ee ens HARVARD UNIVERSITY LIBRARY MUSEUM OF COMPARATIVE ZOOLOGY ay, iy 7 io AZT Oe een ‘ a a¢ Hi ¥ hE : ‘ ni G4 hn, , ur " nH q ae ; ik ue } id s } . ae) er ey 7a heed 4 iz > % "' > o ade? i: : ‘ r ‘ ” Re RSET OARS PASS CYe'y h A eg ei DAs iA cA, ay bY : y - ret a : wl, 4] : rmihy ‘ ; ya i ; A. (" hfe pt eS. 1; ; ie Ce eialy, em ha i Lil if iAg, We . at j ' fis ' fi si u 2 : . we ; ' ’ ‘ bi wy, AD dey Ne wee Pur Ais deg a ; Dm,” Ve, 4 ak Aah i it oe F [ " L% yy P a 7" ‘ ; if . i? ‘ 7 . ' wh, i , . ) L ’ | ye Ua : " whit mJ ry whi : i fd rae Tt Wa) PS. ee ? Ve Ty iy ¥ Pad 'f +. ‘ : (Gs iis ely) A j I a i) ike | 4 4 : me iy WY ths . J 1 3; f A) vie %) fan's ey es ; x { ne ih | : > hi i i” 1 \ 1 14 \ h , { Pi Vi vistee ee ht aN ia, ats ms ; sa) cy, 7A ‘Ok i PURER Jp hy ie) Voki iy if rite) ae 4 ct hy Fhe MUS. COMP. Z00L. LIBRARY MAY 16 1945 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 co 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 i iy al erres 2 oa (ie, ae et at % bei de weil <>" ' Wei, r mie eee E : i cae lee G RAN, t * «. a ree ae Tig ‘ aad Sei ihe Tagg , i, agi an oak = ell ah apie oe Eh a hile SD agen me Se lhe ae Weas ahs 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 BY 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. 18 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 “gehen, EE ae 56 ht PLATE holes te ae ee N ae ye TRANSACTIONS OF THE SAN DIEGO SOCIETY OF NATURAL HISTORY VotuME X, No. 2, pp. 25-60, plates 3-4, figs. 1-5 Zoolteay o® a \ JAN 141944 Bee LLL) NEW MOLLUSKS FROM THE ROUND MOUNTAIN SILT (TEMBLOR) MIOCENE OF CALIFORNIA BY A. Myra KEEN Stanford University SAN DIEGO, CALIFORNIA ~ PRINTED FOR THE SOCIETY DECEMBER 30, 1943 mn of Comps, > iy Ka Zooteay £8, 97 YAN 14 1944 ri LIBRARY ~ KRY Z Tem A { Z (Uy 4@ ROUND MTN NS pp Vij. Tr XN 2 eee \ \rs - -\\ \ - aN \ \ a Nee Ve 2 ND \ x» x heed OP ic Se Re N A- mS Ri »~ \ » O QUATERNARY TERRACES Kern River SERIES x D CHANAC SANTA MARGARITA | D 4 Pe ES RouND MouNTAIN SILT siete lel alee atebcletsatetal elie OLCESE SAND a ac a) fe} JEweETT SILT a < WALKER BASEMENT COMPLEX TRM Tmu| 4 To? UNDIFFERENTIATED Tr MioceNe 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 ferewecom| ruse | < DEL MONTIAN c < <¥ ea 4 Foe St <3 “ w ° x v~ Ww - = z * ° LJ = > Zz. LUISIAN > EE ns aoe £ uw RE AN < LJ ene ee aly v O ° = of > a= a) > Wa eke as Ser a mo, w = aS a ~ WW w = = x 2 se Ee VAQUEROS OLIGOCENE “KREYENHAGEN’? N EOCENE MH oMENGINE-IONE” MEASURE - OF 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 3] 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 c | EUROPEAN ace | cutr coast | panama |TRIDAD [SANTO Lamaca] KERN raver area | EUROFER “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 ns OAK GROVE oe CHIPOLA BURDIGALIAN MIOCENE OLCESE SAND TEMBLOR FREEMAN-JEWETT AQUITANIAN SILTS PYRAMID HILL SAND OLIGOCENE CHATTIAN ue 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. A 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. 36 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
  • 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 37 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 38 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 " Zeoleay ud OX SgAN 251944 Ligra®* GROWTH IN THE WESTERN BLUE-TAILED SKINK BY 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 IOYIIITG “TLLOIY “9 NOLNITO *que4xe auou 9q er1ey4 4eU4 SNoOTKUe oe om S@ ‘AAOO @ATJOSTAp aNnoOLk AOLIYSeP eseaTqF *zayiz0 3u4 TOF aynzTysqns nok yey, ysenbaer am yotym ‘Adoo yooszaed eB uytmerey nox Sutpues are ay ‘*pesoetdstu 99 esed uo SeutyT doz ee144 9U4 PeY PeTTeU or9Mm 4eY4 ONSST aAO0Qge 34 JO setdoo 4ysartz oyy ‘Surqutad ut zozsze ue ysnoruy, "OUT CATIVN “S$ “ON ‘X “TOA ‘AMOLSIH TWUNLYN JO ALZIOOS OOHIA NYS FHT FO SNOILOVSNVEL FO SENEIdGIOWY OF thH6T ‘OT Arenuep ¥L81 GALVYOdHOONI AMOLSIH TWHNLVWN JO ALHIDOS OSFIC NWS SHL A@ GaivdadO GNVY GaqNNO04 VINUYOALIVD ‘OOTIG NVS Mivd VOWIva WOAASOIA AMOLSIPET TV UAL N gonii cod eotid odd 8 di iwored. soy gitbase dts 68 6b yt A Rs Simi, ph Sa A Pah mt 1ot odutitedia Koy” hl Oh EAR oe aw ga .veos evkjosteb suey “etd: 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 aS 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 < 10 67 5 5 107 96 TABLE 3 Distribution of Ventral Scutes in the Type Series MALES FEMALES laterorepens cerastes laterorepens 78 eR NONMVONM VO WN 58 RBS MOM UD BWP He 29 me Mm UW OW DHA DM KN WNW LO 38 99 100 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 «< Fic. 2. Crotalus cerastes laterorepens. Adult male from Borego Valley, San Diego County, California. eee _— = (‘sajmop “gq puoudey 3G Aq ‘erusosyed ‘AIunoD, eHeduy ‘aBnzoy FT PEA &9S UoIeS “yreg Apueg ie saunp pues ayi ul uaxPI ydeas0.04q ) "saqaeq Aq apeur asaM sea} ]pews ayy “pred 243 Aq syzeul gaddn ay ‘pray ey3 Aq apeul a1aM (jensn ueyi padeys-[ ssa] qoyies) syIEUI JAMO] FY T, ‘qyst4 01 Ia] worz Surpaarsy JapulMmapis ynpe ue Jo xe sy] 9 ALV1g : ; YAGNIMAGIG MANN WINVIY ae a “ Nga! Me ds Fetes re reat a es ty Ps 2 pata i ore Nee is man 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 BY 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. A 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. 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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 the Savanna Region of Central Petén, Guatemala. Univ. Mich., Mus. Zool., Misc. Pubs. No. 29, pp. 1-56. SUMICHRAST, F. 1880. Contribution a |’Histoire Naturelle du Mexique. I. Notes sur une Collection de Reptiles et de Batraciens de la partie occidentale 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. Sci. Bull., Vol. 22, no. 9, pp. 203-205. 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 Sinaloa. Univ. Kan. Sci. Bull., Vol. 24, no. 20, pp. 505-537. 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- tion of a New Species of Sceloporus. Proc. Acad. Nat. Sci., Phila., 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. 109 430 S3ID3dS Y¥V INDYAENL-NON 3O SJONVY ONIMOHS ODIXSAW NYSHLYON ONV S31VLS GALINA NY3LSSMHLNOS JO dvW ree yiLLOBav de G oo 53 35953596353 XN ef we, ° 5 se ¢ : SNLVOSINWA PESES: < Ay, “LLOESV -suedalq sijes0waua’y suegajaa’) SN4e4yIW "9 xKu0a]04% JO S3ID3dS YVINDYIAGNL 40 S3ILITVDO1 ONIMOHS VWOINAWNV TIWYLN3AD OGNV ODIXSAW NYSHLNOS JO dVW eG ee af a ev > wiht Os tury. eA ca bias the fs ae ed * ern t i as 7 © > : ' ' ‘a t = 4.9 5 SY j ie nl : 4 ’ ’ Le f i. : 1 ‘ y ‘ a 4 Le 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 BY 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* Z Bid Ko < 2 g $Y g Ropert T. Moore ( sek 12 19409 K 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. x 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 Common to Transverse Volcanic and Sierra Madre Occidental ALIGRTO TOU EES eee eee area et Kel te ce Sener ee SW en 8 Common to Transverse Volcanic and Sierra Madre Oriental ATC enOOtier sues & Antenne. HE ee ery PR. 9) if Common to Transverse Volcanic and Sierra Madre del Sur PIGUET BOC INCES to teens Oi lara Rie ie Fela ei coe atta k= gO Miel 15 Common to Transverse Volcanic and Chihuahua and no others.................. a Common to Transverse Volcanic and Veracruz and no others.................... 1 Common to Transverse Volcanic and Tamaulipas and no others............. 1 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 Common to Transverse Volcanic and all three Sierra Madres TEMTLOMEC A DOV Ce eee pte NM lee LR nal Mee. ee ont 0 5 ae 4 Pye tag se & ae ae gee Ue ee) MK: : Lv, , - ‘4 ut $ 7 ~~ alin ; - Pay / ; ; = =A ine re ee hele il te ee vse ‘Peay ain x ‘ a Bee oo rig is pea a » . ey We ? pe Peay i PO ee ay rv ae ae Bris va 5 an ve wane’ 4 Aah aes it 3 pene te ait re ee sont: ves i ue ¥ =H hae Ale ier ie Aisbeti hee ane co ie ie er, igre mG am ate 4 ee mere Vee 3 ae buenas ah oye fa 7 eats Be arate ete) 9 Py a4 Pa yy . at A ee: } A net Ti % mat ae, we es = i, SAA Bis 7s aes Raps 1. 2 - iD ys us, | be A a nf oe » Ta Sager vale oe en ee te shan Ei) Heavnh oy nee het aa ii it edi +eawy 4 a ie es ae 7 at Ba als Sere pe. i ¥ ts Be Cy mt RN ’ Q Throats on y esa Pre ti ne ie mys a a Pens wa ee ee | ‘Sia Meade 16 ei endo ent ee ni : : 7 } ‘ga aa ia id mer ye t ee aap , fa i } i es, pee wre io wily iy Ne id in j 7 ihe 7 Ba Sirs. ag aa ch hgh in may wy Ps Meet oe A - “f, 2, S| : ‘Si “1 "ae rey ree or , nA . t . ~ u ’ j ] we r . ; £ ~ ~s 1d . 7 i \ a 4 = “ty 7 =a — : 7 1 a 7)? Ba 4 1 ry 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 20 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. 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 aN a -E AA © 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 ce 3i¢ 30° 29° ie 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). o0omAnN nur ff WN — So Ye yey WwW rR Se Se SS — Or SI Cy => 19. 20. 21 Zz. 2S. 2A: 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. 117° ° o © 3 Ho Hi LIFOI RNI Ad. eae : ARIZONA 33 1.2: 32° 131° 30° ‘ SONORA ' ‘ ef 4 ° 29 128° 27° 26° lose DISTRIBUTION MAP Yet NF Virl= IPeOoCcigaw Gores OF BAJA CALIFORNIA, MEXICO Sy TYPE LOCALITIES ‘ p24" SPECIMENS EXAMINED BY AUTHOR___¢ 2310 & BROKEN LINE SHOWS APPROXIMATE BOUNDARY OF ' RANGE, WHEN KNOWN FROM OTHER THAN TYPE c LOCALITY. \ , FOR NAMES INDICATED BY NUMBERS, SEE LIST ON 23° OPPOSITE PAGE. 116° 5° 114° 13° W2° te 110° E : pga an MU Ooze Sar rainy ” ‘en ml Be "7 ee Eee ee as Ain es ois pnb a mtn mnt by tenn pam ttn Senna math ‘ Be : ™ : my ee a bie ue ey f By sane 9 ricaiinee se er -SnTUR OS SSHIMART peigegtiges ery epernien STAMEXOR TAA» uert WAHT ira: REE. ad “Vcr ees 3% eran we arroge a aut Hus ae TRANSACTIONS OF THE SAN DIEGO SOCIETY OF NATURAL HISTORY VotuME X, No. 15, pp. 269-306, text fig. 1, map THE CHUCKWALLAS, GENUS SAUROMALUS BY 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.) Ua AH 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. 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Ayuo suaundeds p 077-8°98I-9S1 06I-b 6SI—CET LSI-Z'8hI-8El c9I—S'8ST-OSI 981-S €9I-ISI IST-8°6€I-O€1 Cal-e0El=8e €7I-8°STI-ZOT 6cI-F IZI-801 sjenua A SN]PUOANYY AO SHIONdSANG AGNV SHIOddS AHL dO SYALOVYVH’) AO AYVNWNS LSI snsaqo °O I¢ snpiun} *o Il tpuasumoz -o II SNLADA 9 S1pD4jSND gI 42qD € Maqgnv]y Ll BUIAOTS II snpidsty suaudeds fo Joquinyy iS) Ss nS) =) ‘Si aS iS: ZS: Si) SHAW—THE CHUCKWALLAS 303 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. of Cn qi ee 15 at er ogee a eee a oe che nt wats . Ce he a ae! Rare Rie te | —_ oes ied hae’ 8 ve Pe Her be ehh ges ete Lire hy fal, pag 1 raeat ne ee Hieicukeoere es Satna eae eth ek rh hae My « * a De an tet a! 0 : igh aig * 4 —_ a Por ae phat oe ie Wen deceit Ne oe oak tise cats ey: ae i") : = - > OH rs oe ave : hy. ; =, : n "me a BP ei URN ar ics Natal Oe re a aie Waar deed : ee | sy a y yer “vr a) ies Soe ie i j ; ai M — - ne, ae ff. a et Tae ane : =r i ae . : » "2 Dey - 7 , - t. = e : 5 a = & ait ® ; ‘ is; iy mit A i ; ua 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 By 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 BY 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 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 Ww © | NMNwMUYUNKF DK BAK NHK CO 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 OCI esl 9°81 b'07 € 81 0ST £61 £07 0'8I sjods ivy 0'6E Z69 6°79 ¢'89 C'8¢ € 8b 0°79 ¢9¢ — n t+ $2YI10]9 Apog 54 pjnrojs Od 00°T €0'1 €9'T b0'1 LOT 00°T €0'1 lire €0'T SADINIOIAT 0¢ TI c6CI 69'CI I8°Cl 06°71 ara | 86 TI OCG) bIel SPP pe sfuy 00°8 662 80°8 +08 80°8 LI'8 c0'8 c0'8 Soom — ioe) spPiqppadney SINNOTT) HOLOTG GNV ATVOS NVal/] ine O' eb Cor £'°0¢ €9Or 60S 8'Lb Sc b CP 1 6b *0°6€ O1¢ 6 Lb TES 61S C9¢ 91G b'9¢ io) 7) 3S 8 & & an 3 RE Q g ¢ °19".L Sappuaf ‘sppaqua 4 0007 CHIT CHIT C6IC 8°807 Cr6l 9'COl 9°F07 LVL Sa ]PuL ‘s]P4qU2 A A]uo uaurdads wUO x. 0°27 VLC SZC SVAG CLG 0°27 EZ? 8°67 T0¢ SMOL 9)PIS 4PIPIDT SHB P20) PEEPUCD) RIEU D3PA1]20 AJ pyjodxq day can IEG supsayq saidadsqng 384 Subspecies Elegans Blanchardi Philipi Expolita Noctivaga Eburnata 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 KLAUBER—THE GLOssy SNAKE 007-291 COCO ZI T'tc-9'81 D6lst Al SOG AZ CCC 881 661-091 sjods PPL 1°99-€°6¢ 9iC9= 09 SCL-E 9 8°09-C'9¢ b'S9-9'8¢ T'09-0°€¢ ZG-9'99r, $2410]9 Apog suaudads qslIfpNsuy * ‘asues apazenbsaqur ay uTyyM Jey [JIM uoNetndod e ur sjenprarpur ayi Fyey inoqy SLb-T Sb CLE SP b6b-1 Or COP-e bb 0'6b-L9b bes 0S CCS-LOS sappua{ ‘spp pnvoqng SLINNOZ) HOLOTG GNV bIc-l 6b V'CS-Z 6b bes-€'0¢ T0S-1'8¢ SVS=GCS b'8S-9'PS VBS-E PS Sa]PUL ‘spppnvoqny 8°SCC-T'0CT DiSCCaCILCC b €€C—-0' 677 EWC EES ICG C LOCC 661 GAT Cac ILe LACCAISECe sajpua{ ‘s)P4qUa A LOLS TLC LOVES CIC DiCEC=S ONC 9°01 7—-0°Z07 ¢961-9'88T 6 LOC-F 107 WAL Gs Cle $9 ]PUL ‘spPajua 4 ATVOS dO SHONVY ATLLYV NOUALNY AGE 4 PIPIVDT 51D JUIP12IOC) Ppipues) pypuing q DBPA1}I0 NJ 4PIodXg dig UY eetman PLAS sups3ajq saisadsqng 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 5) expolita philipt 6 8 pacata 7 occidentalis noctivaga occidentalis 9 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 BatrD, SPENCER F. 1859. Reptiles of the Boundary, in: United States and Mexican Boundary Survey, etc. (Emory), vol. 2, pp. 1-35. 1859. Report upon the Reptiles of the Route, in: Report of Explorations for a Railway Route, etc. (Whipple), vol. 10, pp. 37-45. BLANCHARD, FRANK N. 1924. A New Snake of the Genus Arizona. Occ. Papers Mus. Zool. Univ. Mich., no. 150, pp. 1-5. 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 County, Calif. Bull. So. Calif. Acad. Sci., vol. 29, part 1, pp. 3-14. 1933. Notes on the Snake Dance of the Hopi Indians. Copeia, no. 4, pp: 219-221. BOULENGER, GEORGE A. 1894. Catalogue of the Snakes in the British Museum (Natural History), Vol. 2, (pp x ate 382. Brown, ARTHUR E. 1901. A Review of the Genera and Species of American Snakes, North of Mexico. Proc. Acad. Nat. Sci. Phila., vol. 53, part 1, pp. 10-110. 1903. Texas Reptiles and their Faunal Relations. Proc. Acad. Nat. Sci. Phila., vol. 55, pp. 543-558. Burt, CHARLES E., and Hoyie, LUTHER W. : 1934. Additional Records of the Reptiles of the Central Prairie Region of the United States. Trans. Kan. Acad. Sci., vol. 37, pp. 193-216. 390 San Dieco Sociery oF Naturat History CocKERELL, THEOpoRE D, A. 1896. Reptiles and Batrachians of Messila Valley, New Mexico. Am. Nat., vol. 30, pp. 325-327. CoNnaANT, Rocesr, and BripGes, WILLIAM 1939. What Snake is That? New York. pp. viii + 163. 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. Acad. Nat. Sci. Phila., vol. 12, pp. 368-374. 1861. Contribution to the Ophiology of Lower California, Mexico, and Central America. Proc. Acad. Nat. Sci. Phila., vol. 13, pp. 292- 306. 1875. Check-list of North American Batrachia and Reptilia. Bull. U. S. Nat. Mus., no. 1, pp. 1-104. 1886. Thirteenth Contribution to the Herpetology of Tropical America. Proc. Amer. Philos. Soc., vol. 23, pp. 271-287. 1887. Catalogue of Batrachians and Reptiles of Central America and Mexico. Bull. U. S. Nat. Mus., no. 32, pp. 1-98. 1892. A Critical Review of the Characters and Variations of Snakes of North America. Proc. U. S. Nat. Mus., vol. 14, no. 882, pp. 589-694. 1896. The Geographical Distribution of Batrachia and Reptilia in North America. Am. Nat., vol. 30, pp. 886-902, 1003-1026. 1900. The Croccdilians, Lizards, and Snakes of North America. Report ot U.S. Nat. Mus. for 1898, pp. 153-1294. Cougs, ELLIoTT 1875. Synopsis of the Reptiles and Batrachians of Arizona, in: Explora- ticns and Surveys West of the 100th Meridian (Wheeler), vol. 5, pp. 585-633. Cow es, RAYMOND B. 1941. Observations cn the Winter Activities of Desert Reptiles. Ecology, vol. 22, no. 2, pp. 125-140. Cow Les, RAyMonp B., and Bocert, CHARLES M. 1944. A Preliminary Study of the Therinal Requirements of Desert Reptiles. Bull. Am, Mus. Nat. Hist., vol. 83, art. 5, pp. 261-296. Dunn, Emmett R. 1928. A Tentative Key and Arrangement of the American Genera of Colubridae. Bull. Antivenin Inst. Am., vol. 2. pp. 18-24. GARMAN, SAMUEL 1883. The Reptiles and Batrachians of North America. Mem. Mus. Comp. Zool., vol. 8, no. 3, pp. xxxi 1+ 185. GRINNELL, JOSEPH, and Camp, CHARLEs L. 1917. A Distributional List of the Amphibians and Reptiles of California. Univ. Calif. Pubs. in Zool., vol. 17, no. 10, pp. 127-208. KLAUBER—THE GLossy SNAKE 391 GUNTHER, ALBERT C. L. G. 1885-1902. Biologia Centrali-Americana. Reptilia and Batrachia. (Refer- ence to Arizona in fasc. 16, Feb. 1894.) 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 California. Bull. Zool. Soc. San Diego, no. 8, pp. 1-93. 1938. Notes from a Herpetological Diary, I. Copeia, no. 4, pp. 191-197. 1939. Studies of Reptile Life in the Arid Southwest. Part 1. Night Collecting on the Desert with Ecological Statistics. Bull. Zool. Soc. San Diego, no. 14, pp. 6-64. 1941a. The Frequency Distribution of Certain Herpetological Variables. Bull. Zool. San Diego, no. 17, pp. 5-31. 1941b. The Correlation between Scalation and Life Zones in San Diego County Snakes. Bull. Zool. Soc. San Diego, no. 17, pp. 73-79. 1943. Tail-length Differences in Snakes with Notes on Sexual Dimotr- phism and the Coefficient of Divergence. Bull. Zool. Soc. San Diego, no. 18, pp. 1-60. LINSDALE, JEAN 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 Sci., vol. 73, no. 8, pp. 197-257. LittLe, EvBert L., Jr., and KELLER, JOHN G. 1937. Amphibians and Reptiles of the Jornada Experimental Range, New Mexico. Copeia, no. 4, pp. 216-222. Marr, JOHN C. 1944. Notes on Amphibians and Reptiles from the Central United States. Amer. Midl. Nat., vol. 32, no. 2, pp. 478-490. Mearns, Epcar A. 1907. Mammals of the Mexican Boundary of the United States. Part 1. Bull. U. S. Nat. Mus., no. 56, pp. xv + 530. Mertens, Ropert, and MULLER, LoRENZ 1928. Liste der Amphibien und Reptilien Europas. Abh. Senck. Naturf. Gesell., bd. 41, 1. 1, pp. 1-62. 392 SAN Dreco Society of Naturat History MosavuEr, WALTER 1935. 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. ORTENBURGER, ARTHUR I., and ORTENBURGER, RUTH D. 1926. Field Observations on Some Amphibians and Reptiles of Pima County, Arizona. Proc. Okla. Acad. Sci., vol. 6, pp. 101-121. PERKINS, C. B. 1938. The Snakes of San Diego County with Descriptions and Key. Bull. Zool. Soc. San Diego, no. 13, pp. 1-66. REYNOLDS, FLETCHER A. 1943. Notes on the Western Glossy Snake in Captivity. Copeia, no. 3, p. 196. RipGway, ROBERT 1912. Color Standards and Color Nomenclature, pp. iv + 43, plts. 1-53. RUTHVEN, ALEXANDER G. 1907. A Collection of Reptiles and Amphibians from Southern New Mexico and Arizona. Bull. Am. Mus. Nat. Hist., vol. 23, art. 23, pp. 483-604. SCHMIDT, Kart P. 1922. The Amphibians and Reptiles of Lower California and the Neigh- boring Islands. Bull. Am. Mus. Nat. Hist., vol. 46, art. 11, pp. 607-707. ScHMipT, Karu P., and Davis, Dwicut D. 1941. Field Book of Snakes of the United States and Canada. New Mork; ppizai 36). ScHMIpT, Kart P., and Owens, Davin W. 1944. Amphibians and Reptiles of Northern Coahuila, Mexico. Zool. Ser. Field Mus. Nat. Hist., vol. 29, no. 6, pp. 97-115. ScHMIpT, Kart P., and SmirH, TARLETON F. 1944. Amphibians and Reptiles of the Big Bend Region of Texas. Zool. Ser. Field Mus. Nat. Hist., vol. 29, no. 5, pp. 75-96. SmitH, Hosart M. 1943. Summary of the Collection of Snakes and Crocodilians made in Mexico under the Walter Rathbone Bacon Traveling Scholarship. Proc. U. S. Nat. Mus., vol. 93, no. 3169, pp. 393-504. SmitH, Hoparrt M., and LEONARD, ARTHUR B. 1934. Distributional Records of Reptiles and Amphibians in Oklahoma. Am. MidI. Nat., vol. 15, no. 2, pp. 190-196. STRECKER, JOHN K. 1915. Reptiles and Amphibians of Texas. Baylor Bulletin, vol. 18, no. 4, pp. 1-82. STULL, OLIVE G. 1940. Variations and Relationships in the Snakes of the Genus Pituophis. Bull. U. S. Nat. Mus., no. 175, pp. vi + 225. KLAUBER—TIHE GLossy SNAKE 393 TAaYLor, Epwarp H. 1929. A Revised Checklist of the Snakes of Kansas. Univ. Kan. Bull., vol. 19, no. 5, pp. 53-62. VAN DENBURGH, JOHN 1897. The Reptiles of the Pacific Coast and Great Basin. Occas. Papers Calif. Acad. Sci., no. 5, pp. 1-236. 1906. On the Occurrence of the Spotted Night Snake, Hypsiglena ochrorhynchus, in Central California; and on the Shape of the Pupil in the Reptilian Genus Arizona. Proc. Calif. Acad. Sci., ser. 3, vol. 4, no. 5, pp. 65-67. 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. VAN DENBURGH, JOHN, and SLEVIN, JOSEPH 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. 1921. A List of the Amphibians and Reptiles of the Peninsula of Lower California, with Notes on the Species in the Collection of the Academy. Proc. Calif. Acad. Sci., ser. 4, vol. 11, no. 4, pp. 49-72. WALLS, GorDON L. 1934. The Reptilian Retina. Am. Jour. Ophthalmology, vol. 17, no. 10, p. 892-915, 1942. The Vertebrate Eye. Cranbrook Institute of Science, Bloomfield Hills, Mich., pp. xiv + 785. Yarrow, HEnry C. 1875. Reports upon the Zoological Collections obtained from Portions of Nevada, Utah, California, Colorado, New Mexico, and Arizona, in: Explorations and Surveys West of the 100th Meridian (Wheel- er), vol. 5, pp. 509-584. 1883. Check List of North American Reptilia and Batrachia, with Catalogue of Specimens in U. S. National Museum. Bull. U. S. Nat. Mus., no. 24, pp. 1-249. 1 eae on 3, He ” pape eae oh mane ifsiiaeoasa 0, “ia z > ie a ER RR Et ora ee PY) Ati er “ | de Maree Ms Sst ae ERE Tih rasta ae! ape ote oi oi Fe) ene es : eee ine if eee tip es teed case eo ae a ee ‘auth Dak oe tas yet ins CA Gl ey 6 J = # ah \ oe ee onye ae Bes ak = Oa Aa Ciwocaies sea Bd ih hit ale Piper ini ae | Bes ee pit ao =H. hires oieeas ni 7S ae i Ai a leases je sao ate - (ob) eh amie) htt Sem) aslees A ste 9. ESS ea sited asre nee Oo oe aa mh ey ls ey. rn gos va > pews a aes Stir scieg = aa “oi bh a ‘i ae c Be i ‘ 7 ei PD- ave es 3 7 ~ s PEM Ste A Wigs Oe aT ies LN Caney ay tal 0a we te: hed gales pee | Diets Wah ah, © Rael Su fee ais: picts WES Kit eo ; ah ge lege dy ian Oe Copan aad Pp alls alt Si seta ni Oe pee | ‘, Cian vs 4 Saye € aa | ers | ni } ae if, oe (nG vapene, ts sre ings » PALES A, Sa Aven Bite 1 pete ee x t fe —s eee . a we - 7 1 a ma “a 7 , 4 9 ; f in 4 i! >, . oR : as KLAUBER—THE GLossy SNAKE PLATE 7 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. KLAUBER—IHE GtLossy SNAKE PLaTE 8 Se ae Re a ‘ ret eS . = & Se re a las SV bbb Saree esis $ 2): a Fig. 1. Arizona elegans eburnata. Adult male from Borego Palm Canyon, San Diego County, California. Photograph by L. C. Kobler. Fig. 2. Arizona elegans candida. From Neenach, Los Angeles County, California. Collected and photographed by C. M. Bogert, American Museum of Natural History. re = S~ -& xi Maa a one _ 2b O70 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 BY 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. : es pa ; oetas oo 1 Panel rine aT wast * i ne » -— 7 oe wap eon rte 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. D 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. E 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. F 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. G 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. H 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 K 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. Kobler.. Ieslie ©, 119, 388, 395, 396. Koster, William J., 388. 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. M 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. N 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. O 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 Pp 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. Q Quail, Desert Mountain, 132. R 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 Vv 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. Ww 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. x Xantusia henshawi, 159. Xenospiza, 227. baileyi, 228. Xiphocolaptes promeropirhynchus sclateri, PPB IANG ny. 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