marie ee areeeeetl Brea Seen Pet LS on — — 0
Common to Jaliscan and Orizaba-

Zempoaltepec, but not in others... — — 0
Common to Tarascan and Otomi, but

NOCMMTOtN ers ese eas Me eRe ye ee — = 2
Common to Tarascan and Aztec, but

NOLMntothers re sw ole tse Ne a at — — 2
Common to Tarascan and Orizaba-

Zempoaltepec, but not in others............... = == 0
Common to Tarascan, Aztec, Orizaba-

Zempoaltepec, but not in others... = — 8
Common to Otomi and Aztec, but

MOGRINGORMERS. ie eer saw 7 ee — — 0
Common to Otomi and Orizaba-

Zempoaltepec, but not in others.............. — — 0
Common to Aztec and Orizaba-

Zempoaltepec, but not in others... — — 5

Common to all five Districts..........................-..- =e = +

234 SAN DrgeGco Society oF NATuRAL History

TABLE 3

Showing Distribution of Forms in the Five Provinces of Central Mexico Which
Have the Closest Ornithological Relationships. No Form is
Shown Unless It Appears in At Least Two and Not
More than Three of the Five Provinces.

irs.Vol. S.iOce. SOrn. S- Sur. Chib.
1. Dendrortyx macroura striatus............ x — — x —
2) Pislortyx: fasciatus). 2.2 x _ — Xx —
3. Meleagris gallopavo mexicana............ x xX — — x
4. Chaetura rutila griseifrons.................- Xx x = = —
De Otus- dso: suttoni2<..).6 2 eee XxX — — xX
6. Amazilia beryllina beryllina................ xX — — — — *
7. Centurus chrysogenys flavinuchus...... x — — x —
8. Centurus hypopolius 2... 2cccecqce- x — — x —
9. Campephilus imperialis 2.20.02... x — —
10. Dryobates villosus jardinit.................. x — — x —
11. Dryobates scalaris azelus.........c00.00---- xX — = X
12. Dryobates scalaris centrophilus.......... x x — —- —
13. Lepidocolaptes leucogaster
lewWeo pasterwn Ace 1a Sth SEO xX x — x —
14. Grallaria guatimalensis
ochraceiventris......2--.! 0-00 xX — — xX
15. Elaenia viridicata jaliscensis................ 4 xX — xX ==
16. Empidonax affinis affinis...000...... x — xX — _
17. Empidonax affinis trepidus................ x — — — » 4
18. Empidonax difficilis occidentalis........ x — — D4 —
19. Empidonax fulvifrons rubicundus...... x Xx — _ —
20. Calocitta formosa formosa...........-..-.-- x = = — —
21. Cyanolyca pulchra mitrata:................. x _- — _ — tf
22. Aphelocoma coerulescens cyanotis..... X ao —- a xX
23. Aphelocoma sordida sordida.............. 4 — xX — 4
24. Psaltriparus minimus iulus 0... x — — — xX
25. Psaltriparus minimus melanotis.......... x == = —_ = RE
26. Certhia americana alticola.................. x — = — — £
27. Heleodytes jocosus jocosus........-...---- x — — x —
28. Thryomanes bewickii murinus............ 4 = == = x
29. Henicorhina leucophrys festiva.......... xX — — X —
30. Toxostoma dorsale dumosum............ x —_ = = x
31. Toxostoma curvirostre celsum............ xX == — — x
32. Turdus migratorius phillipsi......... x == xX = =
33. Turdus migratorius permixtus............ x = — x —
34> Uardus infuscatts 62. ee x _ ae — +
35. Catharus mexicanus mexicanus.......... x — — -— —= ||
36. Catharus occidentalis fulvescens........ x = == x =

For explanation of footnote symbols, see end of table on next page.

Moore—Brotic Provinces oF Mexico 235

TaBLE 3—Continued

TVole SOce. S.Ors SaSury Chih.
37. Catharus auranturostris melpomene.. X |

38. Sialia mexicana mexicana...........-..-.--- xX — xX — —
39. Vireo huttoni mexicanus...........-..------ x — x == =
40. Vermivora superciliosa mexicana........ xX — x = —
41. Dendroica graciae gracide...........-.----- xX X — —_ _
42. Basileuterus culicivorus flavescens...... xX xX a _— ==
43. Basileuterus culicivorus brasherii........ x = §
44. Basileuterus belli clarus.......0..0.0.2.-.--- xX — x x =
45. Basileuterus rufifrons dugesi.............. x — x -—- —
46. Agelaius phoeniceus gubernator........ x X = = =
47. Aimophila humeralis humeralis.......... xX — — x —
48. Aimophila rufescens rufescens..........-. X — ¢t
49. Aimophila ruficeps boucardi.............. xX = x — =
50. Aimophila ruficeps fusca.........-.--------- xX — — x a
d1. Spizella passerina atremaeus............-. x x = = —
52. Spizella passerina mexicana................ x — — x —

* Also in Veracruz Biotic.

+ Probably also in Nayarit-Guerrero Biotic.

Also in Chiapas Highlands Biotic.

|| Probably also in Chiapas Highlands Biotic.

§ Also in Tamaulipas Biotic.

TABLE 4

Showing the Forms Common to the Transverse Volcanic Biotic and the Other
Biotics of Mexico Most Closely Allied to It.

Number

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Common to Transverse Volcanic and Chiapas Highlands and no others... 5
Common to Transverse Volcanic and Tehuantepec and no others................ 0
Common to Transverse Volcanic and Nayarit-Guerrero and no others...... 0

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TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VoLuME X, No. 13, pp. 237-244, map Aucust 31, 1945

PRELIMINARY STUDIES ON THE BLACK-THROATED
SPARROWS OF BAJA CALIFORNIA, MEXICO

2.9 89 8 BY

A. J. VAN ROSSEM Me SBR AZ

Dickey Collections, University of California at Los Angeles Nliar

During recent investigations which concerned primarily the
Black-throated Sparrows of Sonora, Mexico, I took advantage of the
opportunity to determine more fully the subspecific status of the num-
erous specimens in the Dickey collections from the Baja California pen-
insula and islands. To augment our own material a series of specimens
from various critical localities has been borrowed from the Bishop, Huey,
and Willett collections, the Los Angeles Museum, the Museum of Com-
parative Zoology, the Museum of Vertebrate Zoology, and the San
Diego Natural History Museum. The combined total is 243 speci-
mens examined. Additionally, I am under obligation to Laurence M.
Huey for the contribution of notes and material on which he had in-
tended to publish and to James L. Peters for information on the type
of Amphispiza bilineata bangsi.

The results outlined in the present paper are not to be considered
conclusive. Material from many peninsular localities and from many
islands must be collected in quantity and accompanied by field notes
before definite ideas of the intra-specific variations and their geograph-
ical limits are possible. Of the more than a score of islands in the Gulf
of California on which the Black-throated Sparrow is a common, or
even abundant resident, there are fairly adequate, good-plumaged series
available only from San Estéban, Tortuga, Carmen, and Espiritu Santo.
Whether the populations on these respective islands are actually the
most pallid, the darkest, relatively the longest tailed, and the smallest
within the area or indeed within the entire range of the species cannot
be assured until populations which inhabit other islands are studied.

238 San Disco Society oF NATURAL HIsTory

Except for a few localities, the material from the peninsula is even more
inconclusive and all too frequently is in sadly abraded plumage or with |
no indication on the tags as to breeding activity. This latter point is
important, as will be appreciated later on in the paper.

As matters stood when Grinnell’s “Distributional Summation of
the Ornithology of Lower California” appeared in 1928, two races of
the Black-throated Sparrow were recognized from the Baja California
peninsula and adjacent islands. These were Amphispiza bilineata
bangsi from Cape San Lucas north to latitude 26° and Amphispiza bil-
ineata deserticola from latitude 27° northward, an arrangement which
was followed in the Fourth (1931) edition of the “Check-list.” More
recently, the present writer described Amphispiza bilineata tortugae
from Tortuga Island (see 19th Supplement to the Check-list, Auk, 61:
463, 1944), and Amphispiza bilineata cana from San Esteban Island,
the latter politically a part of Sonora and therefore outside the “Check-
list” boundaries. The studies here initiated show that considerable
revision of characters and ranges is necessary. This, so far as is pos-
sible with existing material, is attempted below.

Although the sexes are invariably described as “alike,” such is not
precisely true, for females average distinctly browner, the terminal tail
spotting averages less extensive and the measurements are about five per
cent shorter in wing and tail. While the color differences hold good be-
tween races when compared sex for sex, one cannot always be sure of
racial differences by comparing males of one race with females of an-
other. Measurements on the accompanying table are those of males.

PENINSULAR RACES

Amphispiza bilineata deserticola Ridgway

DesERT BLACK-THROATED SPARROW

Amphispiza bilineata deserticola Ridgway, Auk, 15, no. 3, July, 1898, 229
{separates issued May 13} (Tucson, Arizona).

Characters—Among the races here considered, deserticola is the largest,
and is further characterized by a relatively pale, brownish coloration. The
terminal tail spotting on the lateral rectrices averages distinctly smaller than
in any other race, but this is so variable individually as to be of questionable
use in the determination of single specimens. Throughout the extensive range
in the southwestern United States and northwestern Mexico, there seems to be
remarkably little variation in size or color when specimens of equal wear are
compared.

VAN ROSSEM—BLACK-THROATED SPARROWS 239

Range in Baja California—The northern half of the peninsula (except
in the extreme northwest), south at least to about latitude 28° 20’ on the Gulf
and about 29° on the Pacific.

Remarks.—I can perceive no differences between specimens from within
the above range and specimens from Arizona and California. Five worn ex-
amples from the vicinity of San Ignacio eastward to the Gulf (lat. 27°) are
seemingly like equally worn deserticola in color but are smaller. Fresh material
may alter this determination.

Amphispiza bilineata subspecies

On the Pacific side of the peninsula southward from about latitude 28°
30’, and on the Gulf side from about latitude 26° 30’ there occur Black-throat-
ed Sparrows which are, as is the case with various other species, darker than
those to the north or south. In size they are about intermediate between desert-
icola and bangsi. Birds very similar to these are found also on Cedros and
Natividad Islands, but certain minor differences observed suggest that Cedros
birds in particular may prove to be different. Purely on the basis of measure-
ments I place San José Island (Gulf) birds here also. The only two available
specimens from that island are too faded (late June) to be of much use from
the standpoint of color. Specimens from the area outlined above are too few
in number, from localities too widely scattered, and for the most part in plum-
age too worn to justify the use of any subspecific name at this time.

Amphispiza bilineata bangsi Grinnell
BANGS BLACK-THROATED SPARROW

Amphispiza bilineata bangsi Grinnell, Auk, 44, no. 1, January 5, 1927, 71

(La Paz, Lower [Baja} California, Mexico).

Characters.—Distinctly smaller than deserticola in all dimensions, save
possibly those of the bill. Tail, proportionately slightly shorter. Coloration
slightly, but definitely, grayer and darker, and terminal tail spotting more ex-
tensive. For comparison with the still smaller race of Espiritu Santo Island,
see posted.

Range—The Cape region, north on the Pacific side to the vicinity of
Magdalena Bay and on the Gulf side to La Paz.

Remarks.—Although Grinnell ascribed a “slightly paler” coloration to
bangsi as compared with deserticola, he doubtless was misled by the post-mortem
pallor of the Frazar series (Museum of Comparative Zoology) on which the
race primarily was based. Recent material is slightly darker and grayer than
deserticola. I am not prepared to admit the “larger bill” of bangsi, since mea-
surements of the two races are almost identical in this particular. Proportion-
ately, though, this character might be considered valid in minor degree. It is
unfortunate that the type of bangsi was selected from La Paz instead of from
the Cape proper, for not only the type (wing, 64; tail, 57 mm.) but the two
other specimens available from there have slightly longer wings than Cape

(Continued on page 242)

240

San Disco Society oF Natura History

MEASUREMENTS AND BREEDING MontHs oF RAcES SHOWN ON Map

Ref.

Letter

Nex SS CHuMwrOW OZ eH AS eA a Ooo eS

Subspecies

deserticola

carmende

bangsi

sanctissima
Not determined
tortugae

cana

Ay. wing
length

66.8
66.3
67.0
66.0
67.0
65.0
65.1
66.6
66.5
64.0
67.0
63.0
63.0
63.7
Female
64.0
64.0
63.0
61.7
61.0
61.0
62.0
62.7
60.0
62.0
63.1
64.0

Ay. tail

length
62.6
62.0
63.0

63.0
62.0
61.7
63.0
62.0
60.0
63.0
60.0
60.3
60.0
Female
61.0
61.0
60.0
60.3
56.5
56.3
56.5
57.0
54.7
58.0
58.5
60.0

Spec.

measured

ip)
NO N | & Se RB WwW BS KSB PY N OO KF FY KS K CO

—"
aA w

Found
breeding

May-July
Apr.-July
Apr.

Apr.

Feb.-July

Feb.

113° 12° ithe 110° 109°
A x \
@ 1
. |
34 ras |
CALIFORNIA {
? ARIZONA |
\ ‘
33° eS |
o-oo \
Fa a a at '
x ( =e a
> “Sa e :
pT
rs cad ene
3r o Cc \
yi x
& oO &
e° ve, ree
30° a) e& -
> le ~
oo SONORA
| = xe Nl
2s oO :
J *y &
Vv ~ i
sy y go ve
28 A ahaa an °
vi
, VN Ne
I o
Oo0Oor*
are e
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© Yy
35 SN Oo iG “nN
=~ > dQ oe) =P
LOCALITY RECORDS ZL { 41 ip cN
OF 2 2 Zz
INEATA ay ole
se{. AMPHISPIZA BIL Vv er
@ 256-2: -DESERTICOLA v2 &
Se BANGSI v o8w
§----------- - CANA Tw NG
aae|. “8------------ TORTUGAE \
f Che eae oer CARMENAE
—— SANCTISSIMA
& SUBSPECIES vU
Bl 0 Pages aa NOT Vv
aa “V7 DE TERMINED y
|
117° 116° 15° 114° 113° 12° me? 110° 109°

117° 16° 15° 114°

DISTRIBUTION OF THE RAcEs OF Amphispiza bilineata IN BAJA CALIFORNIA

For explanation of letters, see table on opposite page.
Symbols near islands refer to closest island.

335

3i¢

30°

29°

26°

25°

24°

242 SAN Disco Society oF NaAtTurRAL History

specimens and in this respect are intermediate toward the mid-peninsula race.

The closest comparison between the Cape and Magdalena Plain series
(14 specimens) and that from the Magdalena Bay Islands of Santa Margarita
and Magdalena (21 specimens) fails to disclose any tangible morphological
difference. A physiological difference, on the basis of existing evidence, is,
however, shown by the fact that the peninsular birds breed in October while
those on the Bay islands breed in February. Mr. Huey informs me that
throughout the Cape region and north to El Refugio and Buena Vista on the
Magdalena Plain streaked young, mostly just on the wing, were very common
in October and early November, 1941. He collected two of these, one at La
Paz on November 8, and one at El Refugio on November 14. The first is
“bob-tailed;” the second, although scarcely fully grown, had started the post-
juvenile molt. It may be noted that this fall breeding season explains in large
part Frazar’s failure to collect young birds or to note any breeding season (see
Brewster, Birds of the Cape Region, 1902), for he arrived on the peninsula
January 26, too late by some three months for this activity. At the time of
my own visit to Santa Margarita Island (February 27—-March 1), young on the
wing were abundantly in evidence and an occasional late nest with eggs was
found. Bryant (Catalogue of the Birds of Lower California, 1889), had prev-
iously noted this breeding season on Magdalena and Santa Margarita Islands.
The same state of affairs is evident on the Gulf side (see under the Espiritu
Santo race) but in this latter case a morphological difference is evident.

INSULAR (GULF) RACES

The insular populations are possessed of notable constancy of characters,
that is to say the characters which distinguish them one from the other and
from the peninsular races are relatively uniform. This, I suppose, is. to be
expected since inbreeding with little or no infusion of outside stock would act
to fix or to accentuate the characters inherent in the initial occupants. I can-
not conceive that “natural selection,” in the sense of protective coloration, or
of adaptation to special environments has played any significant role in the
divergences shown by these races. All of the islands are essentially similar to
adjacent points on the peninsula both in character and vegetation, nor are there
predators not common to all islands alike, except that a Bassariscus occurs on
some of the larger ones.

Amphispiza bilineata cana van Rossem
SAN ESTEBAN BLACK-THROATED SPARROW

Ampbhispiza bilineata cana van Rossem, Trans. San Diego Soc. Nat. Hist., 6
no. 14, November 28, 1930, 223 (San Esteban Island, Gulf of California,
Sonora, Mexico).

Characters.—The palest and grayest of the races within the Gulf area. Size
slightly but uniformly smaller than deserticola and tail spotting more exten-
sive.

Range.—San Estéban Island.

VAN ROSSEM—BLACK-THROATED SPARROWS 243

Remarks—San Estéban Island, although politically a part of Sonora, is
Baja Californian in its avifaunal relationships and for that reason is included
in the present paper. I have previously called attention to the fact (Condor,
44: 184-5, 1942; further paper in press), that San Estéban races (with one
exception) are similar to or identical with those from more southern parts of
Baja California and not with those of the areas immediately east and west.
The present instance is no exception, for in grayness of coloration cana stands
out conspicuously from deserticola of the peninsula and still more so from
pacifica of Tiburén Island and the southern Sonora mainland. Nineteen speci-
mens have been examined.

Amphispiza bilineata tortugae van Rossem
TORTUGA BLACK-THROATED SPARROW

Amphispiza bilineata tortugae van Rossem, Trans. San Diego Soc. Nat. Hist.,
6, no. 14, November 28, 1930, 222 (Tortuga Island, Gulf of California,
Lower {Baja} California, Mexico).

Characters —Darkest of the described races of Amphispiza bilineata and
no comparison with any of the other Gulf area forms is necessary. Dorsally it
is darker than Amphispiza bilineata bilineata of the lower Rio Grande valley
and slightly more slaty (less brownish), while the lateral under parts are more
extensively and deeply colored than in any other race.

Range.—Tortuga Island.

Remarks.—This dark-colored race, the dark extreme of the species, stands
out all the more conspicuously because of its contrast with the pale brownish
deserticola of the adjacent part of the peninsula and the pale grayish cana
of San Estéban Island to the north. No Black-throated Sparrows are known
to occur on San Pedro Martir, the only island intervening between Tortuga
and San Estéban.

Through some cause now obscure, but which most likely is connected in
some way with an abundant, year-round food supply, the density of the pop-
ulation on Tortuga surpasses anything in my experience with the species. At
the time of my two visits there, in March at the beginning of the breeding season,
and in January when no breeding activity was evident in any of the specimens
collected, there was no noticeable difference in numbers, which certainly ran into
many hundreds on this tiny island of a little over two square miles in area.
Thirty-three specimens have been examined.

Amphispiza bilineata carmenae subsp. nov.
CARMEN BLACK-THROATED SPARROW

Type—RMale, adult, no. 50,416 Dickey Collection; Salinas Bay, Carmen
Island, Gulf of California, Baja California, Mexico, January 21, 1932; collected
by A. J. van Rossem, original number 14,079.

Characters—Nearest in color to Amphispiza bilineata bangsi of the ex-
treme southern portion of the peninsula, but averaging slightly grayer in series.
Tail longer than in bangsi and equal to or only very slightly shorter than the

244 SAN Digeco Society oF NATURAL History

wing, instead of, as in all other described races, decidedly (4-6 mm.) shorter.
Measurements of the type are wing, 61.5; tail, 60.5 mm.

Range.—Carmen Island.

Remarks.—Were it not for the different wing and tail proportions, I would
not suggest a distinctive name for the Carmen Island birds even though in
series they average slightly grayer and paler than bangsi. At one time I was
inclined to place them as nearest to cana, in spite of the intervening, dark
tortugae and the fact that San Estéban and Carmen islands are nearly 200
miles apart. Twelve specimens have been examined. For measurements of the
seven males, see the accompanying table. The five females average in wing,
58.8; in tail, 57.4 mm.

Amphispiza bilineata subspecies

Two specimens from San Francisco Island (a breeding pair in the Dickey
collection) do not differ, so far as I can see, from bangsi of the peninsula, save
that they have notably long, slender bills, attenuate throughout their length
and with the culmen outline slightly concave. This small island which lies near
the southern end of San José Island is only about one and a half square miles
in area but is well populated by Black-throated Sparrows and the collection of
a good series of specimens would present no difficulties.

Amphispiza bilineata sanctissima subsp. nov.
EspirIru SANTO BLACK-THROATED SPARROW

Type.—Breeding male adult, no. 29,936, Dickey Collection; Espiritu
Santo Island, Gulf of California, Baja California, Mexico, March 10, 1930;
collected by A. J. van Rossem, original number, 12,652.

Characters.—Smallest of the races of Amphispiza bilineata with propor-
tionally and actually the largest bill. Very similar to bangsi in coloration, but
averaging very slightly darker in series.

Range—Espiritu Santo Island.

Remarks.—Although separated from the mainland only by some five
miles of channel, this island is known for several endemic vertebrates including
the remarkable Lepus insularis. At this time it appears quite inexplicable why
the Black-throated Sparrows on Espiritu Santo breed in March while those on
the immediately adjacent part of the peninsula (bangsi) breed in October. The
same state of affairs is found in the Magdalena Bay area (see antea), although
without, in the latter instance, any discernible morphological differences be-
tween the two populations. Eleven specimens have been examined. Wing and
tail averages of the nine males are found on the accompanying table. Bill mea-
surements of these nine males are: exposed culmen, 11.2; depth of bill at base,
6.0 mm. Comparative measurements of 23 male bangsi are 10.6 and 5.9 mm.,
respectively.

TRANSACTIONS
OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY

VOLUME X, No. 14, pp. 245-268, map

THE POCKET GOPHERS OF BAJA CALIFORNIA,
MEXICO, WITH DESCRIPTIONS OF NINE
NEW FORMS

BY,

Laurence M. Huey
Curator of Birds and Mammals, San Diego Society of Natural History

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY
AucustT 31, 1945

COMMITTEE ON PUBLICATION
U. S. Grant, IV, Chairman

L. M. KLAuBER CLINTON G. AspotrT, Editor

THE POCKET GOPHERS OF BAJA CALIFORNIA,
MEXICO, WITH DESCRIPTIONS OF NINE
NEW FORMS

BY
BUGIS
Laurence M. Huey
Curator of Birds and Mammals, San Diego Society of Natural History

INTRODUCTION

To one who is interested in mammal ecology and taxonomy, the
pocket gophers (Genus Thomomys) of the Pacific southwest offer an
intriguing subject of study. Early workers found such apparently diverse
characters in specimens from scattered geographical localities, that to
the majority of them they originally assigned full specific rank. Slowly,
‘as intervening regions were explored and closer connections were proven,
certain species were found tu be subspecifically related to distant animals
that had also been named as full species. Gopher populations living
almost as unbroken chains from moist coastal regions, through gradient
mountain passes, to arid deserts were disclosed. Thus the gophers of
a large part of California were found to be linked into a huge closely
related group, all races of the single species, Thomomys bottae. E. A.
Goldman offered a list of 75 races in his paper “Pocket Gophers of the
Thomomys bottae Group in the United States” (Proc. Biol. Soc.
Washington, vol. 48, pp. 153-158, 1935). The gophers of Baja
California were beyond the geographical scope of his paper, though he
did correct the allocation of Thomomys martirensis, by recognizing it
as T. bottae martirensis, and stated that all other gophers so far named
from Baja California had, in his opinion, been properly placed under

T. bottae.

In 1941, Emmet T. Hooper published a complete list of all names
applied to the genus Thomomys known to April 7 of that year (“Type
Localities of Pocket Gophers of the Genus Thomomys,” Misc. Publ.
Mus. Zool., Univ. Michigan, no. 52, pp. 1-26, 1 map, 1941). This
paper, while not indicating relationships of the species or races, arranges
the type localities in geographical order and gives the citation of each
original description. It is thus of decided value to the taxonomist.

248 SAN Disco Society oF NATURAL History

In Hooper’s list, 13 named gophers are shown to have their type
localities in Baja California. However, this was not the complete list
of gophers inhabiting the peninsula, since two of the forms that he
mentions have their type localities in southern California and their
range extends across the International Boundary. With type localities
in California, 47 gophers are listed. Probably this State has been more
intensively explored biologically than any other section within the range
of the genus, while, by contrast, Baja California, a region almost as
interesting faunally and zonally as California, is but little known. This
is particularly true of the northern half.

The gophers inhabiting this section of Baja California offer a
problem of unusual complexity, by reason of the exceptionally varied
terrain and climatic conditions of the region. In that part of the 750-
mile long peninsula which lies north of the 30th parallel of latitude—
an area approximately 200 miles in length by 125 miles in width—are
to be found the highest, lowest, driest, wettest, hottest and coldest
portions, as well as that with the most equable temperature, in the entire
peninsula. From the summits of pine-clad peaks within this area it is
possible to view, in one direction, mesquite-bordered washes, amid
scorched rocky hills and gleaming sandy deserts, and, in the opposite
direction, vast chaparral-covered slopes that stretch for miles in unbroken
expanse to the ocean. Apart from the intensive agricultural develop-
ment of the Imperial Valley, whose limits are defined by the gravity
flow of water from the Colorado River, the hand of man has done
little, as yet, to mar primeval conditions. Ranches are few and evidences
of cultivation but slight. Hence human agency has played almost no
part in increasing, decreasing or disrupting the ranges of the natural
gopher population.

Although there have been several gophers heretofore named from
northern Baja California, it has long been apparent that confusion has
existed and that specified ranges have been incorrect. For example, Bailey
in his revision of the Genus Thomomys (North American Fauna, no. 39,
pp. 1-136, 8 plates, 1915) endeavored to clarify the situation by extend-
ing the range of one race (T. bottae nigricans) over a vast region in
the peninsula and placing another species (T. aphrastus) in synonymy
under it. This broad allocation was unquestionably due to his not
having sufficient specimens from Baja California, and it has complicated
the situation for subsequent students.

In view of these facts, the writer has, at every opportunity during

Huty—Pocket GOPHERS OF BAJA CALIFORNIA 249

the past twenty-two years, gathered material that he hoped would, at
some future time, help to determine the correct status of Thomomys
in Baja California. What he thought were key points have been visited,
yet much remains to be done that would throw a clearer light on the
subject.

ACKNOWLEDGMENTS

Throughout this period of intermittent exploration the most cordial
cooperation has been extended by the several departments and officials
of the Mexican Government. This has been exemplified not only in
granting necessary permits for the work, but also, on the part of both
civil and military personnel, in expediting the passage of equipment
across the International Boundary at the different ports of entry. For all
these courtesies the writer expresses his appreciation.

He borrowed from the Museum of Vertebrate Zoology, through
the kindness of Drs. E. Raymond Hall and Seth B. Benson, all available
specimens contained in the University of California collections that had
a bearing upon the Thomomys of the region in question. From the
Museum of Zoology of the University of Michigan, through the kind-
ness of Dr. W. H. Burt, he borrowed an important series of gophers
from the Magdalena Bay region. For this generous cooperation, he
wishes to acknowledge his indebtedness.

Factors AFFECTING DISTRIBUTION

In view of the sedentary character of the pocket gopher’s life, it
is interesting to speculate upon the causes of its development into the
considerable number of distinguishable races in a region such as northern
Baja California. The factors responsible there are of course the same
as those elsewhere within the range of Thomomys, but are more
intensified as a result of the extreme diversity of climate and terrain
of this region, as was mentioned above. Climate—wet or dry; topo-
graphy—deserts, hills or valleys; soil texture—sand or loam, rock or
clay; moisture content of the soil and vegetation it supports—all these
have a determining influence in the development of physical characters
of this rodent whose existence is almost wholly subterranean.

There are natural avenues of expansion and there are natural
barriers. The latter may be geological or botanical, visible or invisible.

250 SAN Disco Society oF NATURAL History

Valley floors, with their alluvial soil and watercourses, even though dry
most of the year, are avenues for the expansion of gopher populations.
Hills or mountains are forms of barriers. Thus the higher mountain
backbone, that runs the length of California and northern Baja
California, holds in check some of the populations of gophers that have
developed racial differences on either slope. Yet there are two known
streams of gopher populations living in gradient passes through this
mountain chain in southern California that completely link the
relationship of the gophers in both the desert and coastal areas. By
these links the pallid desert forms previously known as the Thomomys
perpallidus group are shown to be definitely related to the earlier-named
coastal form, Thomomys bottae, and all are now considered to fall into
the bottae group.

The drainage flow westward from the central mountain chain in
northern Baja California has formed many valleys and canyons suitable
for Thomomys habitat, with high, rolling hills between. These areas
of rolling hills are in most cases composed of coarse, almost impenetrable
soil and support a vast cover of dense chaparral—a combination which
proves to be an almost insurmountable obstacle to gopher expansion.
Two natural factors here act as a check: (1) the hard compact soil in
which burrowing is exceedingly difficult, and (2) the fact that the
chaparral offers little or no food for gophers. The several plant species
found in the dense growth of shrubs, growing in soil of low fertility,
are not acceptable as food by gophers and do not permit the development
of annual plants or root-bearing species attractive to gophers. Hence
we have a botanical barrier.

Grinnell and Hill in their account of “Pocket Gophers (Tho-
momys) of the Lower Colorado Valley” (Journ. Mamm., vol. 17, no.
1, pp. 1-10, 1936) discuss briefly the effect of a great river flowing
through an arid desert and the arresting influences of rocky barriers
upon the gopher populations where these barriers reach the river’s edge
along its course. Yet beyond the river’s influence of seeping moisture
within the soil, colonies of Thomomys are to be found in seemingly
uninhabitable localities. One such, mentioned in the present paper, is
in northeastern Baja California, and many more populations, similarly
isolated, are to be found over the deserts in the mid-section of the
peninsula.

The question arises as to how these isolated colonies can exist
within such inhospitable desert surroundings. There must be present

HurY—Pocket GOPHERS OF BAJA CALIFORNIA 251

sufficient moisture near enough to the surface for gophers to live and
also enough to sustain continuous plant life for gopher food. Probably
rock formations beneath the surface of the soil exert their influence by
retaining or concentrating moisture to form subsurface reservoirs or
basins. This moisture can come from one or two sources: either from
the scant, irregular precipitation which occurs on the desert, or from
underground flow or seepage originating in distant regions where
rainfall is greater. Thus geological barriers lying beneath the surface
may be responsible for remote gopher colonies that, by isolation, evolve
into differentiated races.

Thomomys bottae has reached the southernmost parts of the
peninsula of Baja California, though the population is very irregular
or spotty over a greater part of the intervening arid desert sections.
It would be interesting to speculate on the way this region was inhabited.
Many theories could be explored, but as yet enough factual data are
not available for sound discussion. As elsewhere, both geological and
botanical conditions have exerted their influence in providing the
fundamental requirements of a gopher’s existence, namely suitable soil
conditions and food.

MATERIAL EXAMINED

For the purposes of this work the writer examined 647 specimens
of Thomomys, of which 188 were from 19 localities situated in the
southernmost half of San Diego County, California. These localities
represent a cross-section from the Pacific Ocean to the Colorado Desert
and include the habitat of three forms of Thomomys whose ranges extend
into Baja California.

The specimens collected in Baja California number 459 and were
taken in 36 localities over the length of the peninsula, but mainly north
of 27° latitude. Of these, the greater portion are from the biologically
complex western slopes north of latitude 30°. Thirty-four specimens,
collected from six different important localities, were borrowed from
the Museum of Vertebrate Zoology, University of California, and 12
specimens, collected at Stearns Point in Magdalena Bay, were borrowed
from the Museum of Zoology, University of Michigan.

Nine new forms are herewith described and one previously-named
race revived. Thus 24 different named forms are found to occupy the
peninsula. That this list is, in all probability, not complete, may be

252 SAN Disco Society oF NATURAL History

readily inferred from the large areas, especially the southern coastal
section of the Vizcaino Desert and the east-central gulf sections of the
peninsula, that are not yet explored.

LIST OF SUBSPECIES

Thomomys bottae albatus Grinnell
IMPERIAL VALLEY POCKET GOPHER

Thomomys albatus Grinnell, Univ. Calif. Publ. Zool., vol. 10, p. 172, June 7,

1912.

Type locality West side of Lower Colorado River at old Hanlon Ranch,
near Pilot Knob, Imperial Co., California.

Range In Lower California—Irrigated section of region south of Interna-
tional Boundary to El Major, east to the Colorado River and west to the limits
of gravity water on the eastern base of the Cocopah Mountain Range.

Specimens examined.—A single specimen from 20 miles east of Mexicali,
Baja California, Mexico, is the only representative of this race from Baja Cali-
fornia in the collection of the San Diego Society of Natural History, though
an abundance of specimens are at hand from the California side of the Bound-
ary, along the Colorado River near the type locality (see Trans. San Diego
Soc. Nat. Hist., vol. 9, no. 15, p. 72, 1939). Grinnell and Hill (Journ. Mam-
malogy, vol. 17, p. 3, 1936) list specimens from five different localities, all in
the true delta of the Colorado River or in irrigated lands close by.

Thomomys bottae lucidus Hall

LAGUNA SALADA PocKET GOPHER

Thomomys bottae lucidus Hall, Proc. Biol. Soc. Wash., vol. 45, p. 67, 1932.
Type locality—Las Palmas Canyon {=2 miles east of Gaskill’s Tanks,
mesquite association, fide J. E. Green, collector of type}, 200 feet altitude, west
side of Laguna Salada north of 32° N. latitude, Baja California, Mexico.
Range.—Type locality, as above. This race is one of those pioneer colonies
confined to an extremely limited area.
Specimens examined—Two, the type and only other known specimen
from same locality.

Thomomys bottae cunicularius’ subsp. nov.
Pattie BAsIN PockeT GOPHER

Type.—From Los Palmitos (western end of Pattie Basin), on the south-
eastern base of the Sierra Juarez (desert slope), latitude 31° 44’ N., longitude

1Cunicularius, Latin, “a miner,’ used as referring to a tunnel or passage under-
ground,

Hury—Pocket GOPHERS OF BAJA CALIFORNIA 253

115° 36’ W., Baja California, Mexico; no. 12,182, collection of the San Diego
Society of Natural History; adult male, collected by Laurence M. Huey, Nov-
ember 29, 1936.

Characters—Compared with Thomomys bottae lucidus, whose range lies
to the northward, T. 6. cunicularius is decidedly grayer, with buffy suffusion on
sides and not light colored, showing its entire relationship to be with the coastal
forms and not with those of the desert, T. 6. lucidus or T. b. albatus. Nor does
it show cranial characters that tend towards the two desert races, though the
skulls are less massive than those of its westward relations, a character that does
offer some suggestion in the direction of T. 6. lucidus, which from the geo-
graphical position of its range would be anticipated. Compared with T. 6. juar-
ezensis,? T. 6. cunicularius is lighter, more grayish in color and has a more
fragilely boned skull with heavier molariform teeth. The bullae are more round-
ed and less elongated than those of T. 6. juarezensis and the pterygoid pro-
cesses are structurally much more fragile. T. 6. cunicularius is also smaller, both
in vertebral length and cranial size.

Measurements—Type: Total length, 215; tail, 68; hind foot, 28; ear,
4. Skull (type): Greatest length, 38.4; spread of maxillary arches, 22.7; length
of nasals, 13.3; interorbital constriction, 5.6; alveolar length of upper molar
series, 8.0.

Range.—Known only from the type locality, as above.

Specimens examined.—Three, the type and only two other known speci-
mens from the same locality.

Thomomys bottae nigricans Rhoads

Tawny Pocket GOPHER

Thomomys fulvus nigricans Rhoads, Proc. Acad. Nat. Sci. Philadelphia, p. 36,

1895.

Range in Baja California Specimens best referable to, though not typical
of, this race in Baja California are to be found along the International Bound-
ary from Nachoguero Valley westward to El Valle de Las Palmas, thence south
to Las Cruces, which lies some 15 miles inland from Ensenada. This area em-
braces a great hilly section in the form of a triangle with its apex at Las
Cruces and its base along the International Boundary. The range of T. 6.
nigricans does not at any point reach the sea coast.

Remarks.—The gopher population of the coastal slope of northwestern
Baja California and southwestern California presents a complicated problem.
According to Bailey (N. A. Fauna, no. 39, p. 57, 1915) the range ascribed
to T. b. nigricans extended from the San Jacinto Mountains in California,
southward along the coastal slope to a point on the mainland of Baja California,
opposite Cedros Island. Yet within this great tract are to be found wide dif-
ferences of plant associations, from pines to palms and mesquites, and areas as-
signable to three life zones.

Witch Creek, where the type specimen of T. 6. nigricans was collected,

2Newly described in this paper.

254 SAN Disco Society oF NATURAL History

lies at an elevation of 2675 feet. It is in a valley filled with live oaks, situated
amid the broad chaparral belt of the coastal foothill slope. This type of coun-
try, with oak-filled valleys surrounded by chaparral, is found westward from the
higher pine-clad mountains of San Diego County and southward into Baja
California, where, from the area about Ensenada, it tapers towards the Sierra
San Pedro Martir, disappearing in the mid-section of that range. Its zonal
character is mainly Upper Sonoran. It is to such an association that the
range of T. b. nigricans, in the estimation of the writer, is restricted.

Within a series of 42 topotype specimens of this race examined by the
writer, a great amount of variation in two important characters was found. These
were the color of the pelage and the shape of the bullae. Neither was stable.
The most dependable cranial characters, and those on which the name of T. b.
nigricans may be said to stand, were the angular flare of the zygomatic arch
from the axis of the skull and the parallel position of both jugals, when speci-
mens of similar age were compared. However, when the writer studied speci-
mens taken in other associational surroundings, especially where Lower Sonoran
conditions prevailed, he found definable differences which included stabiliza-
tion of one or both of the irregular characters observed in the topotype series.
Several new races have therefore been named in this paper from the area that
was ascribed by Bailey to T. b. nigricans.

A similar situation was encountered by Grinnell in the north and des-
cribed in his paper “Distribution in Pocket Gophers of the Thomomys bottae
group in Northern California and Southern Oregon” (Univ. Calif. Publ. Zool.,
vol. 40, no. 11, pp. 403-416, 1935). Here, too, Grinnell found that he could
not match with topotypes gophers taken in different sections of a large range
that had been assigned by Bailey (N. A. Fauna, no. 39, p. 47, 1915) to T. 6.
leucodon. His investigation revealed that several recognizable races were living
within restricted associational conditions, but that wide margins of intergrading
populations were to be found between. A different type of territory from Baja
California was of course involved, but many of the conditions were analogous
and Grinnell’s statement that forested areas were void of gopher life points to
the effect of a “botanical barrier,” such as the present writer has described.

Specimens examined.—2 from Ballena, San Diego County, California;
42 from Witch Creek, San Diego County, California (type locality); 3 from
Santa Ysabel, San Diego County, California; 1 from North Peak, Cuyamaca
Mountains, San Diego County, California; 15 from Laguna Mountains, San
Diego County, California; 9 from Nachoguero Valley, Baja California;? 6
from south end of Valle de Las Palmas, Baja California; 8 from Las Cruces,
Baja California.?

Thomomys bottae affinis subsp. nov.

JACUMBA PocKET GOPHER

Type.—From Jacumba, San Diego County, California; no. 14,083, collec-
tion of the San Diego Society of Natural History; adult male; collected by

3From Museum of Vertebrate Zoology, Berkeley, California.

Huey—Pocket GOPHERS OF BAJA CALIFORNIA 255

Laurence M. Huey, May 9, 1940.

Characters—Thomomys bottae affinis is uniformly lighter in pelage color
than either T. 6. nigricans or T. b. juarezensis. In fact the series of 31 speci-
mens available to the writer, of this closely allied form, shows greater uniform-
ity of color through the various ages of the specimens than do similar series of
either of the other two mentioned forms. Cranially T. 6. affinis differs from
both T. 6. nigricans and T. 6. juarezensis in having the angular development
of the zygomatic arches more accentuated. They flare at almost right angles
from the axis of the skull of a fully mature specimen. Those of T. 6. nigricans
and T. b. juarezensis are obtuse by comparison. The auditory bullae of T. b.
affinis are rounded and inflated, resembling those of T. 6. juarezensis rather
than those of T. 6. nigricans, although the greater size of the skull makes this
character appear more conspicuous. The race T. b. affinis could hardly be pro-
posed without taking into account the race Thomomys bottae puertae, whose
range is in the desert foothills some 30 miles to the northwest of Jacumba. It
inhabits La Puerta (or Mason) and San Felipe valleys, San Diego County,
California, and bears close relationship to T. 6. nigricans; but the characters
that set it apart do not in any way show affinity with T. b. affinis.

Measurements —Type: Total length, 235; tail, 70; hind foot, 30; ear, 5.
Skull (type): Greatest length, 42.3; spread of maxillary arches, 25.4; length
of nasals, 14.5; interorbital constriction, 6.7; alveolar length of upper molar
series, 8.7.

Range.—Jacumba Valley on both sides of the International Boundary.

Specimens examined.—31 from Jacumba, San Diego County, California
(type locality). Thomomys bottae puertae: 10 from San Felipe Valley, San
Diego County, California; 38 from La Puerta Valley, San Diego County,
California (type locality) .4

Thomomys bottae juarezensis subsp. nov.
SIERRA JUAREZ Pocket GOPHER

Type.—From Laguna Hanson, Sierra Juarez, Baja California, Mexico; no.
5849, collection of the San Diego Society of Natural History; adult male; col-
lected by Laurence M. Huey, November 26, 1926.

Characters.—Several average characters differentiate Thomomys bottae
juarezensis from its nearest comparable relative, T. 6. nigricans. In pelage color,
it differs in lacking the brighter buffy sides. When viewed in series, T. b.
juarezensis has a more grayish tone, tending towards black. This race, like
T. 6. nigricans, shows considerable individual color variation and, for best ap-
praisal of this character, must be seen in series. When compared in this way,
the darker grayish Hack of T. b. juarezensis is plainly seen, in contrast to the
warmer brownish color of the mass of T. 6. nigricans. Crnially, T. 6. juar-
ezensis has more rounded and more inflated bullae, with a heavier, wider
spreading V of the interpterygoid, than has T. b. nigricans. The braincase of
T. b. juarezensis is wider at the point of greatest constriction between the ocu-

4Six from collection of L. M. Huey.

256 SAN Disco Society oF NaturaAL History

lar fossae, and the rostra are heavier and broader in a larger number of speci-
mens, when compared with T. b. nigricans. Some overlap is to be found in this
latter character, however. The zygomatic arches of T. 6. juarezensis lack the
sharp angular development found in well-aged T. b. nigricans, and the jugals
are circular-appearing rather than parallel. This is the most pronounced and
constant character of the race, and while some sub-adult specimens of T. b.
nigricans show a rounded arch development, the arches grow angular and par-
allel with age, even to the point of excessive width anteriorly. The selection of
similarly-aged specimens of Thomomys for comparison was made by observing
the development of the temporal ridge, which is an excellent age index.

Measurements.—Type: Total length, 245; tail, 73; hind foot, 30; ear, 5.
Skull (type): Greatest length, 40.7; spread of maxillary arches, 24.1; length
of nasals, 12.9; interorbital constriction, 6.3; alveolar length of upper molar
series, 8.5.

Range.—Known from forested area on summit of Sierra Juarez.

Specimens examined.—8 from El Rayo, 53° from Laguna Hanson, both
localities situated close together on the yellow pine clad summit of Sierra Juarez,

Baja California.

Thomomys bottae jojobae® subsp. nov.
SANGRE DE Cristo PocKET GOPHER

Type.—From Sangre de Cristo, Baja California, Mexico, lat. 31° 52’ N.;
long. 116° 06’ W.; no. 6116, collection of the San Diego Society of Natural
History; adult male; collected by Laurence M. Huey, June 20, 1927.

Characters—Compared with Thomomys bottae juarezensis, whose range
occupies the pine crest of the mountain chain directly to the east, T. b. jojobae
is smaller in size, and brighter in color both ventrally and dorsally, when viewed
in series. Cranially, T. b. jojobae has a more fragilely boned skull, with a
more slender rostrum and smaller incisors. Compared with T. b. nigricans,
whose range lies northwestward and extends into the mountainous foothills of
San Diego County, California, T. 6. jojobae is paler and is more buffy over-all.
Cranially, the skull of T. 6. jojobae is lighter or more fragilely boned than that
of T. b. nigricans, and is not angular or rugose. The upper contour of the
skull, when viewed from the side, is less arched and slightly more flattened or
depressed in the ocular region than is that of T. 6. nigricans. Also, the bullae
of T. b. jojobae are more rounded and inflated, lacking any ridging such as is
found in some specimens of T. b. nigricans.

Measurements——Type: Total length, 218; tail, 65; hind foot, 26; ear, 4.
Skull (type): Greatest length, 37.7; spread of maxillary arches, 21.5; length
of nasals, 13.0; interorbital constriction, 6.2; alveolar length of upper molar
series, 8.0.

Range.—Known only in typical form from the western foothills of the
Sierra Juarez in Valle de San Rafael, Baja California.

5Ten from collection of L. M. Huey.
6Jojoba is the local name for the shrub Simmondsia californica, amongst which this
gopher was found in its greatest abundance.

Hury—Pocket GopHEerS OF BAJA CALIFORNIA 257

Specimens examined.—37 from Sangre de Cristo, Baja California (type
locality) .”

Thomomys bottae xerophilus subsp. nov.
SAN Matias Pass PockET GOPHER

Type—From near Diablito Spring, summit of San Matias Pass (be-
tween Sierra Juarez and Sierra San Pedro Martir), Baja California, Mexico;
no. 11,827, collection of the San Diego Society of Natural History; adult male;
collected by Laurence M. Huey, March 29, 1936.

Characters Compared with Thomomys bottae jojobae, the race that oc-
cupies the foothill area of the Pacific slope of the Sierra Juarez to the north-
ward, T. b. xerophilus is slightly smaller in size. In color, it is dark grayish
instead of buffy. Cranially, T. 6. xerophilus has a proportionately shorter,
heavier rostrum with heavier molariform teeth. The braincase, when viewed
posteriorly, is deeper and more rounded—not an elongated ellipse. T. 6. xero-
philus differs from both mountain-top forms, T. 6. juarezensis and T. 6. mar-
tirensis, in being smaller in size and grayer in color, with more diminutive cran-
ial characters over-all. Compared with T. 6. aphrastus (here revived), T. 6.
xerophilus is smaller in size, gray in color instead of brownish, and cranially has
a proportionately heavier, shorter rostrum.

Measurements—Type: Total length, 198; tail, 65; hind foot, 26; ear, 4.
Skull (type): Greatest length, 34.8; spread of maxillary arches, 20.1; length
of nasals, 11.5; interorbital constriction, 6.2; alveolar length of upper molar
series, 7.9.

Range.—Known from San Matias Pass and from the eastern section of El
Valle de Ia Trinidad, at least to Aguajita Spring. Specimens of Thomomys
from the western part of El Valle de la Trinidad are not referable to this
race, but are intergrades which in some examples are not namable. In fact this
western section lies within an area in which at least three forms (T. 6. xero-
philus, T. 6. aphrastus, and T. b. nigricans) converge, and perhaps both marti-
rensis and abbotti from the south also exert some influence. However, the vast
foothill area west of the Sierra San Pedro Martir is but little known, and the
status of the range of the two latter forms is still to be determined.

Specimens examined.—14 from near Diablito Spring, summit of San
Matias Pass, Baja California (type locality) ; 25 from Aguajita Spring, El Valle
de la Trinidad, Baja California.

Thomomys bottae martirensis Allen
SAN PEpro MartTir PocKET GOPHER

Thomomyus fulvus martirensis Allen, Bull. Amer. Mus. Nat. Hist., vol. 10, p.
147, 1898.
Type locality—San Pedro Martir Mountains (alt. 8200 feet), Baja Cali-
fornia, Mexico [ = La Grulla Meadow, Sierra San Pedro Martir, alt 7400 feet,

7Six from collection of L. M. Huey.

258 SAN Dreco Society oF NaturAL History

fide A. W. Anthony, who was joint-collector, with E. C. Thurber, of the type
series }.

Range.—Higher levels of the Sierra San Pedro Martir. Specimens ex-
amined by the writer from two localities.

Specimens examined.—21 from La Grulla, Sierra San Pedro Martir, Baja
California (type locality); 6 from Valladares Creek, Sierra San Pedro Martir,
Baja California.

Thomomys bottae siccovallis subsp. nov.

Et Cajon CANYON Pocket GOPHER

Type.—From El Cajén Canyon, 3200 feet altitude, east base of Sierra San
Pedro Martir, Baja California, Mexico, lat. 30° 54’ N.; long 115° 10’ W.; no.
37,673, collection of the University of California, Museum of Vertebrate Zool-
ogy; adult female; collected by Raymond M. Gilmore, June 3, 1926, (original
no. 369).

Characters—When compared with Thomomys bottae martirensis, T. b.
siccovallis is found to have a close resemblance in color of pelage, but several
cranial characters separate the two races. T. b. siccovallis has a proportionately
shorter, wider rostrum and heavier, more angular zyogmatic arches, but lacks
the flaring tendency of some examples of T. b. martirensis. The bullae are
slightly less swollen than are those of T. 6. martirensis and the molariform
teeth are slightly heavier. The post-ocular foramina are smaller than in any
example of T. 6. martirensis of comparable age available to the writer. This
latter character is accentuated by the heavier zygomatic arches and the absence
of spreading or flaring. Nevertheless, there is evidently close relationship be-
tween these two forms. A wide divergence in both cranial and color characters
separates T. b. siccovallis and T. 6. xerophilus. Cranially, T. 6. siccovallis has
a larger, more heavily boned, rugose skull than has T. 6. xerophilus, while in
pelage color the contrast is even greater, T. b. siccovallis being within the range
of brown tones, while xerophilus is gray.

Measurements —Type: Total length, 209; tail, 59; hind foot, 28; ear, 4.
Skull (type): Greatest length, 37.1; spread of maxillary arches, 21.6; length
of nasals, 13.0; interorbital constriction, 7.0; alveolar length of upper molar
series, 8.8.

Range.—Known only from the type locality, a very secluded canyon on
the desert side of the Sierra San Pedro Martir, Baja California.

Specimens examined.—3 from El Cajén Canyon, 3200 feet elevation, east
base of Sierra San Pedro Martir, Baja California (type locality). All are the
property of the Museum of Vertebrate Zoology, University of California. Two
of these specimens are sub-adults. However, the salient characters that separate
this race are discernible, even in these non-mature specimens.

Thomomys bottae sanctidiegi subsp. nov.
San Dreco Pocket GOPHER

Type——From Balboa Park, San Diego, California; no. 14,886, collection

HurEY—PocketT GOPHERS OF BAJA CALIFORNIA 259

of the San Diego Society of Natural History; adult male; collected by Laurence
M. Huey, December 18, 1941.

Characters—This form differs from topotypical Thomomys bottae bottae
from Monterey, California, in being paler dorsally when viewed in mass series.
Cranially, several average differences exist. T. b. sanctidiegi has more pro-
truding upper incisors and a more slender rostrum. The bullae are slightly
smaller and more irregularly shaped—not full and rounded. In a large pro-
portion of specimens, the area of interorbital constriction is wider in T. b. sanc-
tidiegi than in T. b. bottae. Compared with topotypical T. b. nigricans from
Witch Creek, San Diego County, California, T. 6. sanctidiegi is larger in size
and lighter in color, with heavier-boned skull. Some specimens of T. b. sancti-
diegi, especially males, attain extremely large size, their skulls far exceeding in
rugosity any of those of the oldest adult males of T. b. nigricans ever seen by
the writer.

Measurements—Type: Total length, 240; tail, 70; hind foot, 30; ear, 5.
Skull (type): Greatest length, 43.2; spread of maxillary arches, 27.6; length
of nasals, 14.0; interorbital constriction, 6.7; alveolar length of upper molar
series, 9.1.

Range—As at present known, the coastal strip from the vicinity of San
Diego, California, to the vicinity of Ensenada, Baja California. Specimens
referable to this race have been examined from La Jolla, California, but its
range may well extend to the northward.

Remarks——Thomomys bottae sanctidiegi is another of the closely related
“differentiates,” though the direction of its relationship is towards the nominate
race, T. b. bottae of Monterey, California. However, T. 6. sanctidiegi is not
the southern end of the branch, but a link in the chain which extends south-
ward into the morass of intergrading population that occupies the foothill area
mentioned in the remarks under T. 6. nigricans.

Specimens examined—Thomomys bottae bottae: 10 from Monterey,
California (type locality). Thomomys bottae sanctidiegi: 1 from La Jolla,
San Diego County, California; 3 from Point Loma, San Diego, California; 18
from San Diego, California; 2 from National City, San Diego County, Cali-
fornia; 1 from Spring Valley, San Diego County, California; 1 from Monument
258, International Boundary, San Diego County, California; 5 from 5 miles
south of Monument 258, Baja California;® 20 from Guatay (about 25 miles
north of Ensenada), Baja California.

Thomomys bottae aphrastus Elliot revived subsp.
PERPLEXING POCKET GOPHER
Thomomys aphrastus Elliot, Field Columbian Mus., Zool. Series, vol. 3, pub.
PO D219 OOS:

Type locality—San [ =Santo} Tomas, Baja California, Mexico.
Characters—Although placed in synonymy with T. 6. nigricans by Bailey

8From Museum of Vertebrate Zoology, Berkeley, California.

260 SAN Disco Soctety oF NATURAL History

(N. A. Fauna, no. 39, p. 56, 1915), the present writer believes that this gopher
possesses sufficient diagnostic characters, and has a sufficiently definable range,
to be revived as a valid subspecies of T. bottae. The pelage color, it must be
admitted, offers but little of value for subspecific determination. In a topotypic
series it was found to vary considerably and only towards the center of the
large range occupied by T. b. aphrastus is it at all uniform. However, a few
cranial characters found in the series of topotypes seem to be constant through-
out the range and thus establish the tenability of the race. T. 6. aphrastus does
not attain rugose, angular cranial proportions as does T. 6. sanctidiegi, but
shows a more definite relationship to the smaller, more modified mountain
form, T. b. nigricans. The skull is lightly boned with weak-angled, fragile and
parallel zygomatic arches, lacking even the almost square flare exemplified in
T. b. nigricans. A series of 21 skulls of T. 6. aphrastus, picked for age equality,
from different points of its range, are all found to show a fairly uniform elon-
gation. That is, the rather long braincase with the weak-spreading zygomatic
arches and the slender-appearing rostrum, are peculiar to them all. This char-
acter does not appear in other named races whose ranges are adjacent. The up-
per incisors of the older males have a procumbent tendency which is not found
in nearly related races.

Range—From Santo Tomas in Santo Tomas Valley eastward to the ex-
treme western end of El Valle de la Trinidad, thence south along the foothills of
the Sierra San Pedro Martir at least to Las Cabras. Southward from Santo
Tomas, the range of this race reaches the coast at Johnson’s Ranch, thence over
the coastal plain to or below San Quintin. The area south of Las Cabras and
east of San Quintin plain is still unknown. This latter region is of transitional
character, where the more arid conditions of the dry inland desert converge
with those of the humid coastal belt, and it could easily be the habitat of an
unnamed race of Thomomys. —

Remarks.—Throughout the perimeter of this large range, local populations
of Thomomys show various degrees of relationship toward the surrounding
named races, and wide areas of intergradation are found. It is unfortunate
that Thomomys aphrastus should have been named from Santo Tomas, a geo-
graphical position which proves to be at the extreme northern limit of its now
known range. It was not until sufficient specimens had been accumulated to
define correctly the characters, even though they are slight, that this race could
be revived and a comprehensive range assigned to it. Had the type been
chosen from a more southern locality, the problem would have been much
easier and perhaps the race T. b. aphrastus would not have been relegated into
synonymy.

Specimens examined.—16 from Santo Tomas, Baja California (type local-
ity) ;? 31 from the extreme western end of El Valle de la Trinidad, Baja Cali-
fornia (some non-typical);!° 6 from Las Cabras, Baja California; 14 from
Santo Domingo, lat. 30° 45° N., Baja California.!!

9Nine from Museum of Vertebrate Zoology, Berkeley, California.
10Seven from collection of L. M. Huey.
11One from collection of L. M. Huey.

Hury—Pocket GopHEerS OF BAJA CALIFORNIA 261

Thomomys bottae proximarinus subsp. nov.
CoasTAL Pocket GOPHER

Type.—From Boca la Playa, 16 miles west of Santo Tomas, Baja Cali-
fornia, Mexico (mesa bordering the sea), lat. 31° 32’ N.; long. 116° 38’ W.,;
no. 14,182, collection of the San Diego Society of Natural History; adult
male; collected by Laurence M. Huey, August 15, 1940.

Characters—The series of 17 specimens used in determining this race are
all uniform in color, being dark “Sepia”!? brown dorsally, grading to “Snuff?!”
brown ventrally. They are darker than any topotypical specimens of the con-
tiguous race T. 6. aphrastus. In fact this color character is outstanding when
compared with all other northwestern Baja Californian Thomomys populations.
In size, too, it is smaller, being a rather depauperate race, living on the mesa-
like benches adjacent to the ocean. Cranially, T. 6. proximarinus is similar to
T. 6. aphrastus, though.the skull is uniformly smaller in size, with the zygom-
atic arches having a more arched appearance and tending towards posterior flar-
ing. The incisors of the older male examples show no tendency towards pro-
cumbency and digress from the main cranial axes at near right angles.

Measurements—Type: Total length, 215; tail, 69, hind foot, 28; ear, 5.
Skull (type): Greatest length, 37.7; spread of maxillary arches, 23.7; length
of nasals, 13.4; interorbital constriction, 5.6; alveolar length of upper molar
series, 7.5.

Range.—Known only from the type locality as above.

Remarks.—Were this uniformly dark-colored, small-sized race of gophers
living in an inland valley, where intergradation could branch from several
directions, it would be unworthy of racial separation. However, with inter-
gradation possible in but one direction and with but one race, the assigning
to it of a name seems to be in order. It is also worthy of mention that its
isolated habitat, backed against the sea, has brought forth such characters as
are shown by many island races of either birds or mammals—darker colora-
tion and diminution in size.

Specimens examined—17 from Boca la Playa, Baja California (type
locality).

Thomomys bottae abbotti Huey
Ext Rosario PockET GOPHER

Thomomys bottae abbotti Huey, Trans. San Diego Soc. Nat. Hist., vol. 5, p.

89, 1928.

Type locality—1 mile east of El Rosario, Baja California, Mexico (river
bottom association), lat. 30° 03’ N., long. 115° 48’ W.

Range.—Known only from environs of river bottom association in type
locality and to the southeastward at San Fernando Mission. In all prob-
ability the range of this race will be found coastwise south from El Rosario,
though as yet the region is unexplored.

12See Plate 29, Color Standards and Color Nomenclature, Ridgway, 1912.

262 SAN Drsco Society oF NaturAL History

Specimens examined.—17 from 1 mile east of El Rosario, Baja California
(type locality); 2 from San Fernando Mission, Baja California.

Thomomys bottae catavinensis Huey

CATAVINA PocKET GOPHER

Thomomys bottae catavinensis Huey, Trans. San Diego Soc. Nat. Hist., vol.
7, p. 45, 1931.
Type locality.—Catavifia, Baja California, Mexico, lat. 29° 54’ N., long.
114°°57' W.
Range——Known only from type locality, as above. It is probable that
T. b. catavinensis will be found along the arroyo for some distance both east
and west of the type locality when these regions are properly explored.
Specimens examined.—12 from Catavina, Baja California (type locality).

Thomomys bottae cactophilus Huey
Punta Prieta Pocket GOPHER

Thomomys bottae cactophilus Huey, Trans. San Diego Soc. Nat. Hist., vol.

5, p. 241, 1929.

Type locality—Punta Prieta, Baja California, Mexico, lat. 28° 56’ N.,
long. 114° 12’ W.

Range.—Known from type locality, as above, and the area near the Pacific
coast at Santa Rosalia Bay, Baja California.

Specimens examined.—28 from Punta Prieta, Baja California (type local-
ity); 2 from Santa Rosalia Bay, Baja California.

Thomomys bottae borjasensis subsp. nov.
SAN Borjas PocKET GOPHER

Type.—From San Borjas Mission, Baja California, Mexico, lat. 28° 52’
N., long. 113° 53’ W.; no. 14,491, collection of the San Diego Society of
Natural History; adult female; collected by Laurence M. Huey, October 14,
1941.

Characters.—In color, as compared with Thomomys bottae cactophilus, its
closest relative, T. 6. borjasensis is less buffy, more gray dorsally, yet belong-
ing to the chain of brownish colored gophers. Cranially, T. 6. borjasensis has
a narrower rostrum, squarer, more angular zygomatic arches, shorter, more
rounded braincase, with more rounded, less elongated bullae. As compared
with T. 6. russeolus to the southward, T. b. borjasensis is darker over all.
Insofar as color is concerned, there is a wide gap between the two forms, T. 0b.
borjasensis, as before stated, belonging to the brownish and not the pallid
group in which T. 6. russeolus falls. Cranially, compared with T. 6. russeolus,
the skull of T. 6. borjasensis is more fragilely boned. The rostrum is more
slender, with slightly protruding incisors. The bullae of T. 6. borjasensis are
smaller, less inflated than those of T. 6. russeolus.

HurEY—Pocket GOPHERS OF BAJA CALIFORNIA 263

Measurements—Type: Total length, 210; tail, 65; hind foot, 26; ear, 5.
Skull (type): Greatest length, 35.7; spread of maxillary arches, 21.8; length
of nasals, 12.4; interorbital constriction, 6.3; alveolar length of upper molar
series, 6.6.

Range-—Known only from the type locality, as above.

Specimens examined.—1, the type, from San Borjas Mission, Baja Cali-

fornia.

Thomomys bottae russeolus Nelson and Goldman
SAN ANGEL POCKET GOPHER

Thomomys bottae russeolus Nelson and Goldman, Proc. Biol. Soc. Wash., vol.

22, p. 25, 1909.

Type locality—San Angel, 30 miles west of San Ignacio, Baja California,
Mexico.

Range.—So far as known, the eastern side and northeastern rim of the
Vizcaino Desert.

Specimens examined.—2 from Mesquital, Baja California; 4 from Calmal-
li, Baja California (not typical); 19 from Campo Los Angeles, Baja Cali-
fornia.

Thomomys bottae incomptus Goldman
MAGDALENA PLAIN POocKET GOPHER

Thomomys bottae incomptus Goldman, Proc. Biol. Soc. Wash., vol. 52, p. 29,

1939:

Type locality—San Jorge, near Pacific coast west of Pozo Grande and
about 25 miles southwest of Comondi (altitude 50 feet), Baja California,
Mexico.

Range.—So far as known, the northern half of the vast Magdalena Plain,
with the exception of the very restricted coastal habitat of the following race,
Thomomys bottae litoris.

Remarks.—A series of 17 specimens from Santo Domingo, which is sit-
uated about 25 miles south of San Jorge, the type locality of this race, show
some slight average divergences. A few of the specimens were moulting and
the new pelage, compared with that of a topotypical specimen in like condition,
reveals the Santo Domingo series to be slightly darker in color. However, it
must be admitted that but two topotypes, though fortunately both good adults,
were available, and so limited a number could hardly be expected to demon-
strate a complete account of characters as they are ascribed to this race.

Specimens examined.—2 from San Jorge, Baja California (type locality) ;
17 from Santo Domingo, Baja California, lat. 25° 30’ N.

Thomomys bottae litoris Burt
MAGDALENA Bay PocKET GOPHER

Thomomys bottae litoris Burt, Occ. Papers, Mus. Zool., Univ. Mich., no. 424,
pp. 1-3, 1940.

264 San Dreco Society oF NATuRAL History

Type locality—Stearns Point, Magdalena Bay (west side), Lower Cali-

fornia, Mexico.

Range.—Known only from the type locality as above.

Remarks.—An assemblage of 31 specimens representing the two races from
the Magdalena Plain and Bay sections of the peninsula, Thomomys bottae in-
comptus and T. 6. litoris, shows a remarkable degree of alkaline pelage stain.
The color is reddish or buffish on the older pelage. Similar examples of alkali
stained Thomomys, Dipodomys and Perognathus are known from other des-
ert localities or where the soil in which they live is heavily impregnated with
mineral salts. This factor of stained coloration has been known to mislead
taxonomists into naming specimens so stained. However, in the case at hand,
regarding the Thomomys from the Magdalena Bay district, moulting examples
of each race are present, revealing their true coloration, and slight dorsal color
differences are plainly visible.

Specimens examined.—12 from Stearns Point, Magdalena Bay (west

side), Baja California (type locality).

Thomomys bottae magdalenae Nelson and Goldman
MAGDALENA ISLAND PocKET GOPHER

Thomomys magdalenae Nelson and Goldman, Proc. Biol. Soc. Wash., vol. 22,
p. 24, 1909.

Type locality—Magdalena Island, Baja California, Mexico.
Range.—Known only from the type locality, as above.

Remarks.—Only one specimen of Thomomys from Magdalena Island is
known to exist and it is the type. The writer has not seen this specimen.

In 1909, when Nelson and Goldman named T. magdalenae, the paucity
of material at hand and the fact that this population was confined to an island
led them to describe this gopher as a species. The first ascription of this species
as a race was made by Burt in 1940 when he described T. 6. litoris (Occ. Pa-
pers, Mus. Zool., Univ. Mich., no. 424, p. 3, 1940). In his list of “Specimens
examined,” there was simply entered, without comment: “T. 6. magdalenae:
from Magdalena Island, 1, the type.” He was following the latest concept
of interrelationship of the widespread “bottae” group. In 1943, Goldman evi-
dently recognized the subspecific relationship and listed this animal as T. 6.
magdalenae (Journ. Washington Acad. Sci., vol. 33, no. 5, pp. 146-147, 1943).

Thomomys bottae imitabilis Goldman
La Paz Pocket GOPHER
Thomomys bottae imitabilis Goldman, Proc. Biol. Soc. Wash., vol. 52, p. 30,
1939.

Type locality—lLa Paz, Baja California, Mexico.
Range.—Known only from the type locality.
Specimens examined.—None have been seen by the writer.

Huryr—Pockxet GOPHERS OF BAJA CALIFORNIA 265

Thomomys bottae alticolus Allen

SERRA LAGUNA PocKET GOPHER

Thomomys fulvus alticolus Allen, Bull. Amer. Mus. Nat. Hist., vol. 12, p. 13,

1899.

Type locality —Sierra Laguna (altitude 7000 feet), Baja California, Mex-
ico.

Range.—The higher sections of the Victoria Mountains in the Cape dis-
trict.

Remarks.—Three specimens collected by the writer at a point about 7
miles northwest of San Bartolo, where the lush mountain chaparral from the
north slope of the Victoria Mountains reaches its lowest level, are darker and
smaller than Thomomys bottae anitae and appear to belong to this mountain
race. This allocation is arbitrary, however, as the writer had no material from
the higher reaches of the sierran chain for direct comparison.

Specimens examined.—3 from 7 miles northwest of San Bartolo, Baja

California.

Thomomys bottae anitae Allen
CaPpE SAN Lucas PockET GOPHER

Thomomys fulvus anitae Allen, Bull. Amer. Mus. Nat. Hist., vol. 10, p. 146,
1898.

Type locality—Santa Anita, Baja California, Mexico.

Range.—Lower levels of the Cape district, south of the Magdalena Plain,
excepting the section about La Paz, which is occupied by T. 6. imitabilis.

Specimens examined.—12 from San José del Cabo, Baja California; 1

from Los Barrilos, Baja California (gulf coast).

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SAN Disco Society OF NATURAL History

THe Pocket GopHers OF BajJA CALIFORNIA, MExico

Name

. Thomomys bottae albatus

. Thomomys bottae lucidus

. Thomomys bottae cunicularius
. Thomomys bottae nigricans

. Thomomys bottae affinis

. Thomomys bottae juarezensis
. Thomomys bottae jojobae

. Thomomys bottae xerophilus

. Thomomys bottae martirensis
. Thomomys bottae siccovallis

. Thomomys bottae sanctidiegi
. Thomomys bottae aphrastus

. Thomomys bottae proximarinus

. Thomomys bottae abbotti

. Thomomys bottae catavinensis
. Thomomys bottae cactophilus
. Thomomys bottae borjasensis

. Thomomys bottae russeolus

Thomomys bottae incomptus

Thomomys bottae litoris
Thomomys bottae magdalenae

Thomomys bottae imitabilts
Thomomys bottae alticolus

Thomomys bottae anitae

Type Locality

Pilot Knob, Imperial Co., Calif.

Gaskill’s Tank, Baja Calif.

Los Palmitos, Baja Calif.

Witch Creek, San Diego Co., Calif.

Jacumba, San Diego Co., Calif.

Laguna Hanson, Sierra Juarez, Baja Calif.

Sangre de Cristo, Baja Calif.

San Matias Pass, Baja Calif.

La Grulla Meadow, Sierra San Pedro
Martir, Baja Calif.

El Cajon Canyon, east base Sierra San
Pedro Martir, Baja Calif.

Balboa Park, San Diego, Calif.

Santo Tomas, Baja Calif.

Boca la Playa, 16 mi. w. Sto. Tomas,
Baja Calif.

El Rosario, Baja Calif.

Cataviha, Baja Calif.

Punta Prieta, Baja Calif.

San Borjas Mission, Baja Calif.

San Angel, 30 mi. w. San Ignacio,
Baja Calif.

San Jorge, Baja Calif.

Stearns Point, Magdalena Bay, Baja
Calif.

Magdalena Island, Magdalena Bay,
Baja Calif.

La Paz, Baja Calif.
Sierra Laguna, alt. 7000’, Baja Calif.
Santa Anita, Baja Calif.

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TRANSACTIONS

OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VotuME X, No. 15, pp. 269-306, text fig. 1, map

THE CHUCKWALLAS, GENUS SAUROMALUS

CHARLES E. SHAW

San Diego Society of Natural History

SAN DIEGO, CALIFORNIA
PRINTED FOR THE SOCIETY
Aucust 31, 1945

DISTRIBUTION MAP
OF THE

Genus Sauromalus

S.o.obesus
S.o.tumidus-___
S.o0. townsend
S.australis
S.hispidus

INTERGRADES
Obesus-tumidus

Tumidus - townsendi____

(LEGENDS ARE SHOWN ADJA-
CENT TO, RATHER THAN ON,
THE SMALLER ISLANDS.)

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fo)

\o NEWA DA Een ae ae

SON OR A

THE CHUCKWALLAS, GENUS SAUROMALUS

BY
Cuares E. SHaw* Hes of Compas
} ; eo Zoviogy =“
San Diego Society of Natural History = ae
ade 4g SEP 12 1949
l

INTRODUCTION

Since the publication of Van Denburgh’s “Reptiles of Western
North America” more than twenty years ago, the genus Sauromalus
has received but slight attention from herpetological workers. During
this period important additional material from Arizona, Sonora, and
Baja California has accumulated, which has made possible a better
understanding of relationships within the genus and a more accurate
determination of the distribution of the several species.

This study was begun at the suggestion of Dr. L. M. Klauber and
was at first confined to the species obesus, in an attempt to determine
whether any races were to be recognized within the range of this very
widespread form. Because of certain nomenclatorial questions and the
acquisition of several interesting specimens from Baja California, the
scope of the study was extended to include the remaining species of
the genus. However, the insular species of Sauromalus, with the
exception of ater and townsendi, have been lightly dealt with in this
paper as these are relatively static from a taxonomic point of view and
probably will remain so. More intensive exploration of the islands in
the Gulf of California may perhaps extend the ranges of one or two
of the insular species, but the possibility that new species remain to be
discovered is unlikely. The insular forms have been included here only
for the sake of completeness.

HistoricAL SUMMARY

The genus Sauromalus was established by Duméril in 1856 (Arch. Mus.
Hist. Nat. Paris, vol. 8, p. 536) on the species ater, the type specimen having
been presented to the Museum d’Histoire Naturelle in Paris without locality.

In 1858 Baird (Proc. Acad. Nat. Sci. Phila., vol 10, p. 253) with a series
of specimens collected by the Mexican Boundary Survey and Lieut. Ives’
Expedition, erected the genus Euphryne for the species obesus, which he
described from Fort Yuma, California. For the next sixty-odd years obesus was
referred to the synonomy of ater.

* At present in United States Marine Corps.

DD. SAN DrseGco Society oF NATuRAL History

In 1891 Stejneger described Sauromalus hispidus (Proc. U. S. Nat. Mus.,
vol. 14, p. 409-411) from Angel de la Guarda Island, Gulf of California,
Mexico. Stejneger distinguished hispidus from ater, with which it had been
previously confused, on the basis of the former’s more spinose and coarser
scalation. At the same time this author also observed that a specimen of ater
from Espiritu Santo Island, in the Gulf of California, exhibited certain
differences from the ater of California and Arizona, differences that would have
warranted its distinction at that time had more specimens from the island been
available.

As a result of the collections made by the Albatross Expedition of 1911
to Baja California and its surrounding waters, Dickerson (Bull. Amer. Mus.
Nat. Hist., vol. 41, pp. 463-465) described as new the following species of
Sauromalus: interbrachialis, type locality, La Paz, Baja California, Mexico;
townsend, type locality, Tiburén Island, Gulf of California; and varius, type
locality, San Estéban Island, Gulf of California. In describing interbrachialis
from two additional specimens from Baja California, Miss Dickerson confirmed
Stejneger’s belief that the single specimen from Espiritu Santo Island available
to him should be regarded as distinct from ater of further north in California
and Arizona.

In 1922 Schmidt (Bull. Amer. Mus. Nat. Hist., vol. 46, pp. 640-641),
observing the agreement in the number of ventral scale rows between the type
specimens of ater and interbrachialis, referred the latter to the synonomy of ater,
and revived Baird’s name obesus for the chuckwallas occurring in the south-
western United States and northern Baja California. This author distinguished
obesus from ater on the basis of a greater number of ventral scale rows between
the gular fold and the anus in the former. In further justification of this
revision, Schmidt (loc. cit., p. 640) stated that “it is highly improbable that
the type of S. ater Duméril, collected by Lieut. Jaurés during the circum-
navigating voyage of the frigate Danaide, and presented without locality to
the Museum d’Histoire Naturelle in Paris, could have been collected in
California or Arizona, since Sauromalus does not reach the coast of California.”

Van Denburgh in 1922 (Occas. Papers Calif. Acad. Sci., no. 10, vol. 1,
pp. 97-99) described S. slevini, type locality, south end of Monserrate Island,
Gulf of California, Mexico. This species was distinguished from hispidus, which
it somewhat resembles, by its finer scalation and smaller size.

As a result of the present investigation, I described S. klauberi, (Trans.
San. Diego Soc. Nat. Hist., vol. 9, 1941, pp. 285-288) type locality, Santa
Catalina Island, Gulf of California, Mexico.

To these may be added two new forms described in this paper, Sauromalus
australis sp. nov. from Baja California, and Sauromalus obesus tumidus subsp.
nov. from southwestern Arizona.

VALIDITY OF CERTAIN NAMES

The fact that the type locality of S. ater is not known has been a source
of confusion to previous authors as well as a considerable annoyance in the
present work. Unfortunately the original description of S. ater gives no

SHAW—THE CHUCKWALLAS 273

diagnostic scale counts which would aid in the determination of the origin of
the type specimen. From figure 3a accompanying the original description it
can be seen that the nuchal scales are considerably smaller than the scales on
the postauricular fold and are quite evenly disposed. This precludes the
possibility of the type having come from any of the islands in the Gulf of
California where klauberi, slevini, or hispidus, occur, for these forms have the
nuchals usually at least as large as the postauricular scales; also klauber: has
elongate nuchals interspersed with those of smaller size and is quite distinct
from the other known forms of the genus in this respect.

Other sources of information concerning the type specimen have been
employed in an effort to determine, as well as possible, the probable source of
the type. Perhaps these references give as much, if not more, valuable
information concerning the type specimen than the original description itself.
A redescription is to be found in Bocourt “Mission Scientifique au Mexique,”
pp. 149-151, but again none of the characteristic scale counts, which might
associate it with a more or less definite locality, are given. However, a clue
to its probable origin, namely one of the several islands in the southern part
of the Gulf of California north of La Paz, is given in the following color
description: “Parties supérieures et inférieures de corps d'un brun jaunatre,
avec la téte, les pattes et la base de la queue d’un tiente un peu plus claire; le
tronc est couvert en dessus de petites mouchetures noires.” The yellowish-
brown coloration referred to is present in nearly all of the specimens of ater
and slevini which have been examined.

From “Contribution 4 la Faune Herpétologique de la Basse-California”*
we have the following remarks by Mocquard: “L’un de nos spécimens, un
male, offre en dessus, cormme le type spécifique, une tiente fondamentale tres
sombre, sur laquelle on distingue assez difficilement une bande noire transversale,
immédiatement en arriére de la racine des membres antérieurs.”’ This evidence
of only one band is not infrequent in the chuckwallas from the southern part of
the peninsula of Baja California, as well as those from the islands off the coast.

Schmidt (loc. cit., p. 640) from a photograph of the venter of the type
specimen of ater, determined the ventral scale count to be about 135, a figure
which corresponds excellently with the counts of the insular specimens. With
these three sources of information, I believe that it can be definitely concluded
that the type of ater came from one of the islands in the Gulf of California,
where this species is now known to occur. The ventral count of. 135, as
determined by Schmidt, also fits townsend: and klauberi very well. But klauberi
completely lacks any evidences of banding, being finely spotted instead; and
townsendi is quite definitely banded with 4 solid, dark transverse bars across
the back, a considerable period of preservation not having rendered these
markings less distinct in the two specimens from Tiburén Island which have
been examined. The coloration, as described by Mocquard, agrees excellently
with some specimens of slevini, but the ventral count of 135 is much too high
for that form; in 17 specimens of slevini the extreme range of variation in the
ventral scale count was found to be 107 to 123 and the average 115.8.

* Nouv. Arch. Mus. Hist. Nat. Paris, ser. 4, vol. 1, pp. 302-303, 1899.

274 SAN Dt1gGo SocliETY OF NATURAL History

From the above facts it seems reasonable to assume the type locality of
ater, hitherto unknown, to be one of the following islands in the Gulf of
California: Espiritu Santo, Isla Partida, San Marcos, San Diego, Santa Cruz,
or San Francisco.

Other counts which might be used to ascertain the probable locality of
collection of the type of ater are the number of scales in a whorl around the
tail two head-lengths behind the vent, and the number of scales around the
upper part of the fore limb. Unfortunately present European conditions make
the determination of these counts on the type of ater impossible.

We now come to the question of the validity of the name interbrachialis
proposed by Dickerson for the chuckwallas occurring on Espiritu Santo Island
and the southern portion of the peninsula of Baja California. The application
of the name ater to the chuckwallas from Espiritu Santo Island makes inter-
brachialis invalid for that population. But as the type locality as given by Miss
Dickerson for interbrachialis is stated to be La Paz, there remains the possibility
that it should be applied to the mainland population in the event the latter
should be proven distinct from the insular ater.

There have been available to me 6 specimens of Sauromalus from the
southern part of the peninsula of Baja California. Although a slight degree
of overlap in differential characters is shown between these specimens and 18
ater from the coastal islands of the southern Gulf, I believe the two populations
to be distinct. And there are a number of reasons which seem to prevent the
application of the name interbrachialis to the chuckwallas found on the mainland
of Baja California.

Schmidt (loc. cit., p. 640) remarks that “Two specimens of Sauromalus
from La Paz are included in the collections of the Albatross Expedition.
These are possibly from the island of Espiritu Santo, off La Paz, like the
specimen recorded by Yarrow and Stejneger.” Mr. Schmidt has informed me
by letter that his reasons for doubting the localities of collection of these two
specimens rest on the fact that some of the material was brought back alive to
the New York Zoo, and, as Miss Dickerson also kept a number of living
specimens, some confusion as to the proper place of origin was not unlikely
under such circumstances. This fact together with the agreement in ventral
scale counts between interbrachialis and ater led Schmidt to place the former
in the synonomy of ater.

In connection with the type locality of interbrachialis it is interesting to
note that the paratype was catalogued at the American Museum of Natural
History as having been collected on Carmen Island in the Gulf of California,
although Schmidt gives the locality of collection as La Paz. It should
therefore, be referred to slevini rather than to interbrachialis or ater. Miss
Dickerson made no mention of the source of the paratype in her description.

Dickerson separated interbrachialis from ater (ater being at that time the
name applied to the chuckwallas in the United States as well as the southern
portion of Baja California and the coastal islands) on the basis of its coarser
scalation and because of the pattern of “double” dark transverse bands. Both
the mainland chuckwallas and those from the islands off La Paz have this

SHAW—THE CHUCKWALLAS 22>

double-barred pattern. With regard to the coarser scalation Miss Dickerson
stated the ventral scale count to be 133, a figure which corresponds excellently
with the counts of specimens from the southern coastal islands. In the
chuckwallas from the peninsula I find that the ventral scale count in 6 specimens
ranges from 151 to 186 and averages 163.5, while the counts of 18 ater from
the islands range from 130 to 151 and average 139.7. In addition, Dr. Doris
M. Cochran informs me that the type of interbrachialis has between 40 and 42
scales around the upper or humeral part of the fore limb, a count which fits
most closely the counts of ater, with a range from 35 to 45, compared to 46
to 55 in the mainland specimens. These differences in countable scale series
between the mainland chuckwallas and the insular ater seem to me to be
sufficiently great to warrant the distinction of the former. In view of the
agreement of the scale counts of the type of interbrachialis with the counts of
the insular ater, and also because of the uncertainty of the locality of collection
of the type of interbrachialis, I concur with Schmidt in placing interbrachialis
definitely in the synonymy of ater. I am, therefore, describing the mainland
population from southern Baja California as a new species, Sauromalus australis.

DIFFERENTIAL CHARACTERS

The characters which have been used to distinguish the several species of
Sauromalus in this study are as follows:
A. Pattern and color.
B. Countable scale series.

1. Ventral scale rows between the gular fold and the anus.

2. The number of scales in a whorl around the tail two head-lengths
behind the vent, hereafter referred to as caudals. A head length, as
used here, is the distance from the anterior border of the ear opening
to the tip of the snout.

3. The number of scales around the upper part of the forelimb, here-
after referred to as humerals.

4. The number of dorsal scales in a head length.

C. Ratio of tail to total length.
D. Size.

Pattern is useful in dividing the banded forms into two groups, one
consisting of the forms townsendi and obesus, which have unicolor bands of
dark-brown or black, as opposed to the forms such as australis, ater, slevini,
and hispidus having the centers of the bands invaded by the ground color or
considerably lightened, thus giving a double-banded effect. Of the remaining
species, klauberi is distinguished by its pattern of fine spots, which have the
appearance of being arranged in some semblance of longitudinal lines, while
varius is characterized by its large, irregular dark blotches.

Of the countable scale series, the number of ventral scale rows between
the gular fold and the anus has been most commonly used in distinguishing
the species. Not infrequently it is difficult to arrive at the same count twice
because of the irregularity and discontinuity of the scale rows. Also, when

276 SAN Disco SociETY OF NATURAL HIsTory

making this count, care must be taken to begin the count at the edge of the
anus and not to include the scales which enter a short distance into the vent.

As is the case with all of the countable characters, the ventral scale rows
show an extreme range of variation both individually and geographically.
The mainland forms in particular are the worst offenders in this respect, while
the insular species fall comparatively close about the mean, this perhaps being
due in a large part to their more uniform environment and to the relative
crowding of the population into a small area.

To illustrate this point, insofar as the ventral scale count is concerned,
the following statistical summary is offered. The specimens of tumidus are
from Telegraph Pass, Gila Mts., Yuma County, Arizona. The specimens

of ater and slevini are from all islands where these forms occur.

tumidus ater slevini
Number of specimens il 18 17
Extreme range 132-170 130-151 107-123
Mean 15227 37 115.8
Standard deviation 12.53 6.45 5.76
Coefficient of variation, per cent 8.20 4.62 4.97

The number of caudal scales also shows considerable variation. Previously
this count has been made “around the thickest part of the tail,” but the hind
limbs of some specimens frequently made a count in this region difficult. Also
it seems best to make this count at a definite point two head-lengths behind the
vent, as the thickest part of the tail may be of some extent, thus allowing room
for variation in the count, depending upon the point in this region where the
count is taken.

The humeral count is made at the middle of the upper part of the forearm.
This is a rather difficult count to make, even in large specimens, because of
the irregularity of the rows of scales around the arm. It is, however, of great
value in distinguishing some of the forms.

The number of dorsal scales in a head length is determined by counting
the number of dorsal scales in a head length at a point half way between the
fore and hind limbs on the middorsal line. This character is most important
in separating the two closely related species, hispidus and slevini.

During the early part of this study it was thought that the ratio of tail
to total length would be of considerable significance in distinguishing several
of the species, but this has proven not to be true. Certain of the species average
much shorter tails than others, but the amount of overlap is too great to make
this of any value diagnostically. However, it is interesting to note the
considerable amount of intraspecific variation in this character. There is
apparently no sexual dimorphism in this character.

Size is useful only in the identification of adults of such forms as hispidus
and varius, which often reach a length in excess of 600 millimeters. S. 0.
tumidus has been found to have a greater average adult length than S. o.
townsendi, to which it is closely related, and this character may be of use in
some instances in distinguishing these two forms.

SHAW—THE CHUCKWALLAS 277

While chuckwallas in the southwestern United States and some of the
islands in the Gulf of California are not uncommon and may even be abundant
in certain areas, adequate homogeneous series for a statistical analysis have
been found lacking. Chuckwallas are rather well represented in collections in
this country, but for the most part they are from many scattered points in the
southwest. This lack of material is probably due to a number of factors. One
is the method of Protection employed by these large lizards when seeking to
escape their enemies, by running to a crevice in the nearest available boulder
and there wedging themselves tightly by inflating their lungs, often making
extrication impossible. Another reason, and perhaps the most important one,
is the large size of these lizards and the consequent unwillingness of some
collectors to “waste” often much-needed space and preservative on such relatively
common creatures.

RELATIONSHIPS WITHIN THE GENUS

That the several species of Sauromalus are all very closely related must be
apparent to those having only a slight degree of familiarity with them.
Speciation has apparently centered upon changes in the degree of coarseness
and spinosity of scalation and to a lesser extent changes in pattern. It is
probable that the original center of distribution was what is now the central
part of the Gulf of California and adjacent areas of Sonora and Baja
California. With the advent of the geological disturbances which brought
about the formation of the Gulf of California, the climatic changes resulting
from these greatly altered conditions, and the splitting off of the islands from
the Sonoran mainland and the peninsula of Baja California, speciation began.

The genus may be roughly divided into two groups, one consisting of
the coarse-scaled species, hispidus, slevini, klauberi, ater, and australis, and the
other group of varius, townsendi, tumidus, and obesus, which have relatively
fine and less spinose scalation. It may be assumed that the formation of the
Gulf of California was the original factor in separating the genus into two
divergent lines, which have been carried on to the present. Hispidus and varius
are apparently the oldest forms of the genus, their differentiation occurring
before that of the other species, through the early breaking off, by faulting,
of the islands upon which they now occur, hispidus being derived from a
peninsular population, while varius was developed from a Sonoran population.

Because of their close relationship to hispidus, the forms slevini and
klauberi are apparently the next to have been developed through insular
isolation. These two species are coarse-scaled and quite spinose but not as
much so as hispidus, nor do they attain the large size of that form. Probably
both are derived from the same ancestral population, which also became
separated from the mainland through geological disturbances. It appears that
Santa Catalina Island, which is inhabited by klauberi, became separated from
the ancestral population at an earlier date than did slevini, as klauberi has
lost its transverse bands and has become finely spotted instead. The same
trend in development is shown in slevini, of which two specimens have been
examined, in which the normal complement of four bands has been reduced

278 SAN Dreco Society oF NaturAL History

to two, the remainder of the dorsal surface of the body being sprinkled with
large brown spots.

S. ater has apparently been the last coarse-scaled form to develop through
insular isolation in the series outlined above. It also seems likely that this
species appeared at a relatively recent date, through the splitting off of a
number of coastal islands from the peninsula of Baja California, as it is
only slightly divergent from S. australis, which occupies the southern half of
the peninsula.

Of the relatively fine-scaled forms, varius, townsendi, tumidus, and obesus,
it is deemed probable that varius is the oldest by reason of its unique pattern

of blotches.

S. 0. townsendi, first described from Tiburon Island, has been found to
occur also on the mainland in Sonora. This species intergrades with S. o.
tumidus on the northwestern coast of Sonora, the two having similar scalation
but very distinct adult patterns in the males. S. 0. tumidus has apparently
given rise to S. o. obesus, with which it intergrades in central Arizona and
extreme southeastern California.

The following diagram represents my idea of the phylogeny of the genus:

australis Ae, obesus
Slevin!

tumidus
klauber!

townsend!

hispidus
varius

Ancestral Sauromalus
Fic. 1

SHAW—THE CHUCKWALLAS 279

DESCRIPTION OF SPECIES AND SUBSPECIES

Sauromalus hispidus Stejneger
SPINY CHUCKWALLA
Sauromalus ater Streets, Bull. U. S. Nat. Mus., no. 7, 1877, p. 36.
Sauromalus hispidus Stejneger, Proc. U. S. Nat. Mus., vol. 14, 1891, p. 409.

Type Specimen.—No. 8563 in the collection of the U. S. National
Museum.

Type Locality—Angel de la Guarda Island, Gulf of California, Mexico.

Distribution Angel de la Guarda, Smith, Pond, Granite, Mejia, and
South San Lorenzo islands, Gulf of California, Mexico.

Diagnosis—This is the coarsest-scaled, and, next to varius, the largest
member of the genus. The largest nuchals are equal to, or larger than the
largest plates on top of the head. The scalation of the limbs and tail is
extremely spinose and more or less strongly carinate. Adult specimens are
nearly uniform dark-brown or black above, while juveniles are transversely

banded with dark-brown or black double bands.

Description—Size large, form stout; head and body much depressed, the
former nearly triangular in outline from above and wider than long in adult
males, and longer than wide in females and juveniles. The top of the head is
covered with irregular, rough plates, largest on the frontal and temporal regions,
and becoming tubercular and spinose in the latter area. The nostrils open
upward and outward in a single, rounded, raised plate much nearer the tip of
the snout than the orbit. The superciliaries and supraoculars are small and
juxtaposed, the latter tubercular and occasionally weakly spinose. There is a
series of short, carinate suboculars, which, following the contour of the orbit,
pass upward and backward to the anterior edge of the ear opening, in the form
of more or less carinate plates. The labials are small and juxtaposed. The
rostral plate is represented by four nearly equal hexagonal plates. The
symphyseal plate is rather short, narrow and subtriangular. There are several
series of enlarged sublabials which merge into the relatively coarse and spinose
granular gular scales. There is a prominent gular fold covered with fine,
feebly spinose scales. The ear opening is nearly vertical, with an anterior
denticulation of from two to four enlarged spinose scales, the two central ones
being the longest. There is a prominent lateral neck fold covered by enlarged,
subconical, sharply spinose scales. The nuchal scales are quite large, some
as large or larger than the frontal plates, strongly spinose and grading gradually
into a broad median band of spinose dorsal scales which extends to the rump.
The scales on the lateral fold are enlarged and each scale is provided with a
short stout spine. The ventral scales are somewhat smaller than the median
dorsals and are weakly spinose. There are from 108 to 129 rows of scales
between the gular fold and the anus, averaging 121.4.

Dorsally, the scalation of the fore and hind limbs is extremely coarse,
almost equaling the nuchal scales in size, and more or less strongly carinate
and spinose. The humeral scales vary in number from 31 to 38 and average

35.6. The femoral pores range in number from 13 to 17 and average 14.9.

280 SAN Dreco Society oF NATURAL History

The tail is about equal in length to the head and body, varying from 49
to 53 per cent of the total length. The scalation is arranged in whorls, those
ventrally being smooth and usually non-spinose while dorsally and laterally
they are strongly spinose, especially towards the tip. The tail scalation is more
or less strongly carinate except at the base. There are from 23 to 28 caudal
scales in a whorl, averaging 25.5.

Coloration in Alcohol—Adults: above, the general coloration is a dull
olive-brown or nearly black with occasional evidences of irregular dark markings,
especially between the shoulders and laterally near the insertion of the fore
limbs. The ventral surface is yellowish- or grayish-brown and immaculate. The
gular region of some specimens shows evidences of dark streaks or spots.
Juveniles: the smallest specimen available is a sub-adult 390 mm. in length
from S. San Lorenzo Island. This is nearly uniform dark-brown dorsally, but
shows traces of one transverse band between the shoulders and another across
the rump. The ventral surface is yellowish-brown with faint indications of a
darker spotting. Schmidt (loc. cit., pl. 50, fig. 2) figures a small specimen of
this species which shows four transverse bands on the body and at least four
bands on the tail. The ground color between the bands is light with an
irregular spotting or streaking of darker color.

Remarks.—This species is apparently most closely related to S. slevini,
which inhabits a group of islands further to the south in the Gulf of California.
From slevini, hispidus may be distinguished by its much larger adult size and
coarser, more spinose scalation. In hispidus the number of dorsal scales in a
head length varies from 16 to 21 while the same count in slevini gives from
20 to 28 scales.

Habits—Van Denburgh (loc. cit., p. 101) remarks: “This chuckwalla
was abundant in rocky canyons. They were found by looking for the spiny
tails protruding from under rocks. On Pond Island they carried about grea:
numbers of long sharp spines of a cactus, Opuntia, which grew in scattered
clumps over the island and under which they ran for shelter. Several were
found with spines sticking even into their eyes. All stomachs examined
contained vegetable matter. On Granite Island, a small rock near the north end
of Angel de Ia Guarda Island, many dead chuckwallas were found strewn
about the tops of the osprey’s nests.”

Sauromalus slevini Van Denburgh
MONSERRATE CHUCKWALLA

Sauromalus slevini Van Denburgh, Occas. Papers Calif. Acad. Sci., no. 10,

voll 19225 9.397.

Type Specimen.—No. 50503 in the collection of the California Academy
of Sciences.

Type Locality—South end of Monserrate Island, Gulf of California,
Mexico.

Distribution—Monsetrrate, Carmen, and Coronados islands, Gulf of
California, Mexico.

SHAW—THE CHUCKWALLAS 281

Diagnosis —A medium sized chuckwalla, intermediate in scalation between
S. hispidus and S. ater. From hispidus it may be distinguished by the greater
number of dorsal scales in a head length (20-28) and the smaller size of the
nuchal scales as contrasted with the plates on the frontal region. The ventral
scale rows range from 107 to 123 and average 115.8.

Description —Form stout; head and body much depressed, the former
nearly triangular in outline from above, the latter very broad with a strong
lateral fold. The top of the head is covered with small, smooth, juxtaposed
plates largest on the frontal region. The nostril is pierced in a single rounded
plate much nearer the tip of the snout than the orbit. There is a series of
carinate suboculars much longer than high, which, posterior to the eye, pass
upward and backward to the ear opening in the form of weakly mucronate
tubercles. The rostral is divided into four small scales of equal size. The
labial plates are small and subequal. The ear opening is vertical, or nearly so,
with an anterior denticulation usually of two prominent spinose scales. Below
the infralabials are several series of sublabials which grade into the granular
gular scales. There is a strong gular fold. Immediately behind the ear opening
there is a prominent lateral neck fold bearing many spinose, subconical scales.
Above and slightly forward of the shoulder there is a fold bearing a small
patch of enlarged, subconical scales. The nuchal scales are much larger than
the median dorsal scales and are strongly spinose. Medianly, between the
rump and the shoulders, there is a broad band of enlarged spinose dorsal
scales. Laterally, between the median dorsal scales and the enlarged, stout,
spinose scales of the lateral fold, the scalation is finer, spinose, and slightly
imbricate. The ventral scales are somewhat smaller than the median dorsal
scales, ranging in number from 107 to 123 and averaging 115.8.

Dorsally, the scales of the forelimb are quite large, mucronate and weakly
carinate. The humeral scales vary in number from 30 to 37 and average 33.8.
On the hind limb the scalation is also coarse, being coarsest on the tibial
portion, mucronate and weakly carinate. The femoral pores vary in number
from 12 to 18 and average 14.5.

The tail is depressed at its base and bears numerous whorls of scales,
more or less strongly spinose and carinate everywhere except dorsally and
ventrally at the base, there becoming smooth and very weakly spinose. In four
specimens the caudals range in number from 22 to 23. The ratio tail to total
length varies from .524 to .565.

Coloration in Alcohol—Dorsally, the ground color of the body is
yellowish-brown to dark-brown, the variation depending upon the age of the
individual, adult specimens being nearly uniform dark-brown with faint irregular
streaks and spots of darker brown or black faintly apparent. The top of the
head is dark-brown or nearly black, becoming yellowish-gray on the sides.
Younger specimens have a lighter brown ground color and are provided with
from two to four prominent dark-brown double bands across the back.

Ventrally, the ground color is yellowish-brown. The gular region is spotted,
streaked or marbled with dark-brown. Above, the tail is yellowish or olive-
brown; below, a somewhat lighter brown.

282 SAN Disco Society oF NATURAL History

Remarks.—While its coarse and spinose scalation indicates affinities with
hispidus, slevini, on the basis of geographical grounds, together with similarities
in scalation, shows a close relationship with klauberi, which inhabits Santa
Catalina Island, only a few miles to the east of Monserrate Island.

Habits.—Nothing is known concerning the habits of this species, although
they are presumably essentially the same as those of the other members of the
genus.

Sauromalus klauberi Shaw
SPOTTED CHUCKWALLA

Sauromalus klauberi Shaw, Trans. San Diego Soc. Nat. Hist., vol. 9, no. 28,
1941, p. 285.

Type Specimen.—No. 6859 in the collection of L. M. Klauber.
Type Locality.—Santa Catalina Island, Gulf of California, Mexico.
Distribution—Confined to the type locality.

Diagnosis —Scalation moderately coarse, being intermediate between that
of S. slevini and S. ater. The ventral scales number from 128 to 132 in the
three known specimens. There are no transverse body bands, the dorsal surface

of the body being sprinkled with small brown or black spots instead.

Description —Form stout; the head and body much depressed, the latter
broad with a strong lateral and gular fold. On top, the head is covered with
small plates, largest on the frontal region. The supraoculars and superciliaries
are small and juxtaposed. The nostril is pierced in a single rounded plate with
raised edges, much nearer the tip of the snout than the orbit. The rostral is
vertically divided into four small scales of equal size. The labial plates are
small and subequal. The suboculars are short and carinate and, following the
contour of the orbit, pass upward and backward to the anterior edge of the
ear opening in the form of nearly round, feebly spinose plates. The ear opening
is large and nearly vertical, with an anterior denticulation consisting of two
adjacent strongly spinose scales bordered by two much smaller scales of the
same shape. Immediately behind the ear opening is a prominent lateral neck
fold bearing large, subconical, spinose scales interspersed with smaller scales
of the same shape. The symphyseal is subtriangular and longer than wide.
Below the infralabials are several series of sublabials which merge gradually
with the very small and feebly spinose granular gular scales.

The nuchal scales are irregular in size, there being large, subconical,
spinose scales scattered among much smaller scales of the same shape. The
largest nuchals are somewhat smaller than the largest scales of the postauricular
fold. Above the shoulder there is a group of enlarged spinose scales. There is
a middorsal band of enlarged spinose scales extending from between the
shoulders to the rump. Between the shoulders, this band of enlarged middorsal
scales is bordered on each side by elongated and strongly spinose scales,
irregularly distributed among those of much smaller size. Laterally, the scales
are much reduced in size, imbricate, and spinose, becoming larger on the
prominent lateral fold. The ventral scales are smaller than the median dorsals,

SHAW—THE CHUCKWALLAS 283

imbricate, and feebly spinose. The number of scale rows between the gular
fold and the anus ranges from 128 to 132 and averages 130.3.

Dorsally, the scales on the fore limbs are large, spinose and weakly
carinate; below, spinose to a slight degree and greatly reduced in size. The
humeral scales vary from 36 to 39 and average 37.6. On the hind limbs the
dorsal scales are also quite coarse, those of the tibial portion being largest,
weakly mucronate, and carinate. The femoral pores range in number from 13
to 16 and average 14.3.

The tail is depressed at its base. The scalation is arranged in whorls of
square or rectangular scales which are strongly spinose everywhere except at
the base, where dorsally and ventrally they are only feebly spinose. There are
24 caudal scales in two specimens on which this count was made.

Coloration in Alcohol—In adult specimens the ground color above is
dark, becoming nearly black on the head. On the ground color of the dorsal
surface of the body irregular black markings are present, but no tendency
toward a banded arrangement is indicated. In a juvenile specimen the ground
color of the dorsal surface of the body is gray, well sprinkled with small brown
spots which have some semblance of arrangement into longitudinal rows. The
gular region is spotted or streaked with brown. The chest is streaked with
reddish-brown and the belly is immaculate, except along the edges, where small
brown spots are faintly in evidence. The tail is uniform gray, dark-brown or
greenish-brown, except at the base, where it may be flecked with yellow.

Remarks.—S. klauberi is most closely related to S. slevini, which occurs
on Carmen, Coronados, and Monserrate islands, only a few miles to the north
and west of Santa Catalina Island. Apparently klauberi became isolated from
the ancestral population through geological processes which occurred at an
earlier date than those which separated the islands on which slevini occurs, thus
allowing sufficient time for development of such a distinctive character as the
lack of transverse body bands. In general, the scalation is finer than that of
slevini, especially on the limbs, and the heterogeneous arrangement of the
nuchals and the elongated anterior dorsal scales are also peculiar to this form.

S. slevini may possibly be undergoing a similar evolutionary development
at the present, for specimens have been examined which lacked the normal
complement of four transverse body bands, two anterior bands being present
and the remainder of the dorsal surface of the body sprinkled with large
brown spots.

Habits—Klauberi, like the other species of the genus, is herbivorous.
Material taken from the large intestine of the three specimens has been identified
as follows: leaflets of Cercidium floridum; fruits of Euphorbia sp.; parts of a
spikelet of Festuca sp.; and leaves of Acacia greggii. The intestine of the type
specimen also contained 14 unidentifiable hard-shelled seeds somewhat
resembling a pinon nut in shape, but smaller.

Parasites —The large intestine of each of the three specimens contained
a large number of nematodes which, according to Dr. L. R. Penner, formerly
of the Research Hospital of the San Diego Zoo, represent two or more new
species of the genus Alaeuris.

284 SAN Disco SoctETy OF NATURAL History

Sauromalus ater Duméril
Espiriru SANTO CHUCKWALLA

Sauromalus ater Duméril, Arch. Mus. Hist. Nat. Paris, vol. 8, 1856, p. 536.

Sauromalus interbrachialis Dickerson (part), Bull. Amer. Mus. Nat. Hist.,
vol. 41, 1919, pp. 463-64.

Sauromalus ater Schmidt (part), Bull. Amer. Mus. Nat. Hist., vol. 46, 1922,

pp. 640-41.

Type Spectmen.—In the collection of the Museum d’Histoire Naturelle,
Paris.

Type Locality—Not definitely known but undoubtedly one of the several
islands in the southern part of the Gulf of California where this species is
known to occur.

Distribution—Espiritu Santo, Isla Partida, San Francisco, San Diego,
Santa Cruz, and San Marcos islands, Gulf of California, Mexico.

Diagnosis.—Similar to S. australis, inhabiting the adjacent peninsula, but
distinguished from that form by its somewhat coarser scalation with corre-
spondingly lower scale counts. The number of ventral scale rows between
the gular fold and the anus varies from 130 to 151, averaging 139.7; humeral
scale count ranging from 35 to 45 and averaging 40.2.

Description—Form stout; head and body depressed, the former nearly
triangular in outline from above and as wide or wider than long in adult males,
and longer than wide in adult females and juveniles. Above, the head is
covered with small, rounded, non-imbricate plates, largest in the frontal and
parietal regions. The supraoculars and superciliaries are small and juxtaposed.
The nostril is pierced in a single rounded plate with raised edges directed
upward and slightly backward, much nearer the tip of the snout than the orbit.
There is a series of large, weakly carinate suboculars which, posterior to the
eye, pass slightly upward and backward to the anterior border of the ear
opening in the form of enlarged tubercles. The rostral is occasionally entire
but is more frequently represented by four hexagonal scales of equal size. The
symphyseal is long and subtriangular. The ear opening is vertical, or nearly so,
with an anterior denticulation consisting of two to five long, spinose scales.
Behind the ear opening there is a prominent crescent-shaped neck fold which
is covered with many tubercles or subconical spines. The lips are bordered by
short, subequal plates. Below the infralabial plates are several series of sublabials
which become successively smaller as they merge with the granular gular scales.
There is a prominent gular fold.

The nuchal scales are somewhat larger than the largest median dorsal
scales and much smaller than the plates on the frontal region. The nuchal
scales may be tuberculate and spinose or simply flattened with an obtuse
posterior spine. There is a broad series of enlarged, feebly spinose, median
dorsal scales which are rectangular in shape, extending from the nuchals to the
rump. Laterally, the scalation is much reduced in size, spinose and slightly
imbricate. The scales on the strong lateral fold are nearly as large as the
median dorsals and are provided with a short, stout spine. The ventral scales

SHAW—THE CHUCKWALLAS 285

are slightly smaller than the median dorsals and are occasionally feebly spinose.
There are from 130 to 151 rows of scales between the gular fold and the anus,
averaging 139.8.

Dorsally, the scalation of the fore limb is coarse and obtusely spinose.
Below, the scales are very much reduced in size and are granular. The number
of humeral scales varies from 35 to 45 and averages 40.2. On the hind limb
the scalation is quite coarse, the dorsal tibial scales being largest, mucronate
and weakly carinate. Below, much reduced in size. The femoral pores vary
in number from 17 to 21 and average 18.7.

The tail is depressed at its base and ranges from 50 to 55 per cent of
the total length. The scalation is coarse and arranged in whorls, the posterior
three-fourths being strongly keeled and spinose, the anterior one-fourth relatively
smooth and less strongly spinose dorsally and ventrally, becoming feebly spinose
laterally. There are from 24 to 33 caudal scales in a whorl, averaging 28.3.

Coloration in Alcohol—The ground color of the dorsal surface of the
body is dull yellowish-brown or occasionally grayish-brown. The top of the
head is dark-brown, usually becoming somewhat lighter on the sides. There
are four broad transverse bands across the back. In juveniles a fifth band is
usually present on the nape. The centers of the transverse bands are invaded
by the light ground color, the dark-brown or black anterior and posterior borders
presenting a double-barred effect. Middorsally, between the bands, the ground
color is yellowish and is spotted with brown or black. The gular region and
the chest are a dull gray or brown and more or less obscurely marbled, streaked,
or spotted with brown or black. The venter is grayish- or yellowish-brown
spotted with brown laterally. There are four or five dark-brown bands encircling
the tail with areas of yellowish-brown between. The limbs are grayish- or
yellowish-brown spotted with dark-brown.

Variation. —The following is a summary of the countable scale series in
this species :

Ventrals Caudals Humerals
Number of specimens 18 9 17
Extreme range 130-151 24-33 35-45
Mean 139.7 28.3 40.2
Standard deviation 6.45 3.95 2.59
Coefficient of variation, per cent 4.62 14.0 6.44

It will be seen that the coefficient of variation is relatively small in this
species, compared to such mainland forms as townsendi and obesus, thus
indicating narrower limits of dispersion for these characters and making them
more reliable diagnostically. The specimens from which the above counts
were obtained were from all islands where this species occurs, as sufficient
material from a single island was not available.

Remarks.—S. ater is apparently most closely related to S. australis, which
inhabits the southern part of the Gulf coast of the peninsula of Baja California.
Ater, however, has a more coarse and spinose scalation than australis and in
this respect shows an approach to slevini, especially in the spinose character of
the nuchal and post-auricular regions. The majority of specimens of ater

286 SAN Disco Society oF NATurRAL History

differ in general coloration from australis in that they are yellowish-brown.
However, LMK 3855 from Espiritu Santo Island has a grayish-brown ground

color, in which respect it is similar to the mainland australis.

A specimen of Sauromalus, CAS 51465, from San Marcos Island, some
150 miles north of Santa Cruz Island, the northernmost of the southern group
of islands where ater occurs, is here referred to this species, as the ventral scale
count of 140 agrees with those of the specimens from the islands further
south. This makes for an erratic distribution, as the islands on which klauberi
and slevini occur destroy the geographical continuity of the distribution of ater
by inserting themselves between Santa Cruz and San Marcos islands. The
occurrence of this species in such widely separated areas seems to support the
idea that ater has but recently become differentiated from a peninsular form
by the splitting off of the several islands on which it now occurs. However,
the single specimen from San Marcos Island seems to have a more oval shaped
head in outline from above and a more truncate snout than the other specimens
of ater which have been examined, and additional specimens from that locality
may possibly disclose further differences which would warrant its distinction
as a separate species.

Sauromalus australis sp. nov.
PENINSULAR CHUCKWALLA

Sauromalus ater Mocquard (part), Nouv. Arch. Mus. Hist. Nat. Paris, ser.
4yvolt 1, 1899) 53302:

Sauromalus obesus Schmidt (part), Bull. Amer. Mus. Nat. Hist., vol. 46,
1922, p. 641.

Type Specimen.—An adult male, No. 30170 in the collection of L. M.
Klauber; collected at San Francisquito Bay, Baja California, Mexico, July 30,
1938, by Robert S. Hoard. The following five paratypes have also been
examined: LMK 30168, Loreto; LMK 30169, 33 mi. north of Canipole; CAS
53710, Agua Verde Bay; SDSNH 17707, Comondi; SDSNH 17708, La
Paz.

Specimens which undoubtedly represent this species are recorded by

Mocquard (loc. cit.) from Santa Agueda, San Ignacio, and Mulegé.

Diagnosis.—Scalation moderately coarse, being intermediate between that
of S. ater and S. 0. obesus, but somewhat finer and less spinose than that of
ater. From obesus it may be distinguished by the transverse bands, the centers
of which are invaded by the lighter ground color, the dark-brown or black
anterior and posterior borders giving a double-barred effect. The ventral
scale rows vary in number from 151 to 186 and average 163.5. The humeral
scales range in number from 46 to 55 and average 49.0.

Description of the Type-—Form stout; head and body depressed, the
former with a greatly swollen temporal region which curves rather abruptly
inward to a narrow, pointed snout. Above, the head is covered by small,
smooth, juxtaposed plates, largest on the frontal and parietal regions and
becoming tuberculate and feebly spinose in the latter area. The nostril is

SHAW—THE CHUCKWALLAS 287

pierced in a single rounded plate directed upward and outward, much nearer
the tip of the snout than the orbit. The superciliaries are small and juxtaposed.
There is a series of carinate suboculars, which, posterior to the orbit, pass
upward and backward to the anterior border of the ear opening. The rostral
is vertically divided into four scales of equal size and shape. The labial plates
are all short and subequal. There are several series of sublabials which merge
into the finely granular gular scales. There is a well-defined gular fold. The
ear opening is nearly vertical, with an anterior denticulation of four strongly
spinose and elongate scales.

Medianly, the nuchal scales are roughly triangular in outline and are
provided with a short obtuse spine. Laterally the nuchal scales become
subconical or tuberculate and more conspicuously spinose. Posterior to the ear
opening there is a prominent lateral neck fold which bears strongly spinose
subconical scales. There is a median band of enlarged dorsal scales. Anteriorly,
between the shoulders, these are rather sharply spinose, becoming less so
posteriorly. Laterally, the scalation is finer, becoming coarser and more spinose
on the strong lateral fold. The ventral scales are smaller than the median
dorsals, smooth, imbricate and feebly spinose. There are 151 rows of scales
between the gular fold and the anus.

On the fore limb the scalation is coarse, obtusely spinose and weakly
carinate. There are 46 humeral scales. On the hind limb the scalation is
coarsest on the tibial portion, carirate and obtusely spinose. The scales of the
dorsal surface of the foot are sharply spinose. The femoral pores number
18-19.

The tail is depressed at its base and covered with whorls of rectangular,
carinate and more or less strongly spinose scales, except at the base where
dorsally and ventrally they are smooth and only very feebly spinose. The
caudal scales number 36. The ratio of tail to total length is .558.

The ground color of the top of the head is dark-brown, becoming yellowish
on the frontal region. The scales on the parietal region are tipped with gray.
Immediately anterior to the ear opening there is a dark-brown blotch. The
labials are gray or white. The gular region is gray with irregularly disposed
streaks or spots of dark-brown. Dorsally, the ground color of the body is a
light-gray with indications of four transverse bands of black or reddish-brown
which do not meet on the median line. The band between the shoulders is
most prominent and is spotted or streaked with gray through its long axis,
thus giving a double-barred effect. Between the transverse bands are small spots
of brown or black. The chest and the undersides of the forearms are gray,
spotted or streaked with brown or black. The tail is an immaculate yellowish-
brown with faint indications of four darker encircling bands. This color
description is of the type specimen as preserved in alcohol.

V ariation.—As in the other mainland forms of Sauromalus a considerable
amount of variation in both coloration and scalation is exhibited by S. australis.
SDSNH 17708 from La Paz possesses only two transverse bands, like some
specimens of slevini which have been examined. The band between the
shoulders is indicated by the black spotting and streaking which overlies a

288 SAN Disco Society of NATURAL History

dark-gray ground color. The band posterior to this is evident only as a
closer grouping of many small black spots.

Usually the coloration of australis differs from that of ater in being gray
instead of yellowish-brown, although CAS 53710 from Agua Verde Bay has the
coloration of the majority of the specimens from the islands.

The following is a summary of the variation in countable scale series in
this species, as well as a comparison of these counts with those of ater:

australis ater
Ventrals 151-163.5—-186* 130-139.8-151
Humerals 46-49.0-55 35-40.2-45
Caudals 32-34.7-37 24-28.3-33

Kemarks.—S. australis is clearly most closely related to S. ater. It may
be distinguished from ater, however, by its generally finer scalation and by its
lack of the degree of spinosity achieved by ater. It seems probable that australis
is the end form of a once widespread species that became differentiated into
the species now found on the islands in the Gulf of California.

The relationship between australis and obesus is uncertain at this time
because of the lack of material from between San Francisquito Bay on the
south and the U. S.-Mexican boundary on the north. However, when more
collecting has been done in this region, the ranges of these two forms will
probably be found to overlap with no intergradation present, for they are
apparently quite distinct from one another despite the overlap shown in
countable scale series.

Habits—Presumably essentially the same as the other members of the
genus.

Sauromalus varius Dickerson
PIEBALD CHUCKWALLA

Sauromalus Townsend, Bull. Amer. Mus. Nat. Hist., vol. 35, 1916, p. 428.
Sauromalus varius Dickerson, Bull. Amer. Mus. Nat. Hist., vol. 41, 1919,
ps 404: :
Type Specimen.—No. 64441 in the collection of the U. S. National

Museum.

Type Locality—San Estéban Island, Gulf of California, Mexico.

Distribution —Confined to the type locality.

Diagnosis —This very interesting species may be most easily distinguished
from the other members of the genus by its large size and by its peculiar pattern
of large irregular reddish-brown blotches on a yellowish ground color.

Description —Size large, adults sometimes reaching a length in excess of
600 millimeters. Head and body much depressed, the latter very broad. From
above, the head is nearly triangular in outline, broader than long in adult males
and longer than broad in females and juveniles. The top of the head is covered

* The outer figures indicate the extremes; the central figure the mean.

SHAW—THE CHUCKWALLAS 289

with smooth, irregular plates, largest on the frontal and parietal regions,
and becoming tubercular in the latter region. The nostrils open upward and
somewhat backward in a single oval, raised plate, much nearer the tip of the
snout than the orbit. The superciliaries and the supraoculars are small and
juxtaposed. There is a series of short, smooth suboculars which, following the
contour of the orbit, pass upward and backward to the anterior border of the
ear opening. The labials are quite small and juxtaposed. The rostral plate is
divided into four subequal hexagonal plates. The symphyseal is long and
narrow. There are several series of enlarged sublabials which merge with the
granular gular scales. There is a prominent gular fold. The ear opening is
nearly vertical with an anterior denticulation, usually of two short and bluntly
spinose scales. There is a lateral neck fold posterior to the ear opening covered
by very small tubercular or subconical scales. The nuchal scales are not greatly
enlarged; they are short, and tubercular or subconical, and only very feebly
spinose, grading into a median band of relatively large feebly spinose dorsal
scales which extend to the rump. Laterally the scalation becomes finer and
somewhat granular with short blunt spines. The scalation on the strong lateral
fold is somewhat enlarged and is bluntly spinose. The ventral scales are smaller
than the median dorsals and may be feebly spinose, especially on the chest.
There are from 150 to 165 rows of scales between the gular fold and the anus,
averaging 158.5 in 11 specimens.

Dorsally, the scalation of the forelimbs is relatively coarse, the individual
scales being as large or larger than the largest nuchal, weakly spinose and very
faintly carinate. Below, they are much reduced in size and are granular. The
humeral scales range in number from 52 to 58 and average 54.3. On the hind
limb the scalation of the femoral portion is relatively fine, increasing in size on
the tibial part of the leg but not quite as coarse as the dorsal scalation of the
forelimbs, occasionally weakly carinate and feebly spinose. Below, the scales
are much reduced in size and weakly spinose. There are from 15 to 18 femoral
pores, averaging 16.1.

The tail ranges in length from 50 to 56 per cent of the total length. The
scalation is arranged in whorls of smooth and weakly spinose scales, except
dorsally towards the tip where they become more sharply spinose and faintly
carinate. The caudal scales range in number from 30 to 35 and average 32.0.

Coloration in Alcohol—The ground color is usually yellowish or
occasionally orange-brown with irregular blotches and spots of reddish-brown
or black on the tail and body. Generally, the snout, chin, and the top of the
head are black or nearly so.

Remarks.—The ancestry of varius cannot be traced directly to any living
species of the genus. Apparently this form became isolated at a very early
date from a mainland population in Sonora, which later gave rise to townsendi,
and its long period of separation has allowed sufficient time for the development
of such distinctive characters as its irregular blotching, large size, and relatively
fine, smooth scalation.

Habits—Regarding this lizard Van Denburgh (1922b, p. 103) says:

“These huge lizards were abundant in the dry washes and small rocky canyons

290 SAN Disco Society oF NaturaL History

of San Estéban Island. Here they lived under rocky ledges and piles of lava.
Numerous droppings about the mouths of their dens, and often their protruding
tails, made it easy to find them. They were easily captured by pulling them
out of their retreats by their tails, and made no attempt to bite when caught.
Five were found in a compact mass in the center of a patch of Opuntia.”

Schmidt (1922, p. 643) states: “Large numbers of the big spotted lizards
of the species, as well as of Ctenosaura hemilopha, were conspicuous, and were
secured by pulling them out from under the rocks where they took refuge, or
by turning over the rocks. Dr. J. N. Rose, who was a member of the party,
has kindly identified the stomach contents of three specimens. He writes:
‘The contents of two stomachs are entirely made up of the flowers of
Pachycereus pringlei Britton and Rose. The third stomach is also largely filled
with this cactus flower, but also contains numerous small leaflets of some
leguminous plants, probably some Cercidium’.”

These large lizards, when on display in an outdoor pit at the San Diego
Zoo, apparently do very well on a diet of lettuce, bananas, tomatoes, apple,
cabbage, and the yellow blossoms of the Palo Verde trees which grow in
the pit.

Sauromalus obesus townsendi Dickerson
SONORAN CHUCKWALLA

Sauromalus ater Belding (part), West Amer. Sci., vol. 3, no. 24, 1887, p. 97.

Sauromalus townsendi Dickerson, Bull. Amer. Mus. Nat. Hist., vol. 41, 1919,
p. 464.
Type Specimen.—No. 64442 in the collection of the U. S. National

Museum.
Type Locality —Tiburén Island, Gulf of California, Mexico.

Distribution—Tiburon Island and the adjacent Sonoran mainland at
least as far south as Guaymas and east to the vicinity of Hermosillo.

Diagnosis—A small chuckwalla distinguished from S. 0. tumidus by the
lack of reddish coloration in the males and also by somewhat coarser scalation,
with correspondingly lower average scale counts.

Description—Form stout; head and body depressed, the former as wide
or wider than long in adult males and longer than wide in females. The top
of the head is covered with small and more or less convex plates, largest on the
frontal and parietal regions, becoming tuberculate and occasionally weakly
spinose in the latter area. The supracculars and superciliaries are small and
juxtaposed. The nostril is pierced in a single rounded plate much nearer the
tip of the snout than the orbit. The rostral plate is usually vertically divided
into from two to four scales of equal size and shape, although these may be
sometimes united to form a single hexagonal plate much wider than high.
There is a series of weakly carinate suboculars, which posterior to the eye pass
upward and backward to the ear opening in the form of more or less mucronate
tubercles. The labial plates are small and subequal. Below the infralabial plates
are several series of enlarged sublabials which change gradually into the granular

SHAW—THE CHUCKWALLAS 291

gular scales. There is a strong gular fold. The ear opening is vertical, or
nearly so, with an anterior denticulation usually consisting of from two to
four strongly spinose scales. Behind the ear opening there is a lateral neck
fold covered with many large subconical or tubercular scales which are strongly
spinose in most males, but less so in females. The nuchal scales may be
subconical and strongly spinose or simple flattened squares with one corner
projected into a spine. From the rump to the shoulders there extends a broad
median band of spinose dorsal scales. Laterally, the scalation is much reduced
in size, imbricate and spinose, becoming enlarged and strongly spinose on the
prominent lateral fold. The ventral scales are somewhat smaller than the
middorsal scales and may be feebly spinose. The number of scales between the
anus and the gular fold varies from 138 to 157 and averages 148.7 in 11
specimens.

Dorsally, the scalation of the forelimb is quite coarse, spinose and
occasionally weakly carinate; below, much reduced in size and granular. There
are from 37 to 45 scale rows around the humeral part of the arm, averaging
40.4.

Dorsally, the scalation of the femoral portion of the hindlimb is coarse
and obtusely spinose, increasing in size on the tibial portion and usually strongly
spinose and carinate.

The tail varies in length from 49 to 55 per cent of the total length and is
depressed at its base. The catidal scales are arranged in whorls, usually strongly
spinose and carinate everywhere except at the base, where dorsally and ventrally
they become smooth and very weakly spinose. There are from 27 to 30 caudal
scales in a whorl two head lengths behind the vent, averaging 28.5 in 11
specimens.

Coloration in Alcohol_—The top of the head is light-brown to yellowish-
gray. There is a very prominent dark spot just behind the ear. There are from
4 to 5 dark-brown transverse dorsal bands between the nape and the rump,
more or less distinct, and spotted or irregularly streaked with gray. Between
the bands there may be well-defined lines of light-yellow on the grayish or
yellowish ground color. The ground color of the gular region and the belly
varies from yellowish to gray with occasional peppering of fine black spots. The
limbs are brownish or gray with spots of yellowish-brown or gray upon them.
The tail is gray or yellowish-brown with four or five dark-brown or nearly black
encircling rings.

V ariation—The following is a summary of the countable scale series in
the available specimens of this subspecies:

Ventrals Caudals Humerals

Range 138-157 27-30 BUA
Mean 148.7 28.5 40.4
Standard deviation 6.37 69 6.55
Coefficient of variation, per cent 4.29 2.41 16.21

As with the other forms of Sauromalus, there is a noticeable variation in
pattern and coloration. Specimens from the Sonoran mainland have a grayish
ground color with nearly black transverse bands, while those from Tiburén

292 San Disco Society oF NAtTuRAL History

Island have a yellowish ground color and brown transverse bands. Some of
the mainland specimens have the transverse bands reduced to only the faintest
indications and in this respect are similar to some specimens of S. australis
from Baja California.

Remarks.—By reason of its coarse and spinose scalation, townsendi is
apparently quite closely related to S. 0. tumidus. The larger size and usually
brilliant reddish dorsal and ventral suffusion of the adult males of tumidus,
together with the higher average scale counts in this form, will serve to
distinguish it from townsend).

The following is a comparison of scale counts in these two forms:

townsendi tumidus
Number of specimens 11 31
Ventrals 138-148.7-157 132-159.4-190
Caudals 27-28.5-30 29-32 4-37
Humerals 37-40.4-45 39-44.6-50

In the specimens of tumidus and townsendi which have been examined,
there is a quite noticeable difference in size. In 22 adult specimens of tumidus
with complete tails, the average total length was found to be 325 mm., while
5 adults of townsendi with complete tails averaged only 285 mm. in length.

Specimens from the northwestern coast of Sonora in the vicinity of Las
Chollas Point and Punta Penasco are considered intergrades between tumidus
and townsendi, as they have the coloration of the latter and the scalation of the
former.

Localities of Collection—Sonora, Mexico: ‘Tiburén Island (type
locality) ; Cerro Prieta, 3 mi. nw. of Puerto de Lobos; 5 mi. n. of Guaymas;
Miramar, 3 mi. nw. of Guaymas; San Carlos; Guaymas; 1 mi. w. of Empalme;
54 mi. sw. of Hermosillo.

Sauromalus obesus tumidus subsp. nov.
GILA CHUCKWALLA

Sauromalus obesus Gloyd (part), Bull. Chi. Acad. Sci., vol. 5, no. 5, p. 106,

19377,

Type Specimen—An adult male, No. 27323 in the collection of L. M.
Klauber; collected at Telegraph Pass, Gila Mountains, Yuma County, Arizona,
June 15, 1937, by L. M. Klauber. The following twelve paratypes are also
available from the same locality: LMK 8613; LMK 27551; LMK 33170-75;
LMK 33224-25; LMK 34141; LMK 35090.

Diagnosis. —A relatively large chuckwalla distinguished from S. 0. obesus
by its coarser and generally more spinose scalation, especially on the limbs and
tail. From S. 0. townsendi it may be segregated by the brilliant red dorsal and
ventral coloration of the males, by its higher average scale counts, and by its
larger adult size.

Description of the Type-——Form stout; head and body depressed, the

former with a greatly swollen temporal region which curves abruptly inward to

SHAW—THE CHUCKWALLAS 293

a narrow, pointed snout. Above, the head is covered by small, smooth,
juxtaposed plates, largest on the frontal and parietal regions and becoming
tuberculate and feebly spinose in the latter area. The nostril is pierced in a
single rounded plate directed upward and outward, much nearer the tip of
the snout than the orbit. The superciliaries are small and juxtaposed. There
is a series of carinate suboculars, which, posterior to the orbit, pass upward and
backward to the anterior border of the ear opening. The rostral is vertically
divided into four scales of equal size and shape. The labial plates are numerous
and subequal. There is a series of sublabials which merge gradually with the
granular and subconical gular scales. There is a well-defined gular fold. The
ear opening is nearly vertical with an anterior denticulation of three strongly
spinose and elongate scales.

The largest nuchals are somewhat smaller than the plates on the frontal
region, somewhat triangular in outline and more or less strongly spinose.
Posterior to the ear opening there is a prominent lateral neck fold which bears
strongly spinose subconical scales. There is a median band of enlarged dorsal
scales. Anteriorly, between the shoulders, these are rather sharply spinose,
becoming less so posteriorly. Laterally, the scalation is finer, becoming coarser
and sharply spinose on the strong lateral fold. The ventral scales are smaller
than the median dorsals, smooth, imbricate, and feebly spinose. There are 132
rows of scales between the gular fold and the anus.

On the forelimb the scalation is coarse, obtusely spinose and weakly
carinate. There are 40 humeral scales. On the hindlimb the scalation is
coarsest on the tibial portion, carinate and obtusely spinose. The scales of the
dorsal surface of the foot are sharply spinose. The femoral pores number 16 on
the left leg, those on the right leg having been injured.

The tail is depressed at its base and covered with whorls of rectangular,
carinate and more or less strongly spinose scales, except at the base where dorsally
and ventrally they are smooth and only very feebly spinose. The caudal scales
number 32. The ratio of tail to total length is .499.

The head, nape, chest, gular region, forelimbs, hindlimbs, and groin are
black with an irregular spotting of yellow on the nape and on the fore limbs.
The middorsal region is suffused with red and generously flecked with yellow
and black spots. The ventral surface is red and irregularly spotted with black.
The tail is an immaculate yellowish-brown with no trace of encircling bands.
This color description is of the type specimen as preserved in alcohol.

Variation —T umidus, like obesus, exhibits considerable variation in both
coloration and scalation. There is a marked sexual dimorphism in the adults.
In specimens from Yuma County, Arizona, nearly all of the males have lost
their juvenile pattern of well-defined transverse bands, except for the prominent
black band across the rump and between the shoulders. In most cases the tail
rings are retained. In the majority of males the head, neck, shoulders, gular
region and forelimbs as well as the hindlimbs, rump and groin are jet-black with
an occasional flecking of gray or white. The ground color of the dorsal surface
of the body is yellowish-gray, well spotted with large amounts of black and
red, the former color becoming more prevalent laterally. The belly is brick-red

294 SAN DrieGco Society oF NaturAL History

spotted with black. The tail is grayish or straw-color with or without indication
of darker rings. LMK 34141 from the Gila Mts. has the red and black on
the dorsal surface of the body arranged into reticulations, making a most striking
and beautiful specimen.

The following color notes describe a live adult male taken at Telegraph
Pass, Gila Mts., Yuma County, Arizona: The head, gular region, chest, neck
and forelimbs together with the hindlimbs, rump and groin are Black.* The
neck, forelimbs and rump are flecked with Deep Olive Gray. The ground color
of the dorsal surface of the body is Deep Olive Gray finely spotted with
Black and Dragon’s Blood Red. The belly is Vinaceous-Rufous finely spotted
with Ivory Yellow laterally. There are four tail rings of Pale Olive Gray with
interspaces of Pale Olive Buff.

The coloration of the females is a brownish-gray with 1 to 4 well-defined
transverse body bands. The bands may be finely spotted with gray, but are
usually a unicolor black or brown with a lighter spotting on the ground color
between. There are usually four dark-brown rings on the tail with yellowish
or brownish interspaces.

In female tumidus, there is a pronounced tendency toward the reduction
of the number of transverse bands from the normal number of four to as few
as one. Chi.A.S. 10133-34 from 4 miles w. of Superior, Arizona, are
excellent illustrations of this trend, the former having only a single band
anteriorly just behind the shoulders, the remainder of the dorsal surface being
finely spotted with brown, while the latter has only two transverse bands. The
majority of the female tumidus also have the transverse bands considerably
widened middorsally and tapering sharply on the sides.

A tendency toward melanism in tumidus is shown by an intergrade from
near Apache Junction, Pinal County, Arizona. In life this specimen was
jet-black in color, except for the tail, which was straw-yellow. Upon preservation
in alcohol the presence of three dark-gray rings on the tail became apparent.
The same condition is also approached in Chi.A.S. 9481 from 4 mi. w. of
Superior, Pinal County, Arizona, except for a grayish spotting of the middorsal
region. Chi.A.S. 9482 is also nearly solid-black except for a lightening of the
middorsal region and a grayish spotting on the gular region and the chest.

In general, the majority of the adult male tumidus have a reddish dorsal
and ventral suffusion, with at least an indication of one or two transverse dorsal
bands, while the females are a drab grayish-brown in color and retain their
transverse bands throughout life.

In scalation the variation is also considerable. The following is a summary
of counts based on 21 specimens from several localities in Yuma County,

Arizona.
Ventrals Caudals Humerals

Range 132-185 29-36 39-49
Mean 156.6 B25 44.0
Standard deviation 13.67 ile7/ll Baz
Coefficient of variation, per cent 8.73 9.29 7.55

* Capitalized colors refer to Ridgway’s Color Standards and Nomenclature, 1912.

SHAW—THE CHUCKWALLAS 295

Tumidus also shows considerable variation in its generally spinose character.
Usually the nuchals and scales on the lateral neck fold are quite spinose as
well as the scales on the tail. Some individuals, however, lack the spinose
development achieved by many of the others and in this respect resemble the
majority of specimens of obesus. Some specimens carry the development of
spines to an extreme. SDSNH 16480, an intergrade between tumidus and
townsendi, from Las Chollas Point, Sonora, has nearly every scale on the body
and tail provided with a sharp spine and in this respect is just as spinose as
hispidus, although lacking the coarse scalation of that species.

Remarks.—S. 0. tumidus is apparently most closely related to S. o.
townsend: in characters of scalation, while it shows a tendency toward obesus
in pattern and coloration. Tumidus, like townsendi, has a relatively coarse
scalation, but its larger size and the distinctive coloration of the males readily
distinguish it from the form occurring further south in Sonora.

Because of the presence of intergrades between tumidus and obesus from
the region about Canyon Lake, Arizona, and from extreme southeastern
California along the western border of the Colorado River, tumidus has been
given subspecific status under Baird’s older name obesus. Although the type
locality of obesus was stated to be Fort Yuma, California, an area of inter-
gradation, the type specimen has the scale counts of a typical obesus. Whether
the type specimen was actually collected at Fort Yuma or in the mountains
to the west is a matter of question, because of the early practice of designating
the shipping point of specimens as the locality at which they were collected.

Localities of Collection—Arizona: Yuma County—Gila Mts. at
Telegraph Pass (type locality), Gila Mts. at Tinajas Altas, Mohawk Mts. at
U. S. 80; Pima County—Black Gap (15 mi. n. of Ajo), Alamo Canyon in
Ajo Mts.; Pinal County—West end of Picket Post Mt. (near Superior), 4

mi. w. of Superior.

Sauromalus obesus obesus (Baird)
GreAT Basin CHUCKWALLA

Euphryne obesus Baird, Proc. Acad. Nat. Sci. Phila., 1858, p. 253.
Euphryne obesa Baird, U. S. Mex. Boundary Survey, vol. 2, Rept., 1859, p. 6,

pl. 27.

Sauromalus ater Cope, Bull. U. S. Nat. Mus., no. 1, 1875, p. 47.
~ Sauromalus obesus Schmidt (part), Bull. Amer. Mus. Nat. Hist., vol. 46,

1922, p. 641.

Type Specimen.—No. 4172 in the collection of the U. S. National
Museum.

Type Locality —Fort Yuma, California.

Distribution.—Southeastern California, southern Nevada, southern Utah,
northern Baja California, and Arizona north of the line Yuma—Casa Grande—
Canyon Lake.

Diagnosis.—Similar in size, coloration and lepidosis to S. 0. tumidus, but
with finer and usually less spinose scalation throughout, especially on the limbs

296 SAN DtgeGo Society oF NaturaL History

and tail. The ventrals range in number from 156 to 220 and average 186.8.
The humerals vary from 46 to 68, averaging 54.7.

Description—Form stout; the head and body much depressed, the latter
with a strong lateral fold. The head is wider than long in adult males and
longer than wide in females and juveniles. The top of the head is covered with
small, smooth plates, largest on the frontal and parietal regions. The supra-
oculars and superciliaries are small and juxtaposed. The nostril is pierced in
a single rounded plate much nearer the tip of the snout than the orbit. The
rostral plate is usually divided vertically into two scales of equal size and shape
although these may occasionally be united to form a single hexagonal plate
much wider than high. There is a series of weakly carinate rectangular
suboculars, which, posterior to the eye, pass upward and backward to the ear
opening in the form of tubercles. The labial plates are small and subequal.
Below the infralabials there are several series of sublabials which merge gradually
into the granular gular scales. There is a strong gular fold. The ear opening
is vertical, or nearly so, with an anterior denticulation of from two to four
strongly spinose elongate scales. Behind the ear opening there is a prominent
neck fold bearing tubercles or subconical spines. The nuchal scales are
occasionally strongly spinose but more often may consist of a flattened scale
with one corner projected into a weak spine. A broad median band of enlarged
and weakly spinose scales extends from between the shoulders to the rump.
Laterally, the scalation is much finer, becoming eniarged and spinose on the
lateral fold. The ventral scales are smaller than the middorsal scales. Between
the gular fold and the anus there are from 156 to 220 rows of scales, averaging
186.8.

Dorsally, the scalation of the forelimb is relatively coarse, occasionally
carinate and weakly spinose. There are from 46 to 68 rows of humeral scales,
averaging 54.7. Below, the scalation is granular and much reduced in size.

The scalation of the dorsal surface of the femoral part of the hindlimbs
is coarser than that of the corresponding part of the forelimb, more or less
obtusely spinose and carinate; below, much reduced in size.

The tail ranges from 48 to 54 per cent of the total length. The scalation
consists of rather small, bluntly spinose, carinate scales, except dorsally and
ventrally at the base, where it is only weakly spinose and usually smooth. The
scalation becomes coarser and more spinose and carinate towards the tip of
the tail. There are from 30 to 42 scales in a whorl two head lengths behind
the vent, averaging 35.9.

Variation—In obesus coloration and scalation are quite variable, both
geographically and individually. There is also sexual variation. Generally
the coloration of the adult males consists of a black head, neck, shoulders,
chest and limbs. These may either be solid back or more or less profusely
spotted with gray or white. The remainder of the body may range in color
from a nearly unicolor black to a light-gray or red. In most adult males some
traces of the transverse bands are usually present, but there are frequent
exceptions to this. Females are usually grayish in color with darker head and
limbs, and usually retain the juvenile pattern of transverse body bands. There

SHAW—THE CHUCKWALLAS 297

may be small and infrequent spots or dashes of orange or red dorsally on the
body, but the females lack the general reddish suffusion found in most of the

males from western Arizona and eastern California.

Many adult males of obesus from Arizona and extreme eastern California
exhibit the same type of coloration as reported for tumidus, that is, with head,
chest, neck, and legs black and occasionally spotted with gray, the remaining
dorsal surface of the body having a yellowish or grayish ground color, well
suffused with red and spotted or reticulated with black. The belly is a uniform
red with more or less dense black spotting. The reddish suffusion seems to be
most prevalent among the large males from Arizona and southern Nevada, and
westward into eastern California a short distance, although in specimens from
the latter area the red is usually not so extensive nor as brilliant. In specimens
from western Imperial County and the eastern slope of the Coast Range in
San Diego and Riverside counties, the reddish coloration is lacking. Specimens
from the Palm Springs region of Riverside County are without the reddish
suffusion, while directly across the Coachella Valley in the Little San Bernardino
Mountains the reddish coloration suddenly appears. LMK 31900, a handsome
male from Fargo Canyon, Little San Bernardino Mts., has a straw-yellow
ground color, dorsally overlain with vivid red reticulations and a fine spotting
of black. The belly is also red, spotted with black. The top of the head,
gular region, chest and limbs are gray, spotted with black. The tail is straw
color without any traces of rings.

A series of specimens from the Mountain Springs region of Imperial
County and the Borego Desert area of San Diego County, California, shows
that the adult males are quite similar in coloration to the adult males of
tumidus from the Gila Mts., Yuma County, Arizona, except that the area

between the head and hindlimbs is a light-gray with or without traces of cross
bands.

Some specimens from Inyo and San Bernardino counties are quite dark
and show tendencies toward melanism like the obesus—tumidus intergrade from
Apache Junction, Arizona. LMK 34218, from 6 mi. nw. of Panamint Spring,
Inyo County, has a jet-black ventral surface, while a large part of the dorsal
surface of the head, body and limbs is also black, except for a grayish spotting.
The tail is yellowish-gray with three darker bands.

The juvenile coloration in obesus, as well as in tumidus, is quite different
from that of adults and the pattern is much more clearly defined. The
following notes were made from a specimen 115 millimeters in length from
Coyote Mountain, Imperial County: The top and sides of the head are Wood
Brown becoming Fuscous on the occipital region. There is a Black spot
immediately anterior to the ear opening. The nape is Blackish Mouse Gray
with a short median dash of Apricot Orange. The gular region is Deep
Mouse Gray. The belly is an immaculate Pale Olive Gray. Dorsally, the
ground color of the body is Prouts Brown. There are three transverse bands
of Chaetura Black between the shoulders and the rump. In the interspaces
between the bands are middorsal streaks of Pale Yellow-Orange. On either
side of these streaks there are several subcircular spots of the same color.

298 SAN DteGo Society OF NATURAL History

Laterally the ground color is Light Pinkish Cinnamon, with scattered spots of
Pale Smoke Gray. The dorsal surface of the fore limb is Mouse Gray
becoming Light Vinaceous Cinnamon on the toes. The dorsal surface of the
upper part of the hind limbs is Deep Mouse Gray irregularly marked with
Blackish Mouse Gray. On the tail the first ring is Saccardo’s Umber with
Fuscous-Black anterior and posterior edges. The remaining three tail rings are
Black. The interspace between the first and second bands is Pale Yellow-
Orange, the other two being White.

The most complete account of color changes as influenced by light and
temperature in this lizard is that of Dr. Sarah R. Atsatt (1939, pp. 249-50).
This author’s conclusions follow: “Experimentally, Sauromalus obesus parallels
Phrynosoma in its responses to temperature and light. At high temperatures
the light phase appears irrespective of illumination; at low temperatures the
dark phase appears irrespective of darkness. At moderate temperatures the
lizard is dark when subjected to bright illumination and in the light phase
when subjected to darkness.

“The speed of the reactions in Sauromalus is much like that of Phrynosoma.
At 28-33° C. in bright light, the process of darkening begins in from five to
fifteen minutes and in thirty minutes shows marked progress but often is not
complete. At higher temperature, however, the change is rapid—one individual
changed from medium dark to light in ten minutes in darkness at 41-38° C.
For lower temperatures, 20-25° C., the change is less for a given length of
time, and at these temperatures the paling process rarely reaches completion. If
the animals have been at a low temperature (below 20° C.) overnight, and
are put in a moderate temperature of 32° C., Sauromalus, because of its bulk,
takes a longer time than Phrynosoma to reach the temperature at which
illumination is the controlling factor. Sauromalus begins to respond to high
temperature around 35° C. and reaches the very light phase only at 40° C.
and higher.

“The change in the distinctness of the tail bands, recorded by Stejneger
(1893, p. 174) as due to intensity of light may also be due to change in
temperature. The change recorded by Van Denburgh (1922b, p. 89) is
definitely in response to excitement.”

The variation in scalation is also quite considerable in obesus. The
majority of specimens of obesus lack the degree of spinosity achieved by tumidus,
but occasional specimens of obesus have been found in which spines were quite
well developed on the nuchal region, the postauricular fold and on the caudal
scales. Obesus generally lacks the well-developed keels present on the scales of
the dorsal surfaces of the limbs of tumidus.

The variation in scale counts found in 40 specimens of obesus from the
Mountain Springs region of Imperial County and the Borego Desert area of
San Diego County, California, is shown below:

Ventrals Caudals Humerals

Range 165-215 30-41 50-61
Mean 189.3 36.9 54.5
Standard deviation 10.58 2.58 3.06

Coefficient of variation, per cent oh) 6.97 5.63

SHAW—THE CHUCKWALLAS 299

Habits —The following notes on the habits of obesus will undoubtedly
apply equally as well to the closely related tumidus.

Obesus, like the other species of the genus, is herbivorous. An examination
of the stomach contents of specimens has revealed the blossoms and leaves of
the following plants: Phacelia sp.; Franseria dumosa; Encelia farinosa; Eriogo-
num sp.; Ditaxis lanceolata; Larrea divaricata; Fouquieria splendens; Lotus
strigosus; Cryptantha sp.; Chaenactis sp.; Salvia columbariae; Tropidocarpum
gracile; Dalea fremontiu; Leptosyne bigelovii; Amsinckia tessellata; Sphaeralcea

munroana; Ephedra viridis; and Euphorbia polycarpa.

In the Borego area of San Diego County two specimens were found in a
crack in a large boulder, from the base of which led a well-traveled path to an
ocotillo (Fouquieria splendens) about fifty feet distant across the sandy bottom
of a dry wash. The stomachs of these two chuckwallas were filled largely with
the brilliant red blossoms of the ocotillo, showing that chuckwallas do not
hesitate to leave their rocky retreats in search of food.

Near Kingman, Mohave County, Arizona, a chuckwalla was observed
perched in the top of a bush about two feet high, eating with slow and deliberate
movements the yellow blossoms. Another specimen in the same attitude was
observed and collected near Quartzsite, Yuma County. Neither of these
specimens was bothered by my presence and I approached to within twenty
feet of them before shooting.

The best time for collecting chuckwallas is early in the morning just after
they have emerged from the rocks to sun themselves and to eat. Specimens
may be collected by shooting or by extracting them from the rock crevices in
which they seek shelter. The former is by far the easiest method but it is not
always a simple matter to get within shooting range. They may be shot while
in the rock crevices although the narrow opening of the crevice, together with
the nearness of the muzzle of the gun, concentrates the shot so that a badly
mutilated specimen is usually the result.

If a live specimen in good condition is sought, the chuckwalla may
usually be extricated from the crevice, in which it tightly wedges itself by
inflating the lungs, by tapping it gently on the nose. This annoyance usually
causes it to back out of the crevice far enough so that it may be seized by the
base of the tail or a leg. It is said that the Indians, who used these lizards for
food, secured them by puncturing them with sharpened sticks and then
withdrawing the lizard from the crevice. Using a heavy, sharpened wire, I tried
this technique on a specimen at Borego Mountain in San Diego County. The
lizard was punctured eight or nine times all along one side but there was no
appreciable deflation.

Practically nothing is known of the breeding habits of this or any other
species of Sauromalus. A specimen taken 7 mi. w. of Townes Pass, Inyo
County, during the middle of May, contained nine eggs which were apparently
about ready to be laid. Bogert (1930, pp. 6-7), who collected fourteen chuck-
wallas at Lovejoy Buttes, Los Angeles County, on May 11 and 12, 1929, found
six eggs in an average sized female. This author also states: “In three cases
male and female found side by side in cracks were apparently mating.”

300 SAN DreGo Society oF NATURAL HIstTory

Occasionally specimens are found DOR in such places as at Sentenac
Canyon, San Diego County, and at Telegraph Pass, Yuma County, Arizona.
This would indicate that chuckwallas do not confine their activities to rather
limited areas, such as a particular pile of rocks, but at times probably travel
considerable distances in search of food or mates. Neither do they confine
themselves to the boulder strewn slopes of desert hills and ranges proper, for
if sufficient rocky debris is available they are occasionally found nearly a mile
from the nearest mountains, as in the vicinity west of the Black Mountains in
Mohave County, Arizona, where a large amount of rocky debris is found on the
plain which slopes away from the base of this range. In this locality chuck-
wallas were found in scattered piles of rocks together with Callisaurus v. ventralis
and Dipsosaurus d. dorsalis which inhabit the open, sandy areas of the desert.

The following remarks by Bogert (/oc. cit.) are of interest: “In the early
part of April, 1928, a large adult male chuckwalla was collected at Lovejoy
Buttes. It had just come out of hibernation and was very thin. After about
a month and a half of captivity it was liberated May 26, 1928, at a spot twenty
feet from the crack in which it had originally been discovered. Then on May
11, 1929, nearly a year later, the identical specimen was re-collected when found
between two rocks possibly twelve feet from the crack where it had been
originally found. It was easily identified by means of a peculiar scar on its
back. Also it was much thinner than any other of the fourteen chuckwallas
collected on that week-end.”

Localities of Collection—CattForNia: Inyo County—7 mi. s. of Saline
Valley, 7 mi. w. of Townes Pass, 5 mi. sw. of Townes Pass, Emigrant Spring,
Wildrose, Pleasant Canyon in Panamint Mts., Darwin (also 7 and 9 mi.
east), 1 mi. n. of Little Lake, Little Lake, 17 mi. n. of Trona, Slate Mts.,
Shoshone; San Bernardino County—Saltwells Valley, 6 mi. nw. of Spangler,
Deadman’s Point, Cave Springs, Victorville, Oro Grande, Barstow, 14.6 mi. ne.
of Barstow, Odessa Canyon in Calico Mts., Minneola, Cat Mountains, Baxter,
s. end Soda Lake Mts., 7 mi. sw. of Baker, Silver Lake, 5 mi. sw. of Ivanpah,
Sacramento Mts. (28 mi. w. of Needles), Piute Mts. (16 mi. w. of Needles),
Essex, 8/2 mi. nw. of Essex, 5 mi. s. of Lavic, 4 mi. s. of Lavic, Hector, Lucerne,
Twenty-nine Palms, 10 mi. ne. of Blythe Junction, Rabbit Dry Lake; Kern
County—17 mi. e. of Inyokern, Red Rock Canyon (20 mi. n. of Mohave),
Red Hill; Los Angeles County—Lovejoy Buttes, Peck’s Butte; Riverside
County—Snow Creek, 4 mi. w. of Garnet, Palm Springs, 5 mi. e. of Palm
Springs, 1 mi. sw. of Palm Springs, 5 mi. se. of Palm Springs, 4 mi. nw. of
Palm Springs, Tahquitz Canyon, Palms to Pines Highway at 3100 feet, Dos
Palmas Springs in Santa Rosa Mts., 5 mi. ne. of Edom, Cottonwood Springs,
near jct. Dead Indian and Grapevine Creeks, Coral Reef Canyon, Murray
Canyon, Fargo Canyon in Little San Bernardino Mts., Berdoo Canyon in Little
San Bernardino Mts., Pushawalla Canyon in Little San Bernardino Mts., Shaver
Well, Hidden Spring Canyon, Rice, Riverside Mt., Desert Center, 16 and 8
mi. e. of Desert Center, Chuckwalla Mountains; Imperial County—Pilot Knob,
Ogilby, American Girl Mine, Chocolate Mts., Fort Yuma (type locality),
near Imperial Dam, Picacho Landing, 12 mi. sw. of Palo Verde, Beal Well,

SHAW—THE CHUCKWALLAS 301

Hanlon Ranch, Coyote Mts., foot of Mountain Spring Grade, Myer’s Creek
Bridge, Mountain Springs; San Diego County—Agua Caliente Hot Springs,
Mason Valley, Box Canyon, Sentenac Canyon, Yaqui Well, San Felipe Wash,
The Narrows, Tubbs Canyon, Coyote Canyon, Borego Mountain, 4 mi. s. of
Benson’s Dry Lake. Arizona: Mohave County—Hackberry, Kingman, Gold-
road, between Oatman and Goldroad, Oatman, 1, 3 and 4 mi. sw. of Oatman,
16 mi. ne. of Topock, Chemehuevis Mountains; Yuma County—8 mi. w. of
Brenda, 5 mi. w. of Brenda, 2 mi. e. of Hope, Dome Rock Mts. (8 mi. w. of
Quartzsite), Quartzsite, Castle Dome, Kofa Mts.; Coconino County—Below
Indian Gardens in Grand Canyon; Maricopa County—Wickenburg, 7 mi. sw.
of Wickenburg, Tempe, Stewart Mt. (near Mesa), Maricopa Mts. at U.S. 80;
Yavapai County—5 mi. n. of Wickenburg, near Congress Junction. Nevapa:
Clark County—W. bank of Colorado River (1/6 mi. north of California line),
Searchlight, Boulder City, island in Lake Mead, Boulder Wash, Las Vegas
Wash, 12 mi. w. Las Vegas, 68 mi. nw. Las Vegas between Indian Springs
and Amargosa, 9/2 mi. s. of Dead Mts., 2/2 mi. e. of St. Thomas, Atlatl
Rock in Valley of Fire, 2/2 mi. e. of Hiko Spring, Hiko Spring; Nye County—
Rhyolite, southeast end of Spectre Range. UtaH: Washington County—St.
George, Black Ridge, Silver Reef (21 mi. ne. of St. George); San Juan
County—Aztec Creek; Kane County—Rock Creek, Warm Creek.

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KEY TO THE SPECIES AND SUBSPECIES OF Sauromalu«

. One or more transverse bands dorsally, across body or rump................-..... 4
INowtransyerse body batids presenti: sect. ne ert ce te a tee sees cece 2

Largest nuchal scales equal to or larger than the frontal plates

SO ee a sane HEE RY OF a3 J a tec Ate scot S. hispidus (adult)
(Angel de la Guarda, Smith, Pond, Granite, and Mejia Islands, Gulf of California,
Mexico. )

Largest nuchal scales smaller than the frontal plates.............-.-.-.-0.--.-.-----.-- 3

Dorsal pattern of large, irregular, dark-brown or black blotches on a yellowish
EOUMCNCOL Ofer sie Pens ee eee rate th, Mews Le hr Ae vate St S. varius
(San Estéban Island, Gulf of California, Mexico. )

Dorsal pattern of small, dark-brown or black spots on a gray ground color

I NN wee CEs AE Ne SF eng PEN hee 4 cat al ae Sd a S. klauberi
(Santa Catalina Island, Gulf of California, Mexico. )

= Venttalvscalesrowssusually less\ than 130%... f ens ee ee et 5
Ventral scale rows usually more than 13022. 2. tere 6

Dorsal scales in a head length usually less than 20......S. hispidus (juvenile)
(Angel de la Guarda, Smith, Pond, Granite, and Mejia Islands, Gulf of California,
Mexico. )

Dorsal scales in a head length usually more than 20................-.--.--.---- S. slevini

(Carmen, Coronado, and Monserrate Islands, Gulf of California, Mexico.)

. Transverse body bands with light centers and dark-brown or black borders

Sivingra wacoublerabanded) ef tect src rests 2, cet eee pete ce eect nate 9
Meransverse * Panis sultal COLOt 2 cs oP chee See es ee ees 7
el-lumeralyscalestusually, less thati 50) cetacean pecs eee eames 8
Flumieralsscalesustially. amore: that) 0 eces te cee ee eee S. 0. obesus

(Deserts of southeastern California, southern Nevada, southern Utah, northern Baja
California, and Arizona north of the line Canyon Lake—Casa Grande-Yuma. )

. No reddish suffusion on the dorsal and ventral areas in the adult males;
maximum adult length averaging somewhat shorter than in S. 0. tumidus
atid: loweraverage scale COUntS. <2. .- at! escyeeecee cee ees S. 0. townsendi
(Tiburén Island, Gulf of California, and adjacent Sonoran mainland at least as far
south of Guaymas. )

More or less brilliant reddish suffusion on the dorsal and ventral areas of
adult males; maximum adult length averaging longer than in S. 0. townsendi
andinigher average scaleicoutlts <5. 2ici t,o eee ns S. 0. tumidus
(Arizona south of the line Canyon Lake-Casa Grande-Yuma. )

Maventraliscale towsilo lor motes. ee ee, S. australis
(Central and southern Baja California from San Francisquito Bay south to La Paz.)
Wentraliscale rows usually less thamj45 le 2 ete. 2 ee terete ene S. ater

(Espiritu Santo, Isla Partida, San Marcos, San Diego, San Francisco, and Santa Cruz
Islands, Gulf of California, Mexico. )

304 SAN Deco SocretTy oF NATURAL History

MusEUM ABBREVIATIONS

Where specimens have been referred to by number the following abbrevi-
ations have been used:

CAS California Academy of Sciences
Chi.A.S. Chicago Academy of Sciences

LMK Private collection of L. M. Klauber
SDSNH_ San Diego Society of Natural History

ACKNOWLEDGMENTS

I am indebted to the following individuals and institutions for the loan of
specimens or information relating to particular specimens: Dr. Doris M.
Cochran, United States National Museum; Mr. Joseph R. Slevin, California
Academy of Sciences; Mr. Thomas L. Rodgers, Museum of Vertebrate Zoology,
University of California; Mr. Karl P. Schmidt, Chicago Natural History
Museum; Dr. H. K. Gloyd, Chicago Academy of Sciences; Mr. Charles M.
Bogert, American Museum of Natural History; Dr. Ross Hardy, Dixie Junior
College; Dr. L. M. Klauber, San Diego Society of Natural History; Dr. E.
H. Taylor, University of Kansas; Mr. Robert S. Hoard, American Vice
Consul at Acapulco, Guerrero, Mexico.

Mr. C. B. Perkins has made many valuable criticisms of this paper in
manuscript.

I am especially grateful to Dr. L. M. Klauber for his advice and criticism
concerning this paper and for the many courtesies extended to me during this
study.

BIBLIOGRAPHY

ATSATT, S. R.
1939. Color Changes As Controlled by Temperature and Light in the
Lizards of the Desert Regions of Southern California. Publ. Univ.
Calif. at Los Angeles in Biological Sci., vol. 1, no. 11, pp. 237-276,
plates 8-12, 9 figures in text.

BairD, S. F.
1858. Descriptions of New Genera and Species of North American
Lizards in the Museum of the Smithsonian Institution. Proc. Acad.

Nat. Sci. Phila., vol. 10, pp. 253-256.

BELDING, L.
1887. Collecting in the Cape Region of Lower California. The West
American Scientist, vol. 3, no. 24, pp. 93-99.

Bocourt, F. (with A. Duméril & F. Mocquard)
1870-1909. Mission Scientifique au Mexique. Les Reptiles, pp. 1-1012.

SHAW—THE CHUCKWALLAS 305

BocGert, C. M.
1930. An Annotated List of the Amphibians and Reptiles of Los Angeles
County, California. Bull. So. Calif. Acad. Sciences, vol. 29, part 1,
pp. 3-14.
Cope, E. D.
1875. Check List of North American Batrachia and Reptilia. Bull. U. S.
National Museum, no. 1, pp. 1-104.

Cow es, R. B., and Bocert, C. M.
1936. The Herpetology of the Boulder Dam Region (Nev., Ariz., Utah).
Herpetologica, vol. 1, no. 2, pp. 33-42.
Dickerson, M. C.
1919. Diagnoses of Twenty-three New Species and a New Genus of
Lizards from Lower California. Bull. Amer. Mus. Nat. Hist.,
vol. 41, art. 10, pp. 461-477.

Dumenrit, A.

1856. Descriptions des Reptiles Nouveaux ou Imparfaitement Connus de
la Collection du Muséum d’Histoire Naturelle et Remarques sur
la Classification et les Caractéres des Reptiles. Arch. Mus. Nat.
Hist. Paris, vol. 8, pp. 437-588, pls. XvII-xxIv.

Groyp, H. K.
1937. A Herpetological Consideration of Faunal Areas in Southern
Arizona. Bull. Chicago Acad. Sciences, vol. 5, no. 5, pp. 79-136,
figs. 1-22.
MocQuarp, F.
1899. Contribution a la Faune Herpétologique de la Basse-Californie.
Nouv. Arch. Mus. Hist. Nat. Paris, ser. 4, vol. 1, pp. 297-344.

ScHmIvT, K. P.
1922. The Amphibians and Reptiles of Lower California and the
Neighboring Islands. Bull. Amer. Mus. Nat. Hist., vol. 46, art.
11, pp. 607-707.
SHAW, C. E.
1941. A New Chuckwalla from Santa Catalina Island, Gulf of California,
Mexico. Trans. San Diego Soc. Nat. Hist., vol. 9, no. 28, pp.
285-288.
STEJNEGER, L.
1891. Description of a New North American Lizard of the Genus
Sauromalus. Proc. U. S. Nat. Mus., vol. 14, no. 864, pp. 409-411.

TAYLor, E. H.
1936. Notes on the Herpetological Fauna of the Mexican State of
Sonora. Univ. of Kansas Science Bull., vol. 24, no. 19, pp. 475-503,
pl. 43.
VAN DENBURGH, J.
1922. The Reptiles of Western North America. Occas. Papers Calif.
Acad. Sciences, no. 10, vol. 1, Lizards; vol. 2, Snakes and Turtles,
pp. 1-1028.

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TRANSACTIONS
OF THE

SAN DIEGO SOCIETY OF NATURAL HISTORY

VOLUME X, No. 16, pp. 307-310 Aucust 31, 1945

L3887%

A NEW WOOD RAT, GENUS NEOTOMA,
FROM THE VISCAINO DESERT REGION
OF BAJA CALIFORNIA, MEXICO

LaureNcE M. Huey
Curator of Birds and Mammals, San Diego Society of Natural History

The gradual accumulaticn, in the collection of the San Diego
Society of Natural History, of specimens of Neotoma lepida from the
central section of Baja California, Mexico, has now reached a point where
they can be studied with some understanding of racial development. As
a result, the presence of an unnamed race from the northwestern coast-
al section of the Viscaino Desert region has been revealed. It may be
known as

Neotoma lepida molagrandis subsp. nov.

VISCAINO DESERT Woop Rat

Type——From Santo Domingo Landing (lat. 28° 15’ N.), Baja California,
Mexico (more precisely, at the site of the old well near the edge of a mesa-like
shelf, some 3 miles inland from the landing beach, elevation about 50’) ;*
no. 14065, collection of the San Diego Society of Natural History; adult male;
collected by Laurence M. Huey, April 28, 1940.

Characters.—In size and color this race from the coastal-llano areas of the
northwestern Viscaino Desert resembles Neotoma lepida egressa from the San
Quintin area, but Neotoma lepida molagrandis differs in slightly more pallid

*This is also the type locality of Dipodomys agilis peninsularis (Merriam), since
Nelson records (Mem. Nat. Acad. Sci., vol. 16, p. 30, 1921) that he and Goldman were
camping there on the date the type specimen was collected.

308 SAN Disco Society Or Natura History

pelage. Cranially the differences are well pronounced. N. |. molagrandis has
heavier molars, more inflated and larger bullae, and slightly broader nasals.
There is a sharper, more angular, bend in the posterior ends of the zygomatic
arches. A most conspicuous character is the shape of the infraorbital foramen.
It is as long as that of egressa, but compressed at the upper end, with the walls
more nearly parallel—similar to those of the smaller Neotoma lepida ravida.
This character is not approached in either of the two comparable races, egressa
and ravida.

Compared with N. l. ravida from the mountainous area southeast and
south of the Viscaino Desert, molagrandis is larger in size and has lighter col-
ored pelage. It lacks the black tendencies found in the pelage of ravida from
the type locality, Comondu, which is located in the lava range, Sierra de La
Giganta. Cranially the differences are also well marked. The molars of mola-
grandis are larger, the zygomatic arches are heavier and more wide-spreading,
posteriorly, and the bullae are more elongated and more inflated. In fact, the
whole cranium is heavier, more robust, than is that of ravida.

There seem to be two family chains of Neotoma lepida down the peninsula.
The smaller, more fragile-appearing, desert type, Neotoma lepida felipensis,
ranges southward over the barren deserts along the eastern side of the peninsula,
transferring these characters into the race ravida. The heavier, more robust type
is represented in Neotoma lepida intermedia, found along the brushy areas of
the Pacific slopes of California and northwestern Baja California, and descends
along the Pacific coast of the peninsula through egressa, molagrandis, and
pretiosa (of the Magdalena Plain area), culminating on the coastal regions.
of the Cape in the robust Neotoma lepida arenacea.

Measurements—Type: Total length, 342; tail, 150; hind foot, 35; ear,
30. Skull (type): Greatest length, 44.4; zygomatic breadth, 23.7; interorbital
breadth, 5.5; greatest length of nasals, 17.1; length of palatal bridge, 8.0; alveo-

lar length of upper molar series, 8.4.

Range.—So far as known, the northern and western coastal section of the
Viscaino Desert region of Baja California, Mexico.

Remarks.—Goldman in his “‘Revision of the Wood Rats, Genus Neotoma”
(North American Fauna, no. 31, 1910) assigned extensive ranges to two races
of Neotoma intermedia (now known as N. lepida; see Jour. Mammal., vol.
13, pp. 59-67, 1932) on the mainland of Baja California. One of these was
Neotoma intermedia intermedia. Its range, as given by Goldman, extended
along the Pacific slope of the peninsula from the International Boundary south
to about latitude 28°. Here, leaving the Pacific side to be occupied by other
races, the range of N. i. intermedia was stated to continue southward along the
Gulf slope to the vicinity of La Paz, including the mountainous backbone of
the region. At La Paz an interruption occurred and N. i. intermedia was not
found again until the upper slopes of Sierra de La Victoria were reached. Men-
tion was made by Goldman in his revision that certain divergent characters
occurred in specimens of this southernmost mountain population.

Hurv—New Woop Rar 309

The other race was Neotoma intermedia gilva. The range ascribed by
Goldman to this form in Baja California was on the desert slope from the In-
ternational Boundary, southward along the Gulf slope to the vicinity of lati-
tude 29°, thence “leap-frogging” westward to occupy the Viscaino Desert
region, including the Santa Clara Mountains bordering the Pacific Ocean.

Subsequently, these allocations were altered. Upon re-examination, speci-
mens ascribed to N. 1. intermedia along the mountainous-gulf area from lati-
tude 28° south to La Paz, and those from the Sierra de La Victoria, were de-
termined to belong to two different races and were named respectively N. i. ra-
vida and N. 1. notia by Nelson and Goldman (Proc. Biol. Soc. Washington,
vol. 44, pp. 107-110, 1931).

Following the publication of these new races, Goldman, in the Journal of
Mammalogy (loc. cit.), revised the Neotoma lepida group. In this account he
revived Elliot’s Neotoma bella felipensis (Publ. Field Columb. Mus. Zool. Ser.,
vol. 3, pp. 217-218, 1903), defining the “desert region, east of the San Pedro
Martir Mountains in northeastern Lower California” as its range. Neotoma
lepida gilva was thus restricted, from its previously assigned gulf-desert strip of
Baja California, to a small range near the International Boundary. No mention
was made by Goldman, in this later revision, of the Neotoma occupying the
Viscaino Desert, nor were the ranges of either intermedia or ravida defined as
covering that area.

In 1934, Orr named a race, Neotoma lepida egressa (Proc. Biol. Soc.
Washington, vol. 47, pp. 109-112, 1934) from one mile east of El Rosario,
Baja California, Mexico. This race was found to occupy the “coastal region
of northwestern Lower California from latitude 31° N. south at least to El
Rosario, latitude 30° 03’ N.” In determining the characters of this race, Orr
compared his specimens with specimens of N. [. felipensis from the opposite
side of the peninsula and with those of N. |. intermedia from California to the
northward. He found that the main characters of his newly defined race were
most directly related to those of N. |. intermedia. This is also well demonstrated
by specimens at the present writer’s command.

No mention was made by Orr of his having seen or compared egressa with
ravida, the nearest geographically named race to the southward at the date he
made his study. There is a wide gap shown in a number of the characters of
these two races, but of outstanding interest is the fact that both are dark col-
ored. Specimens of egressa show a grizzled brown-gray cast or sheen in the pel-
age, while ravida lacks this tone and tends more towards black-gray. These dif-
ferences are no doubt the result of environment, since egressa lives in a humid
coastal region where, in places, heavy chaparral abounds, while ravida lives in
volcanic, black lava country, where vegetation is usually sparse.

The race N. 1. molagrandis, named herewith, has some characters that
are intermediate between those of egressa and ravida, but the large molariform
teeth and flaring zygomatic arches are trenchant, placing the relationship of
molagrandis nearer that of the intermedia-egressa, rather than the felipensis-
ravida, line. Geographically this would be expected. However, there is still much

310 SAN Dreco Society Or NAtTuRAL History

to be learned about the Neotoma populations living in the region between the
southern base of the Sierra San Pedro Martir and the northern edge of the
Viscaino Desert.

Specimens examined.—Neotoma lepida intermedia. California, San Di-
ego County: Murray Dam near La Mesa, 3; Mission Gorge, 3; Alvarado Can-
yon near La Mesa, 1; San Diego, 1; Bonita, 3; Witch Creek, 1; mouth of Tia
Juana River, 1. Baja California, Mexico: Valle de Las Palmas (below Tijuana),
1; Valle de La Trinidad, 14; Summit San Matias Pass, 3.

Neotoma lepida egressa. Baja California, Mexico: San Quintin, 4;
Aguaita, 1.

Neotoma lepida molagrandis. Baja California, Mexico: Punta Prieta, 2;
Mesquital, 2; Santo Domingo Landing, 2 (type locality); Calmalli, 1; 12 mi.
e. El Arco, Rancho Miraflores, 1; Santa Gertrudis Mission, 1; San Ignacio, 3
(not typical).

Neotoma lepida gilva. California, San Diego County: La Puerta Valley,
20. Baja California, Mexico: Gaskill’s Tank, near Laguna Salada, 4.

Neotoma lepida felipensis. Baja California, Mexico: San Felipe, 10 (type
locality).

Neotoma lepida ravida. Baja California, Mexico: south end Concepcion
Bay, 7; Comondu, 5 (type locality).

r
P92 ws

TRANSACTIONS
OF THE

SAN) DIEGO, SOGIETY OF NATURAL HISTORY

Volume X, No. 17, pp. 311-398, plates 7-8, fig. 1, map

THE GLOSSY SNAKE, ARIZONA,
WITH DESCRIPTIONS OF NEW SUBSPECIES

LAURENCE M. KLAUBER

Curator of Reptiles and Amphibians, San Diego Society of Natural History

SAN DIEGO, CALIFORNIA

PRINTED FOR THE SOCIETY

Marcu 29, 1946

COMMITTEE ON PUBLICATION
U. S. Grant, IV, Chairman

L. M. KLauBer CLINTON G. AspotTrt, Editor

TABLE OF CONTENTS

Ma beacl ection ee ae sre See ea et ns epee cas ee Maeda: as 2s oa)
Ii bieeSerteall <QUU DORE eter oR cece eee eee ee ecm er 315
Character Variation and Sexual Dimorphism..................-.--.2----------ese0e- eee 316
Wescrimtionmote SUDspeciess tes: es eeseees eee oo ere acter et 319
(Gretn iS taeAlys CO Niches es ere ted eee cee nc ee Se oe eect rs es 320
Aliconanelenams, ele cdi I eninicOtG see ere ee age ee 320
Arizona elecansiblanchards stilasps tOV 2-2 ee = -ee secon ec eee 328
Arizona elegans» prilaps SUDSP. MOVs -o-a---eetece cctee sce eeepeee nana 333
Arrizomndiel eetin sex POU IEA SUIDS pas OV st er nea teeees eo oer eee eee acne 340
Aniconavelegan’s moctivagd subsp: MOVs act cc eee eee 343
Anizona elegans eburmata sissps MOVs c 2... eee toeec een cee ce eee reece 350
AicOnancle gn séCaaOid a), SUDS ps, MOVs este eee 364
Amzonaselesans occiaentdlas Blamenatd eee cee es tee B72,
Arizonaseleganss pacal disubsps MOV. set. en eae ere 379
Phylogeny: 2:2 3 Se ie eae Eas NO aA 381
Palevatonthe oubspecies,olAgszond elegans a ee asthe eee einer 387
PAGAIO WIECH OICL ES ee reel Wn Oe Pe oh eS ee et eS nae: 388
Abbteviations 2.255.265 = Cp ES ee te rk sR IN eS 388
SO mitnatyaee mee oes ot A ere) ee een ol we te Pee oS ee ON 389

Bibliography

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5 (ca)
Oo
Yn

THE GLOSSY SNAKE, ARIZONA,
WITH DESCRIPTIONS OF NEW SUBSPECIES

LauRENCE M. KLAuBER
Curator of Reptiles and Amphibians, San Diego Society of Natural History

INTRODUCTION

The glossy snakes, or, as they are sometimes less suitably called,
the faded snakes, genus Arizona, comprise a monotypic genus inhabiting
the southwestern United States and northern Mexico. Of late years
new methods of collecting, particularly by driving on paved desert
roads at night, have considerably increased the numbers of specimens
in study collections, so that good series are now available from several
localities, and territorial differences may be determined with a greater
assurance of validity. This genus was once thought to be rare, but is
now known to be among the commonest of snakes in some sections of
the southwest, particularly in parts of the Mojave and Colorado deserts.
In Texas the glossy snakes are diurnal or crepuscular; in the western
deserts they are almost entirely nocturnal.

HIsTorIcAL SUMMARY

The genus Arizona, type species elegans, was set up by Kennicott in 1859,
being differentiated from Pituophis by its paired prefrontal scales and smooth
dorsals, whereas in Pituophis the prefrontals usually number 4, and the central
dorsals are always keeled. Both Arizona and Pituophis are characterized by
undivided anal plates and a relatively high number of scale rows.

In 1860 and 1861 Cope described two new subspecies, Arizona jani and A.
lineaticollis. These have paired prefrontals but keeled dorsals, thus breaking
down one of the differences first presumed to segregate Arizona from Pituophis.
This confusion led Cope to assign elegans to the genus Pituophis in 1875 (or
Pityophis as it was then called), and to Rhinechis (an Old-world genus) in
1886. This latter arrangement was continued by some authors until as late
as 1907. Eventually (1894) lineaticollis and jani were properly assigned to
Pituophis by Gunther (jani as a synonym of P. deppei), and Arizona again
became monotypic, although not so recognized by Gunther, who continued
elegans in Pituophis. However, Arizona had been revived for elegans by Bocourt
in 1888 and has been recognized by most herpetologists since, although Boulen-
ger (1894) placed both Arizona and Pituophis in the genus Coluber, a genus

316 San Dreco Soctety oF NATURAL HIstory

notably composite as set up by him, but comprising primarily snakes which today
would be referred to the genus Elaphe.

The genus Rhinechis is now usually considered a synonym of Elaphe by
European herpetologists, e.g., Mertens and Miller, 1928; but in any case
Arizona differs from Rhinechis in having a slightly elliptical pupil (as pointed
out by Van Denburgh, 1906, p. 66), in having an undivided anal plate, and
single rather than paired apical scale pits. Thus present-day herpetologists are
agreed on the recognition of Arizona as a monotypic genus.

Subsequent to the original description of Arizona elegans, only one ad-
ditional subspecies has been recorded. In 1924 Blanchard (p. 1) described
Arizona elegans occidentalis. type locality La Jolla, California, differentiating it
from the eastern form, A. e. elegans, by its reduced scale rows, more numerous
and narrower blotches, less prominent lateral spots, and shorter tail. He con-
sidered southeastern Arizona to be the area of intergradation between the
two subspecies.

Although Azizona is now generically separated from Pituophis, it is recog-
nized that the relationship is close. The hemipenial differences are slight (cf.
p. 355). Walls (1934, p. 899), in his investigations of eye structure, found
Arizona intermediate between such nocturnal snakes as Phyllorhynchus and
Hypsiglena, and certain diurnal genera, particularly Pituophis. He states: “The
diurnal genus Pituophis stands very close to Arizona and is probably the genus
from which the latter was derived. Arizona can thus be thought of as a Pituophis
which has become generically distinct partly as a result of changes accompanying
its tendency toward nocturnality.” It may be noted that Arizona, in scalation,
most nearly resembles the least specialized (and probably most primitive) forms
of Pituophis, P. deppei and P. lineaticollis, which are like Arizona in usually
having only 2 prefrontals (instead of 4 as in most Pituophis), and in having
2 supralabials in contact with the eye.

With much new material available, it now becomes possible to describe
several additional subspecies. Altogether I have had access to some 540 speci-
mens, compared to the 33 available to Blanchard when his study was made.

CHARACTER VARIATION AND SEXUAL DIMORPHISM

As is my usual practice in surveying a genus, I shall first investigate degrees
of dispersion and sexual dimorphism in the largest territorially homogeneous
series available to me. It has been my experience that coefficients of variation
and sexual divergence tend to be rather constant for each character within a
genus; hence if one is able to determine their values in one area, he will be
better able to evaluate any differences found between subspecies, thus to some
extent compensating for inadequate material. At the same time I shall discuss
some of the character peculiarities of Arizona.

The two largest series at hand are from the Narrows-Dry Lake area of
eastern San Diego County, and the Adelanto-—Kramer area of western San
Bernardino County, California. I have found it necessary not to include, as
a part of the first-named series, a considerable number of specimens from be-

KLAUBER—IHE GLossy SNAKE Ail\7.

tween San Felipe Valley and the Narrows. for in this desert foothill region,
separated by only a few miles from the desert beyond, some differences from
the specimens of the desert flats to the eastward are already manifest.

On the characters having sufficient dispersion to be treated statistically, the
data are as follows:

Narrows— Adelanto-
Dry Lake Kramer
Number of specimens, males 25 40
females 16 13
Coefficients of variation, per cent
Ventrals, males 1.41 1.24
females 1.50 1.40
Subcaudals, males 2.83 3.56
females 3.01 3.09
Body blotches 8.66 6.23
Tail spots 13.41 12.80
Coefficients of sexual divergence, per cent
Ventrals —3.98* ~4.99*
Subcaudals 9.62 9.38

We note here a considerable consistence in the coefficients of variation, and
relatively narrow dispersions, particularly in ventrals and subcaudals. In
Arizona the ventral dispersions are 1/2 per cent or below; while the subcaudals
run about 34 per cent or below. In other genera the coefficients of variation in
the ventrals generally average about 2 per cent, and in the subcaudals 5 to 6
per cent. With such local consistency, relatively small differences may warrant
subspecific distinction. For example, homogeneous populations whose mean
ventral counts are separated by 6 per cent (about 12 scutes in Arizona) would
have about 2.3 per cent of specimens overlapping, while even differences of but 9
scutes would involve only 6.7 per cent of overlapping, if we assume normality
of distribution (Klauber, 1941a, p. 5).

The sexual divergence in ventrals and subcaudals is somewhat less in
Arizona than is general among colubrids}, and there is some overlapping between
the sexes. As is usual, the females have more ventrals and fewer subcaudals than
the males.

The variability in blotch counts, always higher than that of ventrals and
subcaudals, is not greatly different from that found in other blotched snakes.

Some qualifications of the characters used in differentiation, as they are
met in Arizona, are as follows:

There is more variation in scale rows than is evident in many genera.
Variations of at least two rows are present in nearly every subspecies, and
ranges of four rows are occasional. While most subspecies lean heavily toward
one count (27, 29, or 31), some are fairly balanced between two. In such

** Minus signs indicate the females are higher than the males.

+ For examples see Bull. San Diego Zool. Soc., no. 18, p. 14, 1943.

318 SAN Dreco Society oF NATURAL HISTORY

cases some sexual dimorphism is evident. For example, in 29 specimens from
the vicinity of San Antonio, Texas, we have the following distribution:

ScaLtE Rows

29 Sil
Males 12 6
Females 3 8

There are 12 chances in 100 of such a disproportionate number of females
with 31 rows having occurred fortuitously, if the scale rows were, in fact, not
sexually dimorphic.

The scale-row criterion divides the subspecies of Arizona elegans into two
groups: 1) e. elegans and e. blanchardi in the east, with 31 rows often present,
although not necessarily in the majority, which may have 29; and 2) the
other subspecies to the west and south in which 27 rows are always in the
majority, although 29 rows may be present in up to 20 per cent or more of
the population.

With respect to the ventral scutes, I have already mentioned the variability
and sexual dimorphism in homogeneous samples. As is so often the case,
this character is of the highest importance in Arizona, for considerable sub-
specific differences are found.

The subcaudals are less important—they seldom serve as useful key
characters, since differences are less and overlaps greater. It is to be noted
that comparatively few specimens have incomplete tails; also, it is less difficult
to tell whether a tail is incomplete than in some genera—Pituophis and Masti-
cophis, for example.

The supralabials do not constitute a useful character for they are ‘too
constant; there is no subspecies yet known in which 8 are not greatly in the
majority. While there is more variability in the infralabials, it is more intra-
than inter-subspecific. Furthermore, the count is often difficult to ascertain with
accuracy, because of the overhang of the last supralabial. The serial number
of the largest infralabial (counting back from the mental), which is easily
ascertained with accuracy, may eventually prove of value in segregating some
subspecies.

The loreals, postoculars, and temporals, although showing an occasional
intrasubspecific variability, are too constant in number to be of importance in
taxonomic work. On the other hand, in two areas there are trends, one quite
strongly accentuated, toward the division of the normally single preocular.

The blotches, both in number and size, are found to be of importance in
classification, for there are easily recognized territorial differences. It is true that
observers may not always agree as to the number of dorsal body blotches in
a specimen because of a lack of bilateral symmetry. Thus some blotches are
diagonal; some are Y-shaped (paired on one side of the body and single on
the other); while still others are confined to a single side of the middorsal line.
Probably the best way to determine the number is to count each side separately
and use the mean. The tail spots are still less accurately countable. for they are

KLAUBER—TIHE GLossy SNAKE 319

often quite ill-defined, especially toward the tip. Frequently in some subspecies
there is a middorsal light streak which splits these spots, and the last few
may be represented only by irregular dark scale-edges.

Equally important with the number, are the relative sizes of the dorsal
blotches and the interspaces which separate them. I have made it a practice
to evaluate these at mid-body, selecting for study the most regular blotches to
be found there. -The transverse extent of a blotch is determined by counting
the scale rows included, taking the dark edge of a blotch as the lateral limit.
If this edge cuts across the middle of a scale, one half is counted on each
side, thus resulting in an even total. The longitudinal extent of each blotch
is measured in scales, end to end, and the interspaces are similarly determined.
By averaging several blotches and interspaces, considerable accuracy can be
achieved in these measurements. Using scales as units of measurement is
preferable to attempting to gauge the relative extents of blotches and inter-
spaces with the eye. These blotch dimensions are found, in a number of in-
stances, to comprise quite adequate key characters wherewith to distinguish
subspecies.

The lateral auxiliary series of blotches are somewhat less useful, although
it is sometimes of interest to determine the lowest lateral rows darkened by the
interblotch ground-color suffusions, and also the lowest marked by the brown
spots which represent the remnants ae obsolescent lateral blotches in the westerly
subspecies.

I have discussed elsewhere* some of the difficulties involved in using tail-
length proportionality in taxonomic work. In the genus Arizona the trouble-
some ontogenetic changes, although present, are of moderate extent. It is
tree that the juveniles have proportionately slightly shorter tails than adults
(the opposite is the case in many genera) the difference being somewhat more
pronounced in the females than the males. But as the differences are relatively
small, and few juveniles are available in any of the series with which I have
worked, J propose in these studies to use mean proportionalities, treating the
sexes separately. Tail length proves to be of considerable importance in this
genus; it rather sharply separates the forms of eastern Arizona and eastward,
ireh are relatively long-tailed, from those found further west.

DESCRIPTION OF SUBSPECIES

I shall now proceed to the descriptions of the new subspecies, together
with summaries of the old (elegans and occidentalis) as newly delimited. It has
been the new material lately become available, rather than any new methods
of approach, which has made these new segregations possible. The additional sub-
species, far from complicating the picture, seem to me to lead to co-ordination
and territorial consistency. Uncertainties and irregularities are manifest only
where the material is still inadequate.

* Bull. San Diego Zool. Soc., no. 18, pp. 5-60, 1943.

320 SAN Dieco SoctETY oF NATURAL HIstory

While intergrades are not yet at hand between all territorially contiguous
pairs of subspecies, the relationships seem so close, and character overlaps are so
evident, that I have considered all the available forms to be subspecies of Arizona
elegans, thus continuing the monotypic status of the genus. As there are many
areas in which Arizona should thrive from which specimens are not yet available,
I should expect additional forms to come to light from time to time. There
are many places where the most fruitful method of collecting (road cruising
at night) cannot be practiced, and it may take years of collecting there to dis-

close the presence of so secretive and nocturnal a form as Arizona. I have in
mind, especially, parts of north-central Mexico, central Baja California, and
some of the islands off that peninsula.

I shall discuss the subspecies in territorial order, proceeding from east to
west.

Genus ARIZONA

Arizona Kennicott in Baird: aS of the Boundary, United States and
Mexican Boundary Survey (Emory), vol. 2, p. 18, 1859. Type species elegans.

The body is moderately slender with the head slightly distinct. The tail
is from 11 to 17 per cent of the length over-all. The tails of the males, in
proportion to length over-all, average about 11 per cent longer than those of the
females. The snout is rather sharp, with a sharply recurved rostral, and the
underjaw deeply inset. The pupil of the eye in life is slightly elliptical, with the
long axis vertical, although often round in preserved specimens. The head plates
are normal. The nasal plates are usually separated below, but not above, the
nostril. There is generally a single loreal, one preocular, and two postoculars;
however, two preoculars are prevalent in some areas. Two supralabials touch the
orbit. The temporals are commonly 2+3 or 2+4, but other combinations are
not infrequent. The dorsal scales are smooth, in 25 to 31 rows at mid-body,
and with faint single apical pits. The ventrals vary from 185 to 241; the anal
plate is undivided; the subcaudals are in a double series numbering from 39
to 63. The ventral scutes have a sexual divergence of about 4 to 5 per cent
in favor of the females, and the subcaudals 9 to 10 per cent in favor of the
males.

The pattern comprises a series of bown blotches on a light-brown, buff,
or cream, ground color. Smaller auxiliary blotches mark the sides. The ventrum
is immaculate, although the lateral marks may touch the outer edges of the
ventral scutes.

Arizona elegans elegans Kennicott

Texas GLossy SNAKE
Plate 7, fig. 1.

1859. Arizona elegans Kennicott in Baird: Reptiles of the Boundary, United
States and Mexican Boundary Survey (Emory), vol. 2, p. 18. Type
specimen USNM 1722; type locality Rio Grande ( = Lower Rio
Grande {River}, Texas). Cotype USNM 4266; locality between

KLAUBER—THE GLossy SNAKE 321

Arkansas and Cimarron (= between Arkansas and Cimarron rivers,
Oklahoma). The cotype is now to be assigned to the new subspecies

blanchard1.
1875. Pityophis elegans (part) Cope, Bull. U. S. Nat. Mus., no. 1, p. 39.

1883. Pityophis catenifer var. deppei (part) Garman, Mem. Mus. Comp,
Zool., vol. 8, no. 3, p. 151.

1886. Rhinechis elegans (part) Cope, Proc. Amer. Philos. Soc., vol. 23, p. 284.

1894. Coluber arizonae (part) Boulenger, Cat. Snakes Brit. Mus., vol. 2, p. 66.
(See remarks on this Boulenger name, under nomenclatorial and
systematic problems.)

1894. Pituophis elegans (part) Gunther, Biol. Cent.-Amer., Reptilia and
Batrachias ip. 125:

1915. Arizona elegans (part) Strecker, Baylor Bull., vol. 18, no. 4, p. 34
(first mention of specimens from best differentiated area near San
Antonio, Texas).

1924 Arizona elegans elegans (part) Blanchard, Occ. Papers Mus. Zodl.,
Univ. Mich., no. 150, p. 3.

Diagnosis —A subspecies characterized by 31 or 29 dorsal scale rows; and
dark, well-defined blotches, longer than the interspaces. Elegans can be segre-
gated from all subspecies except blanchardi by its having 31 or 29 scale rows,
while the other subspecies usually have 27 rows, although they may occasionally
have 29. However, those western subspecies which have the highest frequency of
29 rows (occidentalis, eburnata, and noctivaga), still average well below elegans;
also, they have proportionately shorter tails and differ in various details of
pattern. The subspecies most like elegans is blanchardi, which, however, has
fewer ventrals and more body blotches, on the average, than elegans.

Nomenclatorial and Systematic Problems.—There is some uncertainty with
regard to the exact locality of collection of the type specimen of elegans,
USNM 1722. This is given as Rio Grande (Kennicott in Baird, 1859, p. 19)
and later by Yarrow (1883, p. 108) as Lower Rio Grande. It was collected by
A. Schott, and. as many of Schott’s Texas specimens were collected at Eagle
Pass, Maverick County, it may have come from that vicinity. The ventral count
(male, 208), while low for elegans as found in southern Texas, cannot be
considered sufficiently abnormal to prove that it came from some other
point. The cotype, USNM 4266. from between the Arkansas and Cimarron
rivers, while presumably from Oklahoma, may have come from Kansas or
Colorado. In some ways this specimen would have constituted a more satis-
factory type, since it may be more definitely allocated; but No. 1722 is the first
named of the two, and besides was selected as the type by Blanchard, the first
reviser. Also, it is the specimen shown in the original plate, the cotype being a
female.

Boulenger (1894, p. 66) transferred the species elegans to the genus
Coluber, and, as the name elegans was preoccupied in that genus, he assigned
it the new specific name of arizonae. We now know that the transfer to Coluber

322 SAN Disco SocrETY OF NATURAL History

was incorrect, and in accordance with a decision of the International Com-
mission on Zoological Nomenclature which is expected to be in print shortly,
secondary homonyms so created are not to be regarded as permanent homo-
nyms. Thus elegans remains valid; however, Boulenger did inadvertently describe
the species arizonae which remains to be disposed of. He had two type specimens,
one designated a from Duval County, Texas, the other, 6, from Warners Ranch,
San Diego County, California. Hence arizonae is a composite of two subspecies,
elegans and occidentalis, and in order to avoid future confusion and the possible
invalidating of Blanchard’s occidentalis, 1 designate the British Museum speci-
man a from Duval County, Texas, as the lectotype of arizonae Boulenger, 1894,
thus making it a synonym of elegans Kennicott, 1859.

Within the territory which Blanchard assigned to the subspecies elegans,
there are available today fairly adequate series of specimens from western
Kansas, Oklahoma, and the San Antonio section of Texas. If we compare the
snakes of the most widely separated areas, western Kansas and San Antonio,
we find a further segregation to be possible, the northern specimens being lower
in ventrals and higher in blotch counts than the southern. There is, in fact,
no overlap in ventrals when these limiting populations only are considered, the
Oklahoma specimens being intermediate, as might be expected. To this extent
the geographical picture is quite consistent, but there are complications. First, the
type specimen of elegans is both uncertain as to locality and intermediate with
respect to characters; second, inadequate material from the area lying south of
the Texas Panhandle and west of San Antonio makes it difficult to allocate the
specimens of this territory, for some of the few available are both confusing as
to characters and from indefinite localities. There is an indication that specimens
from the lower Rio Grande Valley average fewer ventral scutes than those from
the vicinity of San Antonio or from the Chisos Mountains. After considering
the factors involved, I have finally decided to assign to elegans the southern
Texas specimens usually having high ventral counts and a low number of
blotches, and to consider the northern form as the new subspecies. This
makes the most consistent geographical arrangement. The northern form I
shall describe as blanchardi after. the late Dr. Frank N. Blanchard, the first
reviser of this genus, who in 1936, learning I was considering the description
of the Colorado Desert snakes as a new subspecies, generously suggested that
[ include in my study the eastern subspecies as well, although he had already
begun gathering data on them. If, later, it be concluded that I have incorrectly
assigned the type of elegans, then it is probable that the southern subspecies
will take Boulenger’s name arizonae. But it seems to me that geographical
consistency is best served by the allocation I have made. Were it not
that it seems best to begin these subspecies descriptions with the type subspecies,
I should have preferred to start with the western subspecies, which, being repre-
sented by more adequate material and uncomplicated by an indefinite type
specimen, present a more assured arrangement.

Material—The discussion of elegans which follows is based upon 39
specimens from Texas, 29 males and 10 females. Only the Bexar—Atascosa—
Comal counties area is adequately represented (by 30 specimens), the rest

KLAUBER—THE GLossy SNAKE 323

being scattered about, including 4 from the trans-Pecos section. Much additional
collecting will be necessary before the territorial trends in this subspecies can
be fully determined. Two specimens from Coahuila are at hand, but they
have not been included in the statistics which follow, since they are only
tentatively assigned to this form. Necessarily the subspecific description which I
present is strongly influenced by the predominance of material from the San
Antonio area.

Description of Subspecies—This is a snake of moderate colubrid propor-
tions, neither heavy bodied nor attenuated. The head is only slightly distinct
from the neck. Viewed from above, the head is wedge-shaped but with a
rounded snout. From the side, the top of the head is slightly convex or
flat; the jaw is deeply inset. The apex is moderately pointed; the forward
part of the snout slants upward, so that the tip is toward the upper end of
the rostral. The eyes are rather small and non-protuberant; the diameter is
about equal to 60 per cent of the distance from the anterior edge of the
orbit to the nostril. The pupil appears round in most preserved specimens but
is slightly elliptical in life.

The longest specimen I have seen, a male from near Somerset, Atascosa
County, Texas, measured 1386 mm. The smallest specimen measured 293
mm., and, having been collected on August 18, must have been but a few days
old.* The tail proportion (ratio to length over-all) varies from 13.6 to 16.6
per cent in the males, and in the females from 13.5 to 14.8 per cent. The
averages are respectively 14.9 and 14.2 per cent. It is believed that more ex-
tensive series would tend to raise the male figure; the sexual difference is
usually a full per cent. The young specimens have proportionately shorter
tails than the adults.

The hemipenis is single and widens outwardly. The inner half is
almost covered with tiny points, although these are less evident on the side
opposite the sulcus. At the middle of the shaft the organ widens rather suddenly
and simultaneously the points increase in size. However, even the longest
probably do not reach a millimeter in length and most are barely half that length,
so they can hardly be referred to as spines. They are very densely set and
are smallest in size along the sulcus.; At the outer end the points change
to reticulated flounces, whose outer margins are edged with tiny points. At the
outer end there is a truncated surface at an angle with the shaft; this contains
an unreticulated groove, which is reached by the sulcus passing over the
outermost part of the organ.

The body is covered with smooth scales, rather narrow and with slightly
rounded ends. The lower lateral rows are increasingly enlarged. Single apical
scale pits are faintly evident on some scales.

* There is another specimen only 195 mm. in length but its condition is such I cannot
be sure it is an Arizona. It has some scale peculiarities which distinguish it from all the others
I have studied.

+ The points in elegans are notably smaller and more plentiful than the spines in
eburnata and occidentalis.

324 SAN Disco Society oF NATURAL HIstory

The scale rows at mid-body are usually 29 or 31, there being some sexual
dimorphism so that the females more often have 31. One specimen from
northern Texas has only 27 rows; it is possible that this may be a philipt
intergrade. The ventrals in the males vary from 208 to 222, interquartile
range 212.3 to 217.1, mean 214.71+.72,* coefficient of variation 1.65 per
cent; the over-all range in the females is from 220 to 232, interquartile range
223.1 to 227.1, mean 225.07+.79, coefficient of variation 1.31 per cent.
The anal is entire. The subcaudals, all divided, vary from 51 to 62 in the
males, the interquartile range being 54.3 to 58.4, mean 56.36+.62, coefficient of
variation 5.47 per cent. In the females the over-all range is 49 to 53 (this
range will no doubt be considerably enlarged when additional specimens shall
have become available), interquartile range 50.7 to 52.5, mean 51.64+.36,
coefficient of variation 2.57 per cent. The coefficient of sexual divergence in
the subcaudals is 8.7 per cent. The terminal scale is considerably elongated,
with a lateral crease; it is seldom difficult to tell whether the tail is complete.

The head scales approach the colubrid normal in size and arrangement.
The rostral is rather sharply curved, both over the top of the snout and
below. It is wider than high. The upper point separates the internasals for about
half their lengths; in addition it contacts the prenasals and the first supralabials.
The internasals are longer than wide; they curve downward to a point in
front of the prenasals. The prefrontals are wider than high and curve well
down over the canthus rostralis to a level with the center of the orbit. The
supraoculars are triangular in shape; they touch the prefrontals at a point
and widen posteriorly, where they contact the parietals and upper postoculars.
The frontal is widest anteriorly and terminates posteriorly at a point which
does not deeply indent the suture between the parietals. The parietals are
the largest of the head scales. They are moderately regular in shape, de-
creasing in size posteriorly. The dorsal scales which border them are somewhat
enlarged, but are irregular. The prenasal is smaller than the postnasal; it is
widest below, while the contrary is true of its fellow. The nostril is a slit at
the upper end of the suture which divides them; it slants forward above. The
suture is not complete above the nostril. The postnasal usually contacts the
first and second supralabials. The loreal is twice as long as wide, and is
highest forward, becoming pointed posteriorly where it abuts the preocular.
The loreal contacts the second and third supralabials. Although divided loreals
are occasional in the species, I have noted none in this subspecies. The pre-
oculars are usually single, although paired in 2 counts out of 78. The
preocular is somewhat wider above than below. The postoculars ordinarily
number 2, and are subequal in size; there are 3 in one count out of 78. The
temporals are usually 2+3 or 2+4, but vary from 1+2 to 3+5. Those in
the first series are slim and elongated, and slant upward posteriorly. The
supralabials are usually 8 but occasionally number 9 (10.5 per cent). The
fourth and fifth contact the eye; the next to the last is always much the
largest. The infralabials vary from 11 to 15, the distribution being 11(2),

* The minus-or-plus sign indicates that the following figure is the standard, rather than
the probable, error of the mean.

KLAUBER—IHE GLossy SNAKE 325

12(10), 13 (39), 14(22), and 15(1). The first pair meets on the median line.
Usually the seventh is the largest of the series, although it may be the
eighth if there are 14 or more in the series. The mental is small and triangular.
The anterior genials are large and contact on the median line; the posterior are
both slimmer and shorter, and, diverging posteriorly, are separated by from
2 to 4 rows of gulars.

Elegans is a brown snake with conspicuous dark-brown blotches down
the back, and additional secondary blotches on the sides, which may or may not
be obscured by a lateral suffusion of brown. It certainly cannot be called

“faded.”

The head is brown or red-brown above, and lighter-brown on the sides,
shading into buff at the supralabials. There is a brown band, darker than the
ground color, on the posterior edges of the prefrontals; also dark-brown spots
on the supraoculars, frontal, and parietals, often concentrated along the sutures
which separate these plates. There is a brown streak passing backward from
the orbit to the angle of the mouth, and a vertical brown mark between the
supralabials immediately below the eye; this tends to widen as the lip is
approached. The lower jaw is usually immaculate, although the posterior in-
fralabials are sometimes marked by a continuation of the postocular dark streak.

On the neck there is usually a pair of parallel dark marks separated by a
light streak. These, the precursors of the dorsal series, may be joined together
anteriorly, posteriorly, or both.

The dorsal blotches occupy considerably more longitudinal space than the
interspaces which separate them. The body blotches vary in number from
42 to 56, interquartile range 46.6 to 51.7, mean 49.11.62, coefficient of
variation 7.7 per cent. The tail spots have an over-all range of 13 to 24, inter-
quartile range 16.0 to 19.9, mean 17.97+.49, coefficient of variation 15.4 per
cent. The body blotches at mid-body cover from 11 to 16 scale rows across
the body, but in most specimens they span either 13 or 15 rows. The
longitudinal extent of the blotches, again measured at mid-body, varies from
2 to 4 scales, end to end, but most specimens extend over 3 to 31/2 scales. The
interspaces are much narrower; most of them are but a single scale in length,
but in a few specimens reach 12 scales. The lateral brown suffusion is usually
carried down to the third or fourth row above the ventrals. The lowest lateral
spots of the auxiliary series extend considerably lower; in 70 per cent of the
specimens they touch the outer edges of the ventrals, and in the others they
reach the first row above.

The body blotches, although generally rectangular or elliptical, with the
major axis transverse, are often quite irregular because the patterns of the
two sides do not exactly match. This results in some diagonal blotches, and
others which are restricted to one side, while still others are Y-shaped, being
double on one side and single on the other.

In addition to the main dorsal blotches, there are, on each side, two or more
alternating series of progressively smaller spots. The first of these series com-

326 SAN Disco Society OF NATURAL HIstTory

prises subcircular spots, alternating with the dorsal series, and is rather regular;
those below are elongated, irregular, and often indefinite.

The main dorsal blotches are dark-brown with darker-brown or black
edges. There is often considerable red-brown in live specimens, but the red is lost
in preservation.

Between blotches the ground color is cream or buff, but this is evident
only on the 3 to 5 middorsal scale rows, for laterally the area between blotches
is heavily suffused or punctated with brown. This darkening occurs par-
ticularly in scale centers; the scale edges are usually light, giving a net-work
effect characteristic of Arizona. The lateral darkening tends to obscure the
outer edges of the middorsal blotches and the lateral series as well; in some
specimens the lateral suffusion is so dense as to blot out completely the side
blotches. This side-darkening increases with age. Juveniles are usually quite clear
between blotches.

A live specimen said to have been from New Braunfels, Texas, exhibited
the following Ridgway (1912) colors: Ground color of the head, Wood
Brown; lateral streaks on the head, Blackish Brown; centers of dorsal body
blotches, Fuscous; edges of blotches, Black; middorsal interspaces, Mikado
Brown centers, with Orange-Cinnamon edges; lateral blotches, Sepia to Black;
centers of scales in lateral interspaces, Snuff Brown to Orange-Cinnamon; light
edges of interspace scales, Pinkish Buff; ventrum Pale Olive-Buff.

Intrasubspecific Trends—The ventral scutes reach a maximum in the
San Antonio area, and the body blotches attain a minimum there, except for
two Coahuila specimens, somewhat doubtfully allocated to this subspecies.
Specimens from the Chisos Mountains are also high in ventrals, but those from
the intervening territory are lower. I have discussed some of the questions

brought up by individual specimens under “Nomenclatorial and Systematic
Problems.”

The two specimens from Coahuila are low in both ventrals and body
blotches. One is a male with 205 ventrals and 55 subcaudals, the other a female
with 211 ventrals and an incomplete tail. The labials and oculars do not differ
from those of Texas elegans. The male has 40 body blotches and 14 tail spots,
the female 43 blotches. The blotches are both wide and long, greatly exceeding
the interspaces as in elegans. Both specimens have 31 scale rows, showing an
affinity toward elegans and blanchardi, rather than philipi or any other sub-
species farther west. Should further collecting in Coahuila continue to produce

specimens as low in ventrals and body blotches as these, recognition of another
subspecies may be justified.

Relationships with Other Subspecies —Elegans intergrades with blanchardi,
but the exact zone of intergradation cannot now be determined for lack of
material. ‘Tentatively I have presumed the line to be the Red River and the
southern end of the Texas Panhandle. To the west there is little doubt that
elegans intergrades with philipi, this being suggested by several specimens having
indefinite localities but presumed to be from the trans-Pecos area of Texas.
The critical area, with respect to this relationship, is Hudspeth and Culberson

KLAUBER—THE GLossy SNAKE 327.

counties, from which no specimens are at present available. A specimen
(MZUM 81984) from Lubbock, Lubbock Co., Texas, having 27 scale rows and
208 ventrals (male), suggests that this part of Texas also may be an area
of intergradation between philipi and elegans. The determination of the relation-
ship of elegans with the specimens from Coahuila and, through them, with
expolita must also await further collecting.

Life History Notes——I have had no personal field experience with this
subspecies. Such first-hand information as I am able to supply on Arizona
will be found under the western subspecies, particularly eburnata, candida, and
occidentalis. I am of the opinion that the several subspecies are much alike in
habits, except that the eastern forms are less completely nocturnal.

Mr. Albert J. Kirn, of Somerset, Texas, has kindly supplied me with
some interesting field notes on elegans, as found in that vicinity. He writes:
“All of the specimens taken by me were found in the daytime; I do not
recall seeing them at night. April 28, 1929, in the afternoon, I came upon
an Arizona trying to swallow a mole (Scalopus aquaticus texanus) in a corn
field. The snake had two coils around the body of the mole, and was doubled
around the end of the mole so that the entire snake was only about 8 inches

long. The snake had swallowed the head and one foot.

“At 6:30 a.m., September 5, 1932, I saw a small coral snake (Micrurus
fulvius tenere) trying to swallow an Arizona; it had the head and a small part
of the body swallowed. The coral snake measured 187s in. and the Arizona
127% in.

“My earliest record for 1944 was Feb. 23, that of 1945, March 4. When I
come upon any of these snakes they usually contract themselves into short
bends or kinks for the entire length of the body. One, for example had 11
kinks. I do not know if this is done for fright or if it is a method of defence;
they never offer to strike. They do not coil, but remain in a traveling position.
This is, of course, in the daytime.

“These snakes are frequently plowed up by farmers when breaking land.
This would indicate that they do not go deep into the ground; probably they

seek refuge in mole, kangaroo rat, or possibly pocket gopher holes.

“Most of the specimens I have secured are from 7 miles southwest of
Somerset. The country roundabout is sandy to very sandy. There are woods
and cultivated fields. The woods, on more sandy land, are ‘black jack’ oak, mixed
with hickory and live-oak. The topography is gently rolling; there is little rock
except in gullies, and some sandstone outcrops. I think Arizona well-dispersed
throughout this region, especially in the more sandy areas.

“One large female I collected, May 31, 1940, measured 42! inches; it
contained 12 eggs. Another laid 9 eggs.”

Elegans, especially as found in the San Antonio area, is the largest and
darkest of the subspecies. There is no doubt that it is more diurnal than the
westerly subspecies.

328 SAN Disco Society oF NATURAL HIstory

This subspecies reaches an altitude of at least 3800 ft. in the Chisos Moun-

tains.

Range.—Texas east of long. 98°, excluding the Panhandle and El Paso

County; also, tentatively, northern Coahuila, Mexico.

Locality Records —

BayLor CouNTY:
Seymour*
LUBBOCK COUNTY:
Lubbock+
Howarp County:
10 mi. e. of Big Spring on US 80
REEVES COUNTY:
Pecos
BREWSTER COUNTY:
Between Marathon and Alpine
3 mi. w. of Government Spring
(Chisos Mts., alt. 3800 ft.)
ComaL County:
New Braunfels
BExAaR COUNTY:
San Antonio (also 15 mi. s.)
Somerset

ATASCOSA COUNTY:

7 and 8 mi. s. of Somerset
(Bexar County )

Benton

Poteet

10 mi. e. and 8 mi. ne. of
Pleasanton

Lytle (also 4 and 7 mi. se.)

DuvaL CouNTY
KLEBERG COUNTY:

10 mi. sw. of Kingsville

Brooks County:

11 mi. n. of Encino

Hipatco County:

Edinburg

Also the following indefinite localities:

Lower Rio Grande (type locality)
Between Pecos River and Rio Grande

CoaHulILa, MExIco

Hacienda las Rusias (Ciudad Muzquiz)*

Cuatro Ciénegast

Arizona elegans blanchardi subsp. nov.
Kansas GLossy SNAKE

1859. Arizona elegans (part) Kennicott in Baird: Reptiles of the Boundary,
United States and Mexican Boundary Survey (Emory), vol. 2, p. 18.

1875. Pityophis elegans (part) Cope, Bull. U. S. Nat. Mus., no. 1, p. 39.
1883. Pityophis cantenifer var. deppei (part) Garman, Mem. Mus. Comp.

Zool., vol. 8, no. 3, p. 151.

1886. Rhinechis elegans (part) Cope, Proc. Amer. Philos. Soc., vol. 23, p. 284.
1894. Pituophis elegans (part) Gunther, Biol. Cent.-Amer., Rept., p. 125.
1924. Arizona elegans elegans (part) Blanchard, Occ. Papers Mus. Zodl.,

Univ. Mich., no. 150, p. 3

1929. Arizona elegans elegans Taylor, Univ. Kan. Sci. Bull., vol. 19, no. 5,
p. 56 (first mention of specimens from Kansas, best differentiated

locality).

* Specimen no longer available, might be blanchardi or intergrade.

+ May be philip: intergrade.

{These are only tentatively assigned to the subspecies elegans.

KLAUBER—THE GLossy SNAKE 329

Type—No. 10393 in the collection of the Natural History Museum,
Stanford University. Collected by J. W. Anderson, July 7, 1942, in Cheyenne
County, Kansas, 13 miles southeast of Benkelman, Dundy County, Nebraska.

Diagnosis —Blanchardi is a large-blotched, dark-colored, subspecies with 29
or 31 scale rows and a relatively long tail. While some western subspecies
(occidentalis, eburnata, and noctivaga) occasionally have 29 scale rows, they
have more blotches and shorter tails. Blanchardi has more ventrals than philipi
or expolita. It is nearest to elegans, from which it differs in having, on the
average, a lower number of ventrals and more blotches.

Description of the Type—The type specimen is a young adult male;
length over-all 752 mm.; tail length 119 mm.; tail proportion 15.8 per cent.

The head is convex on top; it is a little wider than the neck. The snout is
pointed with the lower jaw inset. The pupil is nearly round. The distance
from the anterior edge of the eye to the nostril is almost twice the eye diameter.

The scales of the head are normal. The rostral is hollowed below and is
20 per cent wider than high. It contacts the first supralabials, the anterior
nasals, and the posterior point separates the internasals for more than half
their lengths. The internasals are longer than wide and curve down to a
point in-front of the nasals. The prefrontals are quadrangular in shape, wider
than long, with their post-lateral points curved well down over the canthus
rostralis, to form a long contact with the loreal. The frontal is pentagonal,
narrowing along the supraoculars and coming sharply to a point between the
parietals. The supraoculars are very narrowly in contact with the prefrontals;
they widen posteriorly. The parietals are large and contact more than half
of the posterior edges of the upper postoculars. The postnasal is considerably
larger than the anterior; the slit-like nostril is at the upper end of the diagonal
separating suture, which is not complete above the orifice. Of the supralabials,
the postnasal touches the first and second. There is a single loreal on either
side, about twice as long as high and pointed posteriorly; it contacts the second
and third supralabials. There is a single preocular on the right, and two, of
which the lower is very small, on the left. The postoculars are 2-2 and are
subequal. The temporals are 2+4, 213, the anterior being long and slender.
The supralabials are 8-8; the fourth and fifth touch the eye and the seventh
is much the largest. The mental is small and subtriangular. The infralabials are
13-14, the first on each side in contact with its fellow behind the mental, and
the seventh on the right and eighth on the left much the largest. There are
two pairs of genials, the anterior considerably the longer, and medianly in
contact; the posterior are slightly divergent and are separated by two or three
rows of gulars.

The dorsal scale rows are 29-31-19. The scales are smooth, rather narrow
and with rounded ends. Single apical pits are faintly in evidence on some
scales. The lower lateral rows are the widest. The ventral scales number 201;
the anal plate is entire; there are 57 subcaudals, all divided.

330 SAN Disco SoclrETY OF NATURAL HIstTory

The head is light-brown above and buff on the sides. There is a brown
bar across the posterior edges of the prefrontals, and some dark-brown spots
on the frontal, supralabials, and parietals. There is a wide, dark streak
from each eye back to the angle of the jaw. There is a faint brown mark under
each eye, otherwise the supralabials are unspotted. The lower surface of the head
is immaculate buff. There is a pair of parallel dark blotches on the back of
the head and beginning of the neck.

The dorsal body-pattern comprises a series of 58 rather irregular brown
blotches, with dark-brown or black edges, on a buff background. The irregular-
ity is due to the fact that the two sides do not match, so that some blotches
are on one side only, while others are Y-shaped. The buff of the interspaces is
evident only on the three middorsal scale rows, the others being heavily stippled
with brown (except on their edges) whether within or between the main blotches.
The effect is almost to obscure the !ateral ends of the median blotches, and the
lateral auxiliary blotches are quite submerged in the general brown color of
the sides. At mid-body the blotches are about 2/2 to 3 scales long (end to end),
and 12 scale rows wide, across the back. ‘The interspaces are about one scale
long. The brown suffusion on the sides reaches the fourth lateral scale row
above the ventrals. Some dark spots, the remains of the obsolescent auxiliary
series, reach the ventrals themselves, which are otherwise immaculate cream-
color. The tail spots are so obscured by the general brownish suffusion that
they cannot be counted with accuracy. The unmarked dorsal interspaces form
an almost continuous middorsal light line on the tail.

Summary of Paratypes—Since, in the characters which distinguish blanch-
ardi from elegans, the specimens from farthest north exhibit the greatest diver-
gence, I have selected the type from northwestern Kansas and deem it desirable
to summarize the northern specimens separately as paratypes, since the inclusion
of those from New Mexico and Oklahoma (as is done in the complete sub-
species description) to some extent compromises these differences. Thus I
include as paratypes one specimen from Colorado, one from Nebraska, and
14 from Kansas.* The statistics on these (including the type) are as follows:
There are 12 males and 5 females. The scale rows are 29(10), 30(2), or
31(5). The ventrals in the males vary from 197 to 206, mean 202.3; in the
females from 207 to 213, mean 210.6. The subcaudals range from 55 to 63 (mean
57.3) in the males, and from 51 to 55 (mean 52.5) in the females. The supral-
abials are 8 in all cases, except for 7 on one side of one specimen. The infralabials
are usually 13 (60 per cent) but vary from 11 to 14. The loreals are all single.
There are two preoculars in 16 per cent of the counts, the others being single;

* LMK 35343 Schramm, Yuma Co., Colo.; WSC 41-197, 2 mi. se. of Stratton, Hitch-
cock Co., Neb.; and the following from Kansas: KUJ 2335 Ashland (Clark Co.); KU 3560
Little Salt Marsh (Stafford Co.); KU 3561-2 Elkhart (Morton Co.); KU 20097 near
Syracuse (Hamilton Co.); KU 20785 4 mi. sw. of Duquoin (Harper Co.); KU 20784 1
mi. e. of Sharon (Barber Co.); KU 20793 Meade County State Park, KU 22206 6% mi.
sw. of Meade, AMNH 62848 12 mi. sw. of Meade (Meade Co.); USNM 89239 near
Hutchinson, CHAS 12244 between Stafford and Sylvia (Reno Co.); and NHMSU 9912
and 9913 2 and 1 mi. se. of Dodge City (Ford Co.).

KLAUBER—THE Glossy SNAKE 331

this is a higher percentage of paired preoculars than in any other subspecies
except candida. There are two postoculars. The temporals are usually 2+3, but
may be 2+2 or 214.

The body blotches vary from 51 to 66, mean 56.7, and the tail spots
from 15 to 27, mean 21.5. The blotches at mid-body vary from 11 to 15, but
are usually 12 or 13 scale rows in width across the body. The blotches are
from 2 to 3 scales (end to end) in length; the interspaces are usually 1 or
1% scales in length, but may be as long as 134. Thus the blotches always exceed
the interspaces in extent along the body. The brownish side-suffusion usually
terminates on the first to third row above the ventrals, and the lowest auxiliary
lateral blotches reach the ventrals or the first scale row above. In general these
Kansas specimens are neither as dark, nor with as large or prominent blotches
as are evident in elegans farther south.

The tail proportionality in these paratypes of blanchardi runs from 14.8
to 16.8 per cent in the males (mean 16.1), and 14.2 to 15.6 in the females
(mean 14.9).

Material_—The description which follows is based on 15 specimens from
Kansas, 1 each from Colorado and Nebraska, 2 from eastern New Mexico,

and 17 from Oklahoma, total 36; of these 23 are males and 13 females.

Subspecies Description—tThe scale rows are 29 to 31, about 58 per cent
having the lower number. The ventrals vary from 197 to 215 in the males, the
interquartile range being 201.4 to 207.9, the mean 204.63+1.10, coefficient of
variation 2.34 per cent. The females range from 207 to 222, interquartile range
211.2 to 217.9, mean 214.541.38, coefficient of variation 2.32 per cent. The
subcaudals, all divided, vary from 52 to 63 in the males, interquartile range
54.6 to 58.4, mean 56.53.68, coefficient of variation 4.97 per cent; and in
the females the range over-all is 47 to 55, interquartile range 50.4 to 53.4, mean
91.90=-.72, coefficient of variation 4.4 per cent. The supralabials are rarely
other than 8, there being one 7 and two 9’s out of 66 counts. The distribution
of the infralabials is 11(3), 12(13), 13(38), and 14(5). The loreals are
nearly always undivided; only two out of 62 are split. The preoculars are single
in 79 per cent; paired in the others. The postoculars nearly always number 2,
although in two counts out of 70 there are 3. The temporals are usually
2+3 or 2+4, but vary from 2+2 to 2+6.

The body blotches vary in number from 47 to 66, interquartile range 53.0
to 60.1, mean 56.54+.88, coeflicient of variation 9.22 per cent. The tail spots
range from 15 to 27, interquartile range 18.8 to 22.5, mean 20.65.56, coefficient
of variation 13.03 per cent. The blotches cover from 11 to 15 scale rows,
laterally; usually they are 12 or 13 rows wide. Longitudinally they extend from
1% to 3% scales, end to end; most fall between 2 to 3 scales. The inter-
spaces are shorter; while most of them occupy but a single scale, a few
extend to as much as 1% scales. But the blotches are always of greater extent
than the spaces between. The dark punctations on the sides extend down to the
third to fifth lateral scale rows above the ventrals. The lateral secondary blotches

332 SAN DreGco SocrETy OF NATURAL HIstory

usually touch the ventrals, but in some specimens only extend to the first
row above.

The longest specimen I have seen measured 1168 mm. This was a male;
the longest female measured 1165 mm. These specimens were from Oklahoma;
the Kansas specimens seem to be somewhat shorter. The shortest specimen
measured 290 mm. The tail ratio (to length over-all) varies from 13.6 to
16.9 per cent (mean 15.7) in the males, and 13.8 to 15.6 (mean 14.7) in the

females.

Intrasubspecific Trends——In blanchardi the ventrals increase from the
northwestern limit of the range, southward through Kansas to Oklahoma. The
proportion of specimens having two preoculars is higher in Kansas than else-
where.

The blotches decrease in number from Kansas southward, and the lateral
spots reach lower scale rows in the northern part of the range.

Relationships with Other Subspecies —Blanchardi intergrades with elegans,
possibly along the line of the Red River and the southern end of the Texas
Panhandle, but the zone cannot be located with assurance with the material
presently available. Intergradation with philipi at the headwaters of the Canadian
River in New Mexico is possible; blanchardi is found at least as far west as
Clovis, Curry County (USNM 118532), and 21 miles southwest of Nara
Visa, Quay County (CHAS 10126), New Mexico.

Life History Notes—I have had no field experience with this subspecies.

Taylor (1929, p. 56) reports a specimen having been found crawling
across the road in the daytime. But he believes the subspecies to be strictly
nocturnal and its being abroad in the day very unusual. Two other specimens
were collected at night.

Smith and Leonard (1934, p. 194) found two specimens in sand dunes in
Oklahoma at 10 p.m. Burt and Hoyle (1934, p. 208) found one in sandy,
sage-brush country, while another was in pasture land. The first was a female
with 10 large eggs in the right oviduct and none in the left. Marr (1944, p. 485)
found a specimen attempting to eat a trapped (dead) kangaroo rat (Dipodomys
ordi).

This subspecies reaches an altitude of at least 3500 ft. in southwestern
Nebraska and northeastern Colorado.

Range.—Northeastern Colorado, southwestern Nebraska, western Kansas,
western Oklahoma, the Texas Panhandle, and northeastern New Mexico.

Locality Records. —
COLORADO
YuMA County:
Schramm
NEBRASKA
Hircucock County:

2 mi. se. of Stratton

KLAUBER—TIHE GLossy SNAKE 333

KANSAS

CHEYENNE County:
13 mi. sw. of Benkelman, Dundy
County, Nebraska (type locality)
HaMILTON County:
Near Syracuse
STAFFORD COUNTY:
Little Salt Marsh
RENO County:
Hutchinson, between Staffore
and Sylvia
5 mi. n. of Turon
Forp County:
1 and 2 mi. se. of Dodge City

Morton County:

81 Ranch (near Elkhart)
MeapE County:

6¥2 and 12 mi. sw. of Meade

Meade County State Park
CLaRK COUNTY:

Ashland
BARBER COUNTY:

1 mi. e. of Sharon
HARPER COUNTY:

4 mi. sw. of Duquoin

OKLAHOMA

BEAVER COUNTY:

2 mi. s. of Beaver
Harper County:

2 mi. s. of Kansas line (s. of

Englewood, Kansas)

Woops County:

Alva

Waynoka
ALFALFA COUNTY:

3 mi. e. of Cherokee
Dewey County:

Y2 mi. nw. of Taloga

BECKHAM COUNTY:
6 mi. e. of Erick
WASHITA COUNTY:
5 mi. ne. of Canute
CLEVELAND County:
Norman
GREER COUNTY
COMANCHE COUNTY
Also the following indefinite localities:
Neutral Strip
Between Arkansas and Cimarron
rivers

New Mexico

Quay County:
21 mi. sw. of Nara Visa

Curry County:
Clovis

Arizona elegans philipi subsp. nov.
PAINTED DESERT GLOSSY SNAKE

1896. Arizona elegans Cockerell, Amer. Nat., vol. 30, p. 326.
1900. Rhinechis elegans (part) Cope, Rept. U. S. Nat. Mus. for 1898, p. 862.

1924. Arizona elegans occidentalis (part) Blanchard, Occ. Papers Mus. Zool.,
Univ. Mich., no. 150, p. 1.

Type—No. 34456 in the collection of LMK, collected 10 mi. east of
Winslow, Navajo County, Arizona, by Charles E. Shaw and Carl Engler,
July 29, 1941.

Diagnosis—A subspecies characterized by a low number of ventral scutes
and a relatively long tail. In the latter character it is like elegans and blanchardi,
from which, however, it may readily be distinguished by its lower number of
scale rows; philipi, like the other subspecies farther west, usually has 27 rows,
while elegans and blanchardi have 29 or 31. From noctivaga and the other
subspecies farther west, philipi may be distinguished by its longer tail; thus
it is seen to be a transition form between the eastern and western subspecies—
like the former in tail proportionality and the /atter in scale rows. Only expolita,

334 SAN Dreco Society oF NATURAL History

a southern extension of philipi, occupies a similarly intermediate position, but

expolita has fewer dorsal blotches than philipi.

Nomenclatorial and Systematic Problems—Three paratypes of philip
are available from northeastern Arizona. One, LMK 20990, the first specimen to
be collected in that area, is a juvenile taken by Philip M. Klauber (for whom I
name the subspecies), August 20, 1933, at the crossing of Canyon Diablo with
U. S. Highway 66 (= Two Guns), Coconino County, Arizona. It was this
specimen whose peculiarities led to repeated trips to the Winslow area in
search of further material, but without success until the Shaw-Engler expedition
of 1941. The initial specimen was not selected as the type because it is a juvenile
and, further, has some abnormal head scales not found in any other specimen.
The two other paratypes are a topotype, LMK 34426, also taken by Shaw
and Engler; and PFNM 123, collected one mile from Second Forest (alt. 5500
ft.), Petrified Forest National Monument, Apache County, Arizona, by L. F.
Keller, October 5, 1943.

It was at first thought that this subspecies was restricted to this section
of Arizona, the snakes of central New Mexico and the El Paso area being
assigned to the southern Arizona subspecies, noctivaga. However, tail-length
studies later showed the New Mexico—El Paso snakes to belong with philipz,
rather than noctivaga. But I have seen fit to leave the selection of the type
unchanged, for the snakes of the Painted Desert area are the lowest in ventral
scale counts and are most differentiated from noctivaga in some other characters.
Later, if additional material should justify a further division of philipi, a new
name may be assigned to the southern race.

Description of the Type-——The type specimen is an adolescent male;
length over-all 565 mm.; tail length 94 mm.; tail proportion 16.6 per cent. The
measurements were made before shrinkage in preservative.

The head is flat-topped and rather narrow, so that it is only slightly
distinct from the neck. The snout is pointed and somewhat uptilted. The
pupil is almost round.

The scales of the head are normal. The rostral is wider than high; it is
sharply recurved with the posterior point inserted between the internasals for
about half their lengths. The internasals are longer than wide and have
sharp points between the rostral and prenasals. The prefrontals are quad-
rangular in shape, wider than long, and with their postlateral points curved
down over the canthus rostralis. The frontal is hexagonal, narrowing consider-
ably along the supraoculars and then terminating in a sharp point between
the parietals. The supraoculars touch the prefrontals at a point; they widen
somewhat posteriorly. The parietals contact nearly the entire posterior edges of
the upper postoculars. Posteriorly they are edged by normal dorsal scales.
There is a suture below the nostril, not above; the postnasals are the larger
of the pair. Of the supralabials, the postnasal touches the first and second,
the latter only at a point. There is a loreal on either side about twice as
long as high, touching the second and third supralabials. Posteriorly it is
curved and indents the preocular. On the left side there is an extra scale

KLAUBER—IHE GLossy SNAKE 335

below the loreal which has been cut from a corner of the third supralabial.
There is one preocular and two subequal postoculars. The temporals are
1+3, 1+3. The supralabials are 8-7, the fourth and fifth touching the eye,
and the seventh largest. This refers to the right side only; the left is peculiar
because of an incomplete suture between what should be the third and fourth
supraoculars. The mental is small and triangular. The infralabials are 13-13, the
first on each side being in contact behind the mental; the seventh is much
the largest. There are two pairs of genials; the anterior pair are medianly in
contact. They are longer than the posterior, which are divergent and separated
by several gulars.

The dorsal scale rows are 25-27-19. The scales are smooth, rather narrow
and with rounded ends. They are somewhat enlarged laterally. Single apical scale
pits are faintly evident. The ventral scales number 185; the anal plate is entire;
there are 51 subcaudals, all divided.

The head is light-brown above with dark-brown spots on the frontal
and parietals. The sides of the head are lighter, particularly the supralabials;
there is an indefinite dark streak from the eyes back toward the commissure.
The lower surface of the head is cream-color, unmarked.

The body pattern comprises a series of brown blotches on a buff back-
ground. There are 57 blotches on the body, and 17 on the tail. The blotches are
wider across the body than their longitudinal extent; at mid-body they are
about 2 scales long (end to end), and 12 scale rows wide. The interspaces
measure about 142 scales (end to end). The blotches are dark-edged, some
bordering scales being tipped with black. The interspaces show the buff ground
color only on the three middorsal rows; laterally there is much spotting with
brown, so as almost to obliterate the outer edges of the main blotch series. The
three lowest lateral rows are cream-color, irregularly spotted with dark-brown
or black at scale edges. A few spots even touch the edges of the ventrals, which
otherwise are cream-color. The scales of the maculate lateral rows are usually

lightest on the edges.

Summary of Paratypes—As there is some intrasubspecific variation in
the subspecies philipi, which might eventually lead to a further subdivision, I
shall summarize separately, as paratypes, the three other available specimens
from the Painted Desert area of Arizona, before discussing the subspecies in its
entirety.

The paratypes agree in all essentials with the type, except that the
specimen from Two Guns has some aberrant scales on the head.

The topotype is a young male measuring 405 mm. over-all, with a tail
length of 68 mm. (preserved measurements). The scale rows are 25-25-17. The
ventrals are 185; the anal entire; the subcaudals number 53, all divided. The head
scales are normal. The supralabials number 8-8, infralabials 13-12, loreals
1-1, preoculars 1-1, postoculars 2-2, temporals 2+4, 2+2. There are 56
body blotches and 20 spots on the tail. The blotches are but little lighter mid-
dorsally than laterally; they are edged with black. The blotches are 12 scale
rows in width and are about 2 scales long (end to end) sniddorsally; the inter-

336 San Disco Society oF NATURAL HIstTory

spaces are 142. A postocular dark-brown dash is evident on the side of the
head. The first two paired spots on the neck are relatively short.

The Canyon Diablo specimen is a juvenile female, evidently hatched but
a short time before collection. It measures 249 mm. over-all, with a 35 mm. tail.
The markings are clear and bright, with brown blotches on a light-gray back-
ground. The scale rows are 27-27-19. The ventrals number 195; the anal is
entire; there are 47 subcaudals. The head scales are anomalous in two particu-
lars, the presence of extra loreals and suboculars. This was one of the reasons
why so much effort was made to secure additional specimens from the vicinity,
for it was hoped that the subspecies would be distinctive in head scales, which,
however, proved not to be the case. The particular anomalies are: A small
extra scale above the loreal on the right, and one below on the left; on both
sides the fourth supralabial is divided transversely, and above this suture the
scale is divided vertically, so that there are two suboculars on either side.
There are 7—7 supralabials, and 12-12 infralabials. There is one preocular on
each side and two postoculars. The temporals are 113, 2+3. There are 62
dorsal blotches on the body and 17 on the tail. The blotches are clear and
evenly outlined. They average a little over 2 scales long, end to end, with
interspaces of one scale. The brown postocular streak is clear and even.

The third paratype, PFNM 123, the only specimen from Apache County,
is a female 615 mm. over-all, with a tail length of 89 mm., tail ratio 14.5 per
cent. It has 197 ventral scutes and 46 subcaudals. The supralabials are 8-8,
infralabials 12-13, loreals 1-1, preoculars 1-1, postoculars 2-2, and temporals
2+3, 2+3. The body blotches number 63; the tail spots are too indefinite to
be counted with accuracy. The blotches are 12 scale rows wide and 134 scales
(end to end) long; the interspaces occupy 144 scales. This specimen is con-
siderably darker laterally than the type or other paratypes, the brownish suffusion
being so dark and extensive as virtually to obliterate the lateral secondary
blotches, particularly toward the tail.

Material—The subspecific description which follows is based on the
four specimens from northern Arizona already discussed, together with the
following from other areas assigned to the same subspecies: Santa Fe County
2, Bernallilo County 5, Lincoln County 1, Sierra County 1, Grant County 1,
Hidalgo County 1, Luna County i, Dona Ana County 3, total New Mexico 15;
El Paso County, Texas, 9; Cochise County, Arizona 2; northeastern Chihuahua
1; grand total 31. There are 14 males and 17 females.

Description of Subspecies —This is a snake of medium colubrid pro-
portions, neither slim nor stout. The head is slightly distinct from the neck.
Viewed from above the snout is rounded. From the side, the top of the
head is slightly convex or flat; the jaw is deeply inset. The apex is moderately
pointed; it seems somewhat blunter than in the desert subspecies farther west.
The forward part of the snout slants upward, so that the tip is at the upper
end of the rostral. The eyes are somewhat protuberant; the diameter of the
orbit is equal to about three-fourths of the distance from the anterior edge of

KLAUBER—IHE GLossy SNAKE 337

the orbit to the nostril. The pupil appears round in most preserved specimens
but is slightly elliptical in life.

The longest specimen available, a female from El Paso, Texas, measured
903 mm., the smallest specimen 249 mm. The tail proportion (ratio to length
over-all) varies from 15.2 to 16.9 per cent in the males, and from 13.8 to
15.9 per cent in the females. The averages are respectively 16.1 and 14.8 per
cent.

The dorsal scales are smooth, rather narrow and with rounded ends.
The lower lateral rows are slightly enlarged. Single apical scale pits are faintly
evident on a few scales.

The scale rows at mid-body normally number 27, but 2 specimens out of
31 have 29, and one has 25. The ventrals in the males vary from 185 to
203, the interquartile range is 188.6 to 196.5, the mean 192.57+1.58, and
the coefficient of variation 3.06 per cent. The over-all range in the females
is from 193 to 211, the interquartile range 199.2 to 207.2, mean 203.18+1.44,
and the coefficient of variation 2.92 per cent. The anal is entire. The sub-
caudals, all divided, vary from 51 to 57 in the males, the interquartile range
being 52.2 to 54.3, the mean 53.21+.42, and the coefhcient of variation 2.96
per cent. In the females the over-all range is 45 to 51, the interquartile range
46.7 to 49.0, the mean 47.88+.41, and the coefficient of variation 3.53 per
cent. The coefficient of sexual divergence in the subcaudals is 10.5 per cent.
The terminal scale is conical, with a longitudinal crease.

The head scales are normal in size and arrangement. The rostral is
rather sharply curved, both over the top of the snout and below. It is
wider than high. The upper point separates the internasals for about half their
lengths; it also contacts the prenasals and the first supralabials. The inter-
nasals are longer than wide, curving downward to a point in front of the
prenasals. The prefrontals are wider than high and curve well down over the
canthus rostralis to a level with the center of the orbit. The supraoculars are
narrower anteriorly where they touch the prefrontals; posteriorly they contact
the parietals and upper postoculars. The frontal is widest anteriorly and
terminates posteriorly at a point between the parietals. The parietals are the
largest of the head scales. The dorsal scales which border them are somewhat
enlarged and irregular. The prenasal is considerably smaller than the postnasal.
The nostril is at the upper end of the suture which divides the nasals; it slants
upward anteriorly. The suture is incomplete above the nostril. The postnasal
usually contacts the first and second supralabials. The loreal is twice as long
as wide, and is highest forward, becoming pointed posteriorly where it indents
the preocular. The loreal borders the second and third supralabials. One
specimen out of 30 has divided loreals. The preoculars are usually single,
although paired in 2 counts out of 60; they are somewhat wider above than
below. The postoculars number 2 in all specimens; they are subequal in
size. The temporals are usually 2+3 or 2+4, but vary from 1+2 to 2+6 or
3+5. Those in the first series are attenuated, and slant upward posteriorly. The
supralabials are usually 8, but may number 7 (7 per cent) or 9 (8 per cent).
The fourth and fifth contact the eye; the next to the last is always much the

338 SAN Disco SocreETY OF NATURAL HIstory

largest. The infralabials vary from 11 to 14, the distribution being 11(1),
12(14), 13(34), 14(5). The first pair meet on the median line. Usually the
seventh is the largest of the series, although sometimes it is the sixth. The
mental is small and triangular, with a sharp posterior point. The anterior
genials are large and contact on the median line; the posterior are thinner and
shorter; they diverge posteriorly and are separated by 2 to 4 rows of gulars.

Philipi is a light-brown snake with conspicuous dark-brown blotches down
the back, and smaller secondary blotches along the sides, which may or may
not be obscured by a lateral suffusion of brown. The blotches are distinctly
more evenly-edged than is the case in noctivaga and the other subspecies farther
west.

The head is light-brown above, and buff on the sides, being lightest on
the supralabials. In some specimens there is a faint brown band, darker than the
ground color, on the posterior edges of the prefrontals; in others, particularly
adults, this has been lost. There are dark-brown spots on the supraoculars,
frontal, and, especially, the parietals. These spots are present in greater number
than in the westerly subspecies. There is a brown streak from the eye to the
commissure. The lower jaw is immaculate.

On the neck there is usually a pair of parallel dark marks separated by
a light streak; these are the first of the dorsal series.

The dorsal blotches occupy considerably more longitudinal space than
the interspaces which separate them. The body blotches vary in number from
52 to 72, interquartile range 58.6 to 65.4, mean 62.00+.88, coefficient of vari-
ation 8.1 per cent. The tail spots have an over-all range of 15 to 23, an inter-
quartile range of 17.7 to 20.8, mean 19.25+.47, coefficient of variation 12.0
per cent. The body blotches at mid-body usually cover 11 or 12 scale rows across
the body, but a few span 13 or 15 rows. The longitudinal extent of the blotches,
again measured at mid-body, varies from 11/2 to 242 scales, end to end, but most
specimens measure 2 scales. The interspaces are narrower; most of them are
but a single scale in length, but in a few specimens reach 14% or 112 scales. The
lateral brown suffusion is usually carried down to the third or fourth row
above the ventrals. The lowest lateral spots of the auxiliary series extend
considerably lower; in more than half of the specimens they touch the outer
edges of the ventrals, and in most of the others they reach the first row above.

The body blotches, although generally rectangular or elliptical, with the
major axis transverse, are sometimes irregular because the patterns of the
two sides do not exactly match. This results in a few diagonal blotches, and
others which are restricted to one side, while still others are Y-shaped, that is,
double on one side and single on the other.

In addition to the main dorsal blotches, there are, on each side, about
four alternating series of progressively smaller spots. The first of these series
comprises subcircular spots, alternating with the dorsal series, and is fairly
regular; those below are irregular, and often indefinite.

The main dorsal blotches are brown, with darker-brown or black edges. Be-
tween blotches the ground color is buff, this being evident middorsally and on

KLAUBER—IHE GLossy SNAKE 339

the lowest lateral rows. The ventrum is immaculate. But dorso-laterally the area
between blotches is suffused or punctated with brown. In some specimens the
lateral darkening tends to obscure the outer edges of the middorsal blotches
and the lateral series as well; in a few the lateral suffusion is so dense as to
blot out the side blotches completely. This side-darkening increases with age;
juveniles are usually quite clear between blotches.

Intrasubspecific Trends.—Since philipi occupies areas of considerable
ecologic diversity, it exhibits, as might be expected, some definite character
variations. ‘

The scale rows are slightly higher in the El Paso area, where specimens
with 29 rows are not uncommon. The ventrals are at a minimum in the
Winslow area, showing an increase eastward toward Albuquerque and thence
south to El Paso. The subcaudals and labials have a similar trend.

The blotches also increase in number from Winslow to El Paso. The
lateral blotches reach lower scale rows in northwestern New Mexico specimens
than in the snakes of the El Paso area.

Relationships with Other Subspecies —The possibility of the intergradation
of philipi with elegans and blanchardi has already been discussed under those
subspecies. All of these forms belong to the long-tailed group. Philipi differs
from elegans, first of all, in scale rows; the fact that the snakes of the El Paso
section occasionally have 29 rows suggests that the connection is probably
in trans-Pecos Texas.

There is no doubt that philipi intergrades with expolita, since they differ
only in the number of body blotches, and there is no ecological barrier between
them. But the actual relationship can only be determined when more material
becomes available, particularly between the most southerly philips (MVZ
24388, 1 mi e. of Samalayuca, Chihuahua), and the most northerly expolita
(USNM 46374 from Casas Grandes, Chihuahua). These two localities are
about 115 miles apart. The same lack of material from critical points prevents
a determination of the relationship of philipi with noctivaga. One belongs to
the long-tailed, the other to the short-tailed group. If serious consideration
be given to the separation of Arizona elegans into two species, it is probable
that the division should be made between these two, representing the westerly
representative of the long-tailed group, and the easterly of the short-tailed
forms. Yet intergradation is by no means impossible; it may involve the snakes
between the Tombstone and Tucson areas, or those between Canyon Diablo and
Congress Junction, by way of Oak Creek and Sedona. Arizona has been collected
at Oak Creek; unfortunately the specimens are not now available so that I
do not know which subspecies occurs there, or whether they show any inter-
mediacy between philipi and noctivaga. The gap in characters over either route
is by no means insurmountable, although the specimens from nearest to the
gaps do not exhibit pronounced intergradative tendencies.

Life History Notes——This subspecies is presumed to be quite nocturnal,
particularly during its season of greatest activity. Shaw and Engler collected the

340 SAN Disco SocreTy oF NATURAL History

type specimen at 11:10 p.m., temperature 72° F. Two specimens were collected
near El Paso in the late evening.

This subspecies reaches an altitude of at least 5500 ft. It is found in
semi-desert areas with sparse vegetation, such as those near Winslow and El
Paso, but in other sections inhabits ground with a heavy brush cover.

Range.—The Painted Desert, or Little Colorado River area of north-
eastern Arizona; central New Mexico, from Santa Fe, south and southwest, to
the Mexican Boundary; extreme southeastern Arizona (Cochise County); El
Paso County, Texas; and extreme northern Chihuahua.

Locality Records. —

' ARIZONA
CocoNniINo County: APACHE COUNTY:
Canyon Diablo at US 66 (= Two 1 mi. from Second Forest (Petrified
Guns ) Forest National Monument) (alt.
Oak Creek Canyon (22 mi. s. of 5500 ft.)
Flagstaff ) * CocHIsE County:
Navajo County: 25 mi. s. of Cochise (on US 666)
10 mi. e. of Winslow (type locality) Near Bowie
Shongopovit
New Mexico
SANTA FE County: GRANT County:
Arroyo Hondo (8 mi. sw. of Santa 26 mi. e. of Lordsburg (Hidalgo
Fe) County )
BERNALLILO COUNTY: Hipatco County:
Albuquerque (also 11 mi. n.) Lordsburg
Isleta Indian Reservation (ponds LuNA County:
on east side of Rio Grande) 29 mi. se. of Silver City
3 mi. w. of Rio Grande (near Dona ANA County:
Albuquerque ) Acacia
LINCOLN CouNTY: Las Cruces
Capitan Mountains} Mesilla Valley

SIERRA COUNTY:
Las Palomas
TEXAS

Ex Paso County:

El Paso (also 2 mi. e. and 2 mi. ne.)

Sand Hills ne. of El Paso

Near Tornillo

CHIHUAHUA, Mexico
1 mi. e. of Samalayuca

Arizona elegans expolita subsp. nov.
CHIHUAHUA GLossy SNAKE
1886. Khinechis elegans (part) Cope, Proc. Amer. Philos. Soc., vol. 23, p. 284.

1888. Arizona elegans (part) Bocourt, Miss. Sci. Mex., Rept., p. 676.
1894. Pituophis elegans (part) Gunther, Biol. Cent.-Amer, Rept., p.125.

* Specimen no longer available, subspecies doubtful.
+ Reported by Bogert, 1933, p. 219—probably this subspecies.

+ Location somewhat uncertain.

KLAUBER—IHE GLossy SNAKE 341

1894. Coluber arizonae (part) Boulenger, Cat. Snakes Brit. Mus., vol. 2, p. 66.

1924. Arizona elegans occidentalis (part) Blanchard, Occ. Papers Mus. Zool.,
Univ. Mich., no. 150, p. 1.

Type.—No. 46374 in the collection of the U. S. National Museum. Col-
lected May 27, 1899, by E. W. Nelson and E. A. Goldman at Casas Grandes,
Chihuahua, Mexico.

Diagnosis—A subspecies characterized by a low number of ventrals and
body blotches. It has fewer body blotches than any other subspecies except
elegans and pacata. From the former it may be distinguished by its fewer scale
rows (27 instead of 29 or 31 as in elegans) and fewer ventrals. From pacata
it may be segregated by its proportionately longer tail. While the numbers
of specimens of each subspecies are too few to set limits, it is believed that
expolita is one of the longest tailed of the subspecies, while pacata will prove to
be one of the shortest. Although expolita resembles philipi in having a low
number of ventrals, it has fewer blotches than philipi.

Description of the Type—The type specimen is a young male; length
over-all 510 mm.; tail length 78 mm.; tail proportion 15.3 per cent.

The head is slightly convex on top; it is little wider than the neck. The
snout is pointed and recurved below; the lower jaw is inset. The pupil is
nearly round.

The scales of the head are normal. The rostral is hollowed below and is wid-
er than high. It contacts the first supralabials, the anterior nasals, and the poster-
ior point separates the internasals for slightly more than half their lengths.
The internasals are longer than wide and curve down to points in front of
the nasals. The prefrontals are quadrangular in shape, with their post-lateral
points curved well down over the canthus rostralis. The frontal is pentagonal,
narrowing along the supraoculars and then more sharply between the parietals.
The supraoculars are narrowly in contact with the prefrontals; they widen poster-
iorly. The parietals are large and touch half of the posterior edges of the
upper postoculars. The postnasal is considerably larger than the anterior; the
nostril is at the upper end of the separating suture, which is not evident
above the orifice. Of the supralabials, the postnasal touches the first and second.
There is a single loreal on either side about twice as long as high, touching
the second and third supralabials. There is one preocular on each side, some-
what wider above than below. The postoculars are 2-2 and are subequal. The
temporals are 2+3, 2+4. The supralabials are 8-8, the fourth and fifth touching
the eye, and the seventh much the largest. The mental is small and sub-
triangular. The infralabials are 12-12, the first in contact behind the mental, and
the sixth much the largest. There are two pairs of genials, the anterior con-
siderably the longest, and medianly in contact; the posterior are slightly diver-
gent and are separated by two or three rows of gulars.

The dorsal scale rows are 27-27-17. The scales are smooth, rather narrow
and with rounded ends. The middorsal and lateral rows are the widest. The

342 SAN Dreco Society oF NATURAL HIstTory

ventral scales number 195; the anal plate is entire; the subcaudals are 49,
all divided.

The head is buff above and lighter on the sides. There is a brown bar across
the posterior edges of the prefrontals, and some brown mottling on the frontal
and parietals. There is a wide, dark streak from each eye back to the angle of
the jaw. The lower surface of the head is immaculate white.

The dorsal body pattern comprises a series of 45 rather irregular brown
blotches, with dark-brown edges, on a buff background. The irregularity is due to
the fact that the two sides do not match, so that the blotches are sometimes
diagonal and others Y-shaped. At mid-body the blotches are about 3 scales
long (end to end), and 13 scale rows wide, transversely. The interspaces are
about one scale long. The ground color is most evident on the three middorsal
scale rows, for laterally the scales between blotches are somewhat suffused with
brown. In addition to the main blotches there is an auxiliary series of dark-
edged brown spots on each side; these generally alternate with the main
series. Below the auxiliary series, many scales are irregularly streaked with
brown, although the lowest lateral rows are not suffused with brown. A few of
the brown spots are present on the outer edges of the ventrals, the ventrum
being otherwise white. On the neck there is a pair of dark blotches separated
by a narrow, middorsal light streak. About 9 scales behind the parietals, these

two blotches join to form the first of the dorsal series. There are 16 spots on
the tail.

Summary of Paratypes—Two paratypes of this subspecies are available:
USNM_ 14298 from Chihuahua* and AMNH 4344 from San Diego,
Chihuahua. They agree in all essential particulars with the type, showing the

same differences from the nearest relative, philipi.

The Chihuahua (City) specimen is a young male 572 mm. over-all with
a tail length of 94 mm., ratio 16.4 per cent. The scale rows are 25-27-19,
ventrals 194, anal entire, subcaudals 53, supralabials 8, infralabials 12 on the
left, indeterminate on the right, loreals and preoculars single, postoculars two,
temporals 2+3. The body blotches number 50 and the tail spots 20. The
blotches cover about 13 scale rows laterally, and 3 scales end to end longitud-
inally. The interspaces average 14 scales. The side suffusion reaches the

third lateral above the ventrals, while the lowest lateral spots touch the edges
of the ventrals.

The San Diego (Chihuahua) specimen is an adult female 785 mm. long,
with a 98 mm. tail, ratio 12.5 per cent. The scale rows are 29-27-20, ventrals.
207, anal entire, subcaudals 39, supralabials 9-8, infralabials 12-13, loreals 2-2
(there is a vertical suture in each), preoculars 1-1, postoculars 2-2, temporals
2+3, 2+4. The body blotches number 50 and the tail spots 12. The blotches
at mid-body cover 12 scale rows transversely, and 3 scales end to end longi-
tudinally. The interspaces measure 11% scales. The side suffusion reaches the

* Although catalogued from Chihuahua, it is known that Wilkinson, the collector,
worked in the vicinity of the City of Chihuahua, and Cope, who first reported on the speci-
men, considered that it came from the city or nearby (Cope, 1886, p. 282).

KLAUBER—THE GLossy SNAKE 343

fourth lateral row above the ventrals, while the lowest secondary spots comprise
a row on each side at the outer ends of the ventrals.

Both paratypes are probably somewhat faded. The Chihuahua City speci-
men has obsolete lateral spots obscured by a brown suffusion; in the San
Diego specimen the upper lateral series is subcircular and almost as clear as
the dorsal series.

In addition to the body blotches, of which expolita has fewer than philipi,
it may be observed that, in 5 counts out of 6, the sixth infralabial is the
largest of the series, whereas the seventh is usually the largest in philipi, as is
the case in the other subspecies. It will be of interest to observe whether this
difference will be maintained when additional specimens of expolita become
available.

Relationships with Other Subspecies ——The probable intergradation of ex-
polita with philipi has already been mentioned. That there may be intergradation
between expolita and elegans is also possible; it may be significant that the two
specimens from Coahuila, tentatively assigned to elegans, are low in both ventrals
and body blotches, thus showing a trend toward expolita, from which they differ
in having 31 scale rows, while the latter has 27.

Life History Notes.—One of the three available specimens of this sub-

species had eaten a small mammal.

Range and Locality Records.—This subspecies is only known from three
specimens from western and east-central Chihuahua, Mexico, the localities being

Casas Grandes, San Diego, and Chihuahua (city).

Arizona elegans noctivaga subsp. nov.

ARIZONA GLossy SNAKE

1875. Pityophis elegans (part) Yarrow. 100th Mer. Surv., vol. 5, p. 541.

1882. Pityophis catenifer var. deppei (part) Garman, Mem. Mus. Comp.
Zool., vol. 8, no. 3, p. 151.

1888. Arizona elegans (part) Bocourt, Miss. Sci. Mex., Rept., p. 676.

1894. Pituophis elegans (part) Gunther, Biol. Cent.-Amer., Rept., p. 125.

1894. Coluber arizonae (part) Boulenger, Cat. Snakes Brit. Mus., vol. 2, p. 66.

1900. Rhinechis elegans (part) Cope, Rept. U. S. Nat. Mus. for 1898, p- 862.

1924. Arizona elegans occidentalis (part) Blanchard, Occ. Papers Mus.
Zool., Univ. Mich., no. 150, p. 1.

T'ype—No. 34188 in the collection of LMK. Found crossing the road
8 miles northwest of Owlshead, Pinal County, Arizona (Owlshead is on
U. S. 80, 45 miles southeast of Florence), by Charles E. Shaw and L. M.
Klauber, at 9:20 p. m., May 31, 1941. Preserved June 16, 1941.

344 SAN Disco SocieETY OF NatTuRAL HIstTory

Diagnosis —Noctivaga is a subspecies belonging to the short-tailed group,
with body blotches slightly more extensive than the spaces which separate them,
and usually with 27 scale rows. It has fewer scale rows than elegans or blanchardi,
has more ventrals and a shorter tail than philipi or expolita, and more body
blotches than pacata. From eburnata and candida it may be distinguished by
the relative longitudinal extents of the dorsal blotches and interspaces; in
noctivaga the blotches exceed the spaces between, while the contrary is true
of the others. Although noctivaga is territorially separated from occidentalis by
the interposition of eburnata, they are superficially much alike and I know of
no single character which will invariably separate them. Noctivaga has fewer
ventrals on the average, fewer body blotches, and is usually lighter on the
sides, with less regular secondary spots, and with the lower lateral scale rows
freer both of spots and the brownish ground-color suffusion. It also lacks the
brown marks on the lower labials which usually characterize occidentalis,
although not those from the San Joaquin Valley.

Description of the Type——The type specimen is a young adult male; length
over-all 829 mm.; tail length 106 mm; tail proportion 12.8 per cent. These
measurements were made before the specimen had been hardened in pre-
servative.

The head is slightly convex on top when viewed from the side; from
above it is little wider than the neck. The snout is somewhat pointed and the
lower jaw inset. The pupil appears round in the preserved specimen.

The scales of the head are normal. The rostral is sharply recurved, wider
than high, and convex below. It contacts the first supralabials, the anterior
nasals (rather narrowly), and the posterior point separates the internasals for
half their lengths. The internasals are longer than wide and curve down in
front along the anterior nasals. The prefrontals are quadrangular in shape, with
their post-lateral points curved well down over the canthus rostralis. The
frontal is pentagonal, narrowing along the supraoculars and then coming sharply
to a point between the parietals. The supraoculars are narrowly in contact with
the prefrontals; they widen posteriorly, but do not jut over the eyes. The
parietals are large and touch somewhat over half the posterior edges of the
upper postoculars. The postnasal is considerably larger than the anterior; the
nostril is at the upper end of the separating suture, which is not complete
at the point where the nasals touch the internasals. Of the supralabials, the post-
nasal touches both the first and second. There is a single loreal on the right
side about twice as long as high, touching the second and third supralabials.
On the left the loreal is split vertically. The posterior end of each loreal is
somewhat pointed and indents the lower part of the preocular, there being one
of the latter on each side. The postoculars are 2-2, the lower being somewhat
the smaller. The temporals are 2+3. The supralabials are 8-8, the fourth and
fifth touching the eye, and the seventh much the largest. The mental is small
and subtriangular. The infralabials are 13-13, the first in contact behind the
mental, and the seventh much the largest. There are two pairs of genials, the
anterior considerably the longer and fully in contact medianly; the posterior

KLAUBER—IHE GLossy SNAKE 345

shorter and thinner, slightly divergent, and separated by two to four rows
of gulars.

The dorsal scale rows are 29-27-19, the highest number being found
only for a short distance on the neck. The scales are smooth, rather narrow and
with rounded ends; the lowest lateral row is the widest. There are single apical
scale pits; however, these are quite small and faint. The ventral scales number
213; the anal plate is entire; the subcaudals 49, all divided.

The head is buff, the supralabials being somewhat lighter than the top
of the head. There is a faint brown bar across the posterior edges of the
prefrontals; another along the suture between the parietals, and another at the
outer edge of each parietal. There is a dark streak from each eye back to the
angle of the jaw. The lower surface of the head is immaculate cream-colored.

The body pattern comprises a series of 66 rather irregular brown blotches
on a buff background. The first blotch on the neck is much longer than the
others and is divided longitudinally. The remaining blotches are wider than long
and are frequently irregular, since the two halves on the sides are not exactly
opposite. Often two on one side match with a single on the other, and several
are diagonal rather than transverse. At mid-body the blotches are about 2
scales long, end to end, and 11 scale rows wide, across the body. The inter-
spaces occupy about 1 scale. The blotches are irregularly dark-edged. The
ground color is clearly evident only on the three middorsal scale rows. Out-
side of these the dorsum is suffused with brown, thus somewhat obscuring the
outer ends of the main blotches, but the two lowest lateral scale rows are
clear. In addition to the main blotches there are two auxiliary series of brown
spots on each side. These are quite indefinite, being somewhat obscured by
the brownish lateral suffusion; nor do they synchronize perfectly with the
main series. The lower and smaller comprises merely a series of scattered black
dots, or black scale-edges, the lowest of which touch the second scale row above
the ventrals. The ventrum is immaculate cream-color. There are about 20
irregular spots on the tail, evident mostly as darkened scale-edges. The

posterior spots are divided into two lateral series by a light longitudinal stripe of
the ground color.

Paratype Series—Noctivaga is sufficiently uniform so that I shall not
separately summarize a paratype series, as was done with several of the other
new subspecies, but shall consider all the available specimens to be paratypes.*

*These paratypes ace AMNH 63640 41 mi. nw. of Safford, CHAS 10024 4 mi. n. of
Pima, CHAS 10025 3 mi. s. of Safford (Graham Co.); LMK 27177 13 mi. n. of Tucson,
LMK 27178 4 mi. n. of Sahuarita, LMK 29222 Tucson, LMK 32293 Martinez Hill, LMK
32521 11 mi. s. of Tucson, LMK 33714 2 mi. ne. of Tanque Verde Ranch, LMK 34018
14 mi. n. of Tucson, SDSNH 13724 Tucson, SDSNH 17949 11 mi. n. of Tucson,
SDSNH 17950-1 4% mi. n. of Tucson, SDSNH 17952 2 mi. n. of Tucson, KU 14137 16
mi. s. of Tucson (near Sahuarita), AMNH 3807 Tucson, AMNH 26015 Pima Co.,
AMNH 60561 5 mi. e. of Tucson, AMNH 60669 Rillito, CHAS 11111 and 11228
Marana (Pima Co.); USNM 8002 Camp Grant, LMK 34438 7 mi. se. of Globe, CHAS
9521 6 mi. ne. of Globe (Gila Co.); LMK 21492-3 Picacho, LMK 27180 Florence, LMK
32323 8 mi. w. of Casa Grande, LMK 34104 3 mi. w. of Superior, LMK 34332 Oracle

346 SAN Disco Society OF NATURAL HIstTory

Material—The material for study, made up of the type and paratypes,
is distributed as follows, all being from Arizona: Graham County 3, Pima
County 20, Pinal County 19, Gila County 3, Yavapai County 4, Maricopa
County 19, Mohave County 2, Yuma County 1; total 71. Other specimens from
Yuma County are either noctivaga-eburnata intergrades, or eburnata, and are
included under the latter subspecies. Of the specimens of noctivaga there are
49 males and 16 females, the other 6 being heads or otherwise indeterminate.

Description of Subspecies —This is a snake of ordinary colubrid propor-
tions, neither heavy-bodied nor slim. The head is only slightly distinct from the
neck. Viewed from above the head is wedge-shaped but with a rounded
snout. From the side the top of the head is slightly convex. The snout is
moderately pointed; the underside slants upward, so that the apex is toward the
upper end of the rostral. The eyes are rather protuberant; the diameter of the
orbit is about equal to 70 per cent of the distance from the anterior edge of
the orbit to the nostril. The pupil appears round in most preserved specimens
but is slightly elliptical in life.

The longest available specimen is a male from Phoenix, Arizona, measur-
ing 1051 mm. The smallest specimen measured 272 mm. The tail proportion
(ratio to length over-all) varies from 12.2 to 15.2 per cent in the males,

and from 11.6 to 13.3 per cent in the females. The averages are respectively
13.7 and 12.7 per cent.

The body is covered with smooth scales, rather narrow and with slightly
rounded ends. The lower lateral rows are moderately enlarged. Single apical
scale pits are faintly evident on a few scales.

The scale rows at mid-body are usually 27, but 13 per cent of the
specimens have 29, and one specimen has only 25. The ventrals in the males
vary from 204 to 214, interquartile range 207.0 to 210.6, mean 208.80+.39, co-
efficient of variation 1.27 per cent; the over-all range in the females is from 211
to 224, interquartile range 216.7 to 221.7, mean 219.19+.69, coefficient of varia-
tion 1.72 per cent. The anal is entire. The subcaudals, all divided, vary from 46 to
52 in the males, the interquartile range being 48.1 to 50.1, mean 49.09.21,
coefhicient of variation 2.96 per cent. In the females the over-all range is
43 to 48, interquartile range 44.3 to 46.5, mean 45.40+.42, coefficient of

Junction, CAS 80685 Florence Junction, CAS 80699 4 mi. n. of Ajo, CHAS 9404 10 mi.
nw. of Sacaton, CHAS 9920 5 mi. w. of Superior, CHAS 10290 6 mi. w. of Florence,
CHAS 10291 5 mi. w. of Florence, CHAS 10292 3 mi. s. of Florence Junction, CHAS
10293 1 mi. s. of Florence Junction, CHAS 10294 5 mi. w. of Florence, CHAS 10295 4
mi. s. of Florence Junction, CHAS 11112 25 mi. se. of Florence, CHAS 11113 10 mi.
se. of Chandler (Pinal Co.); LMK 23925-6 Sentinel, LMK 25829 Mesa, LMK 26010
Mesa, LMK 26913 5 mi. s. of Wickenburg, LMK 26943 Cactus Gardens, LMK 32781 Gila
Bend, LMK 34331 Stanwix, AMNH 8352 and 9021 Phoenix, USNM 44259 Phoenix,
CHAS 3082 1 mi. n. of Wickenburg, CHAS 3386 2 mi. n. of Wickenburg, CHAS 3661
4¥2 mi. s. of Wittman, CHAS 12246 Phoenix, SDSNH 15835 Sentinel (Maricopa Co.);
SDSNH 17623 1 mi. s. of Congress Junction, CHAS 2083 4 mi. n. of Wickenburg, CHAS
3084 10 mi n. of Wickenburg, CHAS 3705 6 mi. n. of Congress Junction (Yavapai Co.);
USNM 37112 mesa n. of Ft. Mohave, MVZ Boulder Lake (Mohave Co.); LMK 34526 5

mi. e. of Salome (Yuma Co.).

KLAUBER—IHE GLOssy SNAKE 347

variation 3.60 per cent. The coefficient of sexual divergence in the subcaudals
is 7.8 per cent. The terminal scale is somewhat elongated, with a lateral crease.

The head scales do not diverge from the colubrid normal in size and
arrangement. The rostral is rather sharply curved, both over the top of the
snout and below, where it is hollowed longitudinally. It is about 20 per cent
wider than high. The upper point separates the internasals for about half their
lengths; in addition it contacts the prenasals and the first supralabials. The
internasals are longer than wide and curve downward to a point in front of
the prenasals. The prefrontals are wider than high and curve down over the
canthus rostralis to a level with the center of the eye. The supraoculars are
narrow and elongated; they barely touch the prefrontals and widen posteriorly,
where they contact the parietals and upper postoculars. The frontal narrows
posteriorly; it separates the parietals for a short distance. The parietals are
the largest of the head scales; the dorsal scales which border them are somewhat
enlarged and irregular. The prenasal is much smaller than the postnasal. The
nostril is at the upper end of the suture which divides the nasals; it slants
forward above. The nasals are not separated above the nostril, being connected
by a narrow ridge. The postnasal usually contacts the first and second supra-
labials. The loreal is twice as long as wide, and is highest forward, becoming
pointed posteriorly, especially where it touches the preocular. The loreal
contacts the second and third supralabials. Divided loreals are more frequent
in this subspecies than any other, being present in 10 counts out of 132. The
preoculars are usually single, although paired in 9 counts out of 132. The
preoculars are somewhat wider above than below. The postoculars ordinarily
number 2, and are subequal in size; there are 3 in 2 counts out of 132. The
temporals are usually 2+3 or 214, but vary from 1+2 to 3+6. Those in
the first row are usually elongated, slanting upward posteriorly. The supralabials
are most often 8, but number 9 in 8 per cent of the counts. The fourth and fifth
contact the eye; the next to the last is always much the largest. The infralabials
vary from 11 to 14, the distribution being 11(1), 12(20), 13 (93), 14(10). The
first pair meet on the median line. Usually the seventh is the largest of the
series, although it may be the eighth if there are 14. The mental is small and
triangular. The anterior genials are large and contact on the median line; the
posterior are both slimmer and shorter, and, diverging posteriorly, are separated

by from 2 to 5 rows of gulars.

Noctivaga is a brown snake with dark-brown blotches down the back, and
additional secondary blotches on the sides, which, however, are somewhat
obscured by a lateral suffusion of brown. In general, noctivaga falls between the
usually darker occidentalis on the one hand, and eburnata and candida, which
are lighter, on the other.

The head is brownish on top, and somewhat lighter on the sides, especially
along the supralabials. There is a brown band, darker than the ground color,
on the posterior edges of the prefrontals; also dark-brown spots on the
supraoculars, frontal, and parietals, often concentrated along the sutures
which separate these plates. There is a brown streak passing backward from

348 San Disco Society oF NATURAL History

the orbit to the angle of the mouth; also, there is often a brown mark below
the eye and another at the anterior end of the loreal. The lower jaw is usually
immaculate, although the posterior infralabials are sometimes marked by a con-
tinuation of the postocular dark streak. All the head marks are much clearer
in the young than the adults. .

On the neck there is usually a pair of parallel dark marks separated by
a light streak. These, the first of the dorsal series, may be joined together
anteriorly, posteriorly, or both.

The dorsal blotches occupy somewhat more longitudinal space than the
interspaces which separate them. The interspaces equal the blotch lengths in
only 3 specimens out of 61; in all the others the blotches are longer. This
is sharply different from eburnata and candida, in which the contrary is true
in nearly every specimen. The body blotches vary in number from 52 to 67,
interquartile range 56.2 to 60.8, mean 58.48.42, coefficient of variation 5.9
per cent. The tail spots have an over-all range of 15 to 23, interquartile range
17.1 to 19.6, mean 18.32.25, coefficient of variation 10.3 per cent. The
body blotches at mid-body usually (70 per cent) cover 11 scale rows across
the body. but a few range from 8 to 13 rows. The longitudinal extent of the
blotches, again measured at mid-body, varies from 1% to 2% scales, end to
end, but most specimens cover 2 scales. The interspaces are somewhat narrower;
most of them are 11/ scales in length, but the variation is from 1 to 2 scales.
The lateral brown suffusion is usually carried down to the second to fourth
row above the ventrals. The lateral spots of the auxiliary series usually are not
present below the second scale row above the ventrals.

The body blotches, although ordinarily elliptical, with the major axis trans-
verse. are often quite irregular because the blotches on one side do not fall
exactly opposite those on the other. This results in some diagonal blotches, and
others which are restricted to one side, while still others are Y-shaped.

In addition to the main dorsal blotches, there are, on each side, two
or more alternating series of progressively smaller spots. These are more
perfect anteriorly. The first of these series comprises subcircular, or vertically
elongated, spots, alternating with the dorsal series, and is rather regular;
those below are hardly more than scattered darkened scale-edges.

The main dorsal blotches are brown, with dark-brown or black edges. Be-
tween blotches the ground color is buff, but this is evident only on 3 to 5
middorsal scale rows, and on the lowest lateral rows, for dorso-laterally the
area between blotches is heavily suffused or punctated with brown. This darken-
ing as accentuated on the scale centers; the scale edges are usually light, giving
a net-work effect characteristic of Arizona. The lateral darkening tends to
obscure the outer edges of the middorsal blotches and the lateral series as
well; in some specimens the lateral suffusion is so dense as almost to blot
out the side blotches completely. This side-darkening increases with age;
juveniles are usually quite clear between blotches. The ventrum is immaculate
cream.

The following colors were noted on a live specimen from Mesa, Maricopa
County, Arizona, using Ridgway’s Color Standards, 1912, for comparisons:

KLAUBER—IHE GLossy SNAKE 349

Centers of blotches, middorsally, Sayal Brown; centers of blotches, laterally
Snuff Brown; edges of blotches, Black; center of scales in lateral interspaces,
Verona Brown; middorsal interspaces, Light Pinkish Cinnamon; ventral sur-

face, Pale Olive Buff.

Intrasubspecific Trends—Noctivaga seems to be rather consistent through-
out its range, and it will take larger series than are yet available to determine
the presence of clines in the characters. There appears to be a slight reduction
in the average number of ventrals from west to east, and a small increase in body

and tail spots.

Relationships with Other Subspecies—The possible intergradation of
noctivaga and philipi has already been discussed. In Yuma County noctivaga
undoubtedly intergrades with eburnata. The snakes of the Yuma—Somerton-—
Gadsden area are probably pure eburnata, the only territory east of the Colo-
rado River inhabited by that subspecies, so far as is now known. The area
of intergradation between noctivaga lies to the east of the Gila Mountians,
from Wellton to the vicinity of Sentinel in southwestern Maricopa County.
Here there is a gradual change from the one form to the other.

Along the Colorado River north of Ehrenberg, and as far north as
Boulder Dam, a lack of specimens prevents any exact knowledge as to the
boundary between the subspecies. The few specimens that are available in-
dicate the river to be the boundary, for those on the east side are noctivaga and
those on the west eburnata.

Life History Notes——WNoctivaga, as the name implies, is nocturnal; this
is shown by the following evening times of collection: 8:30, 9:20, 9:33, 9:40,
9:50 (2), 10:10, 10:37, 10:45, 11:00, 11:25, 11:30, 11:40. I have no daytime
collection records. The following air temperatures in degrees F. were noted when
specimens were found alive on the road: 70, 72(2), 78(2), 85(2), 86, 90.
These records were secured by using the scheme of collecting which has
proved so successful in the deserts of the southwest—by driving on paved
roads at night. I think the best success has been attained where the roads are
bordered by creosote bushes—for example, on the mesa above Tucson on the
Florence road, about Florence Junction, and near Wickenburg. Other sur-
roundings where specimens have been found were as follows: Fields (culti-
vated), grass, mesquite, brushy desert, rocky desert.

Ortenburger and Ortenburger (1926, p. 116) found a specimen at 10
p- m. near a temporary mud puddle, through which it attempted to escape,
swimming readily as if accustomed to water.

While noctivaga appears almost white in the glare of auto headlights
against the black background of a paved road, the type specimen was collected
on a dirt road and appeared darker than its background.

As in the other subspecies, the food comprises small mammals and
reptiles; mammal hair was found in one specimen, and lizard scales in another.
A photograph has been seen of a specimen, probably of this subspecies,
which had died in attempting to swallow a horned toad.

350 SAN Dreco Soctety oF NATURAL HIstTory

This subspecies reaches an altitude somewhat in excess of 3500 ft. at
Globe, and elsewhere along the mountain fringes from Kingman southeast

to Safford.

Range.—Arizona west and south of the central mountains, but excluding
the Yuma Mesa and Yuma Desert; also excluding Cochise County.

Locality Records.—

ARIZONA

Monave County:
East shore of Lake Mead (= Boulder

Lake)
Mesa s. of Fort Mohave
Louise

10 mi. w. of Oatman
Yuma County:
Brenda
5 mi. e. of Salome
Maricopa CouNTY:
Stanwix*
Sentinel *
Gila Bend (also 10 mi. e.)
Wickenburg (also 1 and 2 mi. n.,
A imi. e:,. 5) mi. ss and) 3.) 6;
and 10 mi. w.)
4 mi. s. of Wittman
Cactus Garden
Phoenix
Mesa
YAVAPAI COUNTY:
Congress Junction (also 6 mi. n.
and | and 5 mi. s.)
4, 6, and 10 mi. n. of Wickenburg
(Maricopa County )
Gita County:
6 mi. ne., and 6 and 7 mi. se. of
Globe
GRAHAM COUNTY:
Camp Grant
41 mi. nw. of Safford (near Calva)

4 mi. n. of Pima

Thatcher

3 mi. s. of Safford

PINAL COUNTY:

Superior (also 3 and 5 mi. w.)

Florence Junction (also 1, 3, and
4 mi. s.)

Florence (also 5 and 6 mi. w., and
25 mi. se.)

10 mi. se. of Chandler (Maricopa
County )

10 mi. nw. of Sacaton

Casa Grande (also 8 mi. n. and 12
and 6 mi. w.)

Picacho

8 mi. nw. of Owlshead (type

locality )
Oracle Junction
Pima County:

4 mi. n. of Ajo

Marana

Rillito

10 and 11 mi. s. of Oracle Junction
(Pinal County)

Tucson (also 2, 4, 5, 8, 11, and
139mi- ns > imi es anda
and 16 mi. s.)

Martinez Hill

2 mi. ne. of Tanque Verde Ranch

Rita

Sahuarita (also 4 mi. n.)

Arizona elegans eburnata subsp. nov.

Desert GLossy SNAKE
Plate 8, fig. 1

1892. Rhinechis elegans (part) Cope, Proc. U. S. Nat. Mus., vol. 14, no. 882,

p. 638

1897. Arizona elegans (part) Van Denburgh, Occ. Papers Cal. Acad. Sci.,

no. 5, p. 193.

1924 Arizona elegans occidentalis (part) Blanchard, Occ. Papers Mus. Zool.,

Univ. Mich., no. 150, p. 1.

* Noctivaga-eburnata intergrades.

KLAUBER—TIHE GLossy SNAKE Bil

Type—No. 33094 in the collection of LMK. Taken at Bensons Dry
Lake, in eastern San Diego County, California (3 miles west of the Imperial
County Line on highway Cal. 78), by James Deuel. Preserved June 5, 1940.

Diagnosis.—A desert subspecies characterized by small and narrow blotches,
light color, and high ventral scale counts. It differs from all others except
candida in having blotches markedly shorter than the interspaces which sepa-
rate them. Other differences are as follows: From elegans and blanchardi it
differs in having fewer scale rows and subcaudals; from occidentalis in its
lighter color, narrower blotches, and immaculate infralabials and lower |ateral
scale rows; from noctivaga by its narrower blotches; from philipi and expolita
by its greater number of ventrals and relatively shorter tail; and from pacata
by its greater number of body blotches. Eburnata can be segregated from candida
since it has only one preocular, while the Mojave Desert form generally has two;
also, the dorsal blotches of the latter usually engage 9 scale rows compared to 7
in eburnata.

Description of the Type——The type specimen is a young male; length
over-all 671 mm.; tail length 94 mm.; tail proportion 14.0 per cent. These
measurements were made before shrinkage in preservative.

This is a snake of moderate body shape, neither racer-like nor particularly
stout. The head is rather narrow and little distinct from the neck. The snout
is pointed, sloping backward below, with the lower jaw inset. The pupil
is almost round although slightly narrower than high.

The scales of the head are normal. They comprise a sharply recurved
rostral, twice as wide as high, and strongly indented below. The posterior point
separates the internasals for about half their lengths. The rostral contacts the
prenasals as well as the first supralabials. The internasals are longer than
wide, with anterior points curving down over the canthus. The prefrontals are
quadrangular in shape, wider than long, and with their postlateral points
curved well down over the canthus. The frontal is hexagonal, longer than
wide, narrowing somewhat along the supraoculars, and then coming quickly
to a point between the parietals. The supraoculars are narrowly in contact with
the prefrontals; they widen somewhat posteriorly. The parietals are large and
touch over half the posterior edges of the upper postoculars. Medianly
the parietals are edged by normal dorsal scales. The suture between the nasals
is not complete on either side above the nostril, which is near the upper
end of the suture. The prenasal is much smaller than the postnasal; it is
triangular in shape. The postnasal touches the first and second supralabials.
There is a single quadrangular loreal on either side about twice as long as high,
touching the second and third supralabials. It has a lower posterior point
which indents the lower half of the preocular. There are one preocular and
two postoculars, the latter being about equal in size. The temporals are 2+3.
The supralabials are 8-8, the fourth and fifth touching the eye, and the seventh
largest. The mental is small and triangular. The infralabials are 13-13; the
first on each side are in contact behind the mental; the seventh is much the

352 SAN Disco SociETY OF NaturAL History

largest. There are two pairs of genials, the anterior considerably the longest,
and medianly in contact; the posterior are separated by several gulars.

The dorsal scale rows are 27-27-18. The scales are smooth, rather narrow
and with slightly rounded ends. Single apical scale pits are faintly evident on
some scales, particularly caudad. The ventral scutes number 224; the anal
plate is entire; there are 54 subcaudals, all divided.

The head is buff above, with a dark line across the posterior edges of the
prefrontals; there are dark spots on the posterior edge of the frontal, and along
the suture separating the parietals. The sides of the head are cream-colored;
there is a dark streak from the eye back to the last supralabial, and also below
the eye in the suture between the fourth and fifth supralabials. The lower
surface of the head is cream-colored and unspotted.

The body pattern comprises about 76 irregular, and rather ill-defined
light-brown blotches, on a cream-colored background. The blotches are ac-
centuated by dark posterior scale-edges. They are much wider (across the
body) than long. At mid-body they are about 1 scale long, and 7 scale rows
wide. The interspaces entail about 2/2 scales (end to end). On the neck there
is a pair of parallel blotches about 7 scales long. The irregularity of the
dorsal blotches is such that in some places the blotches are diagonal, while in
others there are separate spots on each side. The ground color is most clearly
evident on the three middorsal scale rows; the next 9 rows on either side are
centrally punctated with buff, causing a suffusion which tends to obscure the
lateral edges of the main blotches. Some of this suffusion is faintly evident down
to the third lateral row above the ventrals. Scattered through this suffused
lateral area there are irregular black spots, none of which is large enough to
cover an entire scale. These are the residual posterior edges of what, in the
more easterly subspecies, are secondary lateral blotches. The lowest of these spots
mark the third scale row above the ventrals, the two lowest rows on either side
being immaculate, as is the ventrum. This is cream-colored in preserved speci-
mens, but white in life. The tail spots are highly irregular, and absent dorsally.
There are about 22, but they cannot be counted with certainty.

Summary of Paratypes—There are available from the type locality, or its
immediate vicinity in the Borego area of eastern San Diego County, 40 speci-
mens of this subspecies. Since these exhibit certain differences from the subspecies
as a whole, which shows some variation throughout its range, it is deemed
desirable to designate these as paratypes and to summarize their characteristics
separately.*

* These specimens and their locations are as follows: LMK 4454, 23854, 23914-5,
25437-8, 26914, 26939-42, 27308, 27383, 27405, 29300, 33094-5, SDSNH 17026, and
CNHM 26036 all topotypes from Bensons Dry Lake; LMK 23024, 23774-6, 23852-3,
26814, 29301, 29487, 32035 from The Narrows, LMK 4862 Beatty Ranch (Borego Valley),
LMK 5136, 21108, 21121 Borego Valley, LMK 23773 5 mi. e. of The Narrows, LMK
26732 2 mi. s. of Borego P. O., LMK 27331 3 mi. w. of Bensons Dry Lake, LMK 26056-7
Borego Palm Canyon, UCLA 548 San Felipe Wash (6 mi. w. of Imperial Co. line), and
UCLA 552 1 mi. w. of Imperial Co. line (2 mi. e. of Bensons).

KLAUBER—THE GLossy SNAKE 353

There are 25 males (including the type) and 16 females. Twenty-seven
scale rows comprise the mode, 27 specimens having this number; one has
25, 3 have 28 and 10 have 29. The ventrals in the males range from 218 to 227,
interquartile range 221.0 to 225.2, mean 223.08+.63, coefficient of variation 1.41
per cent; in the females the over-all range is 226 to 241, interquartile range
232.1 to 236.9, mean 234.50+.88, coefficient of variation 1.50 per cent. The
subcaudal statistics are, males, range 48 to 54, interquartile range 50.0 to 52.0,
mean 51.00+.30, coeflicient of variation 2.83 per cent; females, range 44 to
49, interquartile range 45.5 to 47.4, mean 46.43.37, coefficient of variation
3.01 per cent. The supralabials are distributed thus: 7(1), 8(74), 9(7); and
the infralabials 11(1), 12(14), 13 (63), 14(4). The loreals are single in all
specimens; there are two preoculars in 4 counts out of 82 (4.9 per cent) the
rest having single preoculars; all specimens have two postoculars.

The body blotches vary from 58 to 82, interquartile range 64.4 to 72.4,
mean 68.41.93, coefficient of variation 8.66 per cent; the statistics of tail
spots are, over-all range 16 to 26, interquartile range 18.1 to 21.7, mean
19.90.43, coefficient of variation 13.4 per cent.

The paratypic series well illustrates the outstanding pattern characteristics
of this subspecies—the light color, narrow and short dorsal blotches, wide inter-
spaces and unmarked lower lateral scale rows. The variations in width of the
dorsal blotches at mid-body, measured by the number of scale rows covered,
is as follows: 6(5), 7(26), 8(7), 9(3). The longitudinal extent of the dorsal
blotches is usually a single scale (21 out of 42 specimens) although it may
reach 144 or 1/2, and in two specimens attains 2 scales (end to end). The
interspaces usually measure 2 scales (end to end), but vary from 134 to 3.
The brownish side suffusion generally terminates on the second or third scale
row above the ventrals; in no case does it reach the lowest row. Also, the
obsolescent auxiliary lateral blotches, represented in this subspecies by darkened
scale tips, usually end on the third row above the ventrals, although the lowest
row reached varies from the second to the fifth.

The tail proportion varies between 12.2 and 14.5 per cent in the males,
and from 11.0 to 13.3 per cent in the females. Most adult males fall between
12.8 and 14.2, and the females from 12.2 to 12.7 per cent.

Material_—In addition to the type and paratypes from eastern San Diego
County, totaling 41 specimens, the following are available: California, San
Diego County 44 (including some occidentalis intergrades) , Imperial County 13,
Riverside County 51, San Bernardino County (east of the Mojave Daan.
together with the Twentynine Palms-Morongo section, several of which are
candida intergrades) 14; Nevada, Clark County 8, Nye County 1; Utah, Wash-
ington County 4; eine) Yuma County 13 (induding some noctivaga inter-
grades); Sonora 2; grand total 191. There are 125 males and 62 females, the
others being hones or indeterminate.

Description of the Subspecies—This is a snake of ordinary colubrid pro-
portions. The head is slightly distinct from the neck. Viewed from above the

354 SAN Disco Society oF NATURAL History

head is wedge-shaped with a rounded snout. From the side the top of the head
is slightly convex or flat; the jaw is deeply inset. The apex is moderately
pointed; the forward part of the snout slants upward, the tip being toward the
upper end of the rostral. The eyes are somewhat protuberant; the diameter
of the orbit is about 70 per cent of the distance from the anterior edge of the
orbit to the nostril. The pupil appears round in most preserved specimens but
is somewhat vertically elliptical in life.

The longest specimen I have had, a female from Mecca, Riverside County,
measured 1147 mm. The smallest specimen measured 213 mm. The tail pro-
portion (ratio to length over-all) varies from 12.0 to 15.5 per cent in the
males, and from 11.0 to 14.1 per cent in the females. The averages are respec-
tively 13.7 and 12.6 per cent. The young specimens have slightly shorter tails,
proportionately, than the adults.

In studies of tail length (Klauber, 1943, p. 5) I gave the following
equations for Arizona (Table 9):

Eburnata* males Ti — Onl S715 les
females J = 0.131L — 5.53

Candidat males .T = 0.143L — 2.63
females T = 0.133L — 3.40

where T is the tail length and L the length over-all, both in millimeters.

It will be observed that the equation for the ontogenetic curve of male
eburnata is inconsistent with the others, in that the proportionate tail length is
greater in the juveniles than the adults, whereas in the others, the constant term
being negative, the contrary is true. After noting trends in other subspecies in
the course of the present study, I am of the opinion that the particular sample
available to me produced an erroneous result, and that the regression line in
male eburnata probably falls close to

i—O0N4E — 2.0

Measurements were made of the head lengths of a number of specimens
of eburnata, and it was found, as is usually the case, that the proportionate
size of the head decreases slightly with age. A straight line fairly well defines
the relationship with the length over-all, the equation being

H = 0.022L + 6.5
where H is the head length and L the length over-all, both expressed in milli-

meters.

The hemipenis is single, although somewhat expanded distally. The inner
half is almost covered with tiny points, although less so proximally and on the
side opposite the sulcus. At about the middle of the shaft there is a sudden
transition to spines which are both much longer and heavier. These completely
encircle the shaft, except that they remain fine and delicate along the sulcus.
They are densely set and are not in regular rows; about 20 comprise a complete

* Listed as Arizona e. occidentalis I.
+ Listed as Arizona e. occidentalis II.

KLAUBER—THE GLossy SNAKE 355

ring. Distally there is again a diminution in size, so that the smallest spines
at the outer end are about equal in size to those on the proximal area of the
shaft. At the outer end the organ widens and is truncated irregularly, the end
comprising a hollowed surface, at a slight angle with the main axis. This outer
end is covered with reticulated flounces. The sulcus turns over the highest
part of the outer end and then terminates in a small bare depression. The
transition from points to flounces is quite sudden. The sulcus takes a slightly
diagonal course up the shaft. Specimens of occidentalis show no differences
from eburnata.

Pituophis catenifer exhibits the following differences in hemipenes from
Arizona elegans eburnata: In Pituophis the proximal half of the shaft is quite
smooth, instead of being almost covered with fine points as in Arizona; at the
outer end in Pituophis there is a greater tendency toward bifurcation, and the
smooth area within the cleft is relatively more extensive than in Arizona.

The body is covered with smooth scales, rather narrow and with rounded
ends. The lower lateral rows are somewhat enlarged. Single apical scale pits
are present, but they are so faint as to be located with difficulty.

The scale rows at mid-body are usually 27, but are occasionally 29 (15
per cent), and in one specimen 25. The ventrals in the males vary from 208
to 228, interquartile range 216.5 to 222.6, mean 219.54+.41, coefficient of
variation 2.07 per cent; the over-all range in the females is from 220 to 241,
interquartile range 229.0 to 233.4, mean 231.21+.62, coefficient of variation
2.05 per cent. The anal is entire. The subcaudals, all divided, vary from 47
to 59 in the males, the interquartile range being 50.3 to 53.4, mean 51.82+.21,
coefficient of variation 4.36 per cent. In the females the over-all range is 43
to 54, interquartile range 46.1 to 49.4, mean 47.76.33, coefficient of variation
5.18 per cent. The coefficient of sexual divergence in the subcaudals is 8.2
per cent. The terminal scale is conical, with a lateral crease; it is not difficult
to tell whether the tail is complete. Incomplete tails are not particularly frequent
as compared to Pituophis, for example.

The head scales are normal in size and arrangement. The rostral is rather
sharply curved, both over the top of the snout and below. It is considerably
wider than high. The upper point separates the internasals for half their lengths;
in addition it contacts the prenasals and the first supralabials. The internasals
are longer than wide; they curve down in front of the prenasals. The prefrontals
are wider than high; they curve down to a broad contact with the loreals. The
supraoculars narrowly contact the prefrontals; in addition they touch the pre-
oculars, postoculars, frontal, and parietals. The frontal is widest anteriorly and
terminates posteriorly at a point which does not deeply indent the suture
between the parietals. The parietals are the largest of the head scales. They
are subtriangular in shape, decreasing in size posteriorly. The dorsal scales
which border them are irregularly enlarged. The prenasal is smaller than the
postnasal. The nostril is at the upper end of the suture which divides the
nasals; it slants forward and upward. The suture is not usually complete above
the nostril, being blocked by a ridge which connects the two nasals. The post-
nasal usually contacts the first and second supralabials. The loreal is about

356 SAN Dreco Socrety oF NATURAL History

twice as long as wide, and is highest anteriorly, being pointed sub-posteriorly
where it indents the preocular. The loreal contacts the second and third
supralabials. One specimen has a divided loreal on one side, and another has
no loreal on one side, these being the only 2 aberrants out of 382 counts. The
preoculars are usually single, although paired in 11 counts out of 376 (excluding
possible candida intergrades). The preocular is wider at the prefrontal than the
loreal contact. The postoculars ordinarily number 2, and are about equal in
size; there are 3 in 10 counts out of 382. The temporals are usually 2+3 or
2+4, but vary from 1 to 4 in the first row and 2 to 5 in the second. Those
contacting the postoculars are elongated, and slant upward posteriorly. The
supralabials are usually 8, but occasionally number 7 (3 per cent) or 9 (7 per
cent). The fourth and fifth contact the eye; the next to the last is always
much the largest. The infralabials vary from 11 to 15, with a mode of 13, the
distribution being 11(9), 12(84), 13(246), 14(31), 15(1). The first pair
meet on the median line. Usually the seventh is the largest of the series,
although occasionally it is the sixth. The mental is small and triangular. The
anterior genials are large and contact on the median line, while the posterior
are both slimmer and shorter, and, diverging posteriorly, are separated by from
2 to 4 rows of gulars.

Eburnata is the lightest of all subspecies of Arizona; it is a buff or cream-
colored snake with brownish transverse marks on the back, and additional spots
on the sides, which may be rather obscured by a lateral suffusion of light-brown
punctations.

The head is light-brown or clay-colored above, the sides being somewhat
lighter. There is often a brown band on the posterior edges of the prefrontals,
but in many specimens it is quite faint. There are usually a few brown spots
on the supraoculars, frontal, and parietals, often concentrated along the sutures
which separate these plates. There is a brown streak passing backward from
the orbit to the angle of the mouth, but in many specimens the posterior part
may be absent, or represented by a spot on the last supralabial. Sometimes the
suture between the supralabials immediately below the eye is darkened with
brown. The lower jaw is immaculate.

On the neck there is usually a pair of parallel dark marks separated by a
light streak. These, the precursors of the dorsal series, are rarely joined together
anteriorly or posteriorly.

The dorsal blotches occupy much less longitudinal space than the inter-
spaces which separate them. The body blotches vary in number from 55 to 83,
interquartile range 64.3 to 72.8, mean 68.54.46, coefficient of variation 9.2
per cent. The tail spots have an over-all range of 15 to 29, interquartile range
18.6 to 22.1, mean 20.35+.20, coefficient of variation 12.9 per cent. The body
blotches at mid-body cover from 6 to 10 scale rows across the body, but in
most specimens they span 7 to 9 rows, 7 being in the majority. The longitudinal
extent of the blotches, again measured at mid-body, varies from 1 to 2 scales,
end to end, but most specimens extend from 1 to 1% scales. The interspaces
are always wider (except in occidentalis or noctivaga intergrades) ; most of them
are from 11 to 2 scales in length, but in a few specimens reach 22 or even 3

KLAUBER-—I HE GLossy SNAKE 357)

scales. A lateral brown suffusion is present, usually down to the second to
fourth row above the ventrals. The lowest lateral spots of the auxiliary series
seldom touch any row below the second, and in many cases the third or fourth
is the lowest marked. The relative freedom from marks on the lower lateral
scale rows is characteristic of this subspecies.

The body blotches, although generally comprising short bands, with the
major axis transverse, are quite irregular, in a considerable proportion of the
total, because the spots on the two sides do not synchronize. This results in
some diagonal blotches, and others which are restricted to one side while still
others are Y-shaped, double on one side and single on the other.

In addition to the main dorsal blotches, there are, on each side, several
alternating series of progressively smaller spots. In many specimens these
lateral spots approach obsolescence, and are represented only by irregular

darkened scale-edges.

The main dorsal blotches are brown, with dark-brown edges; the darker
borders are rather irregular. Between blotches the ground color is cream, but
this is evident only middorsally and laterally, for elsewhere there is a brownish
stippling or suffusion which colors the sides and much reduces the contrast
between the blotches and ground color. This darkening is evident particularly
in scale centers; the scale edges are usually light, giving a net-work effect
characteristic of Arizona. There is much variation in the extent of the suffusion
dorsally; in some specimens the dorsum is as much colored as the sides, but in
others the three middorsal rows may be almost unmarked, so as to constitute
a light central streak. Usually, at the tail, the light streak takes precedence
over the main dorsal blotches, dividing them into bilateral series. In any case,
the tail blotches are often so irregular that they cannot be counted with accuracy.
The lowest lateral rows in this subspecies are usually immaculate, differing in
this regard from occidentalis, in which they are usually both suffused and
spotted by the lowest side blotches. The suffusion is less evident in young
specimens, on which both the head and body marks are clearer.

A live specimen from Borego Palm Canyon, San Diego County, exhibited
the following colors, using the designations of Ridgway’s Standards, 1912:
Ground color of the head, Avellaneous; postocular streak, Army Brown; centers
of dorsal body blotches, Avellaneous; edges of dorsal blotches (quite narrow),
Bone Brown; scales of interspaces, middorsally, Pinkish Cinnamon, with White
outer edges; lateral blotches, Sepia, (these are really lines, comprising the
obsolescent remains of blotches, rather than true blotches); centers of scales
in lateral interspaces, Orange Cinnamon; edges of scales in lateral interspaces,
White; ventrum, White. The pupil is almost round, although slightly com-
pressed, vertically; the iris is golden. The eye has a considerable freedom of
horizontal rotation; it can be directed toward the front or rear. The tongue is

black, with whitish tips.

Intrasubspecific Trends—In the subspecies eburnata the ventral scutes
reach a maximum in the Borego area of eastern San Diego County, declining
both to the north and east. But a contrary trend is evident in the subcaudals,

358 SAN Dreco Society oF NAtTuRAL History

which tend to increase in Riverside, as compared to San Diego County; and
the tendency continues northward across the eastern Mojave Desert into southern
Nevada and southwestern Utah. The infralabials are also higher in Riverside
County.

The blotches, both body and tail, are highest in the Coachella-Palm Springs
area of Riverside County, falling off to the north, east, and south. But the
blotches are most highly differentiated from the other subspecies in the Borego
area of San Diego County, for here they are narrowest both along and across
the body. Likewise, more of the lower lateral scale rows are unmarked.

The tail length ratio increases from the type locality toward Riverside
County, and remains higher into Nevada and Utah. The Nevada-Utah snakes
are less highly differentiated from occidentalis than are the specimens of the
western Colorado Desert. It is possible that the availability of additional material
from Utah might validate other differences, and thus substantiate the further
segregation of the snakes of that area into a new subspecies.

Relationships with Other Subspecies—I have already mentioned the inter-
gradation of eburnata with noctivaga in eastern Yuma and western Maricopa
counties, Arizona. Intergradation with candida will be discussed under that
subspecies.

Eburnata intergrades with occidentalis along the eastern slope of the San
Jacinto and Peninsula ranges in Riverside and San Diego counties, California.
Intergradation evidently occurs wherever the mountain ranges are low enough
for Arizona to cross. In the San Gorgonio Pass the intergrades are found in
the vicinity of Cabazon. In eastern San Diego County the specimens found
in the San Felipe Valley are intermediate and are to be deemed intergrades;
and even as far out on the desert as Yaqui Well they are not yet typical
eburnata. In Warners Ranch the snakes are typical occidentalis. Farther south,
at Jacumba, they are intermediate, favoring slightly the desert subspecies. The
same intergradation no doubt occurs across the Sierra Juarez in Baja California,
Mexico, but no specimens are available from the east side of the mountains in
that territory.

Life History Notes—Eburnata occupies the largest area of any of the
truly desert subspecies of Arizona. It is found in a variety of surroundings,
from an almost barren desert, or sand dunes, to places where the brush cover
is quite dense. While not absent from rocky areas, it does not prefer them, and
is less plentiful in such situations than on the sandy flats. Some of the surround-
ings where this subspecies has been observed have been as follows:

Orchard
Cultivated field
Grass

Light brush
Heavy brush
Cactus

m= WHR Ve

KLAUBER—IHE GLossy SNAKE 359

Mesquite 2
Joshua trees 1
Rocks 1
Rocky desert 1
Brushy desert 21
Sandy desert 8

Total 47

Mosauer (1935, p. 20) found a specimen in sand dunes at 10 p. m. after
following the tracks for 200 feet. Cowles and Bogert (1944, p. 285) consider
Arizona a true burrowing snake, which flows in and out of loose desert soil
with little difficulty. I believe that eburnata takes refuge either by burrowing,
or in mammal holes, which are plentiful in most desert areas. The late Frank
Stephens plowed out a specimen at La Puerta, February 10, 1923. Cowles
(1941, p. 134) reports a specimen excavated by a scraper February 20, which
he judges to have been concealed just below the surface of the ground. At
San Felipe on March 11, 1929, one was found beneath a stone.

Eburnata reaches an altitude of about 2800 feet in southwestern Utah and
in eastern San Diego County. But the most typical habitats are at lower levels,
especially in the Salton Basin, where this subspecies occurs down to 240 feet
below sea level. In the Imperial Valley, eburnata does not seem to have been
greatly affected by irrigation; it has neither been driven out, as have such desert
forms as Crotalus cerastes laterorepens and Chionactis occipitalis annulatus, nor
has it greatly increased in numbers as has been the case with Pituophis catenifer
affinis (Klauber, 1939, p. 52).

That eburnata is nocturnal can be readily shown by a list of the hours when
specimens have been encountered alive on the road:

Number of

Time Specimens
6:00 to 6:29 p.m. 1
6:30 to 6:59 1
7-005 to 97329 5
PEN co), Jiao) 5
8:00 to 8:29 7,
8:30 to 8:59 9
9:00 to 9:29 6
9:30 to 9:59 5
10:00 to 10:29 6
10:30 to 10:59 6
11:00 to 11:29 2
11:30 to 11259 2
3:30!to ~3259 a: m. 2.
Total Sy/

360 SAN Disco Society OF NATURAL History

The only specimen found in the daytime was lying in a bush at Las Arenas,
San Felipe Valley, San Diego County, at 3:15 p. m. This was an occidentalis
intergrade.

Linsdale (1940, p. 243) reports a specimen motionless on the ground a
half hour before sunset. Cowles (1941, p. 133) mentions a specimen basking
in the sun near Indio, California, in December. Cowles and Bogert (1944,
p. 285) found Arizona to be secretive in captivity, appearing on the surface only
rarely and for short intervals. There was an avoidance of light. Nevertheless,
a large gravid female was found abroad in the Coachella Valley at 8 a. m. on
July 7, 1940, in full sunlight, although the temperature probably exceeded 100
degrees F. in the shade. This individual was evidently seeking a place to deposit
eggs; they were laid the following day and eventually hatched.

The air temperatures when eburnata has been observed active on the road
at night have been as follows:

Temperature Number of
Deg. F. Specimens
64-65 1
66-67
68-69
70-71
72-73
74-75
76-77
78-79
80-81
82-83
84-85
86-87
88-89
90-91

Total

Eburnata does not have to withstand quite as low temperatures in its season
of maximum activity in the Colorado Desert, as does candida in the western
Mojave Desert. From seasonal and mileage statistics (Klauber, 1939, p. 44), it
is believed that eburnata is most active in late May or early June, earlier in the»
lower Colorado Desert (the Salton Basin) than in the higher, eastern Mojave
Desert. I should place the optimum temperature at about 80 degrees F. The
best time for collecting is immediately after darkness has fallen in the earlier
spring, but later in the evening in June or early July. Activity declines sharply
in mid-summer, although it may be considerable just before dawn; we have
not made really adequate tests in these hours.

Cowles (1941, p. 132) expresses the opinion that eburnata is one of the
most cold-tolerant snakes in southern California; if continuous hibernation
exists, it is probably of short duration, possibly only from the middle of Decem-
ber through January. Cowles and Bogert (1944, p. 285) found that eburnata

KLAUBER—IHE GLossy SNAKE 361

would issue from the ground, but for short periods only, at temperatures as
low as 59 degrees F. They concluded that 66-68 degrees probably constitutes
the normal lower limit for voluntary surface activity. These figures refer to
body, not air, temperatures. The critical temperature was found by these
authors to be from 106 to 109 degrees F., and the putative lethal temperature
109 to 111 degrees. Recently-born young suffered at temperatures from 5 to 9
degrees F. lower.

In order to determine the importance of eburnata as a component of the
total snake population in at least one area (the Borego), I have tabulated all
specimens recorded alive or dead, and whether during day or night traveling,
along the road between Scissors Crossing (San Felipe Valley) via Sentenac
Canyon, Yaqui Well, and The Narrows, to Bensons Dry Lake, in eastern
San Diego County. This includes a section of the desert foothills and the
desert itself; it involves a desert valley (San Felipe), a rocky gorge with some
permanent water (thus accounting for the garter snakes), a wide, sandy, dry
wash, and a stretch of desert with scattered brush, cactus, and sand. The total
length of this sample strip is 21 miles. The snakes which have been recorded
are listed in Table 1.

TABLE 1.

Snakes Encountered on the Road from
Scissors Crossing to Bensons Dry Lake

Number of
Subspecies Specimens Per Cent
Phyllorhynchus decurtatus perkins 27 32.8
Chionactis occipitalis annulatus 181 21.9
Arizona elegans eburnata 70 8.5
Leptotyphlops humilis cahuilae* 55 6.7
Crotalus cerastes laterorepens 49
Rhinocheilus lecontei clarust 43

Masticophis flagellum piceus 39
Hypsiglena ochrorhyncha ochrorhyncha 29
Lichanura roseofusca roseofusca 29
Lampropeltis getulus californiaet 16
Crotalus ruber 16
Trimorphodon vandenburghi 13
Thamnophis hammondu 3)
Tantilla eiseni transmontana 3
Crotalus mitchellii pyrrhus 2
Salvadora hexalepis hexalepis 2
Pituophis catenifer annectens 2
Pituophis catenifer affinis 2

* Including Aumilis-cahuilae intergrades.
+ Including lecontei-clarus intergrades.
+ Ringed phase.

362 SAN Disco Society oF NATURAL History

We find here a greater variety of species than exists in the Mojave-Ade-
lanto-Kramer area (see p. 371), since that portion of the western Mojave contains
no rocky section corresponding to the Sentenac Canyon or The Narrows, which
account for the presence of such forms as Leptotyphlops, Hypsiglena, Lichanura,
Trimorphodon, and Tantilla. Other differences evident between the two tables,
is the high prevalence of the two little snakes Phyllorhynchus and Chionactis
in the Colorado Desert, as compared to their relative rarity in the western
Mojave; and the virtual absence of the usually common gopher snakes
(Pituophis) in the Borego area. As a matter of fact, P. c. annectens is quite
common somewhat farther west, and P. c. affinis is equally prevalent in the
Imperial Valley to the east. This intermediate area is evidently not favored
by either subspecies.

As to eburnata itself, it is seen to be the most common of the larger snakes
in the Borego area, being exceeded in numbers only by the two little ground
snakes, the leaf-nose and the shovel-nose.

Eburnata feeds principally on lizards and small mammals, remains of both
having been found in a number of specimens. One rather small individual
contained a full-grown Uta stansburiana hesperis. J. R. Slevin found a specimen
near Yuma which contained a Dipsosaurus. I have one in my collection which
had eaten a leaf-nosed snake. C. B. Perkins, at the San Diego Zoo, found that
captive specimens preferred lizards, especially Coleonyx, to mice, of which they
seemed to be afraid. Mosauer (1935, p. 20) fed captive specimens on Uma
and Callisaurus.

Eburnata is usually a peaceful snake, although rarely one will bite when
first picked up. It is generally slow-moving and specimens found on the desert
roads at night seldom escape. But I have one field note to the effect that a
large specimen on the road at Bensons Dry Lake at 7 o’clock in the evening,
with the air temperature 91 degrees F., was very lively and quite hard to catch.
One individual at Little Morongo Canyon was seen to raise its head and look
about in the manner so characteristic of the racers.

Specimens have been found containing 4, 5, 6, 7, and 9 eggs. The smallest
of these mothers measured 750 mm. We have not yet been successful in hatch-
ing this species at the San Diego Zoo. Cowles and Bogert (1944, p. 285)
report a female which laid 23 eggs; these hatched in 68 days.

Range—The desert areas of extreme southwestern Utah, southern Nevada,
the eastern and southeastern desert areas of California south of central Inyo
County (but excluding the extreme western Mojave Desert), northeastern
Baja California, Mexico, southwestern Yuma County, Arizona, and north-
western Sonora, Mexico.

Locality Records.—
UTAH

WASHINGTON COUNTY: Watercress Spring
1 mi. e. of St. George Near Bloomington

KLAUBER—THE GLossy SNAKE 363
NEvADA
Nye County: Indian Springs Ranch (42 mi. nw. of
9% mi. s. of Oak Spring (alt. Las Vegas)
4400 fr.) Boulder City (also just outside at
CLARK COUNTY: alt. 2000 ft.)
Kaolin Boulder City to Las Vegas road
Near mouth of Virgin River (at alt. 2400 ft.)
2 mi. s. of Ripley Bracken
Garnet Erie
Jean
CALIFORNIA

SAN BERNARDINO COUNTY:

2 mi. n. of Topock
Sacramento Mountains
Goffs

6 mi. n. of Vidal
Yucca Station

5 mi. e. of Baker

7 mi. w. of Ludlow
34 mi. e. of Barstow

Indio (also 1 and 9 mi. s. and 7 mi.
nw.)

3 mi. up Little Morongo Canyon

5 mi. nw. of Wide Canyon Junction

Ferguson Ranch (Coachella Valley)

Coachella (also 1 mi. n. and 4 and
8 mi. s.)

Thermal (also 3 mi. w.)

Mecca (also 1 mi. w.)

Box Canyon (near Mecca)

Daggett Salton

Nebo : North shore Salton Sea
Barstow* (also 1 mi. n.) Oasis (also 2 mi. se. and 14 mi. n.)
Mace Date Gardens

Lenwood US 99 at Imperial County Line
Hodge (also 3 mi. sw.) Whitewater

Wild Palm Springs RR. Station
Helendale Garnet (also 4 mi. n.)

Bryman Edom

Deadmans Point Dry Camp

5 and 13 mi. e., and 13 mi. ne. of Myoma

Victorville
3 mi. n. of Box S Spring
Twentynine Palms (also 14 and 18

Palm Springs (also 4,5, 10 and 13 mi.
n., and 2, 5, 6, and 7 mi. nw.)
Cathedral City (also 3 mi. s. and 3

mi. w.) mi. se.)
West end of Morongo Valley Junction US 99 with Palms to Pines
Highway

RIVERSIDE COUNTY:

Blythe (also 4 mi. s., and 7 and
16 mi. w.)

West end Granite Mountains (18 mi.
w. of Freda, San Bernadino
County )

2 mi. e. of Hopkins Well

Desert Center

Hayfield

5 mi. e. of Shavers Summit

7 mi. n. of Twentynine Palms
Junction

Cabazon} (also 3 and 5 mi. e.t)

SAN

Indian Wells (also 5 mi. w.)

Dieco County:

Collins Valley

Clark Dry Lake

Beattys Ranch (Borego Valley)
(also 3 mi. e.)

Borego Springs (also 2 mi. s.)

Borego Palm Canyon

Borego (abandoned townsite, also
called San Felipe, near Halfhill
Dry Lake)

Sentenac Canyon (also 2 mi. e. and 1
mi. w. of the bridge)

* Some of the succeeding localities in the San Bernardino County list may represent

candida-eburnata intergrades; these localities are along the Mojave River, between Barstow
and Victorville, from the Apple and Lucerne valleys, and near Twentynine Palms.

+ Occidentalis-eburnata_ intergrades.

SAN Dreco Society oF NATURAL History

San Dieco County (Continued) :

Yaqui Well (also 2 and 4 mi. w., and
3 mi. e.)

The Narrows (also 1, 2, 3, 4, 5, mi.
e., and 1, 2, 3, 414, and 5 mi. w.)

Bensons Dry Lake (type locality)
(also 2, 3, 4, and 6 mi. w., and
2 mi. e.)

San Felipe Wash (1 and 6 mi. w. of
Imperial County Line)

La Puerta (—Mason Valley)

Vallecito

Carrizo Spring

Banner* (also 2 and 3 mi. e.*)

Scissors Crossing* (also 1, 2, and 3
mi. e.* )

Cigarette Hill* (San Felipe Valley)

Top of Sentenac Canyon*

Las Arenas Ranch * (San Felipe
Valley)

ARIZONA
YuMA County:

Yuma (also 9 mi. e.)
2 mi. n. of Somerton
Dublin (also 2 mi. w.)
17 mi. w. of Wellton
Tacnat (also 4 mi. e.)
Pembroke+t

Near Mountain Spring*
Jacumba*

IMPERIAL COUNTY:

Boulder Park*

Mountain Spring*

Coyote Wells

Plaster City (also 3 mi. e.)

Dixieland

Seeley (also 5 mi. s.)

Travertine Rock

Seaview (also 3 mi. s.)

12 mi. e. of Bensons Dry Lake (San
Diego County )

Kane Spring

6 mi. n. of Mt. Signal P. O.

Holtville (also 2 mi. e.)

Date City

East edge of sand hills on US 80
(about 13 mi. w. of Winterhaven )

Winterhaven (also 2, 3, and 11 mi.
w.)

50 mi. e. of Yumat (= 2 mi. e. of
Colfred )

Mohawk (also 2 mi. e.})

Kimt

Stoval+

1 mi. w. of Aztect

Sonora, Mexico
Punta Penasco.

Arizona elegans candida subsp. nov.

WESTERN-MoyaAve GLossy SNAKE

Plate 8, fig. 2.

1930. Arizona elegans occidentalis (part) Bogert, Bull. Sou. Cal. Acad. Sci.,
vol. 29, part 1, p. 5.

Type—No. 34191 in the collection of LMK. Collected at Kramer Hills
(6 miles south of Kramer Junction on US 395), San Bernardino County,
California, by James Deuel, June 16, 1941.

Diagnosis —Candida is a subspecies characterized by its light color, nar-
row dorsal blotches, and the high frequency of paired preoculars. It differs
from all subspecies except eburnata in having dorsal blotches which are uni-
formly shorter (along the body) than the interspaces which separate them.
From eburnata it differs in its high proportion of paired preoculars, and some-
what wider and fewer dorsal blotches, which usually engage 7 scale rows in

* Occidentalis-eburnata intergrades.
+ Eburnata-noctivaga intergrades.

KLAUBER—THE GLossy SNAKE 365

eburnata and 9 in candida. Also, the latter has fewer ventral scutes on the
average, although there is considerable overlapping.

Description of the Type—The type specimen is an adult male; length
over-all 823 mm.; tail length 112 mm.; tail proportion 13.6 per cent. These
measurements were made before the specimen had set in preservative.

The head is slightly convex on top; it is little wider than the neck. The
snout is moderately pointed and the lower jaw inset. The pupil is almost round.

The scales of the head are normal. The rostral is sharply recurved and
somewhat wider than high. It contacts the first supralabials, the anterior nasals
(rather narrowly), and the posterior point separates the internasals for about
half their lengths. The internasals are longer than wide and curve down in front
to the nasals. The prefrontals are quadrangular in shape, with their post-lateral
points curved well down over the canthus rostralis. They are indented by the
upper points of the postnasals. The frontal is pentagonal, narrowing along the
supraoculars and then more sharply between the parietals. The supraoculars are
narrowly in contact with the prefrontals; they widen posteriorly. The parietals
are large and touch somewhat over half the posterior edges of the upper
postoculars. Posteriorly they are rather pointed. The postnasal is considerably
larger than the anterior; the nostril is at the upper end of the separating suture,
which is not complete above the orifice. Of the supralabials, the postnasal
touches only the first. There is a single quadrangular loreal on either side about
twice as long as high, touching the first to third supralabials. The posterior edge
of the loreal is substantially vertical on the right and almost so on the left.
There are two preoculars; the upper is both higher and wider. The postoculars
are 2-2. The temporals are 2+3, 2+5. The supralabials are 8-8, the fourth
and fifth touching the eye, and the seventh much the largest. The mental is
small and subtriangular. The infralabials are 13-13, the first in contact behind
the mental, and the seventh much the largest. There are two pairs of genials,
the anterior considerably the longer, and medianly in contact; the posterior are
slightly divergent and are separated by three rows of gulars.

The dorsal scale rows are 27-27-19. The scales are smooth, rather
narrow and with rounded ends. The middorsal and lateral rows are the widest.
There are single apical scale pits; however, these are quite small and faint. The
ventral scales number 214; the anal plate is entire; the subcaudals 50, all divided.

The head is buff above and lighter on the sides. There is a dark-brown bar
across the posterior edge of the prefrontals; also posteriorly on the frontal, and
along the parietal suture. There is faint evidence of a vertical dark mark below the
eye, and an obsolescent dark streak from the eyes back to the angle of the jaw.
The lower surface of the head is immaculate cream-colored.

The body pattern comprises a series of 61 rather irregular brown blotches
on a buff background. The irregularity is due to the non-matching of the two
sides; sometimes the blotches are diagonal, at others the lack of synchronization
is such that there are separate rows on the two sides. These blotches are much
wider than long. At mid-body they are about 1 scale long, and 9 scale rows
wide. The interspaces are about 244 scales (end to end) long. The blotches
are dark-edged. The ground color is clearly evident only on the three middorsal

366 SAN Dreco Society OF NATURAL History

scale rows. Laterally the dorsum is suffused with brown, but the two lowest
lateral rows are clear. In addition to the main blotches there are two auxiliary
series of brown spots on each side. These are quite indefinite and do not match
perfectly with the main series. The lower and smaller comprises merely a series
of scattered black spots. The lowest touch the third scale row above the ventrals.
The ventrum is immaculate cream-color. There are about 18 indefinite spots
on the tail; these are evident on the sides, rather than dorsally.

Summary of Paratypes—As this subspecies varies slightly in the different
parts of its range, I shall summarize the specimens which are considered para-
types, these being from along highway US 395 south from Kramer Junction to
Adelanto and a little beyond, all within a few miles of the type locality, Kramer
Hills, and all in San Bernardino County, California.* There are in this series
(including the type) 40 males and 13 females, together with one extra head.

The statistics of scutellation are as follows: The scale rows are nearly
always 27 at mid-body, although there is one specimen with 26 and another with
29. The ventral scutes in the males range from 209 to 220, interquartile range
212.4 to 216.0, mean 214.20+.42, coefficient of variation 1.24 per cent; the
corresponding figures in the females are, over-all range 220 to 232 (including
one aberrant individual 8 scutes higher than any other), interquartile range
220.6 to 224.8, mean 222.69+.86, coefficient of variation 1.40 per cent. The
subcaudals in the males range form 47 to 55, interquartile range 49.6 to 52.1,
mean 50.83.29, coefficient of variation 3.56 per cent; females, range 44 to 49,
interquartile range 45.2 to 47.1, mean 46.15+.39, coefficient of variation 3.09
per cent. The supralabials are normally 8 but are occasionally 9 (6 out of 107
counts). The infralabials vary from 11 to 14, the distribution being 11 (4),
12 (43), 13 (54), 14(6). The loreals are single in all but one count out of 107,
this one being split. In 41 counts the preoculars are single; in 67 they are
paired. Thus, in this character, which is typical of candida, 62 per cent are
positive. The postoculars are 2 in 107 counts, and 3 in one. The temporals are
usually 2-3, but may vary from 142 to 2+5 or 374.

The body blotches range from 55 to 73, interquartile range 59.9 to 65.1,
mean 62.50.53, coefficient of variation 6.23 per cent; and the tail spots over-
all range 14 to 24, interquartile range 17.1 to 20.3, mean 18.72+.33, coefficient
of variation 12.8 per cent. The body blotches usually cover 9 scale rows across
the trunk, but occasionally occupy 10, or rarely, 8. The longitudinal extent of

*The paratypes are as follows (A—Adelanto, KJ—=Kramer Juntion, KH=Kramer
Hills; the figures are miles; all specimens are LMK unless otherwise stated): 27251 16 n.
A, 28846 KH, 28848 8 n. A, 31700 KH, 31766 5 n. KJ, 31917 12 s. A, 31940 8 s. KJ,
31941 3 n. A, 31942 3 n. A, 31959 20 s. KJ, 33323 6 sw. A, 33795 KH, 33812 6n. A,
33826 3 s. KJ, 33832 8 s. KJ, 33888 5 s. KJ, 33975 7 s. KJ, 33977-8 10 s. KJ, 33979 12 s.
KJ, 33980 8 n. A, 33981 4s. A, 33982 14 n. A, 33983 8 s. KJ, 34017 7 s. KJ, 34019 8 s.
KJ, 34164 5 s. KJ, 34165 6 s. KJ, 34184 1 w. KJ, 34185 1 e. KJ, 34186 11 s. KJ, 34187 16
n. A, 34189 3 s. KJ, 34190 16 n. A, 34192 5 s. KJ, 35093 4s. A, 35107 13 n. A, 35109
3m. A, 35149) KM, 35150 A, 35151 NJ, 35536) 7 s. KJ, 39537 9%s, NJ, 35594 2 ins AS
35654 6¥2 s. KJ, AMNH 60570 6 w. A, MCZ 44846-7 A, MVZ 39604 7 sw. A, MVZ
39605 3 s. A, MVZ 39606 5 s. A, MVZ 39607 7 sw. A, MVZ 39643 5s. A.

KLAUBER—THE GLossy SNAKE 367
the blotches is 1 to 14% scales, while the interspaces measure 1/2 to 242 scales
(end to end). In only one specimen of the paratype series do the body blotches
equal the interspaces in longitudinal extent. The brownish side-suffusions
usually reach down to the second or third lateral rows above the ventrals, and
the lateral spots generally touch the same rows. This subspecies is the lightest
and has the smallest spots of any except eburnata.

The tail ratio is from 12.8 to 14.7 per cent in the males, and 11.6 to 13.1
in the females. The adult males average about 14.0 and the females 12.5 per
cent.

Material —In addition to the type and paratypes (40 males and 13
females, all from San Bernardino County) the following specimens, all from
California, are available: San Bernardino County 2, Los Angeles County 13
(several may be considered occidentalis-candida intergrades) , Kern County 14,
Inyo County 3; grand total 85. There are 65 males and 19 females, the other be-
ing a head only.

Description of Subspecies —This is a snake of average colubrid proportions,
neither particularly stout nor slim. The head is slightly distinct from the neck;
viewed from above it is wedge-shaped but with a rounded snout. From the
side the top of the head is convex; the snout is advanced and the jaw deeply
inset. The snout is moderately sharp; the forward part slants upward, so that
the tip is toward the upper end of the rostral. The eyes are rather protuberant;
the diameter of the orbit is equal to about 80 per cent of the distance from the
anterior edge of the orbit to the nostril. The pupil appears almost round in
most preserved specimens but is slightly elliptical in life.

The longest specimen I have seen, a male from 5 miles east of Lancaster,
Los Angeles County, measured 870 mm. The smallest specimen measured
245 mm. The tail proportion (ratio to length over-all) varies from 12.6 to 15.3
per cent in the males, and in the females from 11.6 to 13.3 per cent. The
averages are 13.9 per cent in the males and 12.5 in the females. The young
specimens have proportionately shorter tails than the adults to a small degree
(G.p> 354).

The body is covered with smooth scales. The middorsal and lower lateral
rows are slightly enlarged. There are single apical scale pits, but they are not
evident on all scales, and in any case are quite inconspicuous.

The scale rows at mid-body are usually 27, but 3 specimens out of 84 have
29, and one has 26. The ventrals in the males vary from 208 to 220, the inter-
quartile range is 212.3 to 216.1, the mean 214.22+.35, and the coefficient of
variation 1.32 per cent. The over-all range in the females is from 220 to 232,
the interquartile range 221.2 to 225.6, the mean 223.37+.75, and the coefficient
of variation 1.46 per cent. The anal is entire. The subcaudals, all divided, vary
from 47 to 55 in the males, the interquartile range being 49.7 to 52.1, the mean
50.92.22, and the coefficient of variation 3.47 per cent. In the females the
over-all range is 44 to 49, the interquartile range 45.4 to 47.2, the mean
46.25.30, and the coefficient of variation 2.88 per cent. The coefficient of

368 SAN Dreco Socrety oF NATURAL HIstory

sexual divergence in the subcaudals is 9.6 per cent. The terminal scale is some-
what elongated, and has a lateral crease.

The head scales follow the usual colubrid pattern. The rostral is rather
sharply curved, both over the top of the snout and below, and is longitudinally
concave on the underside. It is wider than high. The upper point separates the
internasals for approximately half their lengths; it also contacts the prenasals
and the first supralabials. The internasals are longer than wide, and curve
downward to a point between the prenasals and rostral. The prefrontals are
wider than high and curve down over the canthus rostralis to a broad contact
with the loreals. Rarely the prefrontals are fused into a single plate. The
supraoculars are widest posteriorly; they contact the prefrontals rather narrowly.
The frontal is widest anteriorly; it does not deeply indent the suture between
the parietals. The parietals are the largest of the head scales. They are rather
regular in shape, narrowing posteriorly. The dorsal scales which border them
are somewhat enlarged, but are irregular. The prenasal is much smaller than
the postnasal. The nostril is at the upper end of the diagonal suture which
divides the nasals; it slopes forward above. The suture is usually not complete
above the nostril. While the postnasal most often contacts the first and second
supralabials, in some specimens it touches only the first. The loreal is twice as
long as wide, and is highest forward; posteriorly where it contacts the preocular
the end is almost vertical, especially in those specimens which have 2 preoculars.
In the others the end may be somewhat pointed, as in the other subspecies. The
loreal contacts the second and third supralabials. One specimen out of 86 has
a divided loreal on one side. The preoculars are usually paired in this subspecies
(its outstanding characteristic) , 108 out of 172 counts, or 62.8 per cent having
2. When there are 2 preoculars, the upper is the larger. The postoculars ordi-
narily number 2, and are subequal in size; there are 3 in two counts out of 172.
The temporals are usually 213 or 2+4, but vary from 112 to 2+5 and 314.
Those in the first series are usually long and thin, and slope upward posteriorly.
The supralabials are usually 8, but occasionally number 9 (7 per cent), or even
10 (0.6 per cent). The fourth and fifth touch the eye; the next to the last is
always much the largest. The infralabials vary from 11 to 15, the distribution
being 11(6), 12(58), 13(96), 14(9), 15(1). The first pair meet on the
median line. Usually the seventh is the largest of the series, but not infre-
quently the largest is the sixth. The mental is small and triangular, with concave
sides. The anterior genials are large and contact on the median line; the posterior
are both slimmer and shorter, and, diverging posteriorly, are separated by from
2 to 4 rows of gulars.

Candida is a blotched snake, with brown transverse blotches on a cream-
colored background. In some specimens the blotches are somewhat obscured by
a lateral brownish suffusion.

The head is buff or light-brown above, with somewhat lighter sides, espe-
cially at the supralabials. There is a brown cross-band on the posterior edges of
the prefrontals, a pair of spots at the rear end of the frontal, a mark on each
supraocular, where it contacts the postocular, and several spots, or a brown line,
on the suture between the parietals. There is a brown streak or bar from the

KLAUBER—IHE GLossy SNAKE 369

orbit to the angle of the mouth; also a dark spot or vertical line below the eye.
There may be a few spots in the loreal-nasal region. The lower jaw is immaculate.

On the neck there is usually a pair of parallel blotches edged with darker,
and separated by a light streak. These are the precursors of the dorsal series.

The dorsal blotches occupy considerably less longitudinal space than the
interspaces which separate them. The body blotches vary in number from 55 to
73, interquartile range 60.3 to 65.6, mean 62.91+.43, coefficient of variation
6 24 per cent. The tail spots have an over-all range of 14 to 24, interquartile
rang 17.0 to 20.2, mean 18.61+.26, coefficient of variation 12.6 per cent. The
body blotches at mid-bedy cover from 8 to 11 scale rows across the body, but
in most specimens they span 9 rows. The longitudinal extent of the blotches,
again measured at mid-body, varies from 1 to 2 scales, end to end, but most
specimens cover only 1 to 1% scales. The interspaces are wider; most of them
are 2 scales in length, but the range is from 1% to 3 scales. The lateral brown
suffusion is usually carried down to the second or third row above the ventrals.
The lateral spots of the auxiliary series sometimes touch the scale row next to
the ventrals, but more often do not extend below the second or third row.

The body blotches are elongated transversely, and are often quite irregular
because the patterns of the two sides do not exactly keep step. This results im
many diagonal blotches, and other half-blotches which are restricted to one side.

In addition to the main dorsal blotches, there are, on each side, several
alternating series of progressively smaller spots. The first series is sornetimes
moderately regular and clearly evident; but those below are virtually only ir-
regular darkened scale edges.

The main dorsal blotches are composed of brown punctations, with dark-
brown or black edges. These edges are often irregular and imperfect. Between
blotches the ground color is cream, but if this is evident at all dorsally it is only
on the 3 middorsal scale rows, for laterally the area between blotches is heavily
suffused or stippled with brown. This darkening occurs particularly in scale cen-
ters; the scale edges are usually light, giving a characteristic net-work effect. The
lateral darkening tends to obscure the outer edges of the middorsal blotches and
the lateral series as well. With respect to the side suffusion there is a considerable
variation between specimens, for in some it is relatively light, whereas in others
it is quite dark. In some individuals the middorsal rows are clear, while in the
others the suffusion crosses the dorsum. Where the central light streak is evident,
it is accentuated posteriorly; on the tail it often splits the central spots into two
parallel series. This is the case in all the western subspecies, but is more prev-
alent in candida.

Young specimens usually lack the interblotch suffusion, the dorsal and
lateral spots, and the head marks also, being clearer than in the adults.

Intrasubspecific Trends.—Candida is so territorially concentrated that no
important character trends are evident from the material thus far available.

Relationships with Other Subspecies—Candida inhabits the Antelope Val-
ley, and the Mojave Desert west of the Mojave River and the approximate line
Barstow-Randsburg; it is also found in Inyo County, at the foot of the Sierras

370 SAN Disco Society oF NATURAL HIstory

as far north as Lone Pine. Intergradation with eburnata occurs along the eastern
border of this area, but just where is as yet undetermined. The specimens of
Apple and Lucerne valleys are evidently eburnata, yet two specimens from the
Morongo Valley and west of Twentynine Palms have the paired preoculars typi-
cal of candida. For the present I have considered them intergrades.

Intergradation between candida and occidentalis occurs along the desert
slope of the San Gabriel Mountains in Los Angeles County, at such points as
Humphrey, Vincent, and Valyermo. Presumably intergrades will also be found
on the southeasterly slope of the Tehachapis in Kern County, but I have no
specimens from that region.

Life History Notes.—Candida is a resident of the high, wind-swept, western
Mojave Desert, an area of sparse brush, Joshua trees at the higher levels, and
loose sandy soil. Much of it has an altitude of about 2600 ft. (Mojave 2733,
Palmdale 2669, Adelanto 2877, Kramer 2490). Most specimens were collected
at night when the character of the surrounding country could not be ascertained,
but the classification would usually be light brushy desert. The following addi-
tional surroundings have been observed: Heavy brush, medium brush, Joshua
trees, and sandy desert. It reaches an altitude somewhat in excess of 4000 ft.
(1 mi. east of the summit of Walker Pass).

Times of collection have been noted as follows:

7:00' to 7:29) p. m:
7:30"to 7259
8:00 to 8:29
8:30 to 8:59
9:00 to 9:29
9:30 to, 9:59

10:00 to 10:29
10:30 to 10:59
11:00 to 11:29
330 tos 3-59 a.m:

SS _
fella pomeie sn ey MOON StS

Total

Of course, these collecting records reflect not only the times of activity of
the snakes, but the collectors as well. It is my judgment that in the spring,
when the desert cools quickly after sunset, the snakes seek refuge early; in
summer they may be active all night, and then, in fact, the temperatures may
be most suitable just before dawn. But on the cold and windy spring nights,
one may readily infer from the presence of live snakes early in the evening, and
DOR’s later, that the time of maximum activity is soon passed. At any rate,
the nocturnal character of this desert snake is not to be doubted, for I have yet
to find a specimen abroad in the daytime. Of the 44 snakes whose time of
collection was recorded, the earliest was 7:15 p. m. (May 10, 1941, about one-
half hour after sunset).

The maximum seasonal activity I should place at about June 10, somewhat
later than that of eburnata in the Colorado Desert, which, at a lower altitude,
warms earlier.

KLAUBER—THE GLossy SNAKE 371

The air temperatures when candida was collected were as follows:

56-57 deg. F. 1

NI
4
SK OO OCF AUW ND NAW OO WR WO

Total 38

No doubt this table unfairly represents the higher temperatures, for I have
done almost no collecting in the Mojave in the summer. At any rate, this does
indicate the surprisingly low temperatures at which snakes may remain active.
I remember picking one up at 10:45 on a cold and, as usual on the Mojave,
windy spring night; the air temperature was 60° F., and the snake was so stiff
and dull it was presumed to be a traffic casualty, but the next morning it was
quite well and lively.

In the section of the desert which it inhabits, candida is the second com-
monest snake, as is shown by the following table of specimens recorded by
myself and associates in the Mojave-Palmdale-Adelanto-Kramer area. This table
includes both live and DOR specimens, and both day and night collecting, thus
giving a representation to both diurnal and nocturnal forms.

Number of Specimens Per Cent

Crotalus cerastes cerastes Si 27,
Arizona elegans candida 95 157
Rhinocheilus lecontei lecontei* 89 14.7
Pituophis catenifer deserticola is) 13.1
Crotalus scutulatus scutulatus 78 12.9
Masticophis flagellum piceus 60 9.9
Chionactis occipitalis occipitalis 39 6.4
Lampropeltis getulus californaet 20 Bye
Phyllorhynchus decurtatus perkinsi 8 13
Salvadora hexalepis hexalepis 6 1.0

605 100.0

20 inclodes* some Runnels

+ Ringed phase.

Bi/92 SAN Disco Society oF NaturAL History

Although it is believed that candida prefers lizards, several specimens con-
taining mice (one a pocket mouse), or mammal hair, have been noted.

One specimen 856 mm. long contained 10 eggs; another 661 mm. in
length, 4 eggs.

Although Arizona rarely endeavors to bite, a specimen which had been
injured did so.

Range.—The Antelope Valley and extreme western Mojave Desert, in-
cluding the desert areas of southwestern Inyo County, southeastern Kern County,
northeastern Los Angeles County, and western San Bernardino County,

California.
Locality Records.—

CALIFORNIA
Inyo County: Humphreyt
10 mi. s. of Lone Pine Littlerock+
Brier Fort Tejon Road+ (between Pallet
Haiwee Creek Road and Griffin Road)
Coso Junction Valyermot (also 2 mi. s.)
Little Lake Pecks Butte (= Piute Butte )
Emigrant Checking Station* (Death Lovejoy Springs
Valley National Monument) Llano (also 10 mi. e.)
KERN COUNTY: San BERNARDINO County:
Brown 4 mi. n. of Red Mountain
3 mi. ssw. of Inyokern Kramer
1 mi. e. of Summit of Walker Pass Kramer Junction (also 3 and 5 mi. n.,
Searles Station I ave, Boy 34 SoCs 7, Os A@, itl.
Randsburg (also 5 mi. w.) 1228s eande20 minis sandenl
3,5, 9, and 14 mi. w. of Amargo mi. w.)
Boron (also 6 mi. w.) Kramer Hills (type locality) (also 8
Mojave (also 2 mi. n., and 3, 5, 6, 7, mi. s.)
and 8 mi. e.) Jimgrey
5 mi. n. of Muroc Adelanto (also 2, 3, 6, 8, 13, 14, and
Los ANGELES CouNTY: 16 mi. n., 2, 3, 4, 5, and 12 mi.
Neenach (also 3 mi. w.) s., 5, 6, and 7 mi. sw., and 6 mi.
Between Neenach and Fairmont w.)
Lancaster (also 5 and 15 mi. e.) 5 mi. sw. of Victorville
Denis 4 mi. s. of Hesperia
Palmdale Phelan (also 5 and 6 mi. n.)

Vincent}

Arizona elegans occidentalis Blanchard
CALIFORNIA GLOssy SNAKE

Plate 7; fig. 2.

1894 Coluber arizonae (part) Boulenge:, Cat. Snakes Brit. Mus., vol. 2, p.
66. (See remarks under elegans, p. 321.)

1897 Arizona elegans (part) Wan Denburgh, Occ. Papers Cal. Acad. Sci.,
No: >, ps 193:

1900 Khinechis elegans (part) Cope, Rept. U. S. Nat. Mus. for 1898, p. 863.

* Specimen not available; may be eburnata or an intergrade.
+ Occidentalis intergrade.

KLAUBER—THE GLossy SNAKE 373

1924 Arizona elegans occidentalis (part) Blanchard, Occ. Papers Mus. Zool.
Univ. Mich., no. 150, p. 1. Type specimen USNM 54372; type
locality La Jolla, San Diego County, California.

Diagnosis —Occidentalis is a subspecies inhabiting a territory having a
considerable ecological variability and therefore it is subject in itself to con-
siderable variation. It is characterized by a darker color than the other western
subspecies. It may be distinguished from elegans and blanchardi by its usually
having 27 scale rows compared with 29 or 31 in the eastern forms. Also, its
blotches average higher in number and are narrower across the body. Occidentalis
has more ventrals and a proportionately shorter tail than philip: or expolita, and
more blotches than pacata. It is darker in color and with lower spots and
brownish suffusions on the sides than eburnata or candida. Although territorially
separated from noctivaga by eburnata and candida, it is most like the Arizona
form. However, noctivaga has fewer ventrals on the average, is usually lighter
on the sides, and the lower lateral scale rows are freer of spots, as is also true
of the infralabials.

Material—The description of the subspecies occidentalis, as newly re-
stricted, is based on the following material: Baja California 11; California, San
Diego County 38, Riverside County 7, Los Angeles County 3, San Bernardino
County 5. These are all from the coastal areas of these counties. In addition
there are the following from the San Joaquin Valley section: Kern County 17,
Fresno County 8, San Benito County 1, San Joaquin County 2; grand total 92.
There are 49 males and 40 females, the rest being indeterminate.

Description of the Subspecies—This is a snake of normal colubrid pro-
portions, neither heavy-bodied nor attenuated. The head is only slightly distinct
from the neck. Viewed from above the head is wedge-shaped but with a rounded
or somewhat blunt snout. From the side the top of the head is convex; the jaw
is deeply inset. The apex is slightly pointed; the forward part of the snout slants
upward, so that the tip is toward the upper end of the rostral. The rostral is
sometimes slightly raised above the adjacent scales. The eyes are moderately
protuberant; the diameter of the orbit is about equal to 34 of the distance from
the anterior edge of the orbit to the nostril. The pupil is slightly elliptical.

The longest specimen, a female from La Jolla, San Diego County, measured
1132 mm. over-all. The smallest specimen is 248 mm. in length. The tail pro-
portion (ratio to length over-all) varies from 12.6 to 14.7 per cent in the males,
and from 11.4 to 13.7 per cent in the females. The mean ratios are 13.6 and
12.5 per cent in the males and females, respectively. The young specimens have
somewhat shorter tails proportionately than the adults.

The body is covered with smooth scales, rather narrow and with slightly
rounded ends. The lower lateral rows are increasingly wider. Single apical scale
pits are present, but they are small and inconspicuous.

The scale rows at mid-body are usually 27 but there are 29 in 20 per cent
of the specimens examined. The ventrals in the males vary from 207 to 223,
the interquartile range is 211.8 to 216.7, the mean 214.2454, and the coefficient
of variation 1.72 per cent. The over-all range in the females is from 215 to 231,

374 SAN Dreco Soctety oF NATURAL HIstTory

the interquartile range 220.2 to 225.8, the mean 223.00+.67, and the coefficient
of variation 1.86 per cent. The anal is entire. The subcaudals, all divided, vary
from 47 to 54 in the males, the interquartile range being 49.1 to 51.4, the mean
50.26+.25, and the coefficient of variation 3.30 per cent. In the females the
over-all range is 43 to 50, the interquartile range 45.2 to 47.8, the mean
46.49=.35, and the coefficient of variation 4.27 per cent. The coefficient of
sexual divergence in the subcaudals is 7.8 per cent. The terminal scale is slightly
elongated and creased; it is not sharply pointed in the adults.

The head scales follow the colubrid normal in size and arrangement. The
rostral is sharply curved, both over the top of the snout and below. It is 20
per cent wider than high. The upper point of the rostral separates the inter-
nasals for less than half their lengths; in addition it contacts the prenasals and
the first supralabials. The internasals are longer than wide and curve downward
to a point in front of the prenasals. The prefrontals are wider than high and
curve well down over the canthus rostralis to a level with the center of the orbit,
making a broad contact with the loreal. The supraoculars make a narrow con-
tact with the prefrontals; they widen posteriorly, where they contact the
parietals and upper postoculars. The frontal is widest anteriorly; it terminates
posteriorly at a point which indents the suture between the parietals for
about 14 of their lengths. The parietals are the largest of the head scales.
They are moderately regular in shape, narrowing posteriorly. The dorsal scales
which border them are somewhat enlarged, as compared to the succeeding
dorsals. The prenasal is smaller than the postnasal. The dividing suture slants
forward above; the nostril is at the upper end. The suture curves sharply
above the Abeta and usually terminates in such a way that the nasals remain
connected by a narrow isthmus. The postnasal contacts the first and second
supralabials. The loreal is more than twice as long as wide, and is highest
forward, becoming pointed posteriorly where it contacts the preocular. The
loreal borders the second and third supralabials. Divided loreals are present
in only 2 out of 180 counts in this subspecies. The preoculars are usually single,
although paired in 6 counts out of 180; they are wider above than below, where
they are indented by the loreals. The postoculars ordinarily number 2, and are
about equal in size; there are 3 in 6 counts out of 180. The temporals are
usually 213 or 2+4, but vary from 1+2 to 2+6 and 3+4. Those in the
first series are longer than wide and slant upward posteriorly. The supralabials
are usually 8; the dispersion in the available specimens is 6(1), 7(6), 8(157),
9(11), 10(1). When there are 8 the fourth and fifth contact the eye; the next
to the last is always the largest of the series. The infralabials vary from 11 to 15,
the distribution being 11(3), 12(28), 13(118), 14(24), 15(1), mean 12.95.
The first pair meet on the median line. The seventh is much the largest of the
series. The mental is small and triangular, with concave sides. The anterior
genials are large and contact on the median line; the posterior are both thinner
and shorter, and, diverging posteriorly, are separated by from 2 to 5 rows of
gulars.

Occidentalis is a brown snake marked by darker-brown blotches down the
back, and additional series of smaller blotches on the sides.

KLAUBER—THE GLossy SNAKE 375

The head is medium-brown, although the upper labials are lighter. There
is a brown band, slightly darker than the ground color, on the posterior edges
of the prefrontals; also, irregular dark-brown spots on the supraoculars, frontal,
and parietals. These spots tend toward obsolescence in adults. There is a
brown streak from the orbit to the angle of the mouth; also, there is often a
brown spot below the eye. There is usually a small brown spot on each first
infralabial, and posteriorly the sutures between the infralabials are spotted with
brown. These marks on the lower jaw are characteristic of this subspecies and
will often aid in segregating it from the desert subspecies noctivaga, eburnata, and
candida; however, they are often absent in specimens from the San Joaquin
Valley.

On the neck there is usually a pair of parallel dark marks separated by a
light streak. These, the first of the dorsal series, are much longer than those
which follow.

The dorsal blotches in this subspecies are about equal to the interspaces
which separate them. The body blotches vary in number from 51 to 75, inter-
quartile range 59.3 to 66.1, mean 62.70+.55, coefficient of variation 8.1 per
cent. The tail spots have an over-all range of 13 to 25, an interquartile range
of 16.7 to 20.0, with a mean of 18.33.29, and a coefficient of variation of
13.3 per cent. The body blotches at mid-body cover from 7 to 13 scale rows
across the body, but in most specimens they span 9 to 11 rows. The longitudinal
extent of the blotches, again measured at mid-body, varies from 1 to 3 scales, end
to end, but most specimens extend from 11/2 to 2 scales. The interspaces are
sometimes slightly wider, sometimes narrower, than the blotches; most of them
are from 1% to 2 scales in length. The lateral brown suffusion is usually
carried as far as the first to third row above the ventrals. The lowest lateral
spots of the auxiliary series usually touch the ventrals or the first lateral row,
although rarely these spots do not extend below the second or third row. Some-
times there are a few spots well out on the ventrals, which, otherwise, are buff.

The body blotches, although generally elliptical, with the major axis trans-
verse, are often quite irregular because the patterns of the two sides do not
exactly match. This results in some diagonal blotches, and others which are
restricted to one side, while still others are Y-shaped, double on one side and
single on the other.

In addition to the main dorsal blotches, there are, on each side, at least
two alternating series of progressively smaller spots. The first of these series
comprises vertically elliptical spots, alternating with the dorsal series; it is rather
regular and in most specimens as well defined as the main series. The next
series below are hardly more than dark scale marks in many individuals.

The main dorsal blotches are brown, with dark-brown or black edges.
Between blotches the ground color is buff, but this is evident only on the
middorsal scale rows, and in many specimens not even here, so general and
dark is the interblotch suffusion, often, in fact, so dark as to leave little contrast
between blotches and interspaces. This darkening occurs particularly in scale
centers; the scale edges are usually light, giving a net-work effect characteristic
of Arizona, which is more marked in some territories, the San Joaquin Valley

376 SAN Disco Society oF NATURAL HIsTorRY

for example, than others. The interblotch suffusions are accentuated with age;
the spots and blotches, both on the head and body, are more clearly evident in
the young. The middorsal lightening sometimes affects, not only the ground-
color suffusion, but the blotches themselves, particularly toward the tail, where
the dorsal series may be split by a central light streak.

A live juvenile specimen from Lakeside, San Diego County, showed the
following Ridgway (1912) colors: Dorsal blotches, Seal Brown; dorsal inter-
spaces, Avellaneous to Wood Brown; ventral surface White and somewhat
translucent.

Intrasubspecific Trends.—In occidentalis the ventral scutes are highest in
number in the snakes of coastal San Diego County, declining slightly both to
the north and south. The subcaudals, on the other hand, show an increase
toward the north, as do the infralabials as well.

The body blotches and tail spots are highest in number in San Diego
County, decreasing to the north and south. Also, the blotches are larger both
in width and length, in the San Diego snakes, thus showing the greatest diverg-
ence from eburnata. Both in the San Joaquin Valley, and at the southern end
of the range in Baja California (about lat. 30° 30’ N.), the blotches are
relatively smaller and more frequently exceed the interspaces in extent. There
is less ground-color suffusion on the sides, and the lateral spots are not carried
so low as in the San Diego County snakes, nor are the spots on the infralabials
as prevalent. Thus, in all these pattern characteristics, warmer and drier condi-
tions have produced, in occidentalis, a trend toward the desert form eburnata.
Or possibly this statement should be revised to suggest a retention of the desert
characteristics.

The tail proportionality in occidentalis increases slightly from south to
north,

Relationships with Other Subspecies—As I have already pointed out in
discussing those subspecies, occidentalis intergrades with both eburnata and
candida on the desert slopes of the coastal mountains.

Life History Notes——The differences in daily activity which are found in
Pituophis, between the coastal and desert forms, the former being largely
diurnal and the latter nocturnal, are not so evident in Arizona; for the coastal
subspecies occidentalis is almost, if not quite, as nocturnal as its desert congeners,
eburnata and candida. This is evident from the rarity of its discovery in the
daytime, whereas the frequency of DOR’s indicates that it is not uncommon in
the territory. I have never found a specimen abroad in the daytime, the earliest
being 6:25 p. m. at La Posta. L. M. Huey found one at 9 p. m. at Valle de la
Trinidad, Baja California. Linsdale (1932, p. 377) reports a specimen active
at 6 a. m., just as the sun was coming up. Occidentalis is so much darker than
eburnata and candida, particularly from a lateral view, that it is relatively difh-
cult to see at night. Thus, we have had nothing like the success, in collecting
this subspecies, that has attended our desert efforts; however, it must be admitted
that slow driving for snakes on the coastal side of the mountains is seldom tried.
In the lower San Joaquin Valley, where summer temperatures are high, occiden-
talis is lighter than along the coast, and may be hunted using the desert scheme.

KLAUBER—IHE GLossy SNAKE BF

Here I have taken live specimens on the road at 8:00, 8:50(2), 8:55, 9:06, and
10:45 p.m. The air temperature was as low as 60° F. Judging from my own
collecting experiences, both day and night, I should say that, with the possible
exception of the Bakersfield area, occidentalis represents a much smaller part of
the total snake population than that represented by eburnata and candida in
some desert areas.

There is some evidence that Arizona is not uniformly distributed in the
coastal territory of southern California, but occurs in colonies, for DORs
have been found again and again in the same localities—for example, La Costa,
Mission Valley, and Warners Ranch—whereas other places having the same
characteristics seem uninhabited by this snake. However, in San Diego County,
it is known to occur in all ecological zones, from the coast to the mountain foot-
hills; although in the mountains themselves it is absent, or at least very rare.
I know of no locality of collection higher than Warner Springs (alt. 3132 ft.).
About 60 per cent of the specimens have been from the coastal area, some on
the cliffs immediately above the surf. The relative frequency of surroundings
where specimens have been found is shown in the following list:

Grass (uncultivated) 22

Field (cultivated) 17

Light brush
Chaparral
Barren field
Orchard
Sand

Rocks

eRe Ww RNIN

Total 62

This list indicates a definite preference for open areas, for much of the
territory hunted in has a dense brush cover.

Seasonally, Arizona seems to be a trifle later than the other snakes of
the area, for June slightly exceeds May as the peak month, whereas the contrary
is true of most of the other species.

Specimens of Arizona are occasionally plowed out, showing that they
hibernate at a comparatively shallow depth. I have records of specimens being
plowed out on Feb. 10, 23, and 24. Paul Breese found one under a foot of
sand in a vineyard at Alta Loma, San Bernardino County. L. H. Cook found
one under a board; another was disclosed by turning over a rock, while a third
was buried in sand. Specimens in captivity bury themselves readily in loose
soil or sand.

Lizards seem to be the preferred food of this subspecies, particularly the
two common forms Uta stansburiana hesperis and Sceloporus occidentalis
biseriatus, there being several records of both of these forms being found in
collected specimens. L. H. Cook placed a newly captured snake in a bag with
several lizards. Two utas were taken, one head first, the other tail first.
Mammal hair has been found in several specimens. Cook (1930, p. 158)
Hanley (1943, p. 145) and Reynolds (1943, p. 196) have reported on ocd

378 San Disco Society OF NATURAL HIstTory

habits in captivity. Reynolds states that sparrows were eaten (subspecies of snake
according to the new classification not given). C. B. Perkins tells me that
at the San Diego Zoo lizards are always more readily accepted than mice by
captive specimens.

A specimen 731 mm. long from Oilfields, Fresno County, contained 7
eggs. The hemipenial characteristics are similar to those of eburnata.

A specimen bitten by a small rattlesnake died the next day.

Range—The San Joaquin Valley in California, south from central San
Joaquin County to the Tehachapi Mountains; and coastal and cismontane
southern and Baja California from Los Angeles County south to San Quintin
(lat. 30° 30’ N.). Intergrades with candida on the desert side of the San
Gabriel Mountains, and with eburnata on the desert slopes of the San Bernar-

dino, San Jacinto, and Peninsula ranges.

Locality Records —

CALIFORNIA

SAN JOAQUIN COUNTY:
Corral Hollow Creek (7 mi. ssw.
of Tracy)
Corral Hollow Creek (3 mi. e. of
Tesla)
San BENITO COUNTY:
Panoche Creek (2 mi. se. of
Panoche)
FRESNO COUNTY:
Fresno
3 mi. w. of Conejo
Oilfields
Coalinga (also 2 and 4 mi. e., and
10 mi. ne.)
KERN County:
Famosa
Dow
Saco
Rio Bravo
Rosedale
McKittrick (also 3 mi. n.)
Fellows
Midoil
10 mi. ne. of Taft
Maricopa (also 10 mi. e.)
Pentland
Tupman
North shore Buena Vista Lake
5 mi. w. of Stevens
Old River (also 7 mi. w.)
Greenfield (also 6 and 7 mi. s.)
Reed Station

Bena

Ilmon

Emigdio Station (also 3 mi. e. and
3 mi. w.)

* May be candida intergrade.

Wheeler Ridge P. O. (also 4, 5, 7,

and 9 mi. n.)
Grapevine (also 12 mi. n.)
Los ANGELES COUNTY:
3 mi. se. of Gorman*
Castaic

Saugus (also 1 mi. n., 2 mi.

and 6 mi. ne.)

7 mi. n. of Newhall

4 mi. nw. of Sunland.

Verdugo Hills

Alhambra

9 mi. e. of Azusa
SAN BERNARDINO COUNTY:

Ontario

Alta Loma

Rialto

San Bernardino

Muscoy

Highland Junction

San Bernardino Mountains
RIVERSIDE COUNTY:

Mira Loma

Riverside

West Riverside

Lakeview

San Jacinto

Andersons

Perris

Elsinore (also 7 and 9 mi. ne.)

Sedco

Wildomar

Murrieta

Temecula

nw.,

KLAUBER—IHE GLossy SNAKE 379

OrANGE COUNTY:
El Toro
Galivan
San Juan Capistrano
Coast Royal
San Disco County:
San Mateo Creek (at US 101)
San Onofre
Stuart
Between Oceanside and Carlsbad
La Costa
Leucadia
Encinitas
Cardiff
Solana Beach
La Jolla (type locality)
Pacific Beach
Murphy Canyon
Rosedale
Grantville
Mission Valley
San Diego

Otay

Tia Juana

Monument 258 (International Boun-
dary at Pacific Ocean shore)

Bonsall

Hodges Dam

Poway

Mussey (also middle of Mussey
Grade)

Mission Gorge

Santee

Lakeside

Lakeview (Johnstown)
Ballena

Ramona (also 3 mi. w.)

Warners Hot Spring

Warners Ranch

3 mi. w. of Warners Ranch House
(=3 mi. n. of San Felipe)

Pine Valley

La Posta

Campo

BajA CALIFORNIA, Mexico

Tijuana
Ensenada
Punta Banda

Santo Tomas

Valle de la Trinidad
San José (lat. 31° N.)

San Quintin

Arizona elegans pacata subsp. nov.
PENINSULA GLossy SNAKE

Type—No. 17652 in the collection of the San Diego Society of Natural
History. Collected Nov. 16, 1941, by Frank F. Gander, at Santo Domingo
(lat. 25° 30’ N.), Baja California, Mexico.

Diagnosis —A subspecies characterized by a low number of body blotches
(39 in the holotype), subcircular in shape, as compared with the other western
subspecies (occidentalis, eburnata, candida, philipi, and noctivaga) which
ordinarily have 50 or more rectangular blotches, averaging close to 60. Other
subspecies having low numbers of dorsal blotches are elegans and expolita.
Pacata differs from the first in having 27 rather than 29 or 31 scale rows; and
from the latter in having a proportionately shorter tail and fewer subcaudal
scales.

Description of the Type—The type specimen is an adult male; length
over-all 789 mm.; tail length 94 mm.; tail proportion 11.9 per cent.

This is a snake of moderate body shape, neither racer-like nor particularly
stout. There is a prominent vertebral ridge. The head is flat-topped, rather
narrow, and little distinct from the neck. The diameter of the eye is about 60
per cent of the distance from its anterior edge to the nostril. The pupil is
slightly higher than wide.

The scales of the head are normal. They comprise a sharply recurved

380 SAN Disco Society OF NATURAL History

rostral, considerably wider than high, and with the posterior point separating
the internasals for about half their lengths. The internasals are longer than
wide and have sharp points between the rostral and prenasals. The prefrontals
are quadrangular in shape, with their lateral edges curved downward over the
canthus rostralis. The frontal is hexagonal, and is longer than wide; it is
wedge-shaped at its terminus between the parietals. The supraoculars are in
contact with the prefrontals; they widen posteriorly. The parietais contact more
than half of the upper postoculars. Posteriorly they are edged by slightly
enlarged dorsal scales. Of the nasals the posterior is the larger; there is no
separating suture above the nostril. The postnasal touches the first and second
supralabials. There is a single quadrangular loreal on either side about twice
as long as high, touching the second and third supralabials; the lower edge
is considerably longer than the upper. The preoculars are 1-1; postoculars
2-2, the lower slightly larger than its fellow. The temporals are 214. The
supralabials are 8-8, the fourth and fifth touching the eye, and the seventh
largest. The mental is small and triangular. The infralabials are 13-12; the first
contact medianly behind the mental; the seventh on the right and the sixth on
the left are the largest. The suture between the fifth and sixth is incomplete on
the left. There are two pairs of genials, the anterior pair considerably the longer
and medianly in contact; the posterior are divergent and are separated by two
slim gulars.

The dorsal scale rows number 27-27-19. The scales are smooth, narrow
and are rather pointed posteriorly. There are faint single apical scale pits.
The ventral scales number 200; the anal plate is entire; there are 43 subcaudals,

all divided.

The head is medium-brown above and somewhat lighter on the sides.
There is a faint evidence of a postocular dark stripe toward the angle of the
mouth. The lower surface of the head is unmarked.

The body pattern comprises a series of 39 subcircular brown blotches.
The blotches are not sharply differentiated from the adjacent dorsum except
that the edges are somewhat dark. In addition, the scales outside the blotches
have their edges lightened to a greater extent than those within, thus causing
the blotches to be accentuated. At mid-body the blotches are about 4 scales
long (end to end), and 11 scale rows wide. The interspaces entail about 112
scales (end to end). Below the blotches on either side there is a secondary
series of smaller round spots which are not very clear, and below these there
are other irregular marks on a background which becomes increasingly light
toward the ventrum. The lowest lateral row on either side is immaculate
cream, as is the ventral surface. There are about 12 spots on the tail; one
cannot be sure of the count since they become indefinite posteriorly.

Remarks.—This specimen is the only one from the southern half of Baja
California of which I have knowledge. The most southerly specimen previously
known was from San Quintin. This leaves a gap of over 400 miles, and I
should be justified in making pacata a full species were the differences greater.
But since the differences seem to be largely in pattern, and because the interven-

KLAUBER—IHE GLossy SNAKE 381

ing territory is of such a character that intermediate specimens are to be
expected, I accord it only subspecific rank.

Additional specimens of pacata will probably show that it averages fewer
ventrals and subcaudals than either occidentalis or eburnata, the territorially
nearest subspecies.

Range.—This subspecies is known only from the type locality, Santo
Domingo (lat. 30° 30’ N.), Baja California, Mexico. No doubt it has an

extensive range in the south-central part of the peninsula.

PHYLOGENY

In differentiating the several subspecies of Arizona, the most consistent
characters—territorially—are those which separate them on an eastern-western
basis, these being tail proportionality and dorsal scale rows. The tail pro-
portionality carries with it differences in subcaudals. The ventrals and body
blotches, while consistent in local populations, this being particularly true of the
former, tend to duplication in widely separated areas, as if influenced by con-
ditions of regional ecology. Thus we have high ventral counts in elegans and
eburnata, and low blotch numbers in elegans and pacata. Higher ventral counts
are correlated with higher temperatures (Klauber, 1941, p. 73); fewer blotches
and darker colors with greater brush cover and a less exclusively nocturnal
existence. All of these considerations suggest that the most fundamental
differences lie in tail length and scale rows, and, of these, I should place tail
proportionality first. Had these two characters divided the subspecies with a
co-ordinated consistency, I should have been tempted to formulate two species;
such an eventual conclusion is still by no means impossible, since as yet it has
not been finally determined (through lack of material from critical areas)
whether philipi (or its southern extension expolita) intergrades or overlaps
with elegans-blanchardi to the east, or with noctivaga to the west. But since
philip: is intermediate, being eastern in tail length and western in scale rows,
I have concluded that it would be most logical, for the present, to continue to
consider Arizona monotypic.

It is my view that the species arose in north-central Mexico, radiating
northward from there. Blanchardi is a northern extension of elegans, and
philipi of expolita. Eburnata constitutes the main western branch, as at present
known, but I hazard the guess that another form will be found in Sonora with
fewer ventrals and larger blotches, from which noctivaga, eburnata and pacata
all radiated, the latter across the Gulf of California. Occidentalis and candida
are obviously off-shoots of eburnata. These conclusions are graphically sum-
marized in figure 1.

Subspecific differences in scale and blotch counts, and tail proportions,
are summarized in tables 2 to 7, inclusive.

382 SAN Disco Society OF NATURAL History

philip!
candida ,
noctivaga blanchardi

occidentalis eburnata
elegans

expolita

Ancestral Arizona

Fig. 1. The Phylogeny of Arizona.

383

KLAUBER—THE GLossy SNAKE

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Expolita
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Candida
Occidentalis
Pacata*

SAN Disco SoclETY OF NATURAL History

Table 3

ExTREME RANGES OF SCALE AND BLOTCH COUNTS

Ventrals, males

208-222
197-215
185-203
194-195
204-214
208-228
208-220
207-223
200

* One specimen only

Ventrals, females

220-232
207-222
193-211
207%
211-224
220-241
220-232
215-231

Subcaudals, males

Subcaudals, females

Body blotches

Tail spots

385

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386 SAN Dteco Society oF NaturaAt History
Table 5
DIsTRIBUTION OF SCALE-Row Counts at Mip-Bopy
Scale Rows
Subspecies 25 26 27, 28 29 30
Elegans 1 16 2
Blanchardi alk 3
Philipi 1 28 2
Expolita 3
Noctivaga 1 57 5 7.
Eburnata Z. 154 7 25
Candida 1 80 3
Occidentalis 1 68 7 16
Pacata 1
Table 6
MEAN Ratios oF Tart LENGTH TO LENGTH OVER-ALL
Subspecies Males Females
Elegans 149 142
Blanchardi 157. 147
Philipi 161 148
Expolita 158 .125*
Noctivaga 137 127
Eburnata 137, 126
Candida .139 125
Occidentalis 136 125
Pacata 1207 _
* Only one specimen; believed to be low.
7 Only one specimen.
Table 7
Mopar BLtotcH DIMENSIONS
: :
$ $8 $
=. = x
359 ee 38
° e ° 2 ~ °
Sus Sus £05
Subspecies
Elegans 14 3 1
Blanchardi 13 2% 1
Philipi ll 2 1
Expolita 14 3 1%
Noctivaga 11 2 1%
Eburnata q 1% 2
Candida 9 1% 2
Occidentalis 9 ly, 1
Pacata 11 a 1,

KLAUBER—THE GLossy SNAKE

A Key To THE SUBSPECIES OF Arizona elegans

Dorsal scales at mid-body in 31 or 29 rows
Dorsal scales at mid-body usually in 27 rows; not over
20 per cent of the specimens with 29 rows

Ventral scutes in the males usually exceed 210, and 221
in the females; body blotches (not including those on
the tail) usually less than 54

Ventral scutes in the males usually 210 or less, and
221 or less in the females; body blotches (not including
those on the tail) usually 54 or more

Ratio of tail length to length over-all in adult males
usually exceeds 15 per cent and 1342 per cent in adult
females

Ratio of tail length to length over-all in adult males
usually less than 15 per cent and less than 13! per cent
in the adult females

Body blotches usually 51 or less
Body blotches usually 52 or more

Body blotches equal or exceed the interspaces in longi-
tudinal extent

Body blotches of less extent longitudinally than the in-
terspaces

Body blotches less than 45
Body blotches 45 or more

Marks on edges of ventrals; often with spots on infra-
labials; color generally darker

No marks on edges of ventrals; infralabials clear, except
occasionally a spot on the last infralabial; color

generally lighter

Lowest lateral spots on the edges of the ventrals or
the row next to the ventrals; color darker

Lowest lateral spots rarely touch a lateral scale row
below the second above the ventrals; color lighter

One preocular; dorsal blotches at mid-body rarely span
more than 7 scale rows

Usually two preoculars; dorsal blotches at mid-body
usually span 9 scale rows

387

elegans

blanchardi

expolita

philipt
6

8
pacata

occidentalis

noctivaga

occidentalis

eburnata

candida

388 SAN DrseGo Society oF NATuRAL History

ACKNOWLEDGMENTS

I am much appreciative of the assistance I have received from the following
individuals and institutions, for the loan of material and many other courtesies:
Mr. Charles M. Bogert, American Museum of Natural History; Dr. Charles
T. Vorhies, University of Arizona; Mr. Gordon C. Baldwin, Boulder Dam
National Recreational Area; Mr. M. Graham Netting, Carnegie Museum; Mr.
Joseph R. Slevin, California Academy of Sciences; Dr. Howard K. Gloyd,
Chicago Academy of Sciences; Mrs. Kay Kapp, Department of Zoology,
Cornell University; Messrs. Karl P. Schmidt and Clifford H. Pope, Chicago
Natural History Museum; Dr. Ross Hardy, Dixie Junior College; Prof. Donald
F. Hoffmeister, University of Kansas; Mr. Albert J. Kirn, Somerset, Texas;
Dr. Howard R. Hill, Los Angeles County Museum of History, Science, and
Art; Mr. Arthur Loveridge, Museum of Comparative Zoology, Harvard Uni-
versity; Mrs. Helen T. Gaige, Museum of Zoology, University of Michigan;
Messrs. Thomas L. Rodgers and Wade Fox, Jr., Museum of Vertebrate Zoology,
University of California; Dr. William J. Koster, University of New Mexico;
Dr. Arthur I. Ortenburger, University of Oklahoma; Dr. Hobart M. Smith,
University of Rochester; Dr. Emmett R. Dunn, Academy of Natural Sciences
of Philadelphia; Miss Margaret Storey, Natural History Museum, Stanford
University; Dr. Raymond B. Cowles, University of California at Los Angeles;
Messrs. William E. Branch and L. Floyd Keller, Petrified Forest National
Monument; Dr. Doris M. Cochran and Dr. Waldo L. Schmitt, United States
‘National Museum; Prof. Ray Moree, State College of Washington; Mr. Stanley
Mulaik, University of Utah.

I was assisted in making scale counts by Mr. Charles E. Shaw, now of
the United States Marine Corps, and Mr. Lawrence H. Cook. Mr. James Deuel
secured for me a considerable number of specimens of candida, with much useful
collecting data.

I am much indebted to Messrs. C. B. Perkins and Clinton G. Abbott for
valuable editorial suggestions; and to Mr. Leslie C. Kobler for the maps and

photographs.
ABBREVIATIONS

The following abbreviations are used for the museums whose specimens are
mentioned by number in the text.

AMNH_ American Museum of Natural History

CAS California Academy of Sciences

CHAS — Chicago Academy of Sciences

CNHM__ Chicago Natural History Museum

KU Museum of Natural History, University of Kansas
LMK Collection of L. M. Klauber

MCZ Museum of Comparative Zoology, Harvard University
MVZ Museum of Vertebrate Zoology, University of California
NHMSU Natural History Museum, Stanford University
PFNM © Collection of Petrified Forest National Monument
SDSNH_ San Diego Society of Natural History

KLAUBER—ITHE GLossy SNAKE 389

UCLA — University of California at Los Angeles
USNM — United States National Museum
WSC Charles R. Conner Museum, The State College of Washington

SUMMARY

The monotypic genus of snakes Arizona, of the southwestern United States
and northern Mexico, is surveyed. Seven new subspecies are described: A.
elegans blanchardi, A. e. philipi, A. e. expolita, A. e. noctivaga, A, e. eburnata,
A e. candida, and A. e. pacata; these, with the previously described subspecies
A. e. elegans and A. e. occidentalis, make a total of nine. Tail length, scale
rows, ventral scutes, and pattern are found to be the most important characters
in segregating subspecies. Ranges and locality records, ecological and field notes
are given. Relationships are discussed, and lines of descent are suggested. A
key is given.

BIBLIOGRAPHY
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BLANCHARD, FRANK N.
1924. A New Snake of the Genus Arizona. Occ. Papers Mus. Zool.
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Bocourt, Firmin (with DumériL, AucusteE H. A., and Mocquarp, EF.)
1870-1909. Mission Scientifique au Mexique. Les Reptiles, pp. 1-1012.
Atlas.

BoGertT, CHARLES M.
1930. An Annotated List of the Amphibians and Reptiles of Los Angeles
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1933. Notes on the Snake Dance of the Hopi Indians. Copeia, no. 4,
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BOULENGER, GEORGE A.
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Brown, ARTHUR E.
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Burt, CHARLES E., and Hoyie, LUTHER W. :
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CocKERELL, THEOpoRE D, A.
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Cook, Lorenzo H.
1930. Note on an Arizona elegans occidentalis Blanchard. Copeia no. 4,
wel DS:
Cope, eae DE,
1860. Descriptions of Reptiles from Tropical America and Asia. Proc.
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1875. Check-list of North American Batrachia and Reptilia. Bull. U. S.
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1896. The Geographical Distribution of Batrachia and Reptilia in North
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Cougs, ELLIoTT
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Cow es, RAYMOND B.
1941. Observations cn the Winter Activities of Desert Reptiles. Ecology,
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1944. A Preliminary Study of the Therinal Requirements of Desert
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1928. A Tentative Key and Arrangement of the American Genera of
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HANLEY, GEorGE H.
1943. Terrarium Notes on Californian Reptiles. Copeia no. 3, pp. 145—
Le

Hupson, GeorcE E.
1942. The Amphibians and Reptiles of Nebraska. Neb. Cons. Bull., no.
24, pp. 1-146.

KENNICOTT, ROBERT
1859. {Description of Arizona elegans} in Reptiles of the Boundary by
Spencer F. Baird (q.v.).
KLAUBER, LAURENCE M.
1924. Notes on the Distribution of Snakes in San Diego County, Cali-
fornia. Bull. Zool. Soc. San Diego, no. 1, pp. 1-26.
1931. A Statistical Survey of the Snakes of the Southern Border of
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1939. Studies of Reptile Life in the Arid Southwest. Part 1. Night
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Soc. San Diego, no. 14, pp. 6-64.
1941a. The Frequency Distribution of Certain Herpetological Variables.
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1932. Amphibians and Reptiles from Lower California. Univ. Calif. Pubs.
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392 SAN Dreco Society of Naturat History

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1938. The Snakes of San Diego County with Descriptions and Key. Bull.
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1912. Color Standards and Color Nomenclature, pp. iv + 43, plts. 1-53.

RUTHVEN, ALEXANDER G.
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SCHMIDT, Kart P.
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KLAUBER—TIHE GLossy SNAKE 393

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WALLS, GorDON L.

1934. The Reptilian Retina. Am. Jour. Ophthalmology, vol. 17, no. 10,
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Yarrow, HEnry C.

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Nat. Mus., no. 24, pp. 1-249.

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Fig. 1. Arizona elegans elegans.
Adult male from 7 miles southwest of Somerset, Bexar County, Texas.

Collected by A. J. Kirn: Photograph by L. C. Kobler.

Fig. 2. Arizona elegans occidentalis.
Adult female from Campo, San Diego County, California. Collected and
photographed by J. R. Slevin, California Academy of Sciences.

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Fig. 2. Arizona elegans candida.
From Neenach, Los Angeles County, California. Collected and photographed
by C. M. Bogert, American Museum of Natural History.

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TRANSACTIONS
OF THE
SAN DIEGO SOCIETY OF NATURAL HISTORY
Vol. X, No. 18, pp. 399-402, fig. 1. March 29, 1946

DATA AND FIELD NOTES ON THE
DESERT TORTOISE

CHAPMAN GRANT
San Diego Society of Natural History

When a law was enacted in 1939 forbidding the sale of the Desert
Tortoise (Gopherus agassizii) in California, the writer took the oppor-
tunity to obtain what data were of interest from Robert Heckley’s stock-
in-trade of these tortoises at Hodge, California, before they were liberat-
ed to conform with the new statute. He made a trip to Hodge on May
16-17, 1939, and recorded the following data from 119 specimens. This
supplements an article by the writer, published in 1936,' which discussed

366 specimens.
NorMAL ASYMMETRY OF THE LEFT GULAR

Of the 119 specimens, 105 of both sexes had the left gular the larger;
8 females had gulars of equal size and six females had the right gular the larger.

This is in agreement with previous examination of this species.

ScuTE ABNORMALITIES

Out of over 500 specimens, the writer has found only two with abnormal
scutes on the plastron. The 119 examined on this trip all had normal plastrons,
but 24 specimens had abnormalities of the carapace. The normal carapace con-
sists of a nuchal, 11 marginals on each side, a caudal, 5 neurals and 4 costals
on each side.

Marginals—Four specimens had 12 on each side; three had 12 on one
side. Of these seven, 2 had other abnormalities. Three specimens had 10
marginals on each side and one, 10 on one side.

Costals—Four specimens had abnormal costals. One had the fourth
right and left costals split, forming two complete scutes, thus totaling 5 on
each side. Three others had the third on right side, the first on right side and
the fourth on left side, respectively, completely split, forming two scutes. In
addition, the first of these three had an abnormal neural.

Neurals.—Six specimens had abnormal neurals. One had the second
and third completely split transversely, forming two extra neurals; another had

1Zoologica, vol. 21, pp. 225-229, 1936.

400 SAN Disco Society of NATURAL History

the third completely split and also an abnormal marginal; another had the
fourth completely split transversely and two others had the fifth similarly split,
one having an abnormal marginal in addition.

Caudal—Two specimens had the caudal completely split longitudinally.

Nuchal.—One had the nuchal fused with the first left marginal; another
had a completely split nuchal, appearing like two horns.

MISCELLANEOUS OBSERVATIONS

The largest male had lost his gulars by accident or mutilation when
young and there had been no replacement of either bone or scute struc-
ture. It measured 43 cm. across the carapace over the curve and 45 cm.
in length similarly measured. Old males have the gulars recurved and
forked or worn chisel-shape, but the epiplastral bones show no differentia-
tion. Gulars of the females are never recurved; gulars of old males are
about 6 cm. long; of females, about half that length.

The writer went to a location close to Barstow where he had prev-
iously studied these tortoises and found four in about half an hour at
the optimum time, about 9 a.m. One small specimen had fresh coyote
tooth marks on the shell.

Two old Cavalry Officers who served at Fort Grant, north of
Wilcox, Arizona, told the writer of finding specimens near the fort
while quail hunting. This substantiates a locality record that appears to
be east of usual records.

The accompanying drawing (Fig. 1) is of a specimen of Gopherus
agassizii, no. 17056, San Diego Society of Natural History, taken by the
writer near Barstow, California, in September, 1935. Five eggs were dug
from a shallow nest hole, one being cracked in the process. Four normal
young hatched within a few days and the fifth, from the cracked egg, is
the subject of this note. The illustration was for a contemplated paper
for which the writer had assembled considerable material to illustrate his
theory that originally turtles had a median dorsal suture which disappear-
ed upon the forming of the neurals by fusion of four scutes each. The
pigmentation, as well as the divided neurals in this specimen, was a part
of the evidence.

The drawing is used here to illustrate the position of this species
within the egg. The bend or folding is between the sixth and seventh
marginals, as seen from top and side views, and traces of the folding per-
sist for several weeks. The side view shows a large amount of yolk which
had not been absorbed.

GRANT—NOTES ON THE DESERT TORTOISE 401

The young turtles shed a thin skin from all parts of the body short-
ly after hatching.

Coker* is of the opinion that scute abnormalities are frequently
caused in sea turtles when the leathery eggs are moved from their original
nest due to a difference of pressure. The writer points out that Gopherus

Fig. 1. Specimen 17056, S. D. S. N. H. Upper: dorsal
view of carapace, showing great scute abnormality, and pigmen-
tation offering evidence of a tendency toward complete mid-
dorsal suture. Lower: lateral view of same specimen showing
unabsorbed yolk and normal point of folding between sixth and
seventh marginals, which allows embryo to conform to the curva-

ture of the egg shell. (Norman Bilderback, del.)

eggs are hard-shelled and not susceptible to warping due to moving and
yet there is about the same proportion of abnormalities as is found in
other genera.

2Journal of Morphology, vol. 21, p. 68, 1910.

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MUS. CORP. Z00L.
LIBRARY

MAY 16 1949

HARVARD
UNIVERSITY

INDEX

Transactions of the San Diego Society of Natural History, Volume X
Titles of papers and new systematic names are in heavy-faced type.

A Desert Subspecies of the Snake Aphelocoma coerulescens cyanotis, 234
Tantilla Eiseni, 71-74 coerulescens sumichrasti, 231
: sordida colimae, 231
A new Race of Kangaroo Rat from sordida sieberii, 231
the Argus Mountains, California, sordida sordida, 234
BL-132 unicolor unicolor, 231

Arca nucleus, 41

Arizona elegans blanchardi, 313-389
elegans candida, 313-389
69-70 elegans eburnata, 313-389
elegans elegans, 313-391
elegans expolita, 313-389

A new Snake of the Genus Sonora,
from Lower California, Mexico,

A new Wood Rat, Genus Neotoma,

from the Viscaino Desert Region elegans noctivaga, 313-389
of Baja California, Mexico, 307- elegans occidentalis, 313-389
310 elegans philipi, 313-389
elegans pacata, 313-389
Abbott, Clinton G. 89, 119, 127, 154, jani, 315
206, 246, 312, 388. lineaticollis, 315
Acacia greggii, 283 Arnold, Lee W. 176,
Acteon boulderana, 36, 42, 58. Arrington, O. N. 119.

t tilis, 42.
mera : Artemis africana, 40.
Aechmolophus mexicanus, 231.

Aeluroscalabates, 204.

Agelaius gubernator grandis, 232.
phoeniceus gubernator, 235.

Asio stygius lambi, 228.

Atlapetes torquatus virenticeps, 232
Atsatt, S. R. 146, 206, 289, 304.
Ahi E206. Aulophyseter morricei, 35.

Aimophila humeralis humeralis, 235.

rufescens pallida, 233 B
rufescens rufescens, 235 Bailey, Alfred M. 228, 230.
ruficauda acuminata, 233
ruficeps boucordi, 235 Baird, Spencer F. 89, 135, 136, 207, 271,
ruficeps fusca, 235 272, 304, 389.
Aldrich, John W. 223 Balcis conchita, 36, 43, 57.
melanella, 43.
Allen, Morrow J. 162, 166, 206 montagui as.
Amazilia beryllina beryllina, 234 rutila, 43.
Amphispiza bilineata antea, 244 Baldwin, Gordon C., 388.

bilineata bangsi, 237-244
fiGaesca ae 338.244 Bancroft, Mrs. Grifhing, 69.
bilineata carmenae, 240-243 Barbour, Thomas, 90.
bilineata deserticola, 238-243

bilineata pacifica, 243 Bartsch, P. 4, 9, 18.
bilineata sanctissima, 240-244 Basileuterus belli belli, 232,
bilineata tortugae, 238-241 belli clarus, 235
culicivorus brasheri, 235
culicivorus flavescens, 235

Anachis watsonae, 36, 42, 57 rufifons dugesi, 235
Anderson, J. W. 329 Beach, John H. 27.

Amsinckia tessellata, 299

404 SAN Disco Society OF NATURAL History

Belding, L. 304, Cassidix palustris, 232.
Bellophis zonatus, 76. Carpodacus mexicanus centralis, 232
Benson SedeBe249 mexicanus coccineus, 232

mexicanus roseipectus, 232

Berry, S. Stillman, 1-24. Catharus aurantiirostris melpomene, 235.
Bilderback, Norman, 401. mexicanus mexicanus, 234.

Birch. Donald C. 26-27, 38 occidentalis fulvescens, 234.
? . ets occidentalis occidentalis, 232

Blainville, H. D. de, 81, 87-89.

Blake, Emmet R. 222, 223, 230.

Catherpes mexicanus mexicanus, 232.

Centurus chrysogenys flavinuchus, 234.

cea a ILIA Bile 8S, SiO; hypopolius, 234.
SO SU nes Cercidium, 290.
Bornia, 38-39. 2
: floridum, 283
Bornia (Temblornia) triangulata, 36, 39, ;
55. Certhia americana alticola, 234.
r jali iss eo
Bocoureene7 7, asl 207927303045 315: eu bar
389. Chace, E. M. 15.
Bogert, Charles M. 74, 88, 119, 120, 176, Chace, E. P. 15.
ae 207, 300, 304-5, 360, 362, 388-9, Chaetura rutila grisefrons, 234.
Chaenactis sp. 299.
Boulenger, George A. 135-6, 152, 203, :
207 BS 321322) 380: Chama dosin, 40.
Brachydactylus mitratus, 135-6, 138, 199, Charina bottae bottae, 83-90.
210. bottae umbratica, 83-89
a bottae utahensis, 83-90
Branch, William E. 388. brachylops, 85-87
Breese, Paul, 119, 377. plumbea, 86
Bridges, William, 390. Childs, T. S. Jr. 35.
Brown, Arthur E. 207, 389 Chilomeniscus cinctus, 116.
Brown, Wilmot, 222. Chionactis occipitalis annulatus, 359-362.

- occipitalis occipitalis, 371
Buccinum strigilatum, 47. : F
Chrysallida communis, 43.

Bulla cylindracea, 44. rotundomontana, 36, 43, 58.
Burns, A. E. 5. Chuckwalla, 269-306.
Burt, Charles E. 76, 81, 207, 389. SE ILETAE Sees
Gila, 292.
Burt, W. H. 249. Great Basin, 295.

Monserrate, 280.

Peninsular, 286.

Piebald, 288.
Cc Sonoran, 290.
Spiny, 279.
Spotted, 282.

Cistothorus platensis tinnulus, 232.

Cacodaphnella delgada, 47.
Cahoon, John C. 228.

Callisaurus, 362 Clapp, G. H. 4.

ventralis ventralis, 300 Cnemidophorus, 116.
Calocitta formosa formosa, 234 Cochran, Doris M. 206, 275, 304, 388.
Camp, Charles L. 120, 207, 390. Cockerell, Theodore D. A. 390.

Campephilus imperialis, 234. Coker, 401.

INDEX TO VOLUME X 405

Coleonyx, 134-215, 362
brevis, 134-215
dovii, 199, 203
elegans elegans, 134-215
elegans nemoralis, 134-215
fasciatus, 134-215
mitratus, 134-215
variegatus abbotti, 134-215
variegatus bogerti, 134-215
variegatus peninsularis, 134-215
variegatus slevini, 134-215
variegatus sonoriensis, 134-215
variegatus utahensis, 134-215
variegatus variegatus, 116, 134-215

Colinus virginianus graysoni, 231.
virginianus nigripectus, 231
virginianus thayeri, 231

Coturnicops noveboracensis goldmani, 231.

Coluber, 315, 321.
arizonae, 315, 321, 343, 372.
zonatus, 76.

Columbella scalarina, 42.

Conant, Roger, 390.

Conover, H. B. 228,230.

Cook, Lawrence H. 119, 377, 388, 390.
Cook, porn H. See Cook, Lawrence

Cook, Sherburne F. Jr. 63.
Cope ENID89, 1209189, 207, 305, 315.

Coues, Elliott, 119, 120, 208, 390.

Cowles, Raymond B. 81, 88, 119, 120,
124, 206, 305, 360, 362, 390.

Crotalus adamanteus, 118.

cerastes cerastes, 91-126, 149, 371.

cerastes laterorepens, 92-126,
359, 361

cinereous, 113.

durissus, 118.

enyo, 118.

pricei, 118.

mitchellii, 118.

mitchellit pyrrhus, 361

molossus, 118

ruber, 361.

scutulatus scutulatus, 109, 113, 371

tigris, 118

viridis, 118

willardi, 118

Cryptanth sp. 299.

Ctenosaura hemilopha 290.

Cyanocitta stelleri azteca, 231
stelleri coronata, 231

Cyanocorax, 227.
dickeyi, 228.
Cyanolyca nana, 231.
pulchra mitrata, 234

Cylichna alba, 45.
aula, 45.
? loismartinae, 37, 44, 57
ramonensis, 45

temblorensis, 37, 44, 45, 57
Cynanthus latirostris propinquus, 231.

Cytherea (Transennella) conradina, 41.

Dalea fremontii, 299.
Dalquest, Walter W. 63.

Data and Field Notes on the Desert
Tortoise, 399-402

Davis, Dwight D. 392.

Dendroica aestiva brewsteri, 116.
aestiva dugesi, 232
graciae graciae, 235

Dendrortyx barbatus, 231.
macroura diversus, 231
macroura griseipectus, 231
macroura macroura, 231
macroura oaxacae, 231
macroura striatus, 234

Derbonne, William, 152, 208.

Deuel, James, 112, 119, 351, 364.
Dickerson, M. C. 272, 274, 275, 305.
Diepenbrock, Alex, 30.

Dipodomys, 116, 131-2, 264.

agilis peninsularis, 307.
mohavensis, 131-2.

mohavensis argusensis, 131-132
ordii, 332

Dipsosaurus, 362.
dorsalis dorsalis, 300

Ditaxis lanceolata, 299.
Ditmars, Raymond L. 208.
Dodd, C. A. 11.

Donax, 36, 39, 55.
rugosa, 39.

406 SAN Disco Society oF NaturaL History

Dosinia, 40.
(Dosinidia) margaritana, 36, 40,
De

Dryobates scalaris azelus, 234.
scalaris centrophilus, 234
villosus jardinit, 234.

Duges, Alfredo, 208

Dumeril, A. 135, 208, 271, 305
Ducan, Howard, 127

Dunn, Emmett R. 208, 388, 390.

Elaenia viridicata jaliscensis, 234

Empidonax affinis affinis, 234.
afhnis trepidus, 234
albigularis axillaris, 231
difficilis inmemoratus, 231
difhicilis occidentalis, 234
fulvifrons rubicundus, 234

Encelia farinosa, 299.
Engler, Carl, 333-4, 339.
Ephedra viridis, 299.

Epiphragmophora, 4.
(Micrarionta) hutsoni, 4, 18.

Epitonium (Ferninoscala) ferminianum,

46.
Equus occidentalis, 128.
Eremarionta, 3, 5, 7-9, 12, 14.
Eriogonum sp. 299.
Escobar, Pat. 128.
Eublepharidae, 135.

Eublepharis, 135-6, 152, 204.
dovii, 135-6, 199.
fasciatus, 135-6, 182.
variegatus, 185, 208.

Eulima californica, 45.
gabbiana, 37, 45, 57.
migrans, 45.

Eulimella gabbiana, 45.
Eumeces skiltonianus, 63-67.

Euphorbia sp. 283.
polycarpa, 299.

Euphryne obesus, 271, 295.
Eutamias, 132.

Evalea elegans, 49.

Ferguson, Glen C. 29.

Ferminoscala durhami, 37, 46, 58
prunicola, 46.
pseudoleroyi, 46.
spathe, 46.
whitei, 37, 46, 58.

Ferriss, J. H. 4, 19.
Festuca sp. 283.

Fitch, Henry S. 78, 81.
Foraminifera, 35.

Fossil, 30, 35

Fouquieria splendes, 299.
Fox, Wade Jr. 388.
Franseria dumosa, 299.
Frazer, M. Abbott, 242.
Frick, Childs, 128, 130.
Friedmann, Hubert, 223.

Gadow, Hans, 135, 199, 208.
Gaige, Helen T., 206, 209, 388.
Garman, Samuel, 390.
Gastropoda, 42.

Gecko, Black Banded, 182.
Central American Banded, 199.
Colima Banded, 195.
Desert Banded, 138.
San Diegan Banded, 154.
San Lucan Banded, 160.
Santa Inez Island Banded, 167.
Sonoran Banded, 162.
Texas Banded, 184.
Tucson Banded, 176.
Utah Banded, 171.
Yucatan Banded, 191.

Geckkonidae, 135.
Geococcyx velox velox, 231.

Geothlypis chapalensis, 232.
nelsoni nelsoni, 232.
speciosa, 232.
trichas melanops, 232.

Gila Monster, 152.
Girard, Charles, 89.
Githens, Thomas E., 120.

INDEX TO VOLUME X 407

Gloyd, Howard, 112, 120, 206, 304-5,
388.

Goldman, E. A., 222-4, 247, 307, 309,
341.

Gopher, Pocket, 131-2, 245-268.
Argus Mountains Pocket, 132.
Cape San Lucas Pocket, 265.
Catavina Pocket, 262.

Coastal Pocket, 261.

El Cajon Canyon Pocket, 258.
El Rosario Pocket, 261.
Imperial Valley Pocket, 252.
Jacumba Pocket, 254.

Laguna Salada Pocket, 252.
La Paz Pocket, 264.
Magdalena Bay Pccket, 263.
Magdalena Island Pocket, 264.
Magdalena Plain Pocket, 263.
Pattie Basin Pocket, 252.
Perplexing Pocket, 259.
Punta Prieta Pocket, 262.
San Angel Pocket, 263.

San Borjas Pocket, 262.

San Diego Pocket, 258.
Sangre de Cristo Pocket, 256.
San Matias Pass Pocket, 257.
San Pedro Martir Pocket, 257.
Sierra Juarez Pocket, 255.
Sierra Laguna Pocket, 265.
Tawny Pocket, 253.

Gopherus, 399.
Gopherus agassizii, 399-401.
Gordon, Kenneth, 78, 81.

Grallaria guatimalensis — ochraceiventris,

234.
Grant, Chapman, 399.
Gray, J. E. 209.
Greenberg, B. 209.

Grinnell, Joseph, 132, 209, 238-9, 250,
254, 390.

Growth in the Western Blue-Tailed
Skink, 61-68

Gunther, Albert C. 203, 209, 315, 391.

Gymnodactylus coleonyx, 135-6, 191.
scapularis, 135-6, 191.

Hall, E. Raymond, 249.

Hallowell, Edward, 93, 121.

Hanley, George H., 377, 391.

Hanson, Harold C. 222-3, 230.

Hardy, Ross, 119, 171, 173, 175, 206,
209, 304, 388.

Harter, Samuel G., 131.

Hartweg, N. 199, 209

Hastula gnomon, 37, 47, 57.
jamaicensis, 47.
homala, 47.

Hay, Oliver P., 130.
Hays, Robert P., 88.

Heleodytes jocosus gularis, 232.
jocosus jocosus, 234.
megalopterus megalopzerus, 232.
megalopterus nelsoni, 232

Helminthoglypta, 6.
(H. fisheri), 5.

Hemipenes, 88, 118, 323.
Hemitheconyx, 136, 204.

Henicorhina leucophrys festiva, 234.
leucophrys mexicana, 232

Hill, Howard R. 74, 81, 88, 119, 206,
250, 388.

Helix, 4, 6, 8, 19.

Hoard, Robert S. 304.
Hoffmeister, Donald F. 388.
Hooper, Emmet T. 247-8.
Hoyle, Luther W. 389.

Huey, Laurence M. 131, 227, 242, 245-
268.

Hypsiglena, 316.
ochrorhyncha ochrorhyncha, 361.

I
Ingles, L. G., 5.

J
Jaeger, Edmund C., 5, 11, 15.
Jan, G:,°89:

Johnson, Murray L., 78, 81.

408 San Dreco Society OF NATURAL HIsToRY

Kapp, Kay, 206, 388.

Keen, A. Myra, 25-60.
Keller, L. Floyd, 388.
Kellogg, Remington, 35.
Kennicott, Robert, 320, 391.

Klauber, Laurence M. 69-126, 133-216,
246, 271, 282, 292, 304, 311-398.

Klauber, Philip M., 119, 334.

Kleinpell, R. M., 35.

Krauskoph, Konrad, 27, 30.

1G;
Lamb, Chester C., 222-3, 228, 230.

Lampornis amethystinus _amethystinus,
231.

Lamporpeltis getulus californiae, 361,
Byles

multicincta agalma, 76-80
multicincta herrerae, 76-80
multicincta multicincta, 76-81
multicincta multifasciata, 76-80
pyromelana, 79
zonata zonata, 76, 81

La Rivers, Ira, 209.
Larrea divaricata, 299.
Lathrogecko, 210.

Lazier, Edger L., 122.
Leonard, Arthur B., 392.
Lepidoblepharis, 204, 210.

Lepidocolaptes leucogaster _leucogaster,
234.

Leptosyne bigelovii, 299.
Leptotyphlops humilis cahuilae, 73, 361.

humilis mumilis, 73
Lepus insularis, 244.
Lewis, Thomas H., 81.
Lichanura, 87.

roseofusca roseofusca, 361.
Linsdale, Jean M., 79, 81, 209, 360, 391.
Bittle; Es ies Jr, 209!

Lizard, Fringe Footed Sand, 116.
W7hineatletiG!

Lockington, W. N., 82.

Lophortyx douglasii teres, 231.
Lotus strigosus, 299.

Loveridge, Arthur, 206, 210, 388.
Lowe, Charles H. Jr., 74, 116, 121.
Lucina nassula, 40.

Lucinisca menuda, 36, 40-1, 56.
nuttalli, 40-41.

Lumholtz, Carl, 226, 288.

Mangelia (Cacodaphnella?) kernensis, 37,
47, 58.
densilineata, 48.
striolata, 47.

Marr, John C., 332, 391.
Martin, Lois T., 27, 38, 44, 53.

Masticophis, 318.
flagellum piceus, 152, 361, 371.

McKee, E. D. 210.
Meek, Seth E., 121, 210.
Megarynchus pitangua caniceps, 231.

Meleagris gallopavo gallopavo, 231.
gallopavo mexicana, 234

Melospiza melodia adusta, 233.
melodia pectoralis, 233

Memmler, Viola H., 63-68.
Menzies, Robert J., 119.
Merriam, C. Hart, 121, 230.
Merriam, J. C., 128-130.
Mertens, Robert, 316, 391.

Micrarionta, 4-6, 18, 19.
avawatzica, 16-17.
borregoensis, 9.
desertorum, 5.
(Eremorionta) avawatzica, 16.
(Eremorionta) borregoensis, 8, 9
(Eremorionta) micrometalleus, 6
harperi, 12.
(Helminthoglypta?) _ micrometall-
eus, 6.
micrometalleus, 6.
ora, 12.
reedi, 9.
rexfordi, 14, 17, 19.

INDEX TO VOLUME X 409

Micrurus fulvius tenere, 327.
Miller, Alden H., 88.

Mitrella communis, 48.
flaminea, 48.
lepta, 48.
lissa, 48.
(Mitrella) anchuela, 37, 48, 57.
(Striomitrella) tenuilineata, 48.

Mitrephanes phaeocercus phaeocercus, 231.
Mocquard, F., 210, 273, 305.

Mohavelix, 6.

(micrometalleus), 2.

Mollusk, 25-60.

Moniliopsis electilis, 37, 49, 57.
graciosana, 49.
incisa, 49,

Moore, Robert T. 217-236
Moore, William, 154.
Moree, Ray, 388.

Mosauer, Walter, 114, 115, 121, 122,
399% 392.

Mulaik, Stanley, 206, 210, 388.
Muller, Lorenz, 316, 391.

Murex attenuatus, 47.
scriptus, 48.

Neochloe brevipennis brevipennis, 232.

Nelson, E. W., 222, 225, 227, 307, 309,
341,

Neotoma bella felipensis, 309.
intermedia gilva, 309.
intermedia intermedia, 308-9.
intermedia notia, 309
intermedia ravida, 309
lepida arenacea, 308
lepida egressa, 307-310
lepida felipensis, 308-310
lepida gilva, 309-310
lepida intermedia, 308-310
lepida lepida, 307-309
lepida molagrandis, 307-310
lepida pretiosa, 308
lepida ravida, 308-310

Netting, M. Graham, 119, 206, 388.

New Mollusks from the Round Moun-
tain Silt (Temblor) Miocene of
California, 25-60

New Occurrences of Fossil Tapir In
Southern California, 127-130.

Noble, G. K., 135, 210.

Nucula (Ennucula) birchi, 36, 41 55.
obliqua, 41.
postangulata, 41
taphria, 41
washingtonensis, 41.

Nyctisaura, 207.

Odostomia (Chrysallida) melanoides, 44.
(Chrysallida) pulcia, 44
(Chrysallida) torrita, 43
(Evalea) andersoni, 37, 49, 57-58
(Evalea) californica, 49

Olivella quadriplicata, 50.
ischnon, 37, 50, 57.
pedroana, 50.
purpurata, 50.

Oliver, J. A., 209.
@lsonseAt As, 275.5355:

On the Generic Relationships of Cer-
tain Californian Xerophile Snails,
1-24

Opuntia, 280, 290.

Oreortyx picta eremophila, 132.

Ortenburger, Arthur I., 181, 210, 349,
388, 392.

Ortenburger, Ruth D., 181, 210, 349,
392.

Otophanes, 227.
mcleodii, 228.

Otus asio sortilegus, 223, 231,
asio suttoni, 234
flammeus flammeus, 231.
guatemalae cassine, 231.
vinaceus seductus, 223, 231.

Osprey, 280.

410 San Dieco Society oF Natura History

Pachycereus pringlei, 290.
Pack Eiea|ee2 10:

Parker, H. W., 189, 210.
Parus sclateri sclateri, 231.
Pelecypoda, 38.

Penner, L. R., 283.

Perkins, C. B., 89, 94, 116, 122, 206,
304, 362, 388, 392.

Perkins, Cyrus S., 71.

Perognathus, 116, 264.

Peters, James L., 82, 223, 237.

Peters, W., 135, 200, 203.

Phacelia sp., 299.

Philortyx fasciatus, 234.

Phrynosoma platyrhinos platyrhinos, 116.

Phyllorhynchus, 152, 316.
decurtatus perkinsi, 109, 361,
Bille

Pilsbry,;\ TH Ax, 3-6,29-417-19)

Pipilo fuscus fuscus, 232.
fuscus toroi, 232
torquatus alticola, 232
torquatus nigrescens, 232

Piranga bidentata bidentata, 232.

Pituophis catenifer, 355.
catenifer afhnis, 359, 361-362
catenifer annectens, 361-362
catenifer deserticola, 109, 371
deppei, 315-316.
elegans, 315, 328, 340, 343.,.
lineaticollis, 315-316.

Pleurotoma elaborata, 49.

Pope, Clifford H., 206, 388.

Preliminary Studies on the Black-
Throated Sparrows of Baja Cali-
fornia, Mexico, 237-244

Psaltriparus minimus iulus, 234.
minimus melanotis, 234

Pseudogonatodes, 210.
Pulogonys cinereus cinereus, 232.
Purpura tubifer, 52.

Pyrgolampros mioperplicatus, 52.

Quail, Desert Mountain, 132.

Rallus longirostris tenuirostris, 231.

Rat, Kangaroo, 131, 132.
Argus Mountains Kangaroo, 132.

Rat, Wood, 307-310
Viscaino Desert Wood, 307.

Rattlesnake, Horned, 93.
Reed) Bred) My 535 Iie 12)
Regulus satrapa aztecus, 232.
Reynolds Fletcher A., 377, 392.

Rhinechis elegans, 315, 316, Sy2il SVaeh.
343, 350, 372

Rhinocheilus lecontei clarus, 361
lecontei lecontei, 371

Ridgway, Robert, 70, 80, 122, 145, 158,
175, 180, 210, 261, 294, 326, 348,
392.

Rixford, Emmet, 15.

Rodgers, Thomas L., 63-68, 73, 81, 88,
119, 206, 304, 388.

Roe, Anne, 98,122.
Rose, J. N. 290.
Roveglia, Pablo, 222.

Ruthven, Alexander G., 85, 90, 195,
211392.

S
Safford, Vernon, 131.

Salpinctes obsoletus notius, 232.
Salvadora hexalepis hexalepis, 361, 371.
Salvia columbariae, 299.

Sanderson, I. T., 195, 211.

Saunders, H. F., 152, 211.

Sauromalus ater, 270-303
australis, 270-303
hispidus, 270-303
interbrachialis, 272, 274-275, 284
klauberi, 270-303
obesus obesus, 270-303
obesus townsendi, 270-303

INDEX TO VOLUME X 411

obesus tumidus, 270-303
slevint #270308
varius, 270-303

Scalopus aquaticus texanus, 327.

Sceloporus, 212, 225.
occidentalis biseriatus, 377.

Schenck, Hubert G., 27, 35.

Schmidt, Karl P., 136, 203, 206, 211,
272-4, 280, 290, 304-305, 388, 392.

Schmitt, Waldo L., 388.

Schuchert, Charles, 219, 220, 230.
Searle, Clyde, 83.

Semeloidea donaciformis, 39.

Senn, A., 34.

Shaw, Charles E., 71, 94, 119, 181,
269-306, 333-4, 339, 343, 388.

Sialia mexicana australis, 232.
mexicana mexicana, 235

Sidewinder, 91-126.
Colorado Desert, 94.

Simmondsia californica, 256.

Simpson, G. G., 98, 122.

Sittasomus griseicapillus jaliscensis, 231.
Skink, Western Blue-tailed, 61-68.
Slater, James R., 81.

Slevin, Joseph R., 73, 80-1, 88, 117, 119,
122, 206, 304, 362, 393. 395.

Smith, Allyn G., 5.

Smith, Hobart M., 74, 136, 185, 188-9,
195-6, 206, 211, 225, 230, 388, 392.

Smith, Maxwell, 53, 136, 211.
Snail, 3-5.

American, 4.
Desert, 3, 5.
Xerophile, 1-24.

Snake, Arizona Glossy, 343.
California Black-headed, 71-74
California Glossy, 372.
Chihuahua Glossy, 340.
Coast Range Coral King, 76.
Coral King, 75-82.

Desert Black-headed, 71
Desert Glossy, 350.
Great Basin Rubber, 86.
Kansas Glossy, 328.

Pacific Rubber, 85.

Painted Desert Glossy, 333.
Peninsula Glossy, 379.

San Pedro Martir Coral King, 76.
Sierra Coral King, 76.

Southern California Rubber, 83.
Texas Glossy, 320.

Todos Santos Coral King, 76.
Utah Rubber, 85.
Western-Mojave Glossy, 364.

Sonora bancroftae, 69-70.
mosaueri, 69.
occipitalis, 116.
occipitalis annulata, 109
semiannulata semiannulata, 69-70.

Sordelli, F. 89
Sonorella, 4-9, 18.

desertorum, 4.

ferrissi, 7.

hachitana, 6.

(Mohavelix) micrometalleus, 6, 7,
20.

wolcottiana, 18

Sonorelix, 2, 8, 9.
aetotis, 9, 15-16.
avawatzica, 2, 9, 16, 17, 20-21
baileyi, 9, 17.
borregoensis, 2, 9, 10, 12-15, 20.
borregoensis ora, 2, 12-14
carrizoensis, 9.
depressispira, 9, 15.
eremita, 9, 17.
harperi, 9, 13, 14.
melanopylon, 9.
ora, 9.
orcutti, 9.
rixfordi, 2, 9, 14-17, 20, 22.

Sparrow, Black-throated, 237-244.
Bangs Black-throated, 239.
Carmen Black-throated, 243.
Desert Black-throated, 238.
Espiritu Santo Black-throated, 244.
San Esteban Black-throated, 242.
Tortuga Black-throated, 243.

Sphaeralcea munroana, 299.
Sphaerodactylus, 210.

Spizella passerina atremaeus, 235.
passerina mexicana, 235

Stebbins, R. C., 122.

Stejneger, Leonhard, 74, 82, 87, 90, 122,
13.62 2 272.298: 3.05.

Stenodactylus variegatus, 135, 138, 184.
Stephens, Frank, 211, 359.

412 San Deco Society oF NaAturAL History

Stewart, R., 42.

Stickel, William H., 70.
Stirton, R. A., 130.
Stock, Chester, 127-130

Storey, Margaret, 81, 88, 106, 119, 206,
388.

Strecker, John K., 190, 212, 392.

Strombiformis albus, 43.
glaber, 45.

Stuart WesiG 195712"
Stull, Olive G., 85, 90, 392.
Sumichrast, F., 212.

Syrnola gracillima, 50.
marylandica, 51.

scandix, 37, 50, 58.

Talityphis, 32-34.
Tanner, Vasco M., 85-7, 90, 119.
Tantilla eiseni eiseni, 71-74

eisenit transmontana, 71-73, 361
eisent utahensis, 72

Tapirs, Fossil, 127-130

Tapirus haysii californicus, 128-130.
haysti haysti, 128.
merriami, 128.

Taylor, E. H., 67, 136, 166, 182, 195,
212, 304-5, 332, 393.

Teinostoma calvertense, 51.
depressum, 51.
nanum, 51.
politum, 51.
sapiellum, 51.
sublimatum, 51.

(Teinostoma?) lens, 37, 51, 57.
Temblornia, 34, 38, 39.

Terebra (Hastula) patuxentia, 47.
luctuosa, 47.

Thamnophis hammondii, 361.
The Chuckwallas, Genus Sauromal-

The Coral King Snakes of the Pa-
cific Coast, 75-82

The Geckos of the Genus Coleonyx,
with Descriptions of new Sub-
species, 133-216

The Glossy Snake, Arizona, with
Descriptions of new Subspecies,
311-398

The Pocket Gophers of Baja Cali-
fornia, Mexico, with Descriptions
of Nine new Forms, 245-268

The Sidewinder, Crotalus Cerastes,
with Description of a new Sub-
species, 91-126

The Subspecies of the Rubber Snake,
Charina, 83-90

The Transverse Volcanic Biotic Pro-
vince of Central Mexico and its
Relationships to Adjacent Pro-
vinces, 217-236

Thomomys, 132, 245-268.
albatus, 252
aphrastus, 248, 259
bottae, 247-260
bottae abbotti, 257-266
bottae affinis, 254-266
bottae albatus, 252-266
bottae alticolus, 265-266
bottae anitae, 265-266
bottae aphrastus, 257-266
bottae argusensis, 132
bottae borjasensis, 262-266
bottae cactophilus, 262-266
bottae catavinensis, 262-266
bottae cunicularius, 252-266
bottae imitabilis, 264-266
bottae incomptus, 263-266 —
bottae jojobae, 256-266
bottae juarezensis, 253-266
bottae leucodon, 254
bottae litoris, 263-266
bottae lucidus, 252-266
bottae magdalenae, 264-266
bottae martirensis, 247-266
bottae nigricans, 248-266
bottae proximarinus, 261-266
bottae puertae, 255
bottae russeolus, 263-266
bottae sanctidiegi, 258-266
bottae siccovallis, 258-266
bottae xerophilus, 257-266
fulvus alticolus, 265
fulvus anitae, 265

INDEX TO VOLUME X 413

fulvus martirensis, 257
fulvus nigricans, 253
perpallidus, 250

Thalurania furcata ridgwayi, 231
Thayer, W. N., 219, 220, 230.
Thryomanes bewickit bairdi, 232.

bewickiit murinus, 234
bewicktt percnus, 232

Thryothorus felix grandis, 232.

Toro Avilés, Mario del, 222

Tortoise, Desert, 399.

Tortrix bottae, 85.

Towhee, Argus Mountains Brown, 132.

Toxostoma curvirostre celsum, 234.
curvirostre curvirostre, 232
curvirostre deflexum, 232
curvirostre vetula, 232
dorsale dumosum, 234
ocellatum, 232

Transennella joaquinensis, 36, 41, 55.
Trimorphodon vandenburghi, 361.
Trochilidae, 223.

Troglodytes brunneicollis nitidus, 232.
Tropidocarpum gracile, 299,

Turbo plicatus, 49.
subulatus, 45.

Turbonilla plicatula, 51.

(Pyrgiscus) bravoensis, 36, 51,
52, 57-58

(Pyrgiscus) dominicensis, 52

(Pyrgiscus) interrupta, 52

(Pyrgiscus) virgo, 52

(Pyrgolampros) halibrecta, 52

(Pyrgolampros) mariposa, 37,
52, 57-58

typica, 51

Turdus infuscatus, 234.
migratorius permixtus, 234.
migratorius phillipsi, 234

Typhis alatus, 33, 53-54
alatus obesus, 33
expansus, 53
latipennis, 53
liguniferus costaricensis, 54
obesus, 53-54
pterinus, 54
(Talityphis) alatus obesus, 54, 56
(Talityphis) costaricensis, 27, 54
(Talityphis) expansus, 53, 56

(Talityphis) lampada, 37, 53-54,
56

(Talityphis) latipennis, 56
(Talityphis) olssoni, 27, 54
(Talityphis) pterinus, 33

Uma, 116, 362.

Uta stansburiana hesperis, 362, 377.
stansburiana stejnegeri, 116

Van Denburgh, John, 82-3, 85, 90, 122,
136) 212, 213) 27122. 280) 289. 305.
593%

Van -Rossem, A. J. 228-230, 237, 244
Venus concentrica, 40.

Vermivora superciliosa mexicana, 235.
Voluta dama, 50.

Vireo gilvus amauronotus, 232.
gilvus eleanorae, 232
huttoni mexicanus, 235.
nanus, 232.

Vireolanius melitophrys goldmani, 232.

Volvulella californica, 54.
cylindrica, 54.
gluma, 37, 54-5, 57.
marylandrica, 54.
oxytata, 54

Volvula oxytata dodona, 55.
rostrata, 34.

Vorheis, Charles T., 152, 180, 206, 213,
388.

Walls, Gordon L., 135, 213, 316, 393.
Warbler, California Yellow, 116.
Watson, Elizabeth A., 27, 38, 43, 53.
Weddle, H. W., 130.

Wellborn, V., 213.

Wenona isabella, 85-87.
plumbea, 85-87.

414 SAN Dreco Society oF NATURAL History

Werner, Franz, 136, 213.
Wettstein, Orto, 136, 213.
White, Robert T., 27, 38.
Willett, George, 19, 223, 237.

Woodall, Harold T., 116, 119.

Woodbury, A. M., 213.
Woodring, W. P., 42, 47, 128.
Wright, A. H., 206.

Xantusia henshawi, 159.

Xenospiza, 227.
baileyi, 228.

Xiphocolaptes promeropirhynchus sclateri,
PPB IANG
ny.

Yarrow, Henry C., 82, 213, 393.
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