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Testi CORY Hn} Na Piatt WO a Mr Pa ehh Ob Ba} vibe Wee bh Wt ete 4a COU TAM NM tea! aie Si Wien Va i w) Wide rey aac + 1 Nis whet aut A Nani Wei it had tegagns Vi a Lak ow vow Seah Gunes wir Als: Wy BAM ye etieh fc TA AA ie i eeKON Wole 9¢ i we we La pen! ¥ C0 et | \, ow i ‘ Y nk tat +) SNK Ki Dat atrea yea wdc beg y vy ai oa ve! CR aL OY ha eray wh ANTI CU UCU RRC ‘ ia} 9.1950 i fui Woe & ia SDA tale IO 4) Minted Wi ' Pe Oe Pe Vil Cia ist WH 4 Caan Wald esas wks} t ‘ PA RC wie Ns 1 TL te es (MN a ao Hh ERO ‘ ite LINAS Nt AM | ‘ van ae University of California Publications in LOOLOGY VOLUME XXI 1918-1926 EDITORS CHARLES ATWOOD KOFOID JOSEPH GRINNELL oa) ear aS 7 wv ee: UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA oO 14, 15. CONTENTS . A Revision of the Micretus californicus Group of Meadow Mice, by HEVOTIUETT UO Ta UNE) eh er ese acs oan oe oe ee ew eee . Five New Five-toed Kangaroo Rats from California, by Joseph (Greve: ees ee eee Lb cs oo a SSA A . Notes on the Natural History of the Bushy-tailed Wood Rats of (Chilindoreramiehs lojyig eoorstey oN” 1D i Co) 0) epee ne ete ae cee ea, a occa cee eee . Revision of the Avian Genus Passerella, with Special Reference to the Distribution and Migration of the Races in California, by TEC SI5 PRON COREL: cette ct cece a Eee eee ei ee et . A Study of the California Jumping Mice of the Genus Zapus, by ASE Ta Ziete GO W.ellllip es ese eh Cer ne eek se pele Ue : . Two New Rodents (Genera Thomomys and Marmota) from the East- ern Border of California, by Joseph Grinnell ...................... . A Study of the Californian Forms of the Microtus montanis Group Ole Mead Ova Mace byeckve min oom yell @ soy een anes eee . A Synopsis of the Microtus mordax Group of Meadow Mice in Cali- THOTT, Jony Uivseauirafedionay VSCOM Ove ee oa aera ce ce ccc ced oes eee ree eee . Behavior of the Leaf-nosed Snake, Phyllorhynchus decurtatus, by Sara TEROOXETAS) ZEUS Ea cast eee cnc ser Reece ae Sessa ra i Pre er ee . A Systematic List of the Mammals of California, by Joseph Grinnell . The Systematic Status of the Mountain Lion in California, by Joseph Grrerlrauavenll, sake “AJ OYSEy ol IDS Coy aes ae ee ene eee ee Bee . New Subspecies of Birds from Patagonia, by Alexander Wetmore .... . Revision of the Genus Lynw in California, by Joseph Grinnell and Joseph Dixon ..... ee Ne ee tee a eee Biol. Surv. Coll., U. 8. Nat. Mus.; Lone Pine, 3700 feet altitude, Inyo County, California ; December 23, 1890; collected by E. W. Nelson; original no. 149. Range.—Owens Valley region of California, east of Sierra Nevada, from Olancha, Inyo County, north to Benton, Mono County ; east to head of Willow Creek in north end of Panamint Mountains (fide Bailey, loc. cit.). Vertical range from 3700 feet up at least to 5400 feet ; zonal range.Lower and Upper Sonoran. (See map.) Total number of specimens examined, 96, from the following locali- ties in California. Inyo County: Lone Pine Creek near Lone Pine, Type specimen.—Female adult ; no. 22 University of California Publications in Zoology — [Vou. 21 42; Kearsarge Pass (Grays), 15; Independence, 30; Laws, 5; Silver Canon (3 miles E Laws), 3. Mono County: Benton, 1. Diagnosis—Size large (hind foot, 21 to 25 mm., condylobasal length of skull in largest individuals, 31.6 mm.) ; second upper molar with open posterior internal lobe or loop; skull long, but narrower and not as strongly ridged as in aestuarinus (ratio of zygomatic breadth to condylobasal length about 57%) ; interorbital region with poorly developed ridges. Colors dark. Color—Mass effect of dorsal area dull sepia, peppered with black, or darkened by an admixture of long overhairs, most pronounced mid- dorsally. In fresh pelage: Hairs of upperparts blackish plumbeous basally, with light tippings varying from pinkish buff to light buff; long overhairs ranging from light seal brown to dark aniline black. The light tips of the hairs on the sides become lighter and more buffy, passing abruptly into color of underparts. Rump may show a stronger infusion of light tipped hairs than rest of upperparts. Underparts pallid neutral gray, occasionally with a slight wash of buffy, the plumbeous of hair bases everywhere showing through at surface. Anal area whitish. Terminal portions of whiskers whitish, with bases darker. Nose usually darker than area in front of eyes. Ears rather large, not concealed by fur, with anterior border darkened by the plumbeous hair bases. Tail distinctly bicolor, dark aniline black above and pallid mouse gray below. Hands and feet covered with pallid mouse gray hairs. In worn pelage: Mass effect dull buffy or grayish (moldy appearance), with little infusion of long dark overhairs. The slate gray hair bases show through and take part in the general coloration. Tail distinetly bicolor. Skull—Kssentially the same length as in aestuwarinus, but narrower and less robust; larger and less angular in old age than in californicus. Dorsal profile with a more uniform convexity than shown by other members of this group. Occiput more abruptly truncate, concealing the condyles for the most part from above. Ventral profile similar to that in californicus. From a dorsal aspect the brain-case is distinetly rounded at sides, and truneated anteriorly by strongly developed postorbital tuberosi- ties, which project winglike over orbital fossae. In old adults there are well developed longitudinal ridges near edges of parietals. The interorbital region is rather broad, with a distinct median suleus, but with interorbital ridges well defined. Lambdoidal ridge similar to that of aestuarinus. Interparietal subrectangular, its posterior border variable, but usually somewhat convex, its lateral extremities rather abruptly truneate. Anteroposterior diameter of interparietal, exclu- sive of median projection, less than one-half of greatest transverse diameter. Frontals occasionally convex posteriorly, but usually truneate or even emarginate. Width of rostrum at narrowest region slightly greater than breadth of interorbital constriction. Least depth behind incisors usually greater than, but sometimes equal to, width in same region. Nasals broad at anterior ends, narrowing uniformly toward posterior ends, but with their terminations emarginate in some individuals while in others rounded and notched. In all skulls examined the nasals were shghtly exceeded by ascending branches of premaxillae. Opening 1918] Kellogg: Microtus californicus Group of Meadow Mice 23 for anterior nares, with vertical diameter about equal to transverse. Anteorbital foramen very variable in appearance; in some individuals narrow and high, with its superior portion scarcely if at all wider than inferior ; in others superior width more than twice inferior width. The plate forming outer wall and continuous with lacrimal bone internally, almost complete superiorly, though absent inferiorly. Oecciput evenly convex, the median depth averaging about 53% of greatest width across lambdoidal ridge of brain-case. Foramen magnum subtriangular, averaging smaller than that of aestuarinus. Paroccipital processes heavy, directed downwards and slightly back- wards, terminating rather bluntly. Their tips extend slightly if at all behind most posterior point on supra-occipital above foramen magnum. The posterior margins of paroccipitals are continued up- wards as well marked ridges and usually unite with lambdoidal ridge near outer extremities of interparietal. Basioceipital with poorly defined median ridge. Auditory bullae large, but shghtly smaller than in aestuarinus, somewhat inflated, and otherwise essentially similar. Palate with well marked palatine sulci, and slightly elevated median ridge. Lateral bridges light, but well developed. Lateral pits of palate distinct. Pterygoids and hamular processes rapidly diverging posteriorly, and with posterior margin of palate notched. Interptery- goid fossa somewhat narrower than in aestuarinus or californicus. Posterior palatine foramina almost obliterated. Incisive foramina long, rounded at both ends and with greatest width in median portion. Central portions of zygomatic arches noticeably widened, and maxillary roots heavier than in those of californicus. Jugal enlarged throughout and expanded anteriorly into a broad plate, which abuts against maxillary root of zygoma. Zygomatie arches conspicuously notched at anterior junction with premaxillae. Mandible robust, heavier than in mariposae. Ridges for masseteres laterales strongly developed. Coronoid process light, its base slightly wider than in mariposae, its extremity rising above level of condyle, and curved backward. Angular process short and heavy, but curved strongly outward. Periphery over base of incisor marked by an obvious protuberance. Base of mandibular foramen decidedly above level of cutting surface of last lower molar. Mental foramen on lateral face situated somewhat below superior surface of diastema. Tceth—In most respects the molariform teeth resemble those of californicus. Anterior loop of M+ ecrescentic. An internal lobe nor- mally present on posterior triangle of M?. Long terminal loop of M2 subtended externally by slight notch and internally by deep re- entrant angle. Externoanteriorly this loop is decidedly crescentic in outline. M;with anterior loop more deeply cut on internal side by reentrant angle than on external side. Transverse loops relatively narrower than transverse width of median portions of molars. Remarks.—So far as is known vallicola is separated from mari- posae by the main ridge of the high Sierra Nevada. Collecting so far has failed to show whether or not vallicola intergrades with kernensis 24 University of California Publications in Zoology — [Vou. 21 in the region of Walker Pass. The specimen from Taylor Meadow listed under kernensis, approaches more closely to vallicola in external coloration than it does to kernensis, but cranial characters place it with the latter. Microtus californicus scirpensis Bailey Amargosa Meadow Mouse Microtus scirpensis Bailey (1900, p. 38). Amargosa River (near Nevada line), Inyo County. Microtus scirpensis, of authors. 25979 2 Sane Biol. Surv. Coll., U. S. Nat. Mus.; Amargosa River (near Nevada line [spring near Shoshone, 1560 feet altitude]), Inyo County, California; February 26, 1891; collected by Vernon Bailey ; original no. 2520. Range.—Known only from a small tule marsh at a spring near Shoshone on the Amargosa River, in eastern Inyo County. Altitude of station 1500 to 1600 feet ; zonal range Lower Sonoran. (See map.) Total number of specimens examined, 7, from the following locality in California. Inyo County: Amargosa River [near Shoshone], 7 (Biol. Surv. Coll., U. S. Nat. Mus.). Diagnosis—Size large (hind foot, 25 to 25.5 mm., econdylobasal length of skull in largest individuals, 31 mm.) ; second upper molar with open posterior lobe or triangle; skull long, lighter than in vallicola, with longitudinal ridges much less conspicuous in old adults (ratio of zygomatic breadth to condylobasal length about 60% ) ; inter- orbital region not developing single median ridge with age, but always with definite suleus between ridges ; colors brighter than in aestuarinus or vallicola. Color—Mass effect on dorsal area much brighter than in valli- cola; exact shade variable due to long dark overhairs, but usually near snuff brown. In fresh pelage: Upperparts with light terminal portions of hairs varying from cinnamon buff to buckthorn brown, inconspicuously lined mid-dorsally with long blackish overhairs. Basal portions of hairs blackish plumbeous. Sides much lighter than middle upperparts, with larger proportion of cinnamon-buff tipped hairs. Upperparts pallid neutral gray, irregularly darkened by ap- pearance at the surface of blackish plumbeous hair bases; anal area nearly pure white. Terminal portions of whiskers whitish; basal portions blackish. Upper and lower lips whitish. Nose more blackish than region in front of eyes. Ears large, partially concealed by fur, with anterior border covered by light tipped hairs of same color as those of upperparts. Tail distinctly bicolor, bone brown above and pallid neutral gray below, though when tail is scantily haired the yellowish scales show through. Young: Very young individuals are almost pure black, excepting that there is a suffusion of hight tipped buffy hairs on sides and on head. With age the blackish cast of the young changes to brownish. Type specimen.—Female adult; no. 1918] Kellogg: Microtus californicus Group of Meadow Mice 2d Skull_—In some respects resembles that of vallicola, in others that of aestuarinus. Dorsal profile rather evenly convex, slightly depressed interorbitally. Width across zygomatic arches greater than in any other form within the group. Occiput rather abruptly truncate, concealing the condyles to some extent when skull is viewed from above. Ventral profile similar to that of vallicola. Brain-ease in outline broadly ovate, rounded at sides and trun- eated anteriorly by poorly developed postorbital tuberosities. In old adults the longitudinal ridges near outer edges of parietals are present and unite with interorbital ridges in frontal region. Inter- orbital ridges variable in appearance but apparently always sepa- rated by distinct suleus. Interparietal somewhat rectangular, with posterior border convex and with outer extremities rather abruptly truneate. Anteroposterior diameter exclusive of median projection less than one-half greatest transverse diameter. Frontals convex or truneate posteriorly. Rostrum rapidly tapering anteriorly; width at narrowest region considerably greater than breadth of interorbital constriction; least depth behind incisors greater than width in same region. Nasals narrowly spatulate, their posterior terminations squarely truncate or ‘emarginate. Ascending branches of premaxillae exceeding nasals and extending to or beyond plane of lacrimals. Vertical diameter of anterior narial passage equal to transverse. Anteorbital foramen variable in outline, with its superior portion in some individuals seareely if at all wider than inferior, though superior width is usually greater than inferior. Occiput evenly convex or flattened, the median depth equal to about 60% of greatest width across lambdoidal ridge of brain-case. Foramen magnum subtriangular. Paroccipital processes directed more downwards than backwards, with tips not applied to bullae. Posterior margins of paroccipital processes continued upwards on supra-occipital as sharp crests, terminating at or below lambdoidal ridge. Floor of brain-ease with sharply defined median ridge. Auditory bullae large, with their posterior borders obliquely truneated. Width of basioccipital at suture between it and basisphenoid equal to one- third its median length. Palate relatively wider than that of vallicola, with shghtly deeper palatine sulei and narrower median ridge. Interpterygoid fossa nar- row, though hamular processes rapidly diverge posteriorly. Poste- rior palatine foramina small though open. Lateral pits fenestrated. Incisive foramina long, wide open, the outer sides nearly parallel throughout their length, and ends rounded. Zygomatie arches diverging rapidly anteriorly, the sides nearly parallel ; upper surface of maxillary root narrower than in vallicola. Jugal somewhat expanded anterior to middle, and deeply mortised into maxillary root of zygoma. Zygomatic arches deeply notched at anterior junction with premaxillae. Mandible robust, with moderate diastema. Ridge for masseteres laterales well developed. Coronoid process heavy, its base wide, its extremity rising to or above level of condyle, though but little curved backwards. Angular process heavy, well developed, curved outward, 26 University of California Publications in Zoology — | Vou. 21 but not extending posteriorly beyond plane of condyle. Peripheral tuberosity over base of incisor lying for the most part within masse- teric ridge, but not conspicuous. Base of mandibular foramen de- cidedly above level of cutting surface of last lower molar. Mental foramen on lateral face situated somewhat below superior surface of diastema. Teeth—Molars both above and below differing from those of cali- fornicus in the general tendency of enamel folding of closed triangles to assume more irregular outlines. Anterior loop of M?# flattened. An internal lobe present on posterior triangle of M?. The enamel folding of M? without any marked peculiarity excepting that terminal loop is rather long and external side rather straight. Lower molars not exhibiting any marked peculiarities excepting that transverse loops are relatively large. Anterior loop of M; deeply notched by internal reéntrant angle. Remarks.—This race has apparently been exterminated within recent years, as the type locality, a small tule marsh near Shoshone, Inyo County, has been burnt over for several consecutive years and is now used for a hog pasture. Assiduous trapping at this point in May, 1917, did not yield a single meadow mouse (fide Grinnell, MS, and Storer, MS). There is a possibility, however, that M. c. scirpensis may still exist south of Shoshone near the town of Tecopa, where there is permanent water and tule marshes. The characters possessed by this race show that it is an isolated form of the californicus group. It is probable that the californicus group had a continuous distribution over the state at one time and that geological changes have resulted in isolating certain stocks from one another. Scirpensis resembles both aestuarinus and vallicola, but neither can be considered as the parent stock. It is a remarkable race because of its occurrence away from the main mountain axes and in an area of extremely high summer temperature. Microtus californicus kernensis, new subspecies Kern River Meadow Mouse Arvicola (Myonomes) riparius, Coues and Allen (1877, p. 173), part. Fort Tejon. Microtus californicus, of authors, part. Type specomen.—Female adult; no. 15779, Mus. Vert. Zool.; Fay Creek, 4100 feet altitude, Kern County, California; July 14, 1911; collected by Tracy I. Storer; original no. 219. Range—Kern River basin, from Taylor Meadow, Tulare County, west to Bakersfield and Fort Tejon, Kern County. Vertical range from 400 feet up to 7000 feet; zonal range Lower Sonoran to Tran- sition. (See map.) . 1918] Kellogg: Microtus californicus Group of Meadow Mice 27 Total number of specimens examined, 33, from the following locali- ties in California. Kern County: Bakersfield, 1; Bodfish, 5; Fay Creek (6 miles N Weldon), 8; Onyx, 10; Weldon, 1; Fort Tejon, 7. Tulare County: Taylor Meadow, 1. Specimens from Fort Tejon are apparently intergrades between typical kernensis and the southern California form, neglectus. Diagnosis —Size large (hind foot, 21 to 25 mm., condylobasal length of skull in largest individuals, 31.4 mm.) ; second upper molar with open posterior lobe; skull long, not angular, and relatively smooth in adults (ratio of zygomatic breadth to condylobasal length about 57%) ; interorbital region not developing a single median ridge with age, but always with definite sulcus between ridges. Color.—Mass effect on dorsal area tawny-olive, darkened by the dark overhairs. In fresh pelage: Upperparts with light terminal portions of hairs varying from clay to cinnamon-buft, very incon- spicuously lined with long, dark, warm sepia overhairs. Basal por- tion of fur everywhere blackish plumbeous. Sides lighter than upper- parts, with light terminal portions of hairs approaching pinkish buff. There is no true line of demarcation between dorsal area and sides, but transition to gray of underparts is rather abrupt. Rump not decidedly brighter than dorsal area. Underparts pallid mouse gray or white, sometimes irregularly darkened by the plumbeous bases of the hairs showing through. Anal area nearly pure white. Ter- minal portions of whiskers much lighter than basal portions. Nose normally not conspicuously darker than area in front of eyes. Ears moderate in size, not concealed by fur, with anterior border darkened by plumbeous hair bases. Tail distinctly bicolor, usually warm sepia above and whitish below, though when tail is scantily haired the yellowish scales show through. Hands and feet covered with pale drab gray hairs. In worn pelage: Duller, with greater effect of dark overhairs. Skull.—Skull characterized by relatively smooth brain-case, short rostrum, and small bullae. Dorsal profile rather evenly convex, with slight interorbital depression. Occiput rounded but concealing for the most part the condyles when skull is viewed from above. Brain-case subovate, its sides rounded and truneated anteriorly by winglike postorbital tuberosities which produce a marked break in contour of skull. Longitudinal ridges along outer edges of parietals, if present, but poorly developed. Interorbital ridges variable in ap- pearance, separated by a shallow suleus of varying width. Inter- parietal subrectangular, its posterior border nearly straight and at the most but little convex, with lateral extremities squarely truneate. The anteroposterior diameter exclusive of median projection more than one-third greatest transverse diameter. Posterior margin of frontals, convex, truneate or emarginate. Rostral region shorter than that in either mariposae or mohavensis. Least depth behind incisors greater than width in same region. Proximal depth of rostrum approximately equal to its median length. Nasals long, narrow, expanded shghtly anterior to the middle, but not attaining the pronounced spatulate shape as in mariposae and moha- vensis. Posterior termination of nasals truneate or emarginate, not extending beyond incisors and exceeded posteriorly by ascending 28 University of California Publications in Zoology (Vou. 21 branches of premaxillae; the latter extend as far as or beyond plane of lacrimals. Vertical diameter of anterior narial opening equal to or exceeding transverse diameter. Anteorbital foramen normally high and narrow. Occiput uniformly convex, the median depth equal to about 59% of greatest width across lambdoidal ridge of brain-case. Foramen magnum subtriangular, but somewhat flattened. Paroceipital pro- cesses directed more backward than downward, but their tips may or may not be applied to the bullae; their posterior margins continued upwards as sharp crests. Floor of brain-case not differing in any essential detail from that in other races of californicus; median ridge present and width of basioccipital at suture between it and _ basi- sphenoid more than one-third of its median length. Auditory bullae small, with more marked depression below meatus than in mariposae and mohavensis. Posterior border of bullae obliquely truncated and somewhat indented. | Palate narrow, with shallow palatine sulei and depressed median ridge. Lateral pits fenestrated. Interpterygoid fossa narrow, the hamular processes of pterygoids rapidly diverging. Incisive foramina long and narrow, with portion slightly posterior to middle expanded. Anterior portion of zygoma strongly diverging; median portion almost parallel with main axis of skull; upper surface of maxillary arm of zygoma narrower than in either martposae or mohavensis. Jugal normally expanded and shallowly mortised into maxillary part of zygoma. Zygomatie arches slightly notched at anterior junction with premaxillae. Mandible smaller than that of mohavensis. Ridge for masseteres laterales well developed. Coronoid process long, its base broad, its extremity rising above level of the condyle. Angular process stout, curved. outward but with posterior border not extending as far as plane of condyle. Peripheral swelling over base of incisor marked by conspicuous tuberosity which lies for the most part within the masse- teric ridge. Base of mandibular foramen situated shghtly above cut- ting surface of last lower molar. Mental foramen situated on lateral face below superior surface of diastema. Teeth—Though in general resembling those of californicus, the transverse loops are relatively shorter and less flattened. An internal lobe generally present on posterior triangle of M?. The enamel fold- ing of M2 is without any especial peculiarity. First lower molar with deep internal and external reéntrant angles which are approximately equal. Transverse loops crescentie. Remarks.—This light colored race inhabits a region of dry and (in summer) warm air. Specimens from the vicinity of Fort Tejon are best referred to this form though in certain cranial characters they approach closely to neglectus. Intergradation between kernensis and neglectus occurs around Piru Creek and Mount Pinos. A speci- men from Bakersfield is darker than the average but nevertheless belongs with kernensis. It is of peculiar interest that three other unrelated forms of 1918] Kellogg: Microtus californicus Group of Meadow Mice 29 animals, Agelaius phoeniceus aciculatus, Eutamias merriami kernensis and Neotoma fuscipes simplex, occupy almost the same area as Microtus californicus kernensis. Furthermore, this seems to be the area of intergradation between Cnemidophorus tigris tigris and Cnemidophorus tigris mundus as well as of Uta stansburiana elegans. and Uta stansburiana hesperis. Other examples could be cited, but the evidence points to the fact that in this region there has been a differentiation of species to form new races in several unrelated groups of animals, some of which have become fully differentiated and others of which have not. Microtus californicus mohavensis, new subspecies Mohave River Meadow Mouse Microtus californicus, Elliot (1904, p. 299), part. Oro Grande. Microtus californicus vallicola, of authors, part. Type specimen.—Male adult; no. 5987, Mus. Vert. Zool.; Victor- ville, 2700 feet altitude, San Bernardino County, California; April 6, 1907 ; collected by J. Grinnell and Walter P. Taylor; original no. 2036. Range.—Known only from a limited area along the Mohave River, in San Bernardino County, from Victorville north to Oro Grande. Vertical range from 2500 feet up at least to 2700 feet; zonal range Lower Sonoran. (See map.) Total number of specimens examined, 31, from the following locality in California. San Bernardino County: Victorville, 31. Diagnosis.—Size large (hind foot, 22 to 25 mm., condylobasal length of skull in largest individuals, 33 mm.) ; second upper molar without posterior internal loop or lobe; skull large, angular and ridged in adults (ratio of zygomatic breadth to condylobasal length about 56%); interorbital region not developing single median ridge with age. Color —Mass effect of dorsal area ranging from Dresden brown to mummy brown, with varying admixture of long dark overhairs. In fresh pelage: Upperparts with hght ends of hairs grading from ochraceous-tawny to cinnamon-buff, faintly varied by the sepia or aniline black overhairs. Sides lighter than upperparts, with lght tipped hairs more nearly cinnamon-buff than those of upperparts. Rump not conspicuously brighter than rest of upperparts. Under- parts French gray, irregularly darkened by the appearance, at the surface, of the plumbeous hair bases. Anal area whitish. Terminal portions of whiskers grayish, bases darker. Nose usually not any darker than area in front of eyes. Ears moderate in size, with anterior border covered with hairs of the same color as light tipped hairs of upperparts. Tail distinetly bicolor, covered to a varying degree above with light seal brown or fuscous black hairs; below pale gull gray. Hands and feet covered with pallid neutral gray hairs. In worn pelage: Mass effect much duller and lighter. Light tipped portions 30 University of California Publications in Zoology — (Vou. 21 of hairs of upperparts approaching light pinkish cinnamon. Plumb- eous hair bases of underparts taking a larger share in the general coloration. A lesser infusion of long dark overhairs. Skull—Best characterized by its large size, its widely spreading zygomata, and narrow interorbital region. Dorsal profile considerably flattened, slightly depressed interorbitally, but with region of greatest convexity in plane with lacrimals. Occiput truneated very obliquely, but not concealing the condyles. Brain-case subquadrate, its sides rounded and truneated anteriorly by well developed postorbital tuberosities. Longitudinal ridges extend along outer edges of par- ietals, continuous posteriorly with lambdoidal ridge and anteriorly with interorbital ridges. Interorbital ridges normally prominent. In all specimens examined there is a distinet suleus between the two, though in some the ridges have nearly coalesced. Interparietal sub- rectangular, its posterior border biconvex to a varying degree, with lateral extremities squarely truncate; anteroposterior diameter ex- elusive of median projection one-half (or less) greatest transverse diameter. Frontals convex posteriorly. A condition of unusual variability is exhibited by the rostral region of this form. Normally the least depth behind incisors is greater than width in same region. Nasals long, strongly spatulate, emarginate or convex posteriorly. Nasals exceeded by ascending arms of premaxillae which may extend beyond plane of lacrimals. Nasals normally declining very little anteriorly. Vertical diameter of ante- rior narial opening equal to transverse diameter. Plate forming outer wall and continuous with lacrimal internally, complete superiorly, though absent, as usual, inferiorly. Anteorbital foramen usually rather wide as compared with its height. Occiput evenly convex or flattened, the median depth equal to about 57% of greatest width across lambdoidal ridge of brain-case. Foramen magnum subtriangular. Paroecipital processes well de- veloped, directed more backward than downward, with their tips not applied to bullae. Their posterior margins are continued upwards as prominent ridges, terminating superiorly on lambdoidal ridge within limits of lateral extremities of interparietal. Floor of brain-case similar to that in other races of californicus, though basioecipital possesses a very sharp median ridge. Width of basioccipital at suture between it and basisphenoid usually about one-third its median length. Auditory bullae large, their posterior borders very obliquely trun- eated. Palate with deep palatine sulei and well developed lateral bridges. Lateral pits fenestrated. Interpterygoid fossa narrow, the pterygoids diverging posteriorly and occasionally with posterior margin of palate acutely notched. Posterior palatine foramina moderate, though not always open. Incisive foramina wide open, slightly wider in medial regian, with ends rounded. Region of greatest zygomatic width slightly anterior to middle, and with upper surface of maxillary arm of zygoma rather wide. Jugal normally expanded and deeply mortised into maxillary root of zy- goma. Zygomatic arches slightly notched at anterior junction with premaxillae. Mandible large and robust. Ridge for masseteres laterales strongly 1918] Kellogg: Microtus californicus Group of Meadow Mice 31 developed. Coronoid process long, its base narrow, its extremity ris- ing but little above condyle and curved backward. Angular process heavy, curved strongly outward, but with its posterior margin lying in same plane as that of condyle. Peripheral process over base of incisor marked by a conspicuous tuberosity which lies for the most part above masseteric ridge. Base of mandibular foramen situated slightly above cutting surface of last lower molar. Mental foramen on lateral face situated but little below superior surface of diastema. Teeth—Among the meadow mice of the californicus group this race 1s distinguished by the shortened terminal loop of M2. Anterior loop of M+ erescentie and relatively wide open. No internal lobe present on posterior triangle of M?. The closed triangles are large, and width of molariform series greater than in any other subspecies. Lower molars similar to those of californicus but much larger. Transverse loop strongly crescentic, proportionally long, though narrow. Remarks.—For some time this race has been included by workers with vallicola. As there is no geographical connection with vallicola, and as characters exhibited by mohavensis are so distinct, it seems best to regard it as a separate subspecies. It occurs in the bottom lands along the Mohave River well out on the Mohave Desert to the north of the San Bernardino Mountains. Microtus californicus neglectus, new subspecies Southern California Meadow Mouse Arviecola edax, Baird (1857, p. 532), part. San Diego. Arvicola riparius, of some authors, part. Microtus edax, of some authors, part. Microtus californicus, of authors, part. Type specimen.—Female adult; no. 19119, Mus. Vert. Zool.; Es- condido, 640 feet altitude, San Diego County, Cahfornia; February 7, 1913; collected by Joseph Dixon. Range—San Diegan faunal district, from Mountain Spring and mouth of Tiajuana River, San Diego County, northwest to Gaviota Pass, Santa Barbara County, and north to Bluff Lake, San Bernar- dino Mountains, San Bernardino County. Vertical range from sea level up at least to 9000 feet ; zonal range Lower Sonoran to Canadian. (See map.) Total number of specimens examined, 103, from the following localities in California. San Diego County: Mouth of Tiajuana River, 2; Chula Vista, 9; San Diego, 1; Dulzura, 3; Jacumba, 4; Mountain Spring, 1; Vallecito, 7; Cuyamaca Mountains, 4; La Puerta Valley, 3; Julian, 2; Witch Creek, 1; Foster, 6; Escondido, 4; Warner Pass, 3; San Onofre, 1. Riverside County: Garnet Queen Mine (Santa Rosa Mountains), 1; Hemet Lake (San Jacinto Mountains), 2; Round Valley (San Jacinto Mountains), 1; Tahquitz Valley (San 32 University of California Publications in Zoology — | Vou. 21 Jacinto Mountains), 4; Jurupa Mountains (7 miles NW Riverside), 3. Los Angeles County: El Monte (San Gabriel River), 2; Oak Knoll (near Pasadena), 1; Linda Vista (near Pasadena), 3; Arroyo Seco (Switzer’s Camp), 1. San Bernardino County: South Fork Santa Ana River (San Bernardino Mountains), 3; Fish Creek, 1; Bluff Lake, 14; Mouth of Reche Cafion (near Colton), 4; Devil’s Caton (San Bernardino Mountains), 1. Ventura County: Mount Pinos, 3; Matilija, 6. Orange County: Santa Ana Cafion, 2. Diagnosis.—Size large (hind foot, 21 to 25 mm., condylobasal length of skull in largest individuals, 32 mm.) ; second upper molar without posterointernal lobe or loop; skull long, broad and heavily built (ratio of zygomatic breadth to condylobasal length about 60% ), the interorbital region not developing a ridge with age; superior sur- face of maxillary root of zygoma wider than that in huperuthrus. Color—Upperparts and sides as a rule brighter than in huper- uthrus. Dorsal area not as strongly mixed with brownish and black- ish overhairs. In fresh pelage: Light tipped hairs of upperparts ranging from buckthorn brown, slightly darkened by long overhairs along middle of back, to cinnamon-buff, brightening nearly to pinkish buff on flanks and sides of neck; basal portions of hairs blackish plumbeous. Long overhairs varying from bright Vandyke brown to blackish. Color of upperparts shades into a lighter cinnamon-buff on sides and rump, with long dark overhairs conspicuously absent. Plumbeous bases of hairs usually visible on sides, and these contribute to the general coloration. Underparts plumbeous, overlaid by white tipped hairs and often slightly washed with buffy in pectoral region. In some specimens this suffusion becomes a rather definite light buff. Anal area pure white. Whiskers mixed white and some shade of dark brown. Upper and lower lips whitish. Nose lighter than remainder of head owing to presence of but few dark overhairs. Ears large, not concealed by fur; anterior border covered with blackish hairs ; posterior border covered with hairs of same shade as light tipped hairs of upperparts. Tail may or may not be distinctly bicolor ; usually the same color above as dark overhairs of upperparts, whitish below. When tail is scantily haired beneath, the yellowish scales show through. Hands and feet washed with grayish white. In worn pelage: Much more grayish buffy, with dark overhairs not conspicu- ously darkening the upperparts. Tips of hairs of upperparts pinkish buff; overhairs dark Vandyke brown. Under parts almost devoid of buffy suffusion, while plumbeous bases of hairs appear lighter in tone owing to a thinning out or wearing down of the hairs. Skull—Separable as a rule from that of huperuthrus by the more spreading and heavier zygomatic arches, the relatively broader ros- trum, and larger bullae. Dorsal profile more evenly convex and not flattened as in huperuthrus. Occiput more obliquely truncated and not rounded, but concealing a large part of condyles, when skull is viewed from above. Ventral profile essentially the same as in huper- uthrus. Nearly all skulls examined characterized by broad brain-ease, its sides rounded and truncated anteriorly by well developed postorbital tuberosities. Longitudinal ridges along outer edges of parietals, con- tinuous posteriorly with lambdoidal ridge and anteriorly with inter- 1918] Kellogg: Microtus californicus Group of Meadow Mice 33 orbital ridges, usually well marked. Interorbital ridges variable in appearance, usually rounded, but with well defined suleus between them. Interparietal variable in outline. In most cases biconvex posteriorly, its sides squarely truneated, but with blunt median pro- jection. From a dorsal aspect the rostrum tapers panne anteriorly. Least depth behind incisors greater than width in same region. Nasals long, spatulate, widening at a point anterior to middle, their posterior terminations emarginate. Ascending branches of premaxillae exceed- ing posterior terminations of nasals and extending as far as plane of lacrimals. Vertical diameter of anterior narial passage about equal to transverse diameter. Anteorbital foramen may be high and narrow or short and broad. Occiput flattened, the median depth averaging about 60% of the greatest width across lambdoidal ridge of brain-case. Foramen mag- num subtriangular. Paroccipital processes usually directed more downwards than backwards, and not normally applied to bullae at their extremities. Posterior borders of these processes continued upwards on supra-occipital as sharp erests. Basioccipital shows a well defined median ridge. Width of basioccipital at suture between it and basisphenoid equal to one-third or more of its median length. Auditory bullae large, strongly inflated, their posterior borders ob- liquely truncated, their inner sides rapidly diverging posteriorly. Palate has well marked palatine sulei and prominent median ridge. Lateral pits of palate fenestrated. Pterygoids and hamular processes rapidly diverging posteriorly. Posterior margin of palate usually rounded. Interpterygoid fossa narrow. Posterior palatine foramina distinct. Incisive foramina variable; usually narrow, slightly ex- panded posterior to middle and rounded at both ends. Zygomatic arches rapidly diverging anteriorly, but with moarerion portion nearly parallel to main axis of skull. Jugal enlarged, ex- panded. centrally, and mortised into maxillary root of zygoma. Supe- rior width of maxillary root of zygoma greater than that in huper- uthrus. Zygomatie arches conspicuously notched at anterior junction with premaxillae. Mandible robust but smaller than in mohavensis. Masseteric ridges well developed. Coronoid process light, its base narrow, its extremity rising above level of condyle and curved backward. Angu- lar process long, curved outward, but not extending posteriorly be- yond plane of condyle. Peripheral tuberosity over base of incisor lying for the most part above masseteric ridge. Base of mandibular foramen above level of cutting surface of last lower molar. Mental foramina on lateral face situated somewhat below superior surface of diastema. Tecth—Kssentially like those of californicus except that closed triangles are relatively wider. Anterior loop of M+ usually ecrescentie, though in some individuals it is flattened anteroexternally. No inter- nal lobe present on posterior triangle of M2, although there is a rudi- mentary posterointernal loop in some specimens. Enamel folding of M2 without any especial peculiarity, though long terminal loop is variable in outline. Inner side of posterior limb of terminal loop may bear an incipient reéntrant angle. 34 University of California Publications in Zoology [Vou. 21 Lower molars not differing in any essential details from those of californcus. Remarks.—This race probably intergrades with mohavensis along the headwaters of the Mohave River. On the north it probably inter- grades with califormcus in Santa Barbara County, though so far no definite area of blending has been found. On the south no loeality of occurrence is known below the California boundary line. Microtus californicus huperuthrus Elliot Lower California Meadow Mouse Arvicola edax, Allen (1893, pp. 184-185). San Pedro Martir Mountains, 8500 feet. (Microtus) edax, Trouessart (1898-1899, p. 564), part. Microtus californicus huperuthrus Elliot (1903a, p. 161). Type locality erroneously given as San Quintin. The real type came from La Grulla. Microtus californicus hyperythrus, Elliot (19036, p. 218); and of authors. Type specimen.—Male adult; no. 10740, Field Columb. Mus.; La Grulla, San Pedro Martir Mountains, Lower California, Mexico: September 10, 1902; collected by Edmund Heller. Range.—Lower California, Mexico, from San Quintin, on the sea- coast, east to La Grulla and Aguaje de las Fresas, in the San Pedro Martir Mountains. Vertical range, from sea level up to at least 8500 feet ; zonal range, from Lower Sonoran to Transition. Total number of specimens examined, 7, from the following locality. Lower California, Mexico: La Grulla (Field Columb. Museum), 7. Diagnosis—Size large (hind foot, 24 to 25 mm., condylobasal length of skull in largest individuals, 32.9 mm.) ; second upper molar with or without open posterior lobe; skull long, lighter than in neglec- tus, and with longitudinal ridges feebly developed even in old adults (ratio of zygomatic breadth to condylobasal length about 57% ) ; inter- orbital region not developing single median ridge with age, but always with definite suleus between ridges. Color.—Mass effect of dorsal area ranging from cinnamon-brown to dull Prout’s brown, more or less grizzled by the long overhairs. In fresh pelage: Upperparts with light tipped portions of hairs varying from cinnamon-buff to cinnamon-brown, rather strongly overlaid with long dark Mars brown overhairs. Basal portion of fur everywhere blackish plumbeous. Sides lighter than upperparts. Underparts pallid mouse gray, irregularly darkened by the showing through of the blackish plumbeous hair bases. Anal area nearly pure white. Ter- minal portions of whiskers much lghter than basal portions. Nose lighter and more grayish than region in front of eyes. Ears large, not concealed by fur, with anterior borders covered with hght tipped hairs, while posterior borders are nearly naked. Tail bicolor, dark aniline black above and mouse gray below, though the yellow scales often show through. Hands and feet covered with pale mouse gray hairs. Skull.—hike that of neglectus in many respects, but differs in that zygomata are less widely spreading, rostrum relatively shorter and bullae smaller. Dorsal profile considerably flattened, with marked 1918] Kellogg: Microtus californicus Group of Meadow Mice 35 interorbital depression. Occiput rounded, concealing the condyles to some extent when skull is viewed from above. While there is considerable variation among different individuals, the brain-ease is usually subquadrate, the sides rounded, and truncated anteriorly by postorbital tuberosities. Longitudinal ridges along outer edges of parietals, when present, but feebly developed. Interorbital ridges variable but always separated by suleus of varying width. Interparietal somewhat rectangular, with posterior border convex and with outer extremities rather abruptly truncate. Anteroposterior diameter exclusive of median projection less than one-half, occasion- ally one-third, greatest transverse diameter. Frontals shallowly convex or truneate porteriorly. Rostrum rapidly tapering anteriorly. Least depth of rostrum be- hind incisors greater than width in same region. Nasals long, spatu- late, their posterior terminations squarely truneate. Ascending branches of premaxillae exceeding nasals and extending to or beyond plane of lacrimals. Vertical diameter of anterior narial passage less than, or at the most equal to, transverse diameter. Anteorbital fora- men variable in outline, usually considerably wider dorsally than in neglectus. Occiput more convex and less flattened than in neglectus, but median depth may equal as much as 58% of greatest width across lambdoidal ridge of brain-case. Foramen magnum subtriangular. Paroccipital processes directed more downwards than backwards, with tips usually not applied to bullae. Posterior margins of paroccipital processes continued upwards on supra-occipital, terminating below lambdoidal ridge in a prominent though blunt erest. Floor of brain-case with poorly defined median ridge. Width of basioecipital at suture between it and basisphenoid equal to one-third of its median length. Auditory bullae smaller than those of neglectus, their posterior borders obliquely truncated, their inner sides rapidly diverging posteriorly. Palate narrower than in neglectus, with deep though narrow pala- tine sulci. Lateral pits of palate fenestrated. Interpterygoid fossa narrow, the hamular processes not markedly diverging as in above. Posterior palatine foramina distinct. Incisive foramina in most speci- mens narrower than in above, slightly expanded anterior to middle, and narrowing at both ends. Zygomata widely spreading, the upper surfaces of maxillary arms of zygoma narrower than in neglectus. Jugal narrower and less sharply mortised into maxillary root of zygoma. Mandible robust, with moderate diastema. Masseteric ridges well developed. Coronoid process long and light, its base narrow, its ex- tremity rising above level of condyle and curved backward. Angular process long, curved outward, but not extending posteriorly to plane of condyle. Peripheral tuberosity over base of incisor lying for the most part above masseteric ridge. Base of mandibular foramen above level of cutting surface of last lower molar. Mental foramina situated on lateral face below superior surface of diastema. Teeth—Molars smaller relatively to size of skull than in neglectus, their enamel pattern well defined, with sharp salient and reéntrant angles. Anterior loop of M+ flattened. An internal lobe may or may 36 University of California Publications in Zoology | Vou. 21 not be present on posterior triangle of M?. Enamel folding of M2 without any especial peculiarity except that terminal loop is usually very long. Outer convex surface of loop marked by slight salient angle just behind point of attachment to second outer closed triangle; inner side of posterior hmb of terminal loop with shallow notch or reéntrant angle. Lower molars not exhibiting any marked peculiarities, and resembl- ing very closely those of californicus, except that posterior transverse loops are relatively wider. Remarks.—Elliot in his original description of this form erron eously gave the type locality as San Quintin, whereas the specimen now labelled as the type came from La Grulla. Later, Elliot (1907, p. 292) corrected this error in a footnote. According to Elliot this race occurs in the weeds and grass in the vicinity of San Quintin and follows the streams up into the San Pedro Martir Mountains where it was found even in the highest meadows. LITERATURE CITED ALLEN, J. A. 1893. On a collection of mammals from the San Pedro Martir region of Lower California, with notes on other species, particularly of the genus Sitomys. Bull. Amer. Mus. Nat. Hist., 5, 181-202. BAILEY, V. 1898. Descriptions of eleven new species and subspecies of voles. Proe. Biol. Soc. Wash., 12, 85-90. 1900. Revision of American voles of the Genus Microtus. U. 8S. Dept. Agric., Div. Biol. Surv., N. Amer. Fauna, 17, 1-88, 5 pls., 17 figs. in text. Bairp, S. F. 1857. Report on Zoology. I, Mammals. U.S. Pac. R. R. Expl. and Sury., 8, xlviii + 757, pls. 17-60, 35 figs. in text. Cours, E., and ALLEN, J. A. 1877. Monographs of North American Rodentia, in Rep. U. 8S. Geol. Surv. Territories (Washington, Govt. Printing Office), 11, i-x, 1-939, 1083-1091. Cougs, E., and Yarrow, H. C. 1875. Report upon the collections of mammals made in portions of Nevada, Utah, California, Colorado, New Mexico, and Arizona, during the years 1871, 1872, 1873, and 1874, in Explorations and surveys west 100th meridian (Washington, Govt. Printing Office), Zoology, 5, 35-129, 969-976. ELuiot, D. G. 1898. List of mammals collected by W. W. Price and assistants, chiefly in California. Field Columb. Mus., Zool. ser., 1, 193-216. 1903a. Descriptions of apparently new species and subspecies of mammals from California, Oregon, the Kenai Peninsula, Alaska, and Lower California, Mexico. Ibid., 3, 153-173, 1 fig. in text. 1918] Kellogg: Microtus californicus Group of Meadow Mice 37 1903b. A list of mammals collected by Edmund Heller, in the San Pedro Martir and Hanson Laguna mountains and the accompanying coast regions of Lower California, with descriptions of ‘apparently new species. Ibid., 3, 199-232, pls. 33-38. 1904, Catalogue of mammals collected by E. Heller in southern California. Ibid., 3, 271-321, pls. 38-49, 1 map. 1907. A catalogue of the collection of mammals in the Field Columbian Museum. Ibid., 8, viii + 694, 92 figs. in text. LE CONTE, J. 1853. Description of three new species of American Arvicolae, with re- marks upon some other American rodents. Proc. Acad. Nat. Sci. Phila., 6, 404-415. PEALE, T. R. 1848. Mammalia and Ornithology. U. S. Expl. Exped. (Wilkes), (Phila- delphia, Sherman), 8, i-xxvi, 17-338, several unnumbered figs. in text. TOWNSEND, C. H. 1887. Field notes on the mammals, birds and reptiles of northern Cali- fornia. Proc. U. S. Nat. Mais., 10; 159-241), pl: 5, 4 figs. in text: TROUESSART, EH. L. 1898-1899. Catalogus mammalium tam viventium quam fossilium. 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E S 895 : : ZEEREE = : crEsees aos = a $3) See = = = - vomomne) ae ee Zs BESeRe See SEE re == aS a a re 1918] MEASUREMENTS (IN MILLIMETERS) OF SELECTED ADULT SPECIMENS OF HACH OF THE RECOGNIZED RACES OF Microtus californicus— (Continued) Kellogg: Microtus californicus Group of Meadow Mice Sees S|] SesS'> JOO} PUI | mND Ww AND NAN N NANANN eSVoey ea eee) EAGT SEY IT, Re ERG Ee LO HLS to) LO SH OSH SH Seis) Sy Soe) ysuz] [FIOL | S9M =O MOnsH Ora rr OMOned eee re bo ih an hee aoe eeipupary giver ors. 4SRs en jowysuey |ReReE eR RRRRE rere re Sesser soos | ORR A OOO qvjourdoddQ | ~oo ~~ OMor Mapemeee er peo ee piowen | sGodeg of each cgay ovy[Nq 7v uN Nea Peco) a eecrits ued joys | COM GO COCO reser re ees eyeped amo, ie Se CS Rass ss UWApead sree | ei eae [eyeediejuT | ~ COCO OH DMDHMA HOnamstOon) a) eee tes Se es # [eriqaorsyuy | COHOD cD cMODODOD 3 ‘3 TSIM ilk oe Reese Ego eS cece. = oyeuossg | WrOoOnN KR RKROO & b oon ih oon hoa re res St ost . quduep ee ee ee 3 [SUN | ODD DD DADO o = Jesuay Jo Eee ee ES S BUI] Ivypise O19 219 OOos s pe Pdieaied Gal eae euiew OWI 3S yySue[ Sane a ee S [eseu-Ov1dIN9N | OOD we Reet e AANN A ANNAN 3 quay «| NO S NOAS eseqo,‘puo om oo 2000 8 Ue ence ese AN ABAA MOanh et (Oy aX Gye Glov ata OF hh ieee a arena eee ee GINGER COMnO aliitee ce ts ere ee te ~ (or) AQ 24 } See Peer a=|-x5) boise (ao is ie ag 8 ios : i 8 : ea a :O c= g S| ay = UO: = 26 = _ Eos a elaer Biz) =O oO geo o 3S n “= G ore Hi : 2 je S| n| a SO 2fl8 Cs SS) = 1) $10) MN : 60 q | Segue is Ce o) aS s ao } aie S45 SQ i.» ~ 3 4 MM era's = RPS = 8 =i see g G08 o8 CA? == on) = BS FS = a HHS UD On ORE eure ee BE Os a £8 GEA 4 Ao #99) || NohoNoO. Moe oD Ye) wqunn | BSPS L Be une | RRSS 8 Ss ’ MmMmNN NX ANN 20087 || 3 mi. S. Covelo, Mendocino Co., Calif... Microtus californicus aestwarinus OO SHLD OD NI SH OD SH SH NANNANNNAN eoscsss9ee°2 DOA SROAG A 1 CO SH O19 OOO ooo°o0°0c°o°o°oo S 219 19 09 CO OH 0 ee al eS ais C2 CO O1DIS HOO CD WAAWKOONO A a ee en ce en hc hc Heber nlite eri HOeN es) RRR RORKKKRER BD OD DAVID AI 09 1D 0D 09 69 69 HH od 09 Sree en cn ce ce a OCotoowwwnn so oon hoe ho hoe he hoe aes Sen fh nh oo a on ho a ae HAOMONT OM 19 19 6 19) © 6 19.19 xt See cen heen en cen Doe ae ce HOOT RDOIDO19 OOK OK DOA BOHR DNGBOD H 09 09 09 09 69 6 HOD HONDHONOID errr CHANONMYOS DAA DWOAAG SaaS Orr OOo Ohh O NANNAANANANNN CAANBPOOSOM WHOA KRADDH NANNANNNAN > © CO 1D HH 0 CO 1 aODSOAGanHoooO ALO OO NN OD 0) 09 09 oaquin Co., Calif......... 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Ors = i] ae ONC Ast ts (a\aS-2 & OM & 5 Aadzee -m o |8e5ep99a2 BR aa Axes: Sa. oo Ss f GLAM a a BOM Se SSC SPEDE S 2.2 GE ia -Os SSeSeSOSs des Bou7~ eee Sageakesa saSSess5e8 Sap ge 2 oS ee ES SAn OOURRES5AnA X98 | "Dd OF OF OF Or OF AMOHONMORNRD Joquinud DOAr~MOOtOS unasnwy | COMOOMNOON MO AABDSHAAOG Microtus californicus huperuthrus 107402|| La Grulla, Lower California, Mex............. MDIDINOOCS HoH Hd 19 HH NANNANNN esoesessce aed ak) LD UD 1D AD 1D 19 19) eceosesessee DOarnow Se ie Anna SH SARA BOHBWNHOD aA AA ne O19 01919 Ronrerene Ranoouon Honea AN Son fhaen ih ae han hee hae hae MODAN DOO & HHHOOCOCO Shan en hee aoe en oe rmRORONSt S019 19.19.1919 dea SERCH COUSIN BOOKN OO oom mots 09 6D 69 6 0 0 OD GUS SSUES) ay BOROOOSO ee heel aon than ioe en ae! Tr N19 09 CN SH HDAAARG FOOFABH DOO O19 10 ANNNANNNANN TAANS OD I~ AADHAAND MNANNANNNN BPODNO OOD NAGBOAaNS COCINCAANANN 107412| 92} La Grulla, Lower California, Mee ee 107392] 2| La Grulla, Lower California, Mex............. 107382| 2} La Grulla, Lower California, Mex............. 107372| 2| La Grulla, Lower California, Mex.............[ 107362| 2} La Grulla, Lower California, Mex............. 107352| 9} La Grulla, Lower California, Mex.............|4 [ Vou. 21 2 From Field Museum of Natural History. —— —" 3. be Si3 rary ~ Vol.17. 1 ce 17. 22. 23. 4, oe 6. UNIVERSITY ‘OF CALIFORNIA PUBLICATIONS—(Continued) On Binary and Multiple Fission in. Giardia muris (Grassi); by Charles Atwood Kofoid and Elizabeth Bohn Christiansen. Pp, 30-54, oi, plates 5-8, 1 figure in text. : Nos. 2 and 3 in one cover. N ovember, 1915 Bm St ny ater fh IRE i es SOR On a New Trichomonad Flagellate, Trichomitus parvus, from the Intestine of Amphibians, by Olive Swezy. ‘Pp. 89-94, plate 12. Nos. 6 and 7. in one cover. December, 1915 OS eee Ra OC a eT im Pe On -Blepharcorys equi, sp. nov., a New Ciliate from the Caecum of the Horse, by Irwin C. Schumacher. Pp. 95-106, plate 13. December, 1915... Three New Helices from deren esp by. 8S. Stillman Berry. Pp. 107-111. ML ADEI BEY, LORD re Bai ccs NN ta cen ee PE NG On Trypanosoma triatomae, a New Flagellate from a Hemipteran Bug from the Nests of the Wood Rat Neotoma fuscipes, by Charles Atwood Kofoid and Irene McCulloch. Pp. 113-126, piates 14-15, February, 1916 .............. The Genera Monocercomonas and Polymastix, by Olive Swezy.: Pp. 127-138, Plates=t-6-17 « Fepruary,= 1916 eo ne ee a Notes on the Spiny Lobster (Panulirus interruptus) of the California Coast, by Bennet M. Allen, Pp. 139-152, 2 figures in text. March, 1916 ............ Notes on the Marine Fishes of California, by Carl L. Hubbs. Pp. 153-169, plates: £o-20s" Maren, (O16. he eS Re ee Sr ee The Feeding Habits and Food of Pelagic Copepods and the Question of Nutrition by Organic Substances in Solution in the Water, by Calvin O, Esterly. Pp. 171-184, 2 figures in text. March, 1916 ...:..... pee ye care ee The Kinetonucleus of Flagellates and the Binuclear Theory of Hartmann, by Olive Swezy. Pp, 185-240, 58 figures in text. March, 1916 22202.0000a0 On the Life-History of a Soil Amoeba, by Charlie Woodruff Wilson. Pp. Dat OR —WIAtes LOO COULy, BOLO sei me ee aie ee Distribution of Land Vertebrates of Southeastern. Washington, by Lee Raymond Dice. Pp. 293-348, plates 24-26. June, 1916 oo... cece The Anatomy of Heptanchus maculatus: the Endoskeleton, by J. Frank Daniel, —Pp. 349-370, pls. 27-29, 8 text figures. December, 1916 . 22.2... . Some Phases of Spermatogenesis in’the Mouse, by Harry B. Yocom. Pp. SFI-SSOe plate SOs” sanuar ye AOU eso: Se os Fee ag ea deh ee Specificity in Behavior and the Relation between Habits in Nature and Reactions in the Laboratory, by Calvin O. Esterly. Pp. 381-392. March, B Wt} Sf AaB Se So GNC Sota U US Oia APNE RNS fe ee Sle SS pO SAU UPS Dea ee The Occurrence of a Rhythm in the Geotropism of Two Species of Plank- ton Copepods when Certain Recurring External Conditions are Absent, by Calyvin-O.-Esterly.: Pp; 393-400.— Mareh;=191 7.) oon Ben On Some New Species of Aphroditidae from the Coast of California, by Christine: Essenberg. Pp. 401-480, plates 31-37.. March, 1917 2.002.220. Notes.on the Natural History and Behavior of Emerita analoga (Stimpson), by Harold Tupper Mead. Pp. 431-438, 1 text figure. April, 1917 _............ Ascidians of-the Littoral Zone of Southern California, by Willlam EB. Ritter and Ruth A. Forsyth. Pp. 439-512, plates 38-46. August, 1917 -................. Index in preparation. Diagnoses of Seven New Mamnials from East-Central California, by J ‘oseph -- Grinnell and Tracy I. Storer. Pp. 1-8. . A New Bat of the Genus Myotis from the High Sierra Nevada of Cali- fornia, by Hilda Wood Grinnell. Pp. 9-10. Nos. 1 and 2 in one cover. August, 1916 .2sco... csc cece scene ceneesesesee . Spelerpes--platycephalus, a New Alpine Salamander from the Yosemite National Park, California, by Charles Lewis Camp. Pp. 11-14. Septem- ve) RAE SSF ch Merce SE ile AUD i Od Ml tes pd VLR Sahn eS E oS ee RNG Anal an A aeg we A New Spermophile from the San Joaquin Valley, California, with Notes ‘on Ammospermophilus nelsoni nelsoni Merriam, by Walter P. Taylor. Pp. 15-20;-1- figure: in-text.) October, 1916: fois eo esis Sort bcccheeae Habits and Food of the Roadrunner in California, by Harold C. Bryant, Pp. 21-58, plates 1-4, 2 figures in text. October, 1916... tc. Description of Bufo canorus, a New Toad from the Yosemite National Park, by Charles Lewis Camp. Pp. 59-62, 4 figures in text. ee 1916...... ve il. 18. m 16. Vol. 21. 1. UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 2 The Subspecies of Sceloporus occidentalis, with Description of a New Form from the Sierra Nevada and Systematic Notes on Other California ~ Lizards, by Charles Lewis Camp. Pp. 63-74.. December, 1916 _..........2...2... Osteological Relationships of Three Species of Beavers, by F. Harvey Holden. Pp. 75-114, plates 5-12, 18 text figures. March, 1917 ................. Notes on the Systematic Status of the Toads and Frogs of California, by Charles Lewis Camp. Pp. 115-125, 3 text figures. February, 1917 ............ A Distributional List of the Amphibians and Reptiles of California, by Joseph Grinnell and Charles Lewis Camp. Pp. 127-208, 14 figures in text. PCT Ae A 5S” Alaa ne RRS a ORO et tae = REA ac Se Se Ante ESO AOE ROSE RE A Study of the Races of the White-Fronted Goose (Anser albifrons) Occur- ring in California, by H. 8S. Swarth and Harold ©. Bryant. Pp. 200-222, . 2 figures in text, plate 13. October, 19U7 noise eee teeenenerreenee A Synopsis of the Bats of California, by Hilda Wood Grinnell. Pp. 223-404, plates 14-24, 24 text figures.. January 31, 1918 oo cece The Pacific Coast Jays of the Genus Aphelocoma, by E. 8. Swarth. Pp. 405-422, 1 figure in text. February 23, 1918 — 20.2. c en es Six New ‘Mammals from the Mohave Desert and nyo Regions of ‘California, by Joseph Grinnell. . Pp. 423-430. . Notes on Some Bats from Alaska and British Columbia, by Hilda Wood j Grinnell. Pp. 431-433, Nos. 14 and 15 in one cover: April, TOTB se ee ed eee Revision of the Rodent Genus Aplodontia, by Walter P. ‘Paylor. Pp... 435- 604; plates 25-29,''16 text figures, May TLS no ace ceousareecesteuacccase The Subspecies of the Mountain Chickadee, by Joseph Grinnell. Pp. 505- BTSs 3S CORe SUE ss IMA, LORS Ss ee Lg sae bap ene esoeeE o Excavations of Burrows of the Rodent Aplodontia, with Observations on the Habits of the Animal, by Charles Lewis Camp. Pp. 517-536, 6 figures in’ text. June, gS BR Se caebecea Rai 2 Mor Some ae ale LC AO Ov RIG Nie FR Index in preparation. . Mitosis in Giardia microti, by William C. Boeck. Pp. 1-26, plate 1. Octo- 11s) Pn Es Wy Cig Aa a re aR na ir hace ear cet PNUD ts Ph Pao neNigeeae a ON EET em An Unusual Extension of the Distribution of the Shipworm in San. Fran- cisco Bay, California, by Albert L. Barrows. Pp. 27-43. December, 1917. . Description of Some New Species of Polynoidae from the Coast of Call- : fornia, by Christine Essenberg. Pp. 45-60, plates 2:3. October, 1917... New Species of Amphinomidae from the Pacific Coast, by Christine sions berg. Pp. 61-74, plates 4-5... October, 1917 + ...2:2 oo ee ck ogee Crithidia euryophthalmi, sp. nov., from the Hemipteran Bug, Euryophthalmus convivus Stal, by Irene McCulloch. Pp. 75-88, 35 text figures. Decem- DOE SORT Se Ns a eR OE Rs Sec Se toe re On the Orientation of Erythropsis, by Charles Atwood Kofoid and Olive Swezy. Pp. 89-102, 12: figures in ‘text. December, 1917 22... knees . The Transmission of. Nervous Impulses in Relation to Locomotion in the Earthworm, by John F, Bovard, Pp. 103-134, 14 figures in text. : January, 1S ee ee eee ee eee ee ee aicg Slaps Nach delta ocea U aa RR AA Ree RE The Function of the Giant Fibers in Earthworms, by Foun F. “Bovara. EDs: 185-144,.1 figure intext. “January, 1918 sc. cae cence oees toca n tebececs ape Rapid Method for the Detection of Protozoan Cysts in ‘Mammalian Faeces, by William C. Boeck. Pp. 145-149. December, 1917 ..................— The Musculature of Heptanchus maculatus, by Pirie Davidson... Pp. 161-170, : 12-figures:in text.. “Miarch,< 1938 ce oi ease crc een botw bc ddacnneenstrncee The Factors Controlling the Distribution of the Polynoidae of the Pacific - Coast of North America, by Christine Essenberg. Pp. a71=288, plates 6-8, & firurés in ‘text, (Aa rch TOU Sse Pa cay capa mn ane ra tr gar omenne meee . Differentials in Behavior of the Two Generations of Shiba democratica ~ Relative to the Temperature of the Sea, by Ellis L. Michael. Pp. 239-298, plates 9-11, 1 figure in text. “March, T9418: o-oo eee rhcees A Quantitative Analysis of the Molluscan Fauna of San Francisco Bay, by E. L. Packard. Pp. 299-336, plates 12-18, 6 figs. in text. April, 1918... . The Neuromoter Apparatus of Huplotes patella, by Harty B. Yocom, Pp. 337-396, plates 14:16. -September, 1918. 5.0)..22.-22 2c k0e 2S . The Significance of Skeletal Variations in the Genus Peridiniwn, by A. L. Barrows. Pp. 397-478, plates 17-20, 19 figures in text. .June, 1918-2002... The Subclavian Vein and its Relations in Elasmobranch Fishes, by J. Frank Daniel.. Pp. 479-484, 2 figures in text. August, 1918 o.oo... A Revision: of the Microtus Californicus Group of Meadow Mice, by Rem- ington Kellogg. Pp. 1-42, 1 figure in text. ° fattened b LB hs Rai SRR Cary CA te PS ee = UNIVERSITY OF CALIFORNIA PUBLICATIONS cS ee Note—The University of California Publications are offered in exchange for the publi- © cations of learned societies and institutions, universities, and libraries. Complete lists of © all the publications of the University will be sent upon request. For sample copies, lists — of publications or other information, address the MANAGER OF THE UNIVERSITY PRESS, BERKELEY, CALIFORNIA, U. 8. A. All matter sent in exchange should be addressed to THE EXCHANGE DEPARTMENT; UNIVERSITY LIBRARY, BERKELEY, » CALIFORNIA, U.S.A, WILLIAM WESLEY & SONS, LONDON Agent for the series in American Archaeology and Ethnology, eee, Geology, Physiology, and Zoology. ZOOLOGY. —W. E. Ritter and ©. A. Kofoid, Editors. Price per volume, $3. 50; beginning with vol. 11, $5.00. 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A Report upon the Physical Conditions in San Francisco Bay, Based upon. the Operations of the United States Fisheries Steamer ‘‘Albatross’’ dur- ing the Years 1912 and 1913, by F. B. Sumner, G. D, Louderback, W. L. — Ne Schmitt, E, C. Johnston. Pp. 1-198, plates 1- 13, 20 text figures. July, = ~ aS) Peper psreer, Sa cean ie oor Geel Me Me ne gs VES @ BLAME NI een on PMU ae MS teats ae 2 - 9, Molluscan Fauna from San Francisco Bay, by HE... Packard. - Pp. 199-452, Ps plates14-60. September, 1918 ron soccer canes tctgenee ns eb ae WEE aa Pen Volume 15, 1915-1916, 360 pages, with 38 plates ......22 pe ir aires Saete, S e ee So git pat a ee Volume 16, 1915- 1917, 522 pages, with 46 plates PS ae MO ee SOR SERRE i Os AA SENS Volume 17, 1916-1918, 545 pages, with 24 plates Bele nt acs: grin eee rex he ee etn ae DO. Vol. 17. 1. Diagnoses of Seven New Mammals from East-Central California, by J oseph Grinnell and Tracy I. Storer. Pp. 1-8. - 2. A New Bat of the Genus Myotis from the High Sierra Nevada of Cali- fornia, by Hilda Wood Grinnell. Pp, 9-10. Nos. 1 and 2 in one cover, August, 1916 witntasentersseanerntnensencedsneenenscnsnaannensten 3. Snelerpes. platycephalus, a. New. Alpine Salamander from the~ Yosemite National Park, California, by Charles Lewis Camp. Pp. 11-14. Septem-~ Boss pee BE SRS pee. Amstel ee ya eae a res Settee pup A elena tei oe bP a oo ae 4, A New Spermophile from the San J oaquin Valley, California, with Notes on Ammospermophilus nelsoni nelsoni Merriam, by Walter P. Taylor. Pp. ae 15-20;-1 figure in “text. October = 1 O06 ae eras wcacceseee te honk aetpca eon ccensctageeaes “06”. 5. Habits and Food of the Roadrunner in California, by Harold Cc. Bryant. See Pp. 21-58, plates 1-4, 2 figures in: text,-~October;, 19165s2:2 a ae ge 35 Rein 6. Description of Bufo canorus, a New Toad from the Yosemite National Park, py Charles Lewis Camp. Pp, 59-62, 4 figures in text. November, 1916... .05 > fi The Subspecies of Sceloporus occidentalis, with Description of a New Form ~from the Sierra Nevada and Systematic Notes on Other California Lizards, by Charles Lewis Camp, Pp. 68-74. December, 1916 ici ---sveecn- 10° SS UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Vol. 21, No. 2, pp. 43-47 March 29, 1919 FIVE NEW FIVE-TOED KANGAROO RATS FROM CALIFORNIA BY JOSEPH GRINNELL (Contribution from the Museum of Vertebrate Zoology of the University of California) Study of the collection of five-toed kangaroo rats (genus Perodipus ) contained in the Museum of Vertebrate Zoology discloses the existence within the state of California of five species not heretofore named. Ability to deseribe these now is mainly due to the successful efforts during the past two years on the part of our field collectors to secure topotype series of those species which had already been described. This topotype material has made it possible to define clearly the main characters of the species as would have been impossible from the published descriptions alone. Perodipus elephantinus, new species Elephant-eared Kangaroo Rat Type.—Male adult, skin and skull; no. 28511, Mus. Vert. Zool. :; 1300 feet altitude, one mile north of Cook P. O., Bear Valley, San Benito County, California; July 9, 1918; collected by Halsted G. White; original no. 2296. Diagnosis —A large sized, very large-eared, long-tailed Perodipus, of moderately dark tone of coloration (about as in P. agilis [Gambel] ) ; skull with very large mastoid and auditory bullae, very narrow supra- occipital and inter-parietal, nasals flaring at ends, and tapering and weakly angled maxillary arch; incisors heavy. Comparisons.—In matter of general size, P. elephantinus is, of all the known species of Perodipus, smaller only than P. ingens Merriam. From this species it differs in that the ear is very much larger, the body smaller, the hind foot shorter, the coloration darker in tone, the skull generally smaller, narrower and more shallow, the jugal more slender, and the maxillary arch tapering and less strongly angled. 44 University of California Publications in Zoology [Vou. 21 From P. tularensis Merriam, P. elephantinus differs in much larger size, much larger ear, darker general coloration, heavier dentition, much larger mastoid and auditory bullae, less strongly angled max- illary arch, heavier malar bar, and in much heavier rostrum, the end of which flares conspicuously. From P. agilis, P. elephantinus differs ereatly in general size (though not in coloration), the ear is very much larger, the rostrum heavier, the nasals flare at their ends, and the bullae are proportionally greater in size. Other details of difference are numerous, but each seems to be correlated more or less directly with one or more of the several features already noted. Measurements—The average measurements of ten adult speci- mens, five males and five females, are as follows: total length 324 mm. ; tail vertebrae 197 mm.; hind foot 46.8 mm.; ear from crown 17.4 mm. ; ereatest length of skull 43.0 mm.; width of skull across bullae 26.3 mm.; greatest length of nasals 15.7 mm.; width of rostrum near end 4.8 mm.; width of maxillary arch at middle 5.2 mm. Distribution——The only material at hand representative of this species is from the type locality, as specified above. This consists of twenty-seven skins with skulls, adults and young, taken July 8 to July 21, 1918, by the Museum field men, Halsted G. White and Richard M. Hunt. So far as ascertained up to the present time this mammal is an exclusive inhabitant of dense chaparral. Remarks —tThe relationship of this form with any one of the other Californian species which have been previously described is not close ; P. elephantinus, in combination of characters, stands quite alone. The very large ear and its shape and manner of folding has suggested the name selected. Perodipus swarthi, new species Carrizo Plain Kangaroo Rat Type—Male adult, skin and skull; no. 14440, Mus. Vert. Zool. ; seven miles southeast of Simmler, Carrizo Plain, San Luis Obispo County, California; May 26, 1911; collected by H. 8. Swarth and W. L. Chandler; original no. 9144, H.S.S. Diagnosis—Most nearly resembles Perodipus tularensis Merriam, from whieh it differs in brighter ochraceous-buff coloration, larger skull, and decidedly more inflated bullae. Comparisons.—This is the most brightly colored, that is, the most clearly ochraceous-buff, of all the species of Perodipus so far known from west of the desert divides. The bullar inflation is near the maximum, relatively to general size of skull. Measurements.—The average measurements of six adult specimens, two males and four females, are as follows: total length 297 mm.; tail vertebrae 175 mm.; hind foot 43.5 mm.; ear, from crown, 11 mm. ; greatest length of skull 41.1 mm.; breadth of skull across bullae 25.8 mm.; greatest length of nasals 15.0 mm.; width of rostrum near end 4.1 mm.; width of maxillary arch at middle 5.1 mm. 1919] Grinnell: New Kangaroo Rats A5 Distribution.—The material at hand representing this species con- sists of seven specimens, all from points near together, about seven miles southeast of Simmler, on the Carrizo Plain, 2000 feet altitude, San Luis Obispo County, California. Remarks—The name is selected as a recognition of the success im general vertebrate field work of Mr. Harry 8. Swarth, of the staff of the Museum of Vertebrate Zoology. Perodipus dixoni, new species Merced Kangaroo Rat Type—Male adult, skin and skull: no. 26805, Mus. Vert. Zool. ; Delhi, near Merced River, in Merced County, California; Mareh 235 1917; collected by J. Dixon; original no. 5580. Diagnosis —A medium-sized kangaroo rat, slightly darker in color than Perodipus tularensis Merriam, but paler than P. streatort Mer- riam. Bullae smaller, and rostrum and dentition much weaker than in either of those species. Comparisons.—Perodipus dixont is conspicuously different from P. tularensis, its neighbor to the south on the floor of the San Joaquin Valley, on comparison of the skulls. The bullae are much less inflated, the nasals are much shorter and narrower, and the dentition is weaker. The supraoccipital and interparietal, however, are quite as narrow, and the relative width and angulation of the maxillary arch are much the same. As compared with P. streatori, the form belonging to the Upper Sonoran zone of the adjacent Sierran slopes, the skull of P. dixoni is much smaller, and again the rostrum and bullae show marked inferiority in volume, though, with the bullae, not to so great a degree as in the case of P. tularensis. Externally P. dixont shows no special peculiarities except in dimensions; the body is smaller than in either of its near neighbors, and its tail is relatively longer. Measurements.—The average measurements of ten adult or sub- adult specimens are as follows: total leneth 283 mm.; tail vertebrae 170 mm.; hind foot 41.1 mm.; ear, from crown, 14.1 mm.; greatest length of skull 37.0 mm. ; breadth of skull across bullae 23.2 mm. ; ereatest length of nasals 13.2 mm. ; width of rostrum near end 3.6 mm. ; width of maxillary arch at middle 4.5 mm. Distribution.—Material is at hand only from the east side of the lower San Joaquin Valley, in Stanislaus and Merced counties, and is represented by localities as follows. Stanislaus County: three miles south of Lagrange, 3. Merced County: near Delhi, 2; five miles north of Snelling, 12; one and one-half miles south of Merced Falls, 1. Remarks.—This new form is named in recognition of the pains- taking quality of the field work carried on by Mr. Joseph Dixon in the interests of the Museum of Vertebrate Zoology. It is through his success in specially searching for kangaroo rats that some of our most valuable specimens have been obtained. 46 University of California Publications in Zoology [Vou. 21 Perodipus leucogenys, new species Pale-faced Kangaroo Rat Type—Male adult, skin and skull; no. 26938, Mus. Vert. Zool.; 5600 feet altitude, Pellisier Ranch, five miles north of Benton Station, Mono County, California; September 20, 1917; collected by J. Dixon; original no. 6371. Diagnosis—A large kangaroo rat, closely similar to Perodipus panamintinus Merriam in size and general appearance, but differing in paler tone of coloration, greater extent of white markings, more heavily haired tail, shehtly smaller bullae, and heavier rostrum. Comparisons —More in detail, Perodipus leucogenys differs from P. panamintinus as follows: less black or dusky and more white about face and ears (white spot above eye larger, cheek whiter, forehead paler, hght patch on upper rim of ear whiter and larger, dark nose markings less distinctly black) ; white bar across thigh broader; tone of coloration more pinkish buff, with less of dusky intermixture down middle of back; tail distinctly ‘crested to nearly or quite halfway from tip to base; white side-stripes of tail broader, in other words, black stripes narrower and the lower one of these playing out short of tip of tail; feet (in fall specimens at least) more densely haired. From P. mohavensis Grinnell, P. lewcogenys differs in darker, less brightly ochraceous-buff body color, generally larger size, bushier tail toward end, relatively heavier rostrum anéd dentition, and broader supra- occipital. Measurements——The average measurements of ten adult and sub- adult specimens, five males and five females, are as follows: total length 301 mm.; tail vertebrae 176 mm.; hind foot 45.2 mm.; ear, from crown, 12.4 mm.; greatest length of skull 40.4 mm.; width of skull across bullae 24.38 mm.; greatest length of nasals 15.4 mm.; width of rostrum near end 4.4 mm.; width of maxillary arch at middle 5.4 mm. Distribution area.—The sagebrush territory southeast of Mono Lake and about the head of Owens Valley, thence southward, at least along the west side of Owens Valley, as far as vicinity of Indepen- dence. Material is at hand as follows. Mono County: Mono Mills, 7300 feet, 10; Dry Creek, 6700-6900 feet, near Mono Lake, 3; Mono Craters, 6600-7100 feet, 3; Pellisier Ranch, 5600 feet, five miles north of Benton Station, 12; Benton, 5639 feet, 6; Taylor Ranch, 5300 feet, two miles south of Benton Station, 29. Inyo County: Farrington Ranch, 4100 feet, west of Laws, 9; Walters Ranch, 3900 feet, two miles north-northwest of Independence, 19. Perodipus monoensis, new species Mono Kangaroo Rat Type.—FK emale adult, skin and skull; no. 27002, Mus. Vert. Zool. ; 5600 feet altitude, Pellisier Ranch, five miles north of Benton Station, Mono County, California; September 21, 1917; collected by J. Dixon; original no. 6384. 1919] Grinnell: New Kangaroo Rats 47 Diagnosis—A member of the ordi group of five-toed kangaroo rats. Differs from Perodipus columbianus Merriam as follows: in paler tone of coloration, which involves much reduction of black or dusky on face, ears, and tail, as well as a decidedly more creamy tint of body color; in larger ear; and in larger and more backward- projecting bullae. Differs from its near neighbor, the altogether distinet P. microps Merriam, in the following characteristics: larger ear, smaller foot, shorter tail, much broader, more sharply angled, and more widely spreading maxillary arches, longer nasals, which are also broader posteriorly, broader interparietal, and more incurved incisors. Measurements.—The average measurements of ten adult and sub- adult specimens, five males and five females, are as follows: total length 233 mm.; tail vertebrae 125 mm.; hind foot 38.7 mm.; ear, from erown, 11.1 mm.; greatest length of skull 36.9 mm.; width of skull across bullae 23.3 mm.; greatest length of nasals 13.7 mm.; width of rostrum near end 3.5 mm.; width of maxillary arch at middle 4.3 mm. Distribution.—The material at hand consists of fourteen adult or nearly adult specimens, all from the sagebrush flats in the extreme head of Owens Valley, in the vicinity of Benton, Mono County, Cali- fornia. Three places are represented: Pellisier Ranch, 5600 feet alti- tude, five miles north of Benton Station; Taylor Ranch, 5300 feet, two miles south of Benton Station; and the old town of Benton, 5639 feet altitude, four miles west of Benton Station. Remarks—Perodipus levipes Merriam, Perodipus microps, and Perodipus monoensis, although roughly similar to one another in external appearance, are specifically distinct; each has excellent eranial characters distinguishing it from either of the others, and no evidence of intergradation geographically is at hand. P. monoensis is apparently most nearly related to P. columbianus, of the north- western Great Basin. The latter form is represented in the collection of the Museum of Vertebrate Zoology, from within the state of Cali- fornia, by four specimens from Vinton, Plumas County. Perodipus columbianus seems not to have been recorded from California pre- viously to this note. Transmitted November 26, 1918. eS eae si is Ff) * a ‘ we: . vo) ahs - e eae, Po ee ? lio if. 4 r 4 I 74 f rar ‘ da . ac hon ns. Pa ee Han, wee UNIVERSITY OF CALIFORNIA PUnLiGatroNn qeeucucay 8. Osteological Relationships of Three Species of Beavers, by F. Harvey Holden. Pp. 75-114, plates 5-12, 18 text figures. March, 1917... BA 9, Notes on the Systematic Status of the Toads and Progs of California, by ~ Charles Lewis Camp. Pp. 115-125, 3 text figures. February, 1917............. 10,.A Distributional List of the Amphibians and Reptiles of California, by Joseph Grinnell and Charles Lewis Camp. Pp. 127-208, 14 figures in text. BES i Pah AY Sn aR etn ime eres eed ad lle A ae MO ca ES Aces Re ae 11. A Study of the Races of the White-Pronted Goose (Anser albifrons) Occur- ring in California, by H. 8S. Swarth and Harold c, Bryant. Pp. 209: 222, 2 figures in text, plate 13. October, 1917 ..ccc.c.ccccsccccecceccscecscecceeceoccsastzeeecesdecee 12, A Synopsis of the-Bats of California, by Hilda Wood Grinnell. Pp. 223-404, plates 14-24, 24 text figures. January 31, 1918 220000 e i 13. The. Pacific Coast Jays of the Genus za RIEL tlh, by H.-S. pfu dant Sn Pp. 405-422, 1 figure in text. February 23,1918 <0... = 14, Six New Mammals from the Mohave Desert — Inyo Regions of California, - by Joseph Grinnell, Pp. 423-430. 15. Notes on Some Bats from Alaska and British Columbia, by Hilda Wood . Grinnell... Pp. 431-433. Nos. 14 and 15 in one cover. April, OA Gs hey) ote 16. Revision of the Rodent Genus Aplodontia, by Walter P. Taylor. Pp. 435- ‘504, plates 25-29,:16 text figures. May, 1918 -2.ooct ici eceectcte eee sy 17. The Subspecies of the Mountain Chickadee, by Joseph Gingell Pp. 505- x OLD, Srbext: Heures; May; BOLG <2 oa a os ee a eee : -18, Excavations of Burrows of the Rodent Aplodontia, with Observations on the Habits of the Animal, by Charles Lewis Camp. Pp. 517-536, 6 figures in text... June, 1918 2... PEE EES aD Ss Aen a Ee Reap, Se ee ME eer Index, pp. 537-545, Re Vol 18. 1, Mitosis in @ariia microti, by William Cc. Boeck. Pp. 1-26, plate 1.° Octo- Sy Het EY Ee RSE aR caer ei RES nS eas eee Msc hs eign IO 2. An Unusual Extension of the Distribution of the Shipworm in San ‘Fran- “cisco Bay, California, by Albert L. Barrows. Pp. 27-43. December, 1917. 8. Description.of Some New Species of Polynoidae from the Coast of Cali- _ fornia, by Christine Essenberg. Pp.45-60, plates 2-3: October, 1917 ...... 4, New Species of Amphinomidae from the Pacific Coast, by Christine Essen: berg. Pp. 61-74, plates 4-5. October, 1917) 2o..co.occ ccc icc ccecscdedecceee 5. Crithidia ewryophthalmi, sp. nov., from the Hemipteran Bug, DuPyiphihahient conyiwus Stal, by Irene ssiuahe gag Pp. tha 88, BP text Agures. Decem- BBP SOE ee Sra eas Se a eA eee ee ee ee a oe 6, On the Orientation of Erythropsis, by Charles Atwood Kofoid and Olive Swezy. Pp, 89-102, 12 figures.in text. December, 1917 sone a eee ae 7. The Transmission of Nervous Impulses in Relation to Locomotion in the Earthworm, by John FE Bovard.: Pp. 103-134, 14 figures in text. January, OG a a ae ae an SE Ee RD ae ee 8, The Function of the Giant Fibers in Earthworms, by 2 ohn F, Bovard. Pp. 22135-144.-1- fioure-in text. January, 1018 We a a. 9. A Rapid Method for the Detection of Protozoan Cysts in Mammalian Faeces, by William C. Boeck. Pp. 145-149. December, 1917... 2... 10. The Musculature of Heptanchus maculatus, by Pirie Davidson... Pp. 151- 170, SOT Oures An: teRe., “Mar Che OLS. se eS A, a eead a acs 11. The Factors Controlling the Distribution of the. Polynoidae of the Pacific ~ Coast of North America, by Christine Essenberg. Pp. 171-238, plates 6-8, o-nguresein text; Waren; S101 8 a atc Sas SE emi 12. Differentials in Behavior of the Two Generations of Salpa democratica Relative to the Temperature of the Sea, by Ellis L. Michael. Pp. 259-298, plates 9-11, 1 figure in text. March, 1918 ooo cece ecto ncecen sect ecere 1s. A Quantitative Analysis of the Molluscan Fauna of San Francisco Bay, by E. L. Packard. Pp. 299-336, plates 12-13, 6 figs. in text. April, 1918... 14, The Neuromotor Apparatus of. Euplotes patella, by Harry B. Yocom, Pp. 837-396, plates 14-16.” September, 19189 ocean aoe ects neee 15, The Significance of Skeletal Variations in the Genus Peridinium, by A. L. Barrows, Pp. 397-478, plates 17-20, 19 figures in text. June, 1918 ...........: 10 2.00 _ UNIVERSITY OF CALIFORNIA PUBLIATIONS— (Continued) 16. “The Subclavian Vein and its Relations in ‘Elasmobranch Fishes, by cs Frank Daniel. Pp. 479-484, 2 figures in text. August, 1918.0... ‘17. The Cerearia of the Japanese Blood Fluke, Schistosoma Japonicum Kat- surada, by William W. Cort. Pp, 485-507, 3 figures in text. | e, 18. Notes on the Eggs and Miracidia of the Human Schistosomes, by William A -W. Cort. Pp. 509-519, 7 figures in text. - Nos. 17 and 18 in one cover. pany) 1919 eat ORR NS i tan ice ere ; Index in preparation. Vol. 19. 1. Reaction of Various Plankton Animals with Reference to their Diurnal Migrations; by.:C.°O. Esterly. PD, 1-83 snc css eect ctcc cn Sendeecteedonen ey 2. The Pteropod Desmopterus Pacificus (sp. nov. ), by Christine Essenberg. Pp...00-88, 2 fisures an stext cso 5 Sa een es capeecs vane dea cepa ae 3. Studies on Giardia microti, by William C. Boeck. Pp. 85- 136, plate 1, 19 fe eae PiOUTES I CORY: eee ee re OO a Neem aa ce aN cee as 4, A Comparison of the Morphology and the Life Cycle of Crithidia with those eat of the Crithidial Stages of Trypanosoma, by Irene A. McCulloch....... (In press). 5 5. A Muscid Larva of the San Francisco Bay Region which Sucks the Blood f of Nestling Birds, by O. E, Plath. Pp. 191-200. February, 1919 —..22..0.... 2s AOS Vol. 20. Studies on the Termites, by C. A. Kofoid and Olive Swezy......2-2.2i0c.0cc (In ney a Vol, 21. 1..A Revision of the Microtus Californicus Group of Meadow Mice, by Rem- ington Kellogg. Pp. 1-42, 1 figure in text. December, 1918 .....2220.0..222... 00 2, Five New Five-toed Kangaroo Rats from California, by. Joseph. 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November, 1916... ap: } UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Vol. 21, No. 3, pp. 49-74, plates 1-3, 3 figures in text December 10, 1919 NOTES ON THE NATURAL HISTORY OF THE BUSHY-TAILED WOOD RATS OF TTT Ms BY ; 2 JOSEPH DIXON (Contribution from the Museum of Vertebrate Zoology of the University of California) CONTENTS PAGE TEraY waeVONG HUGH SS Uae PS apse Na Sar a ee So Ee A So ee eek ENE a nea eee ey ee 49 GROVER CG) SU OFS) OSV OUNCENSS trot Car a re a YR ee er PE eee 50 FHL OS Sill TUNG pete eee aoe me ane erucn ek Sie Ae Se etree 3 fhe SC le te et al 57 IDeScrTer ONDA, chap Meas Leon ses See RAT EL Pl Eee Sees CE nse erate cn OCR iy aoe SBiA ER ERS SE TERE opr 57 PIT Cy ew eANS SC cats OT See ee eae eas ey a ene ge 2 ene ae Beep eee 58 Venere prea Gate leat Ole seen seen eh ete eel. PB AEE 2 eee ool. 58 Avviramal CEN cereal rete Ce ey) aaa eee ae ha om eee ve ns ge ee Re Uae BR eee ere Sioa 59 “GET REANGLEGS): FAATANG IS Ghee SU Res ee eae Ree es MR ne ea 60 TELRUUSIEED eee cee) EY aN i See Rn a ee Oe ee ne Meee Re cee 2 eee, 61 PE=Tiil fo @rsbes te OT epee ee ee es eek OM ee PR AE BI EE IRS a RRA. ea 2. 2 to 62 SEES CIM OMS EAS OMe eae es Ata ean MEA Neate LE Neto Ree CS tO UE Sd arrears 62 (ire OsyvcU nba Oile yO UT Oars 8 Es Se Ee eae a ee Jad ged scenes es ceennsenee 63 Ve AONE CO ROL NG wala | Specs tee eres Deca RE Deg ee ee Se ee eee 64 TEN rat rath ofan ec ee ae eG ea Ne pee Cee rece Batra 20 Stren een Soe ea ee nea in Er 65 TENGYOXG! sa giana Pete AS EUS a ESS ane TEES ae es ce ee ee eh eee eer ee ea Uae ean eee 65 PEER ro ule 01 a pe ee eee Sate ae ae oh es ee ee eR cee 66 BE Cont CNA CS AALS eae eee ee Se eS ce nee es ee 67 Wiener GU Che Case (ees etek eee es et le A Bnd eeeo AR Sees eee er foe. JUN Pea ee een ed 68. INTRODUCTION It is the function of the present paper to place on record such facts as have been learned to date in regard to the habits and associational relationships of the bushy-tailed wood rats occurring in California. The chief available source of this information has lain in the field notebooks and series of specimens contained in the Museum of Verte- brate Zoology. The systematic position and subspecific constitution 50 University of California Publications in Zoology [Vou. 21 of this group of rodents (subgenus Teonoma of genus Neotoma) have already been given full attention by Goldman (1910). This phase of treatment is therefore here reduced to a minimum. Two subspecies of the one species, Neotoma cinerea, occur in Cali- fornia. These are so closely similar to one another in structural features that their natural history may well be treated in one account. But first, some of the diagnostic features of the two races will be set forth in some detail. THE TWO SUBSPECIES IN CALIFORNIA Neotoma cinerea cinerea (Ord) Gray Bushy-tailed Wood Rat Mus cinereus Ord (1815, p. 292). Original description. Neotoma cinerea, Baird (1857, p. 499). Teonoma cinerea, Elliot (1903b, p. 247). Teonoma cinerea acraia Elliot (1903b, pp. 247-248). Original description of supposed new form from vicinity of Mount Whitney, California. Teonoma cinerea acraea, Elliot (1904, p. 280). Teonoma acraea, Elliot (1905, p. 221). Neotoma cinerea cinerea, Miller (1912, p. 201). Type locality —Great Falls, Cascade County, Montana (fide Gold- man, 1910, p. 95). Diagnosis (as compared with the round-tailed wood rats, subgenera Neotoma and Homodontomys).—Size large. Average total length of adults, in millimeters: males 370, females 352; average weight of adults, in grams: males 295.6, females 252.2. Tail long, averaging 43 per cent of total length; average length of tail, in millimeters: males 159, females 152. Tail squirrel-like, bushy; hairs on each side of tail about 30 millimeters long. Skull with rostrum and incisive foramina long; sphenopalatine vacuities usually absent. A persistent musky odor is especially characteristic of this animal. Pelage—Of whole body, dense, long and soft. Dorsal surface, sides of body, and top and sides of tail, covered with soft underfur which is well concealed by the long overhairs. Sole of hind foot normally furred from heel to posterior tubercle (see fig. A). Fig. A. Hind foot of Neotoma cinerea cinerea, to show hairing on sole. This is seen to extend as far forward as the posterior tubercle. In California the regular annual molt of Neotoma cinerea cinerea takes place in September and early October. Although there is nor- mally but one change.of pelage each year, prompt regeneration of 1919 | Dixon: Bushy-tailed Wood Rats of Califorma 51 . pelage evidently occurs at any time when areas become denuded of hair through accident. This is well illustrated in an adult female (no. 24992) taken September 2, in which a triangular area 20 milli- meters in extent on the rump is thickly covered with a short growth of both underfur and overhair between 3 and 4 mm. long. The position of this spot is such that normally it would be one of the last portions of the body to be invested with the new pelage. No other indication of molt could be discovered in this individual, so the loss of pelage at this particular place was clearly due to accident. From an examination of the material at hand, it is apparent that the beginning, progress and termination of molt in Neotoma c. cinera is very similar to that of Peromyscus as described by Osgood (1909, p. 20) and by Collins (1918, pp. 78-79). Stated briefly, the growth of new hair, which is well concealed by the old, begins on the sides and progresses rapidly upwards on each side, meeting at the middle of the back and then inelosing the rest of the body. A small area above the base of the tail and another between the nose and shoulder are the last parts to be clothed with the new hair. (See Collins, 1918, p. 79, figs. 1-8.) The following five specimens have been chosen as best illustrating the time and method of molt in Neotoma cinerea cinered. No. 24993, 9 adult; Bullfrog Lake, 10,600 feet, Fresno County, September 2, 1916. No sign of molt; hairs slate gray to base; hairs on tail much worn. No. 23112, 9 adult; Walker Lake, 8000 feet, Mono County, September 9, 1915. New light buff hairs 5 mm. long on head, shoulders, sides and rump. The new overhairs are entirely black and 8 mm. long. These new hairs are concealed by the longer older hairs and can not be seen until the old hairs are separated and held apart. No. 23115, ¢ adult; Warren Fork, Leevining Creek, 9200 feet, Mono County, September 26, 1915. New hairs on rump and shoulders 8 mm. long. Tail well covered with new hairs 24 mm. long on sides, and with a thick woolly underfur 8 mm. long on top and sides. No. 26352, 2 adult; 3 miles east of Jackass Springs, 7200 feet, Panamint Moun- tains, Inyo County; October 8, 1917. Molt complete except on rump and shoulders where a few hairs are still light buff at base. No. 26353, ¢ adult; 3 miles east of Jackass Springs, 7200 feet, Panamint Mountains, Inyo County, October 8, 1917. Molt complete. New hairs slaty gray to base, except for a few hairs on nape which still show buffy at base. Color—General effect, ashy gray. Fresh pelage (adult, Octo- ber 2): Dorsal surface warm buff, palest on head; back and rump darkened by black tipping of overhairs. Ventral surface, including upper lip, white; dorsal surface of fore and hind feet white; ears faintly margined with white (see pl. 3, fig. 6), and scantily covered with grayish and brownish hairs outside, and brownish or blackish hairs inside. The pelage on the dorsum consists of two distinct types of hairs: a silky type forming the soft underfur, and coarser, longer, less numerous overhairs. A typical individual hair of the underfur is 12 mm. long and usually lacks any black tip although some are of the banded or agouti type with the tip black. The basal portion of such a hair is slate gray and the terminal portion light buff (see fig. B, a). A typical overhair is 22 mm. long and is slate gray at the base and black at the tip (see fig. B, b). A typical hair of the underfur on the 52 Umversity of California Publications in Zoology [Vou 21 sides is slate gray at base with terminal portion white, while the over- hairs are entirely white (see fig. B, c,d). Hair on underparts, includ- ing under side of tail, white from base to tip (see fig. B, e, f). All color terms used are according to Ridgway (1912). Adults in worn pelage sometimes acquire a rusty tinge over the entire dorsal surface. This appearance is due to the light pinkish cinnamon of the terminal portions of the underfur. The oldest adults, judging from cranial characters, are not always the rustiest, and in individuals of comparable age the rusty tinge is much less pronounced in fresh fall pelage than in the worn summer pelage. This change in color could not well be called fading, since the animals are nocturnal and the change is from a duller to a brighter hue. 8mm | black 4mm light buff 5mm | white 14mm \ all while Silke /4mm | Slate Hel z 7mm | slate qra 8mm | Slate gray j a b c d emm| all while roram \ all while e f Fig. B. Diagrams to illustrate coloration of typical hairs in pelage of Neo- toma cinerea cinerea: a, underfur of back; b, overhair of back; c, underfur of side of body; d, overhair of side; e, underfur of under surface of body; f, over- hair of under surface. The juvenal pelage of N. c. cinerea, as illustrated by a series of specimens from the Mount Whitney region and from the White Moun- tains, shows considerable variation in color, some specimens being much paler than others taken at the same time and locality. The underfur and overhairs are both shorter and finer than in the adults. The basal portions of these hairs are slate gray as in adults, but the terminal portion of the underfur is much lighter, being pale pinkish buff. Extensive black tipping of the overhair in certain juvenals obscures the paler underfur and gives the animals a dark appearance as viewed dorsally. In this connection it is interesting to note that certain young examples of cinerea from the Mount Whitney region are indistinguishable from juvenal co-types of Neotoma c. lucida from Charleston Peak, Nevada, this race having been described as being paler than cinerea (Goldman, 1917, p. 111). Cramal characters—The cranial characters and dentition of Neotoma have been so well covered by J. A. Allen (1894) and by Goldman (1910) that it seems unnecessary to deal with this subject in the present paper. It may not be amiss, however, to mention the 1919] Dixon: Bushy-tailed Wood Rats of California 53 fact that skulls of specimens from the Panamint Mountains, Inyo County, particularly those from Telescope Peak, near Death Valley, show some of the features which have been given (Goldman, 1917) as characterizing the race lucida, from Charleston Peak, within sight in e Neotoma cinerea cinerea o Neotoma cinerea occidentalis S 2 fat ios ‘ L eu Cm ayy Pie Set : DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA Fig. C. Map showing stations of occurrence in California of Neotoma cinerea cinerea and Neotoma cinerea occidentalis, as established by specimens examined by the author. Assumed general range of the species, cinerea, outlined. southern Nevada. For instance, in the specimens from Telescope Peak the zygomatic arches lie more nearly parallel to the axis of the skull than in typical cinerea. However, the bushy-tailed wood rats of Telescope Peak are seemingly much more closely similar to cinerea, the Sierra Nevadan form, than to lucida, and they are therefore placed under the name cinerea. 54 University of California Publications in Zoology [Vou. 21 Measurements and weights—In the accompanying table the average weights of twenty-nine adults are given along with the aver- age measurements of body and skull of thirty-nine adults. It is to be noted that the series from the southern Sierra Nevada, inclusive of both sexes, averages smaller than the comparable series from the Yosemite region. i) a? El Tora Cr Son Joaquin Valley \|Sa Jacinto Volley goon na Wash view Mts den Wilmingt | Long |Be. Santa |A Te Son i} Jacinto River Cotton wood wngton Lo remescol Loke an Eogle PD: Mts) Mountains by} qs: d San Pedro through San Jacinto Peak Fig. G. Four cross-sectional profiles of California. The shaded areas indicate the altitudinal distribution in summer of the subspecies of fox sparrow occurring within the state at that season. a. Brevicauda on the Yolla Bolly Mountains; mariposae on Lassen Peak; fulva on the Warner Mountains. b. Mariposae on the Sierra Nevada west of Mount Lyell; monoensis in the Mono Lake region on the east slope of the Sierra Nevada; canescens on the White Mountains. c. Stephensi on Mount Pinos and on the Sierra Nevada west of Mount Whitney ; mariposae on the Sierra Nevada east of Mount Whitney. d. Stephensi on San Jacinto Peak. through the interpolation of unsuitable territory, it never takes place through the interpolation of another race of fox sparrow. Regarding each isolated colony as a separate race (as in a sense they are, of course, though we can not always distinguish between them), and it is evident that any one line of variation extends continuously in a 1920] Swarth: Revision of Avian Genus Passerella 105 given direction, but that appreciable changes in appearance do not occur wherever there is discontinuity of range. Passerella is closely related to Melospiza. Fox sparrows are very like song sparrows, in structure, appearance, habits, and actions. The respective ranges also of Passerella iliaca and Melospiza melodia are suggestive of rather close relationship between the two, for their habitats are in a measure complementary since the two species, though frequently occurring in the same general regions, occupy different life zones. Passerella is mostly in the Boreal zone, descending oceasion- ally into high Transition, Melospiza mostly in the Sonoran zones, though some races ascend to Transition and higher. On the coast of Alaska, at the northern extreme of the range of Melospiza, the general separation of the two does not hold, neither Sonoran nor Transition extending this far north, but at the latitude of Vancouver Island we find the birds thus separated, Passerella on the Boreal mountain tops, and Melospiza in the Transition zone valleys. All of the races of Passerella iliaca are migratory. Some are much more so than others, and in a general way it may be said that the subspecies breeding farthest north migrate farthest south. Passerella iliaca iliaca winters far south of its Boreal nesting grounds, concentrating into a relatively limited area in the South Atlantic and Gulf states. Thus the lines of migration in this sub- species converge, birds from Labrador and Newfoundland traveling southwest and those from Alaska and Mackenzie southeast, toward their common winter home. Birds of the Unalascheensis group move directly southward along the Pacific eoast, each into a more or less definitely circumscribed winter habitat. The six subspecies of this group are arranged in summer, from north to south (from the Alaska Peninsula to Puget Sound) in the following order: wnalaschcensis, insularis, sinwosa, annectens, townsendi, fuliginosa. The subspecies breeding at the northern extreme, wnalaschcensis, insularis and sinuosa, move the farthest south in winter, passing completely over both summer and winter habitats of annectens, townsendi and fuliginosa, and reaching the extreme southern limits of California. Annectens winters mainly on the coast of California in the Santa Cruz and San Francisco Bay regions, passing over the habitats of townsendi and fuliginosa to reach this destination. Townsendi in turn leap-frogs over fuliginosa and winters in numbers as far south as Humboldt Bay, California. Fuliginosa hardly migrates at all. Occasional stragglers reach San 106 University of California Publications in Zoology (Vou. 21 Francisco Bay and a few more winter in the Humboldt Bay region, but the main winter home of this subspecies certainly lies north of the latter point. Undoubtedly, however, it performs an altitudinal migration, moving down from the mountain tops which constitute its summer home, as on Vancouver Island. (See fig. H.) Fig. H. Map showing in somewhat conventionalized form the nature of the migrations performed by the subspecies of the Unalaschcensis group. 1, wnalasch- censis; 2, insularis; 3, sinuosa; 4, annectens; 5, townsendi; 6, fuliginosa. A solid line surrounds the summer habitat; a dotted line surrounds the main winter habitat. The broken lines connecting summer and winter habitats do not necessarily indi- cate migration routes; they are inserted to emphasize the overleaping feature in the seasonal movements. The subspecies occupying the northernmost summer habitats (unalaschensis, insularis, and sinwosa) travel the farthest south. The subspecies occupying the southernmost summer habitat (fuliginosa) is practically resident there. As bearing upon the manner in which these birds migrate, certain facts gathered during several seasons of field work carried on by the California Museum of Vertebrate Zoology on the coast of Alaska and 1920] Swarth: Revision of Avian Genus Passerella 107 British Columbia are worthy of record, indicating a striking absence of migrants representing subspecies that might be supposed to occur commonly. An expedition to the Prince William Sound region, Alaska, covering the migration period in part at least, took no speci- mens of Passerella other than the breeding form, sinwosa. Of two expeditions to the Sitkan district, Alaska, each one covering nearly the entire migration period, both spring and fall, one collected nothing but the breeding fox sparrow, townsendi, the other secured, besides an abundance of townsendi, a single migrating individual of another race, insularis. An expedition to Vancouver Island, covering both spring and fall migration, secured, besides the breeding fuliginosa, one migrating individual of insuwlaris and three of townsend. I have at hand, however, two specimens of wnalaschcensis taken at Ketchikan, on the coast of southeastern Alaska, April 12 and 15, 1916, migrating birds, of course, and proof that some individuals, at least, do pass through this region. On the whole, however, it seems probable that these northern Alaskan fox sparrows do not as a rule travel south- ward by slow stages, passing continuously through the regions directly south of their breeding homes, but rather that departure is made abruptly and by long flights to distant destinations. It may very well be that unalaschcensis, insularis, and sinuosa, with their summer habitats lying so far to the westward, fly southeast over the open ocean for a long distance, the main body of the travelers not reaching the North American mainland until at about the northern boundary of the United States. In the above connection it may be noted that the one migrating example of insularis just referred to as having been collected in the Sitkan district, Alaska, was taken on Kuiu, one of the westernmost islands of the region, and a station where there was also found migrating the Aleutian Savannah sparrow (Passerculus sandwichensis sandwichensis), whose summer home lies in the same general region as this fox sparrow. In the Schistacea group the winter migration involves a moving down from the high mountains, but there is a decided southward passage also. The lines of travel and the respective winter habitats of the several subspecies are not so well understood as is the ease in the other groups; although the material at hand for the present study serves to explain some of the questions involved, there is need of further collecting of specimens and observation of migration at many points. 108 University of Califorma Publications in Zoology [Vou 21 The winter habitats of certain subspecies are still practically unknown, the few winter specimens that have been collected being undoubtedly stragglers from the centers of abundance. Details of distribution are given elsewhere in this paper, under each of the subspecies, but the nature of the several problems may be briefly touched upon here. The winter homes of fulva and canescens are still practically unknown. Megarhynchus appears in numbers in winter in southern California from an unknown breeding ground. Stephensi in departing from its summer habitat in the Sierra Nevada and the higher mountains of southern California does not visit the adjacent valleys, either as a transient or as a winter visitant. It leaves the region absolutely, in a similar manner to the Green-tailed Towhee (Oreospiza chlorura) and the Wright Flycatcher (Empidonax wright) of the same habitat. The latter two species, however, are known as migrants to the southeast, while the route and destination of stephensi, are facts that are still hidden. The manner in which megarhynchus, mariposae, and brevicauda are distributed locally in southern California in winter suggests that the winter homes of the other subspecies may be found by further explor- ing at the proper season those regions whose winter avifaunas are still but slightly known. It may be pointed out, as regards the mountain ranges of southern California, that while we are well acquainted with the summer birds, there are but one or two points where winter collecting or observations have been carried on at all extensively. Judging from the local distribution in winter of certain of the subspecies of Passerella, it may well be that those races whose winter homes are not known are just as definitely delimited, but in sections from which reports covering the proper season are lacking. The accompanying charts (tables 1, 2) are designed to show the winter distribution of the subspecies of Passerella in California. Under each of the several faunal areas are entered the number of specimens examined of each subspecies, taken within the specified areas between October 15 and April 15, and also the percentage of the total Passerella winter population that is formed by each sub- species. The restriction between limited dates is not quite fair, as it throws out a number of specimens of the more southerly wintering subspecies, taken unquestionably in their winter habitats, but some such arbitrary rule seemed necessary to allow for the presence of migrating individuals in the more northern parts of the state. It is also possible that October 15 is a little too early a date to insure the presence of a bird in its winter home in all eases. 1920] Swarth: Revision of Avian Genus Passerella 109 TasBLe 1.—Manner of occurence in winter of the subspecies of Passerella iliaca in the different faunal areas of California. Specimens listed were taken between October 15 and March 15. The figures in the body of the table indicate the number of each subspecies in each faunal area. Column at extreme right indicates total number of specimens ofeach subspecies. Bottom row of figures indicates number of specimens of all subspecies from each faunal area. 2 ) . n = pt > > n 4 #2| 2 a|3 eles ~~ = an S-a Ss 3 o q S 8 = On | OF ° > £ R ioral a 4 |/03|/08 3 FI 3 2 io) = g ~ ca n N 2 ) cD) = 2 A S | Ss/25| 8 |Ba\3Bag| 2] a Boe ta &| S/| a] o > (72 (24) = |Fe(ee/ Oo] 2) 2/2) 2) 214 | ~ ALP RHE Ca Nikee (StS Sel ele ioe eeaual ce ale ee ies Se Heep ll ORS Ie Se || esa eras 3 3 x a a 3 3 oH | a Dn n |G Z op) nN op) 6) oa) 7) 2) 16) PM TAGS. O sesccacneesseo2 a ee 2 2 1 7 12 Pe it altivagans)...c..:....: : 3 2) 4 1 1 3 32 46 P. i. unalaschcensis...... 1 2 8 3 Bo ae 2 1 1 36 2 56 122: shubelspl iat peeereresyeel| 6 tags 3 4 10 3 10 1 3 oss 25 1 ae 60 PB SLNUORAneccsesesceseore 1 12 13 meee Nee 6 28 15 4 4 3 | 124 3 nia | eri 183 Pia; annectens).:.c..:.:--- 1 i: par oes His 28 54 27 i = 1 16 a axa ||| Wz P. i. townsendi... 40 4 1 4 Sie oa weed sess is a, 49 P. i. fuliginosa..... 6 9 1 16 P. i. schistacea... ae ee ae ae 1 Pee Lats see Pie: woes ah ae 14 aoe sa 15 Piet vasnccscreccs ene eens ae ae ne at ee a3 ie a a, oa 4 Bs: baa 4 P. i. megarhynchus...... 3 1 1 ee ah il 1 355 nti i css 82 2 4%, 92 P. i. brevicauda............ a ets zee eS a ill Ye fos A ae oe 24 3 ad 38 P. i. canescens................ _ see ae co Be — be ae a a 3 8 oes 1 9 P. i. monoensis... 8 11 1 20 P. i. mariposae... 2 | Woe ie as 1 Feds ae ace xe Be at ae 8 Bae Be 9 IPereistephenstrcc--s2-e.-- ee one eae ee as =e fr = = ro det 4 see we 4 MOtalstecen nc fetcrcts 6 21 28 ile 46 63 90 71 5 9 5 | 396 12 1 || 770 Taste 2.—Manner of occurrence in winter of the subspecies of Passerella iliaca in the different faunal areas of California; based upon the same specimens as those listed in Table 1. The figures indicate the percentage of the Passerella population of each faunal area that is formed by the indicated subspecies. j 2) i) : 2 om Bes > > n = 22\2¢ aes Ses ~ = gn a-5 8 3 3) | S S | <5, || f5= is) > & R =i || ae Zz Hal Wi tO)as) |) (SO) es = =| s 2 Cle BS 6 rc ZA GS) Bs 2 reals ge 2 | Sclealen| 8 le@elsale |e) 3 | 4) oe |g | eal >» |4e|43| & #6) 85) 0 g =e Sdn Se irae || eg ~ BES Weel ce en ee ae eal Boia irc alee as Eee hee taaee a Iheia ttees SOE {Ie es aatl|| Stas 3 3 = 3 a 3 9 H | na mn n |G Z op} N n 1S) mM 10) op) iS) PR ae area 13st Cag eee tee ccacere occa racer eA reeesc pw vee coca]? acdsee | aaeecs I aeseses SPO leet ellie csnse+| btecces||. ‘esesce i ate) eR oes | eee PSI Althva PANS 2,.c.ciseccasteescesee|| | tozasas ENO her PETG) srr La)» Av sil B31) ceead | h aeeenl lo ecoaee pe Ul ea eae P. i. unalaschcensis.................. A ele At estes tithers|! cece) abieo| ccc PTR tapers 11.1] 20 Qe GAG le soe Wie NSU STIS: feccesccccseescces ce cesess ZR AW ATE: Wd hel a eeds||. | Graces 15.8] 3.3] 13.3] 20 Sano meee 6. SES! Seseucs TREY, (yak boc}: ena tere oanseconenECE ena: ; GAL on eeteal Reese 9.5] 31.1} 21.2) .80 44.4] 60 SIeS|e2br lp cess Paste ADNECCENB hc ccssetecceseseussess AG | ycuseeen|h, acctoe| aera llmneereee 44.4) 60 SS ell Meese cosas 20 ARES | Soest sal\e cee res Pe -ehOWMBENG Tee css sessealle | accel” toeceealt coare™ ON Gee) TEU Geri Foes eee Neck | ale aeella) hcl emacs PEL DAM ORE, eee etece seteccessceea| [om scoters ll ue ececvel |e tieeecsll omcesers TST) zeecsll -cdzeat Tete Bases a caee seal le Ge ceced oC) octal | eereaaae P. i. schistacea... 5 4) Mean | eee | urreeeeree [eee aM RE A te hen) Met mallee mee 2 |" Readers TE TESA Ras ty ee SRE EE ne ero (mene [Wha yrerra| IMM) Manama a eel (ok Meera lt ase eel errs | need ee mnertcen| lancer UL pes lh is ace P. i. megarhynchu................... 50 ZS MEO SIM rastal| » scecte WEG), “Well eel sashes iA WS ee 2029 16).6]) a: eI DT OVUC AUG ee seceres eeeececal lee ectece|tweesealll mcskee|| Seeeedlll | weepers LO een eeecealedeereeal lt. ececses| reeretce (Sat 2155 || Morea IM CANGSCONA eee ee eee rally Be kell ce I een aa cee ated merce cceslas Lecce |(y “ccvse aly ugtaecs PFPA tie coos 100 IPM ONOENSIS alec esse ere | exeecs|| | yaetese lly cesses ATE il Rete Seeeccsl eens || Ucceaee ll meteena|la tetsccs| in cerere DES IP vera: Uses Tedoriyaeat huh oles \s eae sore cere eae | Nae [emeeereD | PR nore BEO! Ge Pea Po x-5| Peel! seen P ceases || Acedeellfrsiiaes 2 r6ll| ewe cs allie ea ees IPAM SLE PHONB crest ececsesccssu| | [eccess|( sasave|' vsees NS eed | A Aree Pee IA nes cere Eee es kill ke SA Pe 14 Uy ee es en 110 University of California Publications in Zoology [Vou. 21 From certain of the faunal areas the numbers of birds examined were too small for the results to have much significance, but there are certain facts in winter distribution that the chart shows very clearly. Attention may be called to the demonstrated preponderance of annectens in the Marin, San Francisco Bay, and Santa Cruz sections, compared with its absence from the Humboldt Bay section directly to the northward, and the limited numbers found elsewhere in the state; the preponderance of townsendi in the Humboldt Bay section, Td Fig. I. Map of southern California indicating the position of the localities designated in the text as (1) the Santa Monica Mountains region, (2) the Pasa- dena region, (3) the Claremont region. and the few individuals taken at other points; and similar restrictions in range shown by certain other subspecies. From the Sierra Nevada, Humboldt, Marin, Santa Cruz, San Francisco Bay, and San Diego regions, there are sufficient winter specimens to warrant some generalizations. From the San Diego region (with which is included the San Bernardino Mountain region) there are enough birds available to permit of analysis of conditions at several localities within this area, such as can not be done elsewhere in the state. Of the 396 winter collected specimens of Passerella from the San Diego region, 326 were taken from three relatively restricted areas, in Los Angeles and San Bernardino counties. These points I have designated as follows (see fig. I): (1) Santa Monica Mountains region (Coldwater Canon, Franklin Canon, Hollywood, ete.); (2) Pasadena region (Arroyo Seco, Millard Canon, Mount Wilson, ete.) ; (3) Claremont region (San Antonio Canon, San Dimas Canon, Lytle 1920] Swarth: Revision of Avian Genus Passerella 1B Creek, ete.). Hach of these three areas includes places that were favorite collecting grounds of certain persons (Santa Monica Moun- tains: J. E. Law, L. E. Wyman, W. B. Judson, H. S. Swarth; Pasa- dena: J. Grinnell, F. S. Daggett, H. S. Swarth; Claremont: W. M. Pierce), and the series of Passerclla are results of years of collecting at these points. I believe, therefore, that these series may be taken as fairly indicative of average conditions within the areas where they were obtained. As to the characteristics of the three areas, the Santa Monica Mountain region is widely different from the other two, while the Pasadena and Claremont regions are essentially alike. The part of the Santa Monica Mountains that is here included is a spur extending eastward from Santa Monica Canon about fifteen miles. These hills are low, nowhere reaching an elevation of 2000 feet, densely brush covered, with very little large timber and very little water. The mountains of the Pasadena and Claremont regions are of a different character, much higher and more rugged, with deep canons containing large streams of water, and with an abundance of timber. TaBLeE 3.—Winter distribution of the subspecies of Passerella iliaca at three neighboring localities in the San Diego region (see map, fig. I). Specimens listed were collected between October 15 and March 15. Los Angeles Pasadena Claremont and Santa Monica and adjacent and adjacent Mountains mountains mountains PSST Cai eats scthcesesccs savssonuseee sees 1 2 2 1A ae Lig: 002s 6 pereereeer eon eee arenes 4 6 17 P. i. unalaschcensis 2 a 26 EPMA SULATIS eters 5 17 1 EIS ASIN OE CONST: is he oe eee 8 9 83 Peer aTINCCLENS seccoeesesessieces eee 10 2 Py SCHIStACER Se iceete sear eeces oes : 2 7 IPP meganhwanChus!ss...cci7.ceee. ec 26 19 26 Js OTO WCAG Aen ceee ces sence seeeackne 18 we P. i. canescens 1 6 PS leyI ON OCDSIS tence saseeareesees sie 4 Uf Pio pI ATID OSAC -c-.cecesekosccitecseseess 2 s 3 IPI Step MeN Seas Msececiss vse sasecesns 2 see 1 4 BG) FS oa eee 68 80 178 There are certain features in the distribution of Passerella in the three sections that are at once apparent (see table 3): the preponder- ance of schistacea in the Claremont region, the restriction of brevi- cauda to the Santa Monica Mountain region, the scarcity of any subspecies of the Unalaschensis group in the Santa Monica Mountain region, the preponderance of insularis in the Pasadena region, and the preponderance of sinwosa in the Claremont region. 112 University of California Publications in Zoology [Vou. 21 The figures given for the southern half of the Sierra Nevada and for the Sierra foothill sections are based in part on material secured by expeditions from the California Museum of Vertebrate Zoology into the Yosemite region, covering every month of the year. The results apply only to this lhmited area, but judging from these figures the subspecies of the Schistacea group form but a small part of the winter population of the Sierra Nevada. I am convinced that there is significance in the results of these tabulations. I believe them to indicate that some forms really have a winter habitat nearly as sharply defined as the summer home, while others show preferences somewhat less marked. One lesson to be learned from this analysis of winter collections is that inasmuch as the extensive collecting of specimens at certain limited spots has yielded definite knowledge of a sort that obviously could not have been obtained in any other way, similarly extensive collecting at places regarding which we are still very ignorant as to the winter avifauna, may be expected to explain many details of distribution in this group of birds of which at present we have no knowledge. In the writer’s opinion, the solution of such complicated questions of relationship and distribution as are presented in the Passerella group lies only in the wholesale but intelligently directed collecting of specimens. In the course of the writer’s field experience he has from time to time noted facts in the distribution and migration of fox sparrows, which, disconnected though they be, are set down here as having some bearing on the subject in general. In a number of years of observa- tion about Los Angeles, fox sparrows were found to be extremely rare in the flat valley in which the city is situated, one or two of the Unalascheensis group, and two altivagans being all that were noted, and these at long intervals. In the low foothills of the Santa Monica Mountains, but a few miles away, certain subspecies could be found in fair numbers regularly every winter. This was also the case in the brush of the Arroyo Seco, a stream issuing from the mountains near Pasadena, fox sparrows being seen regularly in winter in the bordering willow thickets, practically as far as the junction of this stream with the Los Angeles River, at the edge of the city. In years past many southern California valleys were covered with miles of more or less dense chaparral. Much of this land has been cleared for agricultural purposes, but some fairly extensive areas of this nature still remain, the haunts of many typical Californian animal . 1920] Swarth: Revision of Avian Genus Passerella 113 species. These tracts afford shelter for brush-loving birds, and it might be supposed that they would be occupied by the winter visiting fox sparrows; but such is not the case. In many years experience in such surroundings, while quail shooting and bird collecting, the writer has never seen a fox sparrow thus situated. In southern Cali- fornia fox sparrows in general apparently occur but rarely in the Lower Sonoran valleys. The brush covered hills, Upper Sonoran and as far up as low Transition, is the favored winter habitat, with local variations as elsewhere noted. In April, 1912, while the author was collecting at the eastern base of the Sierra Nevada, near Lone Pine, Inyo County, fox sparrows were observed migrating northward. All that were taken were schistacea, the forms that nest in the adjacent mountains, mariposae in the Sierra Nevada, and canescens in the White Mountains, not being detected in the lowlands at all. This fact, and also the non-occurrence of stephensi in the foothills of the mountains wherein the latter breeds, is, I believe, in each case, indicative of rather abrupt arrival at, and departure from, the breeding grounds, with no tarrying in the low- lands nearby ; suggestive of just such migration, in fact, as has been inferred on similar grounds in certain of the Alaskan subspecies. In September, 1916, in the Kings River Canon, in the Sierra Nevada, the first migrating fox sparrow was noted on September 10, and several others were observed during the next few days. Those eolleeted proved to be all schastacea, evidently from some point con- siderably to the northward, and no specimens were taken of mariposae, the race breeding in the Sierra Nevada to the north and east of Kings River Canon. No Passerella was found breeding on the floor of the cahon. A few days later, at Horse Corral Meadow, on the adjacent table land to the southward, fox sparrows of several subspecies were found migrating abundantly. It seemed apparent that there was no migration from the east to the west side of the Sierras, or vice versa, in which the Kings River Canon could be used as a highway, and this was borne out by the manner of occurrence of many species of birds besides the fox sparrows. It was also apparent that migrants were travelling southward in numbers along the line of the Sierras, stopping at favorable places along the higher ridges, but not descending, except rarely, into the deep gorge that cut across their path. 114 University of California Publications in Zoology [Vou. 21 Passerella iliaca iliaca (Merrem) Eastern Fox Sparrow Original description.‘ Fringilla iliaca Merrem, Beytr. besond. Gesch. Vogel, 11, 1786, pl. x. (North America.)’’ (A. O. U. Check-List, 1910, p. 277.) Type specimen.—Not known to exist. Range.—‘‘North America. Breeds in Boreal zones from tree limit in northeastern Alaska, northern Mackenzie, central Keewatin, north- ern Ontario (Moose Factory), and northern Ungava south to central Alberta, northern Manitoba, southern Keewatin, Magdalen Islands, and Newfoundland; winters from lower Ohio and Potomac valleys (occasionally further north) to central Texas and northern Florida; casual on the coast of southern Alaska and in California.’’ (A. O. U. Check-List, 1910, p. 277.) One record from Arizona (see p. 118). Specimens examined.—48 (see list, p. 184). Distinguishing characters—From all subspecies of the Schistacea group, iliaca is distinguished by different proportion of wing and tail lengths (wing longer than tail in iliaca, wing equal to, or shorter than, tail in the Schistacea group), and by the predominance of bright, ruddy coloration. From all subspecies of the Unalaschcensis group, iliaca is distinguished by the much brighter hue of its red markings, by the sharply contrasting red and gray streaking dorsally, and by the heavier and more stubby bill. Within the Iliaca group ‘liaca is distinguished from the one other subspecies, altivagans, by greater general size, and brighter, more ruddy coloration, with dorsal streaking much more sharply defined. Remarks.—There are at hand fourteen specimens of Passerella taken in California that are evidently referable to Passerella tliaca iliaca; a small enough number to allow of individual consideration being given to each. No. 4932, collection of A. B. Howell; male; Pasadena; December 13, 1907; collected by A. Van Rossem. (See Willett, 1912, p. 85.) This bird appears to be absolutely typical of iliaca, being closely matched in all respects by selected specimens from Illinois. It is, however, rather small for a male, though it is marked as such, approaching more nearly the dimensions of some females. No. 867, collection of J. R. Pemberton; female; Big Sur River, Monterey County; December 22, 1903; collected by J. R. Pemberton. (See Pemberton, 1908, p. 50.) Apparently a typical example of iliaca. Very closely similar to the specimen last deseribed, and likewise duplicated by selected eastern birds. No. 384, collection of A. B. Howell; female; Burbank, Los Angeles County; November 11, 1911; collected by A. B. Howell. (See Howell, 1912, p. 41.) Ven- trally indistinguishable from eastern birds, but dorsal coloration generally more reddish. The streaks on the back are broader and consequently less sharply detined, and there is none of the clear gray on head and neck that is apparent on all the eastern specimens used in comparison. Wing bars but faintly indicated. No. 789, collection of J. Eugene Law; sex not determined; Lankershim, Los Angeles County; March 22, 1908; collected by J. Eugene Law. (See Willett, 1912, p. 85.) Closely similar to eastern examples, the main difference lying in 1920] Swarth: Revision of Avian Genus Passerella DOS the duller dorsal coloration of the California bird. The dorsal streakings are less ruddy than in eastern specimens, and less sharply defined. Top of head is gray, washed with brownish, as in many eastern birds, and the sides of neck are as clear gray as is seen in other winter collected specimens of iliaca. Ventrally it is indistinguishable from typical iliaca. Wing bars faintly indicated. ae Me a a net wt vat vat LLPILES IIL SY Gl SE 1 ! l ine Oy 4 © Passerella iliaca iliaca DISTRIBUTION MAP | MUSEUM OF YERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA ~ Ta Tae 7 ra = ora ~ - — a Fig. J. Map showing stations of occurrence in California of Passerella iliaca iliaca, as established by specimens examined by the author. No. 30445, Mus. Vert. Zool.; male adult; Seaside, Monterey County; Decem- ber 26, 1918; collected by H. G. White. Differs from average eastern birds mainly in somewhat darker coloration. Dorsal streaking and wing bars not so strongly indicated as in eastern collected specimens. No. 3233, collection of J. Grinnell; male; Mount Wilson, Los Angeles County ; October 31, 1897; collected by J. Grinnell. (See Grinnell, 1915, p. 130.) This bird, together with the rest of the specimens here listed, is suggestive of iliaca 116 University of California Publications in Zoology [Vou. 21 rather than any other subspecies, but differs from the mode of that race in certain details of markings and color. Dorsally this bird has hardly a trace of streaking, the back being almost uniformly reddish, with a suggestion of under- lying gray about the head and neck. The ventral streakings, though of a brighter, more ‘‘foxy’’ red than in the Unalaschcensis group, are not so red as in typical iliaca. Wing bars are but faintly indicated. No. 4630, collection of J. and J. W. Mailliard; male; San Geronimo, Marin County; January 26, 1901. (See Mailliard, 1901, p. 72.) Close to typical diaca, differing from eastern birds mainly in less contrasted streaking above, and in more blackish markings below. Bill as in typical iliaca, and wing bars conspicuous. No. 4208, collection of J. and J. W. Mailliard; female; Plumas County, ‘‘alt. 4000’’; September 27, 1898. Differing from eastern birds only in being less streaked dorsally. The general appearance of the back is reddish brown, mottled rathern than longitudinally streaked. Wing bars conspicuous. No. 750, collection of A. B. Howell; female; Santa Barbara; January 1, 1911; - collected by J. H. Bowles. (See Bowles, 1911, p. 175.) Ventrally indistinguishable from selected eastern specimens. Back and top of head almost uniformly dull grayish brown, back obscurely mottled with somewhat darker brown. Reddish coloration on rump and upper surface of tail feathers rather paler than in typical iliaca. Wing bars fairly well defined. No. 2003, collection of W. Otto Emerson; male; Hayward, Alameda County; December 12, 1898; collected by W. Otto Emerson. Differs from average eastern examples of iliaca in the darker, less reddish, coloration of the spots below, and in the almost uniformly brown back. The dorsal region is dull brown, faintly mottled with darker brown, and on the head and neck there is a dull brown wash over an underlying grayish color. Wing bars faintly indicated. No. 1811, collection of W. M. Pierce; male; Palmer’s Cafion, Los Angeles County; February 2, 1919; collected by W. M. Pierce. Differs from eastern birds in its generally darker coloration and heavier ventral markings. Upper parts dark grayish brown, washed with reddish; back obscurely streaked with dark reddish. Breast streakings of a brighter reddish color than the back. Wing bars faintly indicated. This bird is fairly intermediate between typical iliaca and unalaschcensis. No. 1810, collection of W. M. Pierce; male; Palmer’s Cafion, Los Angeles County; February 1, 1919; collected by W. M. Pierce. To all appearances this specimen lies between typical iliaca and altivagans. It is larger than average examples of the latter, darker colored and more heavily marked below. Upper parts nearly uniform grayish brown, more grayish on the head, more brownish on dorsum. No dorsal streakings and no wing bars. In color and markings this bird is close to altivagans, in dimensions it is nearer tiaca. No. 2805, collection of F. 8S. Daggett; male; Nicasio, Marin County; Febru- ary 6, 1906; collected by C. A. Allen. Spotting beneath of a darker brown than in any eastern specimen; top of head and back almost uniform dull brown; dorsal region faintly spotted. Wing bars faintly indicated. This bird is about inter- mediate in color and markings between typical iliaca and insularis. The bill is as in iliaca, and on the whole the specimen seems best referred to that subspecies. No. 1180, collection of L. E. Wyman; female; Santa Monica Mountains, Los Angeles County; January 31, 1917. Collected by L. E. Wyman. This bird can be considered an example of iliaca only by reason of the manner in which the series of variants just detailed connects this extreme with the mode of the subspecies. It is in a slight degree further from the type than the specimen last described. Under parts are rather heavily streaked, and with a dark shade of 1920] Swarth: Revision of Avian Genus Passerella AT brown; upper parts almost uniform, a faint suggestion of gray underlying the brown of the head and neck, the back slightly mottled, and the wing bars barely apparent. The character of the bill, however, is closely similar to that in iliaca. From the appearance of the specimens described above several deductions may be drawn. These birds are all to be referred to the subspecies iliaca, but, with two exceptions (no. 4932, coll. A. B. Howell, and no. 867, coll. J. R. Pemberton), they vary from typical iliaca sufficiently to be noticeable if inserted among a series of eastern collected skins. The specimens all resemble iliaca most nearly in the ventral markings ; the greatest variation is evident on the upper parts. These birds all show a tendency toward elimination of dorsal streak- Fig. K. Wings of the two subspecies of fox sparrow of the Iliaca group, natural size. a. Passerella iliaca iliaca, adult male; no, 6088, Mus. Vert. Zool.; Eastford, Connecticut; April 12, 1876. b. Passerella iliaca altivagans, adult male; no. 26039, Mus. Vert. Zool.; Aspen Valley, Yosemite National Park, California; October 14, 1915. ings, with consequent greater uniformity of color on the upper parts. This color in some is reddish, of a darker shade than is seen in typical iliaca, while in others it is decidedly gray. Such variation as appears in the breast streakings tends toward the darker shades of red. In the subspecies iiaca there is, according to Ridgway (1901, p. 387), great variation in color, the extremes being designated by him as the ‘‘gray phase’’ and ‘‘red phase,’’ differences are sufficiently correlated with different regions to suggest the possibility of the eventual recognition of two subspecies. ‘‘The extreme rufous phase seems to be represented only in the Atlantic Coast district, and is altogether wanting in Alaska and other north- western portions of the continent, where only birds representing the eray extreme occur’’ (Ridgway, loc. cit.). However, it is extremely unlikely that the reddish-colored birds occurring in California in win- ter are migrants from the Atlantic Coast, where the red phase of iliaca has its habitat. The probabilities are that the Californian migrants respectively, and these 118 University of California Publications in Zoology [Vou. 21 in question arrive from the northwest together with the other fox sparrows with which they are found associated. My personal convic- tion is that they hail from an intermediate locality, or localities, between the ranges of typical claca of the interior of Alaska and those of some of the Unalascheensis subspecies of the coast. Some of the specimens described in this paper, together with certain variants of insularis and unalaschcensis at hand, form, both in color and propor- tions, a graded series connecting the extremes by such short steps that any dividing line between the two must be arbitrarily drawn. Then, too, the reddish coloration of these birds is of an appreciably darker shade than in the few Atlantic coast winter birds available for comparison, tending more toward the color of the Unalaschcensis subspecies. The more grayish-backed birds may represent a step toward altivagans. I can not agree with the opinion that the character of any of these birds points toward interbreeding of diaca with schistacea (see Grinnell, 1915, p. 130). Beside the California birds described, there are two other specimens from unusual localities. One is a female (no. 2524, coll. J. Hugene Law) taken by A. Van Rossem in Pinery Canon, at 6000 feet, Chiri- cahua Mountains, Arizona, November 6, 1914. This specimen, here placed on record, comprises the first recorded occurrence of the sub- species in Arizona. It is extremely close to typical diaca, differing mainly in its more uniform dorsal coloration, the back being quite erayish, and the dorsal streakings but poorly defined. This may be indicative of intermediateness between ilaca and altivagans. The second specimen is a male (no. 967, Provincial Museum, Victoria, B. C.), collected by John Fannin at Sicamous, British Columbia, September 25, 1893. This bird is absolutely indistinguishable from selected eastern examples of aliaca. The California specimens of iliaca are too few in number to indicate any definite manner of occurrence in any specified area, but it seems unquestionable that some individuals of this subspecies winter each year within the borders of the state. Most of the speci- mens taken are from the San Diego region (where, however, there has been more winter collecting of fox sparrows than anywhere else in California), and as these were all secured between October 31 and March 22, they may be assumed to have been in their winter home. The specimens taken in December in Monterey and Alameda counties, and in January and February in Marin County, indicate the prob- ability of the winter habitat including the whole coastal area north ee ee EE 1920] Swarth: Revision of Avian Genus Passerella 119 to the San Francisco Bay region. The one specimen from the Sierra Nevada of Plumas County, taken September 27, was probably a migrant on its way farther south. It is noteworthy that nearly all the California examples of cliaca were secured by collectors who had gathered specimens for years at the several points where these birds were taken, hence the small number of specimens of this race, compared with those of most of the other wintering subspecies of Passerella, may be taken as fairly indicative of the relative numbers of each actually present. _ Fig. L. Bills of the two subspecies of fox sparrow of the Iliaca group; natural size. a. Passerella iliaca iliaca, adult male; no. 6088, Mus. Vert. Zool.; Eastford, Connecticut; April 12, 1876. b. Passerella iliaca altivagans, adult male; no. 26039, Mus. Vert. Zool.; Aspen Valley, Yosemite National Park, California; October 14, 1915. Besides the specimens here listed there are others recorded from California that I have not seen, as follows: ‘‘ Accidental in California (spec. in Mus. S. I.)’’ (Coues, 1874, p. 161) ; specimen from Poway, San Diego County, January 3, 1888 (W. E. Bryant, 1889, p. 90); specimen from Oakland, December 2, 1892 (W. E. Bryant, 1893, p. 363); Saticoy, December 14, 1872 (Baird, Brewer and Ridgway, 13714; )p..516)). Passerella iliaca altivagans Riley Alberta Fox Sparrow Original description.—Passerella iliaca altivagans Riley, 1911, p. 234. Type specimen.—No. 222832, U. S. Nat. Mus.; male, immature; Moose Branch of Smoky River (about 7000 feet altitude), Alberta; July 31, 1911; collected by J. H. Riley (original number, 2175). Range—In summer known only from a few points in the interior of British Columbia and the adjacent part of Alberta (see map, fig. N). In winter southward at least to extreme southern California. 120 Umiversity of California Publications in Zoology [Vou. 21 Winters mainly in the San Diego region and on the western slope of the Sierra Nevada (see map, fig. M). Specimens examined.—90 (see list, pp. 185-186). Distinguishing characters—Of the Iliaca group (see p. 89). Somewhat smaller than iliaca; size and proportions about as in the subspecies of the Unalascheensis group (see table 4). Coloration more subdued than in iliaca; rather brighter, with more strongly contrast- ing reds and grays, than in any of the Unalascheensis group. Remarks.—While the breeding range of the Alberta fox sparrow cannot yet be exactly defined, it may be said in general terms to be in the interior of British Columbia and in the adjoining portion of western Alberta. The breeding stations afforded by Riley’s type series are the Moose Pass region on the boundary line between British Columbia and Alberta, and Thudade Lake, British Columbia. To these I am now able to add Jasper Park, Alberta, and McGillvary Creek, and Mt. McLean, Lillooet District, British Columbia. The migration route, as indicated by specimens at hand, would be through the western part of the Great Basin, in a path whose width extends from western Montana to central Oregon, and southwestwardly at least as far as extreme southern California. The winter home appar- ently lies almost entirely within the confines of California. There is one specimen at hand (not typical, however) from Government Island, near Portland, Oregon, taken December 15. In California the line of migration seems to le entirely east of the coast ranges. There are no records from these mountains, nor even from the floor of the Sacramento-San Joaquin Valley, immediately to the eastward, while there are specimens, migrants, from many points the whole length of the Sierra Nevada. In all probability, judging from the available data, the winter metropolis of altivagans is in the western foothills of the southern Sierra Nevada, perhaps as far north as Eldorado County, and in southern California west from the eastern bases of the several moun- tain ranges separating the desert and the Pacific slope, with sporadic occurrences in the San Francisco Bay region. It may be that the subspecies winters regularly in the coast ranges from San Francisco Bay southward. There is but one specimen at hand from this region, but there has been practically no winter collecting carried on there. As to dates of arrival and departure in California, a bird taken September 22, at Horse Corral Meadow, in the Sierra Nevada of Fresno County, is the earliest fall migrant, and a specimen from the Sierras of Plumas County, ‘‘alt. 4000 feet,’’ April 24, is the latest in A a 1920] Swarth: Revision of Avian Genus Passerella 121 the spring. The Horse Corral Meadow bird was taken on an expe- dition headed by the present writer, and, as observed by him, was one of migrating northern fox sparrows which had just ai se fp 6 ?? of a ‘‘wave Ix ran aNd h + ¢ =/ iy fb y o Passerella iliaca altivagans DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OP CALIPORNIA tet rrr rm my i= a ra ora > ora Fig. M. Map showing stations of occurrence of Passerella iliaca altivagans in California, as established by specimens examined by the author. The region enclosed within the dotted outline is believed to be the main winter habitat of the subspecies. arrived in a region where we had been collecting birds for some time. It may therefore be regarded as quite accurately representing the arrival of the subspecies there in that year. Passerella iliaca altivagans was described by Riley (1911, p. 234) upon the basis of four young birds, three from the Moose Pass region, 122 University of California Publications in Zoology [Vou. 21 on the British Columbia-Alberta boundary, and one from Thudade Lake, British Columbia. The subspecies is described (loc. cit.) as ‘*Similar to Passerella iliaca schistacea, but middle of the back mars brown instead of mouse gray; wings and tail with more red in the brown (near burnt umber).’’ Of some additional specimens, migrants and winter birds, Riley remarks that ‘‘from typical Passerella iliaca schistacea, to which they have been referred, they are quite distinct, having the mouse gray of the upper parts of that race replaced by brocecoh brown and burnt umber and the spotting of the under parts prout’s brown, not sepia. Apparently there is no difference in size.’’ The A. O. U. Committee (1912, p. 386) denied recognition to this subspecies on the ground that it was ‘‘too near P. 2. schistacea.’’ The form was recognized by Grinnell (1915, p. 180), and some years later Oberholser (1918, p. 186) reasserted the validity of altivagans, affirm- ing its near relationship to iliaca, and its unlikeness to any other race. Most of Riley’s type series has been available for the present study. The three birds collected by him (in July) and deseribed (loc. cit.) as ‘‘nearly adult”’ or ‘‘slightly immature,’’ are, speaking more exactly, molting from juvenal into first winter plumage. The type specimen has reached the most advanced stage in the change, but all three still show large tracts of juvenal feathers. However, there is enough of the new plumage in place to afford a fair basis for comparison with full feathered migrants from points farther south, and thus to estab- lish with fair certainty the color characters of altivagans in its typical form. A specimen in fresh fall plumage (no. 26039, Mus. Vert. Zool.), selected for its close resemblance in comparable parts with the molting type of altwagans, is colored as follows: Upper parts generally mummy brown, most ruddy on dorsum, dullest on rump. Feathers of dorsum with dark centers, giving an obscurely streaked effect. Sides of neck washed with grayish. Upper tail coverts auburn, with narrow edgings of paler brown; exposed portion of rectrices auburn. Under parts white, chin, throat and abdomen nearly immaculate, else- where marked with triangular spots, varying in color from some the same shade as the back, to others suffused with blackish. Exposed portion of wing coverts and remiges auburn, like tail. Median and greater wing coverts slightly tipped with whitish, forming two ill- defined bars. . The above described specimen is an average fall-plumaged example of this subspecies, though there are others at hand showing diverse extremes of dorsal coloration, of grayish. or of reddish. Specimens 1920] Swarth: Revision of Avian Genus Passerella 123 taken in the spring, and consequently in less fresh plumage, are duller, less ruddy, and more grayish than fall birds. From what can be seen of the juvenal plumage of the Moose Pass birds, in this stage the markings are more ruddy than in juvenals of schistacea, which they otherwise most nearly resemble. There are no young of iliaca available for comparison. The large series that I have been able to bring together, though yielding few summer birds additional to those of Mr. Riley, abundantly demonstrates the distinctness of this subspecies, and there is no difficulty in recognizing in the winter birds from California the same race as that from British Columbia described by Riley. The small series from MeGillvary Creek, British Columbia, pre- sents several points worthy of emphasis. It includes five specimens, one in juvenal plumage, two well advanced in the post-juvenal molt, and two in fully acquired winter plumage, whether immatures or adults is not apparent. From the dates at which they were taken (August 10 to 18), and from the fact that one is still in the juvenal plumage, these birds may be assumed to have been taken upon their breeding ground. MecGillvary Creek is, in a direct line, less than ninety miles east of the general line of the coast, much less than that distance east of several deeply penetrating arms of the sea, and may be assumed to represent a point on the western boundary of the habitat of altivagans. It indicates the occurrence of the subspecies as far west as the eastern slope of the ranges most closely adjoining the coast. The birds from this point, though properly referable to altivagans, differ appreciably from the mode of that subspecies. The under parts are of a richer brown, and more heavily streaked, and there is an entire absence of streaking above, producing an evident appearance of intergradation toward the character of the Unalaschcensis sub- species. They can be very nearly matched with some winter birds from California. There is a series of July birds from Mt. McLean, so short a distance east of MeGillvary Creek that the two lots must perforce be of the same race. Yet the worn and faded plumage of the Mt. McLean birds betrays hardly a trace of the brown coloration seen in the MeGillvary Creek fall specimens, and the former are superficially very much like schistacea. However, the breast spottings still show a browner tinge than is seen in that race. The California series as a whole exhibits some little variation in color, though not so much but that the birds can, with all certainty, 124 University of California Publications in Zoology [Vou.21 be placed with altivagans, as now defined. From a style that may be taken as typical of the subspecies, the mode of coloration most frequently seen—pale reddish back, obsoletely streaked with darker red, and underparts moderately streaked—there are two extremes of variation. One is darker colored, dark reddish brown above, heavily marked below, the other predominantly grayish above and but sparsely spotted on the breast. These specimens I take to represent various degrees of intergradation, the reddish colored birds toward the coastal subspecies of the Unalaschcensis group (as shown in the MecGillvary Creek series), the grayish colored ones toward iliaca. It will be noted in these non-typical birds as well as in typical altivagans, that tail is always shorter than wing, as is the case in all fox sparrows outside the Schistacea group. There are a few apparent intergrades of another type, of duller, less reddish coloration and with tail of about the same length as wing. These I take to represent intermediate stages between altivagans and schistacea. Of a series of nine specimens taken by W. M. Pierce in certain ecafons near Claremont, southern California, during the winter of 1918-19, practically all are variants from typical altivagans, appar- ently tending toward some of the darker colored coastal races. One of these birds (no. 1776, col. W. M. Pierce) is closely similar to one of the summer specimens from MeGillvary Creek, British Columbia. Compared with the subspecies of the Schistacea group, altivagans is apparently always distinguishable by the proportionate lengths of wing and tail, wing longer than tail in altivagans, rarely the same leneth, wing usually shorter than tail, in all members of the Schistacea group. In the occasional variant of altivagans in which coloration somewhat closely approaches some examples of schistacea, the wing and tail measurements suffice for identification. In typical altiwvagans the spots on the lower surface are more ruddy than in the Unalasch- censis group, and rather less thickly distributed, leaving more white showing through where the spots are thickest, and more extensive pure white areas on throat and belly. It is quite possible that the name altivagans as now used covers a composite of several recognizable subspecies, as was the case in the former ‘‘megarhynchus’’ and ‘‘schistacea.’’ The breeding birds available are few in numbers and from but two or three scattered points, while as regards the greater part of the interior of the vast northwest nothing is known of the Passerella population. In view of 1920] Swarth: Revision of Avian Genus Passerella 125 the manner in which fox sparrows are distributed over much of the general range of the species elsewhere, locally segregated at scattered points, and considering the variation seen elsewhere in series of breed- ing birds taken from isolated, but relatively close colonies, it would not be surprising if the diverse specimens of altivagans from the winter home of that race, should prove to be representative of addi- tional recognizable subspecies, forming chains of variants comparable to those just mentioned. Riley’s type series from the Moose Pass region was taken at high altitudes (see Riley, 1912, p. 69), and the McGillvary Creek birds were collected at 5000 feet and 7000 feet altitude. These facts point to the probability of altivagans inhabiting in summer the higher mountains, and not occurring in the valleys, just as is the case with the related subspecies to the southward. Under such conditions there is added probability of the existence of distinguishable forms within the general range here ascribed to altivagans. Two skins from Penticton and one from Midway, British Columbia, show a leaning toward schistacea. While from the dates of capture (April 21 and May 5) they may have been migrants at the points where they were taken, it is probably fair to assume that they would have bred somewhere near-by, and that these places therefore lie just north of the boundary line between altivagans and schistacea. In typical form, therefore, altivagans probably does not breed south of the British Columbia-Washineton boundary, nor schistacea north of it. An interesting sidelight on the status of this subspecies is afforded by the comments written upon the labels of many specimens examined from contributing collections. There is hardly a bird collection of any size in California but contains one or more examples of this race, many of them collected years before altivagans was described. These specimens have almost invariably been a source of trouble to their owners, for various names are written and rewritten upon the labels, frequently with question marks appended. Some of these birds have been referred to different authorities for naming, and the labels reflect opinions of their relationships, which, though diversely worded, are mostly in agreement as to the general status of the race. ‘‘Schistacea —not typieal,’’ ‘“‘schistacea X iliaca,’’ ‘‘ilaca—intergrade,’’ are the terms most frequently seen, terms which quite accurately indicate the systematic position of altivagans, and in their unanimity also express the need of, and justification for, the naming of the subspecies. 99 [ Vou. 21 University of California Publications in Zoology 126 ‘psombuny ‘8 fypuasumo, ‘1 fsuaqgoauun ‘9g Snsonus “Gg /sivpjnsu fsisuaoyasnyoun ‘e fsunboaryyo °% -vovryr ‘T ‘suoryemrxordde Ajerour ore Avy, “WoYy} WoaeMjod SotrBpUNod JovxO SUL} BOTpUL SB pooysrepun oq 0} Jou aie sotoedsqns oy} Fo szEzIqVy [e1oues oy} durpuno.ims soul]jNo sy, “AoYyNe oy} Aq poururexe suoumoeds Aq poyst[qeyse sv ‘su0Tyeys SUL -paeiq JO WOT}BI0T BY} ePVorpur spoquidés oy, ‘Uolsal oy} Suryrqeyur Morreds xoF Fo sormadsqns oy} FO syezIqeYy JouUINS SurMoYs ‘TOISUTySVA, PUB VyLoqry Fo syed pue RIQUN[OHN YSyug ‘eysepy Surpnpour “BolLouLy UW1oJsomyyAOU FO dew ‘oN ‘Sly 1920] Swarth: Revision of Avian Genus Passerella 127 Passerella iliaca unalaschcensis (Gmelin) Shumagin Fox Sparrow Original description—Emberiza unalaschcensis Gmelin, 1788, p. 875. Based on ‘‘Unalashka Bunting. Arct. zool. 2. p. 364. n. 232. Lath. syn. II. 1. p. 202. n. 48. Habitat in Unalaschea....’’ Type spectmen.—Not known to be in existence. Range.—In summer, the Alaska Peninsula, Shumagin Islands, and Unalaska Island; in winter, south at least to extreme southern Calli- fornia. Specimens examined.—83 (see list, pp. 186-187). Distinguishing characters—Of the Unalascheensis group (see p. 89). Within this group, of relatively large size, with large bill, and with the minimum of red or brown coloration seen in this section of Passerella; and thus easily distinguished from fuliginosa, townsendi or annectens. Compared with sinwosa, unalaschcensis (in both types of color and bill structure [see postea]) is even less reddish, and with larger bill. Of about the same size as insularis, but with differently shaped bill, and of duller coloration. Remarks.—Unalaschcensis 1s apparently of rather rare occurrence in most parts of California, to judge from the few specimens con- tained in collections. The series of breeding birds at hand also is unsatisfactory, in that but few localities are represented within the probable summer habitat. The variety of characters represented in birds in the winter series is such as to raise questions regarding their proper classification; questions that can not be answered prior to the accumulation of much more information about the summer home of the subspecies, and some additional material from various points therein. California winter birds may be divided into two categories, one comprising birds of pale brownish coloration, rather ashy in general tone, and with long pointed bill; the other, birds decidedly darker in appearance, rather plumbeous in tone, and with short, heavy bill. Extremes of these two types are as widely different as any two recog- nized subspecies within the Unalascheensis group. They are here lumped under wnalaschcensis, first, because of intergradation which links them so closely as to indicate nearer relationship between the two than to any other race, as seems to be shown by the winter speci- mens, and second, because of the present impossibility of a satisfactory understanding of conditions in the summer habitat. Emberiza unalaschcensis Gmelin (1788, p. 875) was based upon the Aoonalashka Bunting of Latham (1782, p. 202), and the Unalashka Bunting of Pennant (1785, p. 364). Ridgway (1901, p. 389, footnote) 128 University of Califorma Publications in Zoology [Vou. 21 comments upon the applicability of the name as follows: ‘‘ Although no Passerella has hitherto been found on Unalaska (unless the Aoona- lashka Bunting of Latham really came from there), it is not at all unlikely that the present form may occur there, at least accidentally. ° | i Wace Wy; ft © Passerella iliaca unalaschcensis as \ ere < Maw c Way ce Ve ‘ Vs DISTRIBUTION MAP P MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIPORNIA wean Ta ro T= ue 1 - ae “— 7 Fig. O. Map showing stations of occurrence of Passerella iliaca unalaschcensis in California, as established by specimens examined by the author. Certainly if any form of the genus does occur on that island it would be the present one rather than the Kadiak form, most of the land birds of the Shumagins and Unalaska being the same, e. g., Lewcosticte griseonucha, Passerina nivalis townsendi, Melospiza melodia cinerea, ete. This form inhabits, on the Shumagins and the peninsula, alder 1920] Swarth: Revision of Avian Genus Passerella 129 thickets. within sight from the town of Unalaska or its vicinity, but they may have done so formerly, or may now occur in other parts of the island. At any rate, it seems better to retain the name wnalaschensis for the present form rather than to give it a new one, since the Emberiza unalaschensis of Gmelin seems almost certain to have been the present No alders now grow on that portion of Unalaska Island form.’’ There is in the Museum of Vertebrate Zoology a Passerella from the island of Unalaska. This specimen (no. 4585, male adult), col- lected by C. L. Hall, is labelled as from ‘‘Ounalaska, Alaska, June 4, 1894.’’ There is no reason to doubt the authenticity of the skin. It is one of a large collection of Alaskan animals that came into the possession of this Museum, all taken by the same man, and all carefully prepared and fully labelled; furthermore, the date of capture of this skin accords with other dates upon specimens, from the same island, of species well known to occur there. These facts, together with other cor- roborative adduced from the collection as a whole, fix the origin of this fox sparrow Fig. P. Bills of specimens of Passe- rella iliaca unalaschcensis, natural size. These birds are from different parts of the range of the subspecies, as it is now defined. Note the short, stubby bill of a, as compared with the long, slender one of b. These structural differences are correlated with differences of coloration. a. Adult male; no. 4585, Mus. Vert. Zool.; ‘‘Ounalaska,’’ Alaska; June 4, 1894. evidence beyond any reasonable doubt, even were there occasion to view it with suspicion. This skin is b. Adult male; no. 90835, Amer. Mus. Nat. Hist.; Muller Bay, Alaska Penin- sula, Alaska; June 1, 1903. unfortunately a rather poorly made specimen, but it is apparently of the plumbeous colored, heavy- billed type above described. é On the other hand, the considerable series of breeding birds from points on the Alaska Peninsula are all of the more ashy colored, slender billed type, also represented in California in winter. These facts point to the further restriction of wnalaschcensis, as a plumbeous colored, heavy-billed bird, breeding on Unalaska Island and possibly some adjacent territory, with the consequent naming of a new sub- species (of ashy coloration and with longer, more slenderly pointed bill), breeding on at least the eastern two-thirds of the Alaska Penin- sula. In the lack of an adequate series of birds from Unalaska it is 130 University of California Publications in Zoology [Vou. 21 undesirable to take any such formal action, and I prefer merely to indicate what may be the explanation of the two different types of birds I have here included under the subspecifie name wnalascheensis. Birds of the Alaska Peninsula type show a range of variation undoubtedly indicative of intergradation with sinuosa, the subspecies next adjoining on the southeast. There are also some individuals apparently inclining toward insularis, of the closely adjacent Kadiak Island. This range of variation, including apparent intergrades as above indicated, as well as the two diverse types of wnalaschcensis, all oceurring among winter birds in California, has added greatly to the difficulty of recognizing the relationships of many specimens there taken. At the outset of this study the proper separation of wnalasch- censis and insularis from the confused series brought together, was one of the most difficult problems encountered, and while the present solution of the question seems logical and fairly satisfactory, it must be borne in mind that it is largely hypothetical, and advanced merely as an apparent explanation of the observed facts. Absolute proof of its correctness awaits the study of additional summer specimens. In the range of variation seen in winter birds, all the Alaska Peninsula type show some trace of brown or rufous. Even the summer birds show it to some degree, especially in the breast spotting. The plumbeous colored type of bird is much less rufous, many of this type presenting a dull slaty effect with scarcely a trace of reddish anywhere. In its southward migration wnalaschcensis, like sinwosa, apparently avoids the coast to a great extent. This is especially noticeable in California, where nearly all the record stations are at inland points until southern California is reached (see map, fig. O). Specimens taken at Helena, Trinity County, February 17, at Drytown, Amador County, December 8 and February 29, at. Lakeport, Lake County, December 13, and at Berkeley, February 15, are apparently indicative of winter visitants at these points, but they are only scattered indi- viduals, the main winter home of the subspecies apparently lying south of Tehachapi in southern California. In the accompanying list of specimens (pp. 186-187) birds of the plumbeous colored type are marked with a superimposed figure (*) above the collection number, and it will be seen that this type of bird in winter is practically restricted to southern California. The paler eolored Alaska Peninsula bird, found in southern California too, 1920] Swarth: Revision of Avian Genus Passerella 131 occurs also in the Sierra Nevada. October 6 is an early date of arrival in the Yosemite region, October 7 in southern California. In the spring unalaschcensis lingers rather late, as shown by two birds taken in Ventura County on May 2 and 3. Unalaschcensis is one subspecies that may prove to winter well south into Lower California, and from the data afforded by the series at hand I am inclined to believe that it does so. The San Pedro Martir es & =< a a> = Fig. Q. Wings of four subspecies of fox sparrow of the Unalaschcensis group, natural size. a. Passerella iliaca unalaschcensis, immature male; no. 26031, Mus. Vert. Zool. ; near Yosemite Point, Yosemite National Park, California; October 30, 1915. b. Passerella iliaca sinuosa, adult male; no. 1535, Mus. Vert. Zool.; Hinchin- brook Island, Prince William Sound, Alaska; June 25, 1908. c. Passerella iliaca townsendi, adult male; no. 495, Mus. Vert. Zool.; Admiralty Island, Alaska; May 4, 1907. d. Passerella iliaca fuliginosa, male; no. 27152, Mus. Vert. Zool.; Blue Lake, Humboldt County, California; December 4, 1916. region, at least, should offer suitable surroundings for fox sparrows in winter. There is no information at hand regarding the winter bird population of those mountains, and it will be of interest to note the nature of occurrence of Passerella in the range. A specimen from Tacoma, Washington, on January 10, and one from Portland, Oregon, November 7, are possibly indicative of winter visitants at these points, surely the extreme northern limits of the subspecies in winter. The Portland bird is not a typical example of unalaschcensis, and is apparently an intergrade toward sinwosa. One from Salem, Oregon, April 24, was probably a migrant. 132 University of California Publications in Zoology [Vou 21 Passerella iliaca insularis Ridgway Kadiak Fox Sparrow Original description.—Passerella iliaca insularis Ridgway, 1900, p. 30. Type specimen.—No. 52475, U. S. Nat. Mus.; male adult; Kadiak, Alaska; May 17, 1868; collected by Ferdinand Bischoff. Range.—In summer, Kadiak Island, Alaska; in winter, south along the Pacific Coast to extreme southern California. Specimens examined.—l00 (see list, pp. 188-189). Distinguishing characters—Of the Unalascheensis group (see p. 89). Compared with fuliginosa, townsendi, and annectens, msularis is of paler coloration, of slightly greater general size, and with a notably larger bill. Compared with sinwosa and wnalaschcensis, in- sularis is of brighter, more ruddy coloration. The bill of imsularis is heavier than in sinwosa, about the bulk of that of wnalaschcensis but differently shaped (see fig. S). A specimen of insularis in juvenal plumage at hand differs from the corresponding stage in the neigh- boring races sinwosa and annectens in just such color characters as are shown in the adults. It is of a brighter hue than the darker red annectens, and decidedly ruddy compared with the relatively grayish young siuosa. Remarks.—Insularis is apparently restricted in summer to Kadiak Island. It has also been tentatively ascribed to Middleton Island, near Prince William Sound (Ridgway, 1900, p. 30), but the two specimens from this island that I have examined, and upon which this claim may have been based, hardly justify this conclusion. These two birds are not exactly alike, but to my mind they both most closely resemble the subspecies sinuwosa (see p. 136). The series of winter birds does not suffice for as explicit an out- lining of the winter habitat as is possible with some of the related subspecies, but there are, nevertheless, certain features of the distri- bution at this season that seem fairly well demonstrated by the data at hand. Judging from the few migrants secured at intervening points, the Kadiak fox sparrow, like the other northern coast forms of Passerella, apparently makes long, direct flights in its migrations to and from the summer home. Onee central California is reached, insularis does not seem to shun the seacoast, as do its near relatives wnalaschcensis and sinwosa; on the contrary, it favors the coastal region and avoids the Sierras. There are no specimens of insularis at hand from any point on the California coast north of Marin County, though many Passerellas have been examined from this section; in Marin County insularis is a fairly common winter visitant, but outnumbered by annectens. In the Sierra 1920] Swarth: Revision of Avian Genus Passerella 133 Nevada, though there are several record stations of capture, the number of individuals of insularis secured forms but a small per- centage of the total number of Passerellas taken. The Pacific slope of southern California apparently forms the main winter home of this sxe" “— ot a - ° Passerella iliaca insularis DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA acne crs cu 7 = ue 7 - “ ~~ Fig. R. Map showing stations of occurrence of Passerella iliaca insularis in California as established by specimens examined by the author. The region enclosed within the dotted outline is believed to be the main winter habitat of this subspecies; in all probability most of the birds winter in the southern half of this region. The general avoidance of the mountains of the interior is in marked contrast to the distribution of the closely related wnalaschcensis and sinuosa. subspecies, and even here there is apparently a local restriction to regions fairly near the coast. Insularis is the most common form of the Unalaschcensis group in the western part of the San Gabriel 134 University of California Publications in Zoology (Vou. 21 Mountains, in the vicinity of Pasadena. To the eastward, in the canons above Claremont and Pomona, it is far outnumbered by sinwosa, and this appears to be the case in the San Bernardino Moun- tains also. There are only a few specimens at hand from that range, and there are no examples of insularis among them. September 23, at Echo, Eldorado County, and September 25, at Hume, Fresno County, form early dates of arrival in the Sierra Nevada. There is one specimen at hand from Lakeport, Lake County, taken as early as September 7. In southern California dates of capture le between October 19 and February 25, though it can hardly be doubted that the birds arrive there earlier and depart later than is indicated by these dates. Farther north there are specimens secured at Stony Ford, Colusa County, March 5, at Oakland, Alameda County, March 14, and at Stockton, San Joaquin County, April 8. Following are dates and places of capture between California and Alaska: Fort Klamath, Oregon, April 13; Westminster Junction, British Columbia, April 27; New Westminster, British Columbia, March 22; Parksville, Vancouver Island, April 26; Howkan, Alaska, April 25; Kuiu Island, Alaska, April 30. These data indicate about the time of the northward migration, but the specimens of insularis thus represented form so small a percentage of the fox sparrows taken in the region that to my mind they should be considered as little more than strays from the regular line of travel. They will serve, perhaps, to indicate dates of passage, but to all appearances the mode of travel is mostly a long, direct flight, rather than a more leisurely series of short flights with frequent stops. One specimen from Clackamas County, Oregon, February 16, possibly indicates the occasional wintering of insularis as far north as this. It is rather singular that among the non-typical winter birds, referred perforce to insularis, the form they most nearly resemble, there should be so many apparently leaning toward iliaca. It is a question whether there can be actual intergradation between the two subspecies, with the ranges of wnalaschcensis or sinuosa interposed between them, and with insularis restricted to an island habitat. The variants referred to usually resemble insularis in size of bill and bright, ruddy coloration, but show more or less obscure mottling or streaking upon the back, more or less distinct wing bars, and rather sharply defined breast markings. They may really be intergrades between wnalaschcensis and iliaca, owing their bright coloring, and a 1920] Swarth: Revision of Avian Genus Passerella 135 consequent superficial resemblance to imsularis, to their iliaca strain, but this is a point that can hardly be determined from winter birds alone. Passerella iliaca sinuosa Grinnell Valdez Fox Sparrow Original description.—Passerella iliaca sinuosa Grinnell, 1910, p. 405. Type specimen.—No. 1593, Calif. Mus. Vert. Zool.; male adult; Drier Bay, Knight Island, Prince William Sound, Alaska; August 26, 1908; collected by J. Dixon; original number 309. ae a Fig. S. Bills of three subspecies of fox sparrow of the Unalaschcensis group; natural size. a. Passerella iliaca insularis, adult male; no. 9655, Mus. Vert. Zool.; Kuiu Island, Alaska; April 30, 1909. b. Passerella iliaca sinuosa, adult male; no. 1591, Mus. Vert. Zool.; Hawkins Island, Prince William Sound, Alaska; June 22, 1908. c. Passerella iliaca annectens, adult male; no. 7198, Mus. Vert. Zool.; Nicasio, Marin County, California; February 21, 1909. Range—In summer the Prince William Sound region (both on the islands and the mainland), Middleton Island, and the Kenai Peninsula, Alaska. In winter south on the Pacific slope at least as far as southern California. In its migrations and winter habitat sinuosa for the most part avoids the immediate vicinity of the coast. The main winter habitat appears to be the eastern part of the San Diegan region, southern California, but a certain percentage of the birds winters in the foothills along the west slope of the Sierra Nevada, and small numbers even as far north as central Oregon. Specimens examined.—365 (see list, pp. 189-194). Distinguishing characters——Of the Unalascheensis group (see p. 89). Bill rather long and slender, not as heavy as in wnalaschcensis and insularis, but larger than in the more stubby-billed annectens, townsendi, and fuliginosa (see fig. S). Coloration rather more reddish than in wnalaschcensis, but decidedly less so than in the brighter colored insularis, and the darker annectens and townsendi. A notice- ably ashy tinge dorsally and on sides of neck. Remarks.—The summer habitat of sinwosa lies between the ranges of unalaschcensis and annectens, and, in physical characters also, 136 University of California Publications in Zoology [Vou. 21 sinwosa occupies an intermediate position between these two subspecies. It is of less slaty, more brownish, coloration than wnalaschcensis, but it is decidedly less brownish than the ruddy-colored annectens to the southward. In bill structure also it is intermediate, with bill smaller than in wnalaschcensis, but rather larger than in annectens. The con- siderable series of sinuosa at hand from Prince William Sound includes many breeding adults, some juveniles, and some adults and immatures in fresh fall plumage. It is thus illustrative of age and seasonal varia- tion, and there is considerable individual variation exhibited also, but nothing that can be correlated with local distribution. That is, birds from the several islands and from the mainland, as well as from the opposite sides of the sound, are all intrinsically alike. (See Grinnell, 1910, p. 407.) Two adult specimens from Montague Island (nos. 1569, 1572, Mus. Vert. Zool.), exhibit, perhaps, the extremes, one of brown, the other of gray coloration as seen in the entire series. Six specimens at hand from points on the Kenai Peninsula are clearly referable to sinwosa. These birds, from the collection of the American Museum of Natural History, were referred to by Chapman (1902, p. 248; 1904, p. 404) under the names annectens and unalasch- censis, the subspecies sinwosa not having been described at that time. Two birds from Middleton Island (nos. 115823, 115824, U.S. Nat. Mus.) appear to me to be clearly referable to sinwosa. These birds, in fresh fall plumage, were taken on August 26, a date sufficiently late to allow of their being perhaps wanderers from some other point, though they are more probably representative of the form breeding upon this island. They are very slightly aberrant from the mode of the Prince William Sound series in that both are a trifle paler colored and have distinctly more buffy under tail coverts, and that one has a slightly larger bill. It is interesting to note that two winter speci- mens from Echo, Eldorado County, California (nos. 88757, 88758, Amer. Mus. Nat. Hist.) duplicate these Middleton Islands birds in every particular. It is not uncommon, of course, to find winter speci- mens so different from the normal type of any of the described sub- species as to be difficult to place satisfactorily, but there are only one or two instances in the large series of fox sparrows here assembled where such birds can be linked up with a summer habitat, as appears to be the case with the two just mentioned. There are many winter specimens at hand from points in Califor- nia, which, though referred perforce to sinwosa, exhibit among them- 1920 | Swarth: Revision of Avian Genus Passerella 137 selves a much wider range of variation than is seen in the series from the summer habitat, showing practically every degree of divergence from typical sinwosa, on the one hand toward wnalaschcensis, on the other toward annectens. These different types of birds, it is to be inferred, hail from appropriately intermediate localities between the summer home of typical sinwosa and the homes of the races they apparently approach in appearance. Whether the whole Passerella population from any one such place is uniformly of intermediate appearance, or whether in such regions a percentage of intermediates occurs among a preponderantly typical population, are questions which can not now be answered. The field work carried on at such points has as yet been insufficient to solve these problems. Among winter birds there are other non-typical specimens also, apparently referable to sinuosa, but showing a trend of variation not to be explained as an approach either to wnalaschcensis or to annectens. There are four birds from Vancouver Island (nos. 1623, 4082, 4091, 4095, Victoria Mem. Mus.) that I have provisionally placed with sinuosa though they are certainly not typical examples of that sub- species. One of these (1623) taken at Victoria on April 29, and probably a north-bound migrant, is, perhaps, to be explained as an intermediate toward annectens; but the other three, winter birds from Departure Bay, present peculiarities of dark coloration and thick bill not to be explained as an approach toward any other neighboring race. It is always possible in such a case that while the superficial appear- ance of the specimen is most nearly like some one race (in this case sinuosa), its real affinities lie elsewhere ; and that its peculiar characters may be due to an intermediateness of distribution in summer at present not understood. There are such large expanses of territory in the Pacific northwest where fox sparrows must occur and where no collecting has been done, and the manner of variation throughout the eroup is as yet so imperfectly known, that it is too much to be expected that the peculiarities of all winter specimens collected be perfectly understood. Aside from these aberrant specimens there is no example of sinwosa at hand from any point between Prince William Sound and northern Oregon, though extensive collecting has been carried on at many places in this region. The inference to be drawn is that travel between summer and winter homes is for the greater part of the distance by one continuous flight with few or no stops. 138 University of California Publications in Zoology [Vou.21 Most California specimens of stnwosa come from inland points. There is evidently very little travel immediately along the coast, and only a small proportion of the birds winters in the coastal region. eee gene mee ln et Ly Passerella iliaca sinuosa Ne iets O = eee O BENy 7 a IES IE \ \ y: = RA S¢ ee MUSEUM OF VERTERRATE ZOOLOGY di i weyo) ll ixYyiess 2ea= AQ . J DISTRIBUTION MAP UNIVERSITY OF CALIFORNIA 1 rd rd rd 7 org rd Fig. T. Map showing stations of occurrence of Passerella iliaca sinuosa in California, as established by specimens examined by the author. The regions enclosed within the dotted outlines are believed to comprise the main winter habitat of the subspecies. The stations outside these areas are points where the specimens of sinuosa taken were migrants, or, if winter visitants, where they formed but a small percentage of the total Passerella population. From Humboldt Bay there are two or three migrating individuals and no midwinter birds; from Marin County, six winter specimens out of a total of 63 Passerellas, and one or two additional migrating ot) ae 1920] Swarth: Revision of Avian Genus Passerella 139 sinuosa; from the Santa Cruz region 28 wintering sinuosa out of 90 Passerellas. On the other hand, from points in the Sierra Nevada there are 23 wintering sinwosa out of a total of 51 Passerellas, and from the San Diegan region, 124 out of a total of 396. - An interesting feature of the winter distribution of sinuosa lies in the manner of its occurrence in the San Diegan region (see table 3). Sinuosa is the predominant winter fox sparrow in the foothills and eafions at the eastern extremity of the San Gabriel Mountains. An extensive series of Passerella from the Wright M. Pierce collection, mostly from San Dimas and San Antonio cafions, is available in the present study, and there are some additional birds from the same general region available from other sources. Out of a total of 178 winter specimens of Passerella from this section, 83 are sinuosa. Only 29 of the remaining 95 fox sparrows are representative of other sub- species of the Unalaschcensis group (unalaschcensis, insularis, and annectens). On the other hand, from the region about Pasadena and in the adjacent mountains, toward the western extremity of the San Gabriel Range, there are but nine wintering sinwosa out of a total of 80 Passerellas. Here insularis is the predominant form. Specimens, to all appearances referable to sinwosa, from Corvallis, Oregon, in January, from Netarts, Oregon, in December and January, and from Mercer, Oregon, in February, are indicative of the wintering of at least occasional individuals as far north as this. There are a few winter birds at hand from scattered points in the interior of northern California: the Siskiyou Mountains, October 31; Clear Lake, December and February; Colusa, January; Eagle Lake, October 27. Undoubtedly, however, the main winter home of sinwosa les in the foothills at the western base of the Sierra Nevada, and in parts of the San Diegan region. The series from Prince William Sound includes specimens taken from June 25 to September 2. While this gives no clue as to the date of arrival on the breeding ground, it seems probable that the latter date, September 2, is close to the time of final departure for the south. Among the birds from California and Oregon there are scattered specimens that were secured upon dates affording some information regarding the time of arrival at the winter habitat and departure therefrom. Of autumnal arrivals in the Sierra Nevada there are birds from Sierra City, Sierra County, September 22; and from Horse Corral Meadow, Fresno County, September 21. This latter date may 140 University of California Publications in Zoology [Vou.21 be accepted with especial confidence as the date of arrival at that point on that particular year (1916), since fox sparrows had been assiduously sought for some time before. There is a specimen from Kunz, Trinity County, taken September 22; from the coast region there are specimens from Fortuna, Humboldt County, September 23, and from Oakland, Alameda County, September 28. The earliest date of arrival from southern California is Strawberry Valley, San Jacinto Mountains, September 28. There are but few specimens at hand from California of later date than February, though occasional captures indicate that the last _ sinuosa does not leave for the north until much later. One bird from Santa Cruz Island was secured on April 27, much the latest date of capture in California shown in the series here available. Other late spring dates are: Newhall, Los Angeles County, March 4, Bolinas, Marin County, March 31, and Eureka, Humboldt County, Mareh 3. From Oregon there are birds from Portland, Multnomah County, March 23, from Salem, Marion County, April 23, and from Warm Spring, Cook County, April 30. Passerella iliaca annectens Ridgway Yakutat Fox Sparrow Original description.—Passerella iliaca annectens Ridgway, Auk, 17, Jan., 1900, p. 30. Type specimen.—No. 170222, U. S. Nat. Mus.; male adult; Yaku- tat, Alaska, June 20, 1899; collected by R. Ridgway. Range.——lIn summer on the coast of Alaska in the Yakutat Bay region; probably extends northwestwardly nearly to the eastern edge of Prince William Sound, southeastwardly about to Cross Sound. In winter restricted almost entirely to the immediate vicinity of the coast in central California, from Marin County south through Monterey County. A few stragglers reach southern California on the coastal slope. Specimens examined.—170 (see list, pp. 194-197). Distinguishing characters——Of the Unalascheensis group (see p. 89). In geographical position annectens lies between sinuwosa and townsendi, and in its characters it forms a link between the two. Coloration is brighter, more ruddy than in s¢nwosa, not so dark as in townsendi; breast spotting less heavy than in townsendi. In fresh fall plumage, general coloration about cinnamon-brown; breeding birds have lost much of the bright, ruddy appearance of the fall plumage, and are duller in appearance. Annectens is of shghtly smaller general size than sinwosa, insularis and unalaschcensis, to the northward, slightly larger than townsendi, to the southward (see table 4). Bull notably smaller than in uwnalaschcensis and insularis, shghtly smaller than in sinwosa (see fig. 8). 1920] Swarth: Revision of Avian Genus Passerella 141 Remarks.—Vigors (1839, p. 19) deseribed Fringilla meruloides, a species that ‘‘was met with at Monterey’’ by the naturalists of H.M.S. ‘‘Blossom.’’ His description is brief, but evidently pertains to a fox sparrow, and to one of the Unalaschcensis group, although it is not sufficiently explicit to be pinned down to any one of the six subspecies within the latter aggregation. Therefore the proper application of the name meruloides rests to some extent upon outside facts, inde- pendent of the written description. Vigors gives no dates of occur- rence, but the time at which the ‘‘Blossom’’ visited Monterey, as ascertained from Beechey’s ‘‘narrative’’ (1831) was in January, and again in October and November, 1827. It was undoubtedly during the second visit (October 29 to November 17) that the natural history collections were made, but either stay was at the proper season to find fox sparrows there. No Passerella is known to breed in that region (see Grinnell, 1902, p. 44). Although Vigors’ description cannot be used to distinguish between the several closely related subspecies concerned, it so happens that at Monterey there is one particular race that occurs in greater numbers than any of the others. The Passerella that winters there in such abundanee is, as verified by com- parison of specimens, the form breeding in the Yakutat Bay region, Alaska, named by Ridgway (1900, p. 30) Passerella itlaca annectens. Collecting carried on at Monterey in December, 1901, by Grinnell (1902, p. 44) resulted in the capture of but the one subspecies; exam- ination of additional specimens from the same general region showed these also to be of the same form. On this basis Grinnell (loc. cit.) pointed out that Vigors’ name, meruloides, could be used with pro- priety for this subspecies, as apparently the only form occurring in the region. With this view the present writer was in perfect accord until a very recent date. When the writing of the present report was begun there were at hand seventeen fox sparrows from the vicinity of Monterey. Of these, fifteen were of the Yakutat Bay subspecies, one was sinwosa and one insularis, a representation of subspecies that bore out Grinnell’s assertions, above indicated. A little later, however, additional material was received that put a different aspect upon the matter. In December, 1918, Mr. H. G. White, of this Museum, spent some weeks collecting birds and mammals in the immediate vicinity of Monterey. As one result of his labors he sent in thirty-one fox spar- rows. Of these, thirteen are of the Yakutat Bay race, sixteen are sinuosa, and there is one specimen each of tliaca and megarhynchus. 142 University of California Publications in Zoology [Vou. 21 Altogether there are at hand forty-six fox sparrows from Monterey, including twenty-six of the Yakutat Bay race, seventeen sinuosa, and one each of dliaca, nsularis, and megarhynchus. The name meruloides can be applied to one particular subspecies only on the assumption that the chances are overwhelmingly against any other race having been taken at the specified locality. As shown by the recent collecting indicated above, while the one form is by far the most abundant in the region, there is at least one other that occurs in numbers. Altogether five subspecies are here recorded from that place. Under the circumstances it seems to me that there can be no absolute certainty as to the bird Vigors had in hand. On the contrary, it seems apparent that there is no alternative but to discard Vigors’ name meruloides as undeterminable, as has been done by Ridgeway (1901, p. 890). That name is replaced by Passerella iliaca annectens Ridgway (1900, p. 30), with type locality at Yakutat Bay, Alaska. There is another angle to the question, in that the winter birds bearing the closest resemblance to Yakutat Bay specimens, both as regards color and bill structure, and making due allowance for seasonal changes in color, are those from Marin County. The Yakutat speci- mens at hand are all midsummer birds, taken a month or so prior to the molt, but I can not conceive that these birds, even in the freshest of plumage, ever presented the bright, ruddy appearance so frequently seen in fall and winter specimens from the vicinity of Palo Alto and of Monterey Bay. These latter seem to present an extreme of differ- entiation within this subspecies, both as regards bright color and increased size of bill. Discovery of a definable summer range of birds with these characters may necessitate the recognition of still another race, with annectens occurring at Yakutat Bay in summer, and mainly in the region immediately north of San Francisco Bay in winter. At present, however, no such division can safely be made, but the birds from the several points indicated must all be lumped as exemplifying the range of variation within the one subspecies. In the series of Passerella here accumulated, annectens is abun- dantly represented from many points near the coast, from Sonoma County south through Monterey County, and by strikingly few speci- mens from other parts of the state. Although this is such a common winter bird in the San Francisco Bay region and in Marin County, to the northward, it is almost entirely absent from the California coast region but a little way farther north. In an extensive series of fox sparrows from the Humboldt Bay region, but one example of aE 1920] Swarth: Revision of Avian Genus Passerella 143 annectens was discovered. This bird, taken September 20, was prob- ably a migrant at this point. Two others specimens, from Logan, Oregon, March 18, and from Tacoma, Washington, November 10, fo) Passerella iliaca annectens \yi Ws © DISTRIBUTION MAP MUSEUM OF YERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA Fig. U. Map showing stations of occurrence of Passerella iliaca annectens in California, as established by specimens examined by the author. The region enclosed within the dotted outline has been satisfactorily demonstrated to be the main winter habitat of the subspecies. indicate at least the occasional appearance of the subspecies at these points, but in all probability it is of uncommon occurrence there. In the winter metropolis, the coastal region of central California, dates of occurrence range from September 25 (Sebastopol, Sonoma County) to April 19 (Hayward, Alameda County). In southern 144 University of California Publications in Zoology (Vou. 21 California, specimens have been taken from October 22 (Arroyo Seco, near Pasadena) to February 14 (San Dimas Cafion). The only inland points where specimens have been taken are Hume, Fresno County (in the Sierra Nevada at 5300 feet), September 26, at Stockton, San Joaquin County, November 28, and at Tower House, Shasta County, March 8, one specimen at each place. Specimens of annectens from Clayoquot, Vancouver Island, September 17, from Kalama, Cowlitz County, Washington, October 18, and from Beaverton, Washington County, Oregon, from February 25 to April 22, are presumably migrants. It is a point worth noting that not only are the lmits of the winter habitat sharply defined and not only are by far the greater number of these birds found in this habitat, but also a large propor- tion of the stragglers found elsewhere are not absolutely typical of the subspecies. Intergrades occur, linking annectens with sinuosa, on the one hand, and with townsendi, on the other, and many of the individuals taken outside the usual winter habitat are of such equivocal character, though referred to annectens as the form they most nearly approach. Passerella iliaca townsendi (Audubon) Townsend Fox Sparrow Original description—‘‘ Plectrophanes townsendi Audubon, Birds Amer. (folio), volume IV, part 85, plate 424, figure 7, engraved, printed and colored by Robert Havell, 1838. ‘«Fringilla townsendi Audubon, Ornithological Biography, vol. V, p. 236, 1839. ‘Shores of the Columbia River.’ ’’* Type specimen.—No. 2874, U. S. Nat. Mus.; female adult ; Colum- bia River, Oregon; February 15, 1836; collected by J. K. Townsend. Range.—In summer the coast region of southern Alaska, on islands and mainland, from Glacier Bay and Lynn Canal south to Forrester, Island; also on the Queen Charlotte Islands, British Columbia. In winter south on the coast of northern California; to Humboldt Bay, abundantly, to the San Francisco Bay region in small numbers. One specimen from Arizona. Specimens examined.—155 (see list, pp. 197-199). * These references are copied verbatim, as given me by Mr. W. H. Osgood, Assistant Curator, Mammalogy and Ornithology, Field Museum of Natural History, from copies in the library of that institution, the works cited not being available for my own personal inspection. I wish here to express my appreciation of his courtesy in supplying me with the needed references. Authors in citing the original description of this subspecies give sometimes one and sometimes the other of the above volumes, or a mixture of the two under one heading (ef. A. O. U. Check-List, 1910, p. 278; Ridgway, 1901, p. 393), a procedure that is apt to cause confusion where the actual circumstances are not understood. 1920] Swarth: Revision of Avian Genus Passerella 145 Distinguishing characters—Of the Unalascheensis group (see p. 89). Compared with the subspecies to the northward, wnalaschcensis, insularis, sinwosa, and annectens, coloration is much darker, more rufescent, spots on breast larger and more crowded. Compared with fuliginosa, to the southward, townsendi is brighter colored, less sooty. General size slightly less, and size of bill appreciably less than in unalaschcensis, insularis, and sinuosa, slightly less even than in annectens (see table 4). An average fall specimen just through the molt, in general coloration of upper parts (head, neck and back) is close to bister. Spring birds are appreciably paler colored, about vandyke brown. Remarks.—The Townsend fox sparrow is a notable example of a bird with a winter habitat nearly as sharply defined as its summer home. In the Humboldt Bay region of California it is an abundant winter visitant, but even here it is evidently restricted to a narrow strip of territory immediately adjoining the coast. Of forty-six fox sparrows examined from the Humboldt Bay region taken during the winter months, forty are townsendi; of sixty examples of townsendi taken in California (migrants and winter visitants), forty-nine are from the Humboldt Bay region. From these figures it is apparent that in California townsendi is closely confined to this particular region; it is also evident that it is the predominant form of Passerella there. The only other fox sparrows I have seen from that section are a few winter visitant fuliginosa, as listed beyond, several migrating stnuosa, and one migrating annectens. The last mentioned subspecies is so abundant a winter visitant to the California coast to the south- ward, in the regions about San Francisco and Monterey bays, as to make its rarity at Humboldt Bay a matter of surprise and of evident significance. Conversely, the subspecies townsend is almost as rare in the winter metropolis of annectens. The winter habitats of the two forms are nearly as sharply defined as the breeding ranges, and there is very little straying on either side. The summer home of the Townsend fox sparrow includes a long stretch of coast in southeastern Alaska, but the bird apparently is not continuously distributed throughout this area. The 1907 Alexan- der Alaska Expedition found this subspecies breeding at Glacier Bay (about the northern limit of the form), and at various points on Admiralty, Baranof, and Chichagof islands (see Grinnell, 1909, p. 232). The 1909 Alexander Alaska Expedition did not find the bird on any of the more southern islands of the Alexander Archipelago, nor at the more southern mainland points touched upon (see Swarth, 1911, p. 93). It is, however, known to be a common summer visitant to 146 University of California Publications in Zoology [Vou 21 Forrester Island, near the southern extremity of the Alexander Archi- pelago, and there is a June specimen at hand from Wrangell Island, where I myself failed to find it. Specimens from Howkan, Long Island, taken March 29 and April 6, were probably newly arrived migrants, but one secured there on April 27 may well have been upon its nesting ground. The available facts, however, all things considered, point to a rather uneven distribution within the known summer habitat. The specimen above mentioned as secured at Howkan (near the southern extremity of the Alexander Archipelago) on March 29, is probably representative of as early a date of arrival upon the breeding grounds as usually occurs. The Townsend sparrow was plentiful on Admiralty Island by April 17, and eggs were found there on May 3 (Grinnell, loc. cit.). The present writer found the subspecies abun- dant at Port Snettisham (on the mainland opposite Admiralty Island) at the end of August; as birds in juvenal plumage were secured the inference is that they were upon their breeding ground. ‘Townsend fox sparrows were seen on the Taku River early in September. The last was noted September 19; as the writer remained in that region until the end of September without seeing another one, that date may be assumed to indicate the final departure for the year (Swarth, loc. cit.) The first arrival in the winter home may be represented by a bird (no. C. 292, Dickey coll.) taken at Fortuna, Humboldt County, California, September 19. There is another specimen at hand (no. 2432, Emerson coll.) from Haywards, Alameda County, California (near the southern limit of the subspecies), taken September 20. The latest date in the spring represented in the available series is of a specimen (no. 21, Dickey coll.) taken at Eureka, Humboldt County, California, on March 3. It is very probable, however, that this sub- species really has not finally departed for the north until some weeks later. In California, as noted, the Townsend fox sparrow is quite closely restricted to the Humboldt Bay region. Additional specimens from seattered points in Sonoma, Marin, Alameda, and San Mateo counties are in some eases typical examples of townsendi; in others they show an obvious tendeney toward annectens. As no stray specimens of townsendi have appeared among the hundreds of fox sparrows exam- ined from southern California, the occurrence of an individual of this subspecies from a point in southeastern Arizona seems doubly strange. There is, however, one specimen at hand (no. 3178, Law coll.) collected 1920] Swarth: Revision of Avian Genus Passerella 147 in Pinery Canon (altitude, 6000 ft.), Chiricahua Mountains, Arizona, November 28, 1914, that so closely resembles typical townsend: that it must perforce be placed in that category. It is a trifle duller colored, less richly brown, than Alaskan specimens, but the difference is shght. The series of townsendi from throughout the breeding range is on the whole notably uniform in appearance, despite the segregation of the birds on different islands. From the northern extreme, Glacier Bay, there are, unfortunately, but two juveniles available. These are not appreciably different from young from other points, and they are unlike the one juvenal annectens at hand, but the material is too scanty for a decision as to the existence of a tendency toward annectens at this point. Birds from Forrester Island, an outlying island near the southern boundary of Alaska, do not depart appreciably from the mode of townsendi, nor, as far as I ean see, do those from the Queen Charlotte Islands, still farther south. A series of the last mentioned was reported upon by Osgood (1901, p. 48), who remarks that the birds ‘‘are not identical with breeding birds from Sitka, and perhaps should be considered intermediate between townsendi and fuliginosa.’’ I do not know the amount of comparative material that was available when that report was written, but in the light of the extensive series now assembled, the Queen Charlotte Islands specimens can not be regarded as outside the range of variation seen elsewhere in the habitat of townsendi. They certainly show no appreciable approach toward fuliginosa, though such variation would not have been surprising. Birds in fresh fall plumage from the Taku River and Port Snetti- sham are notably dark-colored as compared with breeding birds from neighboring islands (see Swarth, 1911, p. 94), and this darkness is also apparent in comparison with a September bird from Sitka. There is a fall specimen at hand taken at another point on Baranof Island (Peril Strait) of the same dark type of coloration as is seen on the mainland birds. The type specimen of Plectrophanes townsendi Audubon has been available for the present study. This is a well made skin, considering the period at which it was collected, and in good condition. Taken late in the winter (February 15) the condition of the plumage would not be noticeably different from birds collected six weeks later upon the breeding ground, and to my eye the coloration of this specimen is not appreciably different from selected specimens from the Alex- 148 University of California Publications in Zoology [Vou 21 ander Archipelago and the Queen Charlotte Islands. There can be no shadow of a doubt as to the applicability of the name townsendi. The only label upon the skin (besides the red ‘‘type label’’) was Oo Passerella iliaca townsendi 4 Passerella iliaca fuliginosa DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA ve ord Fig. V. Map showing stations of occurrence of Passerella iliaca townsendi and P. i. fuliginosa in California, as established by specimens examined by the author. The region enclosed within the dotted outline has been satisfactorily demonstrated to be the only part of California visited commonly by townsendi; it is evidently the southern extremity of the normal winter habitat of the sub- species. Fuliginosa is of little more than casual occurrence in California. presumably attached to it by Professor Baird.. Upon the face of the label the words ‘‘Smithsonian Institution, Washington, D. C.’’ have been crossed out with a pen, and ‘‘Coll. of S. F. Baird’’ written in. Upon the back of the label are written the words ‘‘From Audubons 1920] Swarth: Revision of Avian Genus Passerella 149 Coll. Property of 8. F. Baird.’’ The locality of capture, Columbia River, Oregon, must necessarily be regarded as established by the statement in the original description, rather than that upon the label, which must have been attached at a much later date. Passerella iliaca fuliginosa Ridgway Sooty Fox Sparrow Original description—Passerella iliaca fuliginosa Ridgway, 1899, p. 36. Type specimen.—No. 157611, Biol. Surv., U.S. Dept. Agric. ; male adult; Neah Bay, Clallam County, Washington; June 10, 1897; col- lected by E. A. Preble; original number 319. Range.—Imperfectly known. In summer known to occupy parts of Vancouver Island and northwestern Washington; presumably the mainland coast of southern British Columbia also. In winter non- typical birds occur in small numbers along the coast as far as San Francisco Bay, rarely to southern California. Specimens examined.—s89 (see list, pp. 199-200). Distinguishing characters—Of the Unalascheensis group (see p. 89). Much darker than any of the other subspecies, and also more heavily marked. One specimen at hand in freshly acquired winter plumage is about clove brown on the head, and very slightly more castaneous on the lower back. The lateral under surface of the body and the spots on the breast are dull and sooty, with hardly an indica- tion of reddish. In general size fuliginosa is slightly larger than townsend. Remarks.—This is one of the least known of the subspecies of fox sparrows. Its summer home is in a region where relatively little bird collecting has been done; its migrations at any season are extremely limited, and do not carry it into regions where specimens are much more likely to be taken than in the summer habitat. As a result there are but few skins in collections. Owing to the diversity of appearance exhibited by the specimens, the series here assembled, small though it is, raises certain questions that can not be answered prior to the accumulation of much additional material. The type specimen of fuliginosa (a summer bird from northwestern Washington) possesses a relatively long and slender bill. This is seen also in the more fully grown young birds from Vancouver Island, and again in some winter specimens from Van- couver Island. Other winter birds from the latter place, however, though of dark coloration and with the heavy markings of fuliginosa, have noticeably short, stubby bills. Winter birds from California, though here listed under the name fuliginosa, are so placed by the 150 University of Califorma Publications in Zoology [Vou.21 author reluctantly and with a strong sense of dissatisfaction. They are certainly dark colored, but of a duller tone than in typical fali- ginosa, they are not so heavily marked below (that is, the streakings are less crowded), the under tail coverts are more nearly whitish, and they all have stubby bills. There are no specimens available from the mainland coast of British Columbia, and I have consequently no knowledge of the characters of the birds of that region. The nature of variation that may be assumed to exist at the outskirts of the range of fuliginosa, perhaps showing intergradation between fuliginosa and townsend, perhaps between fuliginosa and altivagans, and fuliginosa and schistacea, is entirely unknown, and until series of specimens are gathered from many points where such intergrades are to be looked for, and the summer habitats of the several subspecies concerned con- sequently platted in much greater detail than is now possible, there is little to be gained by speculation as to the nature and origin of the puzzling winter birds above described. It may be said with all certainty, however, that the California winter visitants here included under fuliginosa were not migrants from northwestern Washington or Vancouver Island. With the exception of one or two specimens from Humboldt Bay they do not even approach typical fuliginosa very closely in appearance; they are placed in that category because their characters are such as to indicate a closer affinity to fuliginosa than to any other form, and probably illustrate intergradation between fuliginosa and some one of the adjacent subspecies. In what little is known of the summer habitat of fuliginosa there are certain apparent inconsistencies of occurrence. Dawson (1909, p. 152) found this subspecies breeding upon the San Juan Islands, Puget Sound, but the present writer did not meet with it at the lower levels of the closely adjacent southeastern portion of Vancouver Island. Here it was discovered breeding in the Canadian Zone of the higher mountains, from 2200 feet altitude upward. Farther north on Vancouver Island, at Nootka Sound, however, it was seen at sea level (Canadian Zone) (Swarth, 1912, p. 62). While Dawson (1909, p- 155) defines the summer habitat in Washington as on ‘‘the north and west slopes of the Olympic Mountains, together with the islands of Lower Puget Sound,’’ a lst of the birds of the Lake Crescent region, Olympic Mountains, by Rathbun (1916) ineludes no mention of Passerella. The present writer collected four specimens of fuliginosa in the summer habitat on Vancouver Island. Three are in juvenal plumage. 1920] Swarth: Revision of Avian Genus Passerella 151 and one in. almost complete first winter plumage. The last mentioned (no. 16253, Mus. Vert. Zool.), in perfectly fresh, unfaded plumage, is of intensely deep brown coloration, about clove brown on the head, and very slightly more castaneous on the lower back. It is much more heavily marked below than is the case in townsendi, the dark markings being so nearly confluent across the breast that in this case it is the white ground color that shows through as interrupted markings. The lateral under parts and the dark spots on the breast are dull and sooty, with but a suggestion of reddish. The three juveniles exhibit some diversity of color. In the darkest of them (no. 16252, Mus. Vert. Zool.) the general coloration is bister, as compared with snuff brown in typical young townsendi; the mark- ings below are so extensive, and the ground color so very little paler than these markings, that it appears almost ‘‘solid’’ brown below. The other two young (nos. 16251, 16254) are less heavily marked and not so dark-colored, but they are still appreciably darker than town- sendi at the same stage. Fuliginosa (speaking now of the non-typical birds previously described) is a rare winter visitant to the northern coast region of California. The earliest arrival noted is one from Fortuna, Humboldt County, September 19; the latest in the spring, one from southeast Farallone Island, May 31. As the subspecies had been represented in loeal collections heretofore only by an occasional specimen (nearly all from the Humboldt Bay region, included among large series of townsendi), it was rather unexpected that in the fall of 1918 there should be a sudden influx of the birds into the San Francisco Bay region. On October 1, 1918, R. M. Hunt, of the Museum of Vertebrate Zoology, noted numbers of fox sparrows in the hills east of the Uni- versity Campus. The following day he returned to the place and collected six of the birds. Four of the six proved to be fuliginosa. On October 18 four more fox sparrows were collected at the same place, and three of them were fuliginosa. Later collecting failed to disclose more of these dark-colored birds. On September 26, 1918, J. Grinnell collected an immature female fuliginosa at Morro, San Luis Obispo County. In the light of these several captures, and con- sidering the absence of the subspecies in the extensive series of fox sparrows available and collected in the San Francisco Bay region during previous years, there seems to be evidence of a rather unusual ‘“‘Wwave’’ of this bird. There are certain other birds, more con- spicuous in actions and appearance (the varied thrush is a good 152 University of Califorma Publications in Zoology [Vou. 21 example) that we know to vary in numbers from year to year, and it may be that among the subspecies of the fox sparrows also there is this question of fluctuating numbers (or even total absence some winters compared with relative abundance during others) to make still more complheated the already involved problem of the migration and distribution of the different forms. One specimen of fuliginosa at hand (no. 27153, Mus. Vert. Zool.), like the Berkeley birds in appearance, was taken at Bear Flat, San Antonio Cafion, San Gabriel Mountains, California, on November 30, 1916. This is by far the southernmost record for the subspecies. Fig. W. Wings of three subspecies of fox sparrow of the Schistacea group, natural size. a. Passerella iliaca schistacea, adult male; no. 9054, Mus. Vert. Zool.; Pine Forest Mountains, Nevada; June 19, 1909. b. Passerella iliaca mariposae, adult male; no. 29089, Mus. Vert. Zool.; Sisson, Siskiyou County, California; May 15, 1918. c. Passerella iliaca stephensi, adult male; no. 3426, coll. G. F. Morecom; Bear Valley, San Bernardino Mountains, California; June 10, 1886. Passerella iliaca schistacea Baird Slate-colored Fox Sparrow Original description.—Passerella schistacea Baird, 1858, p. 490. Type specimen.—No. 5718, U. S. Nat. Mus.; female; ‘‘ Platte river, K. T.’’ (= ‘‘South Fork of Platte River, about 25 miles east of northeastern corner of Colorado, Nebraska’’ [A. O. U. Check-List, 1910, p. 277]) ; July 19, 1856; collected by Lt. F. T. Bryan (original number 131). Range.—The summer habitat is principally in the high mountains of the Great Basin region, north into Alberta, south into Nevada and east probably to eastern Wyoming. In winter south to southern California and (casually, at least) to southern Arizona. Occasional in migration in western Nebraska and Kansas. 1920] Swarth: Revision of Avian Genus Passerella 153 Specimens examined.—98 (see list, pp. 200-201). Distinguishing characters——Within the Schistacea group (see p. 89), schistacea is distinguished from all the other subspecies save canescens by its diminutive bill (see figs. E, F, and table 4). From canescens, schistacea is distinguished by its more slender pointed, less stubby bill, and by color, canescens being more grayish in general coloration, schistacea more brownish. In an average example of schistacea in fresh fall plumage (no. 23203, Mus. Vert. Zool., Blue Canon, Placer County, California, August 31, 1912) the upper parts generally are overlaid with brownish, the general effect being not far from sepia, while the ventral spots, each one blackish centrally, are laterally of a richer brown than the back. By April much of the brownish tinge has disappeared through wear and fading, but direct comparison of specimens of schistacea and canescens shows an appre- ciable difference, still, in this regard. In juvenal plumage schistacea exhibits throughout a brownish suffusion, just such as is seen in the later stages. Remarks.—The type specimen of Passerella schistacea Baird was collected upon the expedition headed by Lieut. Francis T. Bryan in exploration of a road from Fort Riley to Bridger’s Pass in the sum- mer of 1856. The locality of capture of the specimen as entered upon the label is simply ‘‘Platte river, K. T.,’’ the date, July 19, 1856. The itinerary of Lieut. Bryan’s party appears in a report contained among ‘‘The executive documents printed by order of the senate of the United States’’ (1858, pp. 455-457) ; and there is a map showing the route the party traversed, in the Annual Report of the Wheeler Survey (1876, opp. p. 36). In the itinerary, unfortunately, a detailed account of day to day travel begins only on July 23, but by comparison of the less detailed statements just preceding this date, with the table of distances (pp. 485, 486) a fairly accurate idea may be reached of the whereabouts of the party on July 19. On that day, from the above data, they would seem to have been traveling along the South Platte River, probably between Laramie Crossing and Goodale’s Crossing, approximately two hundred miles west of Fort Kearny. This would fix the type locality of schistacea at about the point given by Cooke (1897, p. 107), upon the authority of Dr. T. S. Palmer, and as repeated by the A. O. U. Check-List (1910, p. 277). The latter authority gives it as ‘‘South Fork of Platte River, about 25 miles east of northeastern corner of Colorado, Nebraska.’’ I myself, after studying the documents above mentioned, did not feel able to fix the locality quite so exactly, but it is, apparently, in Nebraska and on the South Platte River, not many miles east of the Nebraska-Colorado boundary line. It should be stated that on the map above cited the 154 Umversity of Califorma Publications in Zoology [Vou. 21 only part of Lieut. Bryan’s route that is clearly labelled as such is the stretch between the South Platte River in northeastern Colorado and the Solomon River, Kansas, the homeward route taken by the party later in the summer.* The type specimen of schistacea is an adult female in badly molting condition, of little value for color comparisons. The subspecies is evidently rare in Nebraska and in the adjacent portions of Colorado and Wyoming, and it is improbable that this bird was breeding at the place where it was taken. From the account in Bryan’s itinerary, above cited, the headwaters of Lodgepole Creek (which empties into the South Platte not far west of the point of capture) would seem a not unlikely breeding ground for Passerella, so it may be that this bird had strayed down that stream, and then down the South Platte. In Baird’s original deseription of Passerella schistacea (1858, p. 490), his comments were applied to a series of specimens that included two or more distinct subspecies, as now understood. As far as the description itself is concerned, both the small-type paragraph headed ‘‘Sp. Ch.’’ and the general comments following, all laying stress upon the great size of the bill, as compared with iliaca, apply to the bird he later called megarhynchus, and not to the small-billed bird for which the name schistacea has been generally used. The definite statement is made, however, that :‘‘The essential characters of the preceding diagnosis are based on a specimen (5118) from the head of the Platte, and collected by Lieutenant Bryan, in 1856.’’ It was a curious slip to have made, especially by as keen an observer as Baird, for it is literally impossible that a comparison of Bryan’s specimen from the Platte with examples of iliaca, could have inspired a diagnosis of the former containing the statement: ‘‘Bill very thick; the upper mandible much swollen at the base.’’ It seems as though the descriptive paragraphs had been based upon Fort Tejon specimens, and the comment upon the Platte specimen inserted at some later time. It may, perhaps, have been put in with an idea of defining the geographic range of the species. However that may be, and however definitely every sentence in the diagnosis applies to the California bird, there is no doubt that the statement above quoted fixes the type of schistacea as being the Platte River specimen. In addition to this there is Baird’s action in an appendix to the volume containing the description of schistacea, where *T am indebted to Dr. T. 8. Palmer, of the United States Biological Survey, for putting me upon the track of the accounts from which the above statements are taken. 1920] Swarth: Revision of Avian Genus Passerella 155 he applies the name megarhynchus to birds from Fort Tejon (1858, p. 925), and specifically mentions the Platte River specimen as the type of schistacea. Judging from the variation exhibited by the birds assembled for this study, it seems possible that the name schistacea, even as here restricted, covers a composite of two or more recognizable subspecies. That such is the case, however, is not capable of proof by the exam- ination either of winter birds alone, or of the few summer birds avail- able, which represent but a limited number of localities. The general summer range of schistacea consists of many scattered high mountain localities, more or less widely separated, and it would not be surpris- ing should some of the isolated colonies thereon prove to be as dis- tinguishably different from normal schistacea as is canescens. The rather extensive series of summer specimens from the Pine Forest Mountains, Nevada, are apparently similar in character to the type specimen of schistacea and to summer birds from the vicinity of Fort Bridger, Wyoming. Schistacea, proper, is thus primarily a bird of the Great Basin. Birds of this type are of distinctly brownish coloration, as compared with canescens (both adults and juvenals), and, in the extreme of differentiation from that race, have a notably longer, more slender pointed bill. In midsummer plumage the pres- ence of the brown coloration is usually obscured by wear and fading, but molting birds from the Pine Forest Mountains show it unmistak- ably in the new-coming feathers. Most California winter taken speci- mens are evidently of this same type, slender-billed, and more brown of color than canescens. There is a small proportion of skins, from various seattered localities, that have this brown coloration carried to an extreme, nearly as much so as in some of the Unalaschcensis group, from which, however, they are readily distinguished by other characters. The capture of one of these extremely rufescent birds at Anthony, Baker County, Oregon (October 11), suggests the possibility of the summer habitat of this type lying somewhat farther to the northward, or perhaps it is an indication of intergradation between schistacea and altivagans. In support of this supposition attention may be drawn to a series of summer birds from Banff, Alberta (coll. Victoria Mem. Mus.). As compared with the Pine Forest Mountains series, the Banff birds are much more heavily marked below, and of appreciably browner coloration. One specimen in particular (no. 1175) is notably rufescent, and with a faint suggestion of streaking above. On the other hand, a specimen from a neighboring locality 156 University of California Publications in Zoology [Vou.21 (no. 100358, Amer. Mus. Nat. Hist., Ptarmigan Lake, near Laggan, Alberta), is not appreciably different in coloration from the Nevada birds. The Alberta specimens are short-tailed, as compared with those farther south, this again being undoubtedly an approach toward altivagans. Schistacea oceurs in California as a migrant and winter visitant, the specimens at hand indicating the occurrence of this subspecies over a large part of the state, though it is not a common bird at any point. The line of migration extends along the Sierra Nevada, with the main winter home in California in the San Diego region. There are many records from the Sierras extending from August to November, and a lesser number during March and April. One individual from Mari- posa County, November 29, might have been supposed to be in its winter home, but Tyler (1918, p. 85), speaking of the occurrence of this subspecies in the Fresno district (south of Mariposa County), asserts that while some of the birds remain until December, they all depart during this month, presumably for points still farther south. There is one winter bird (December 5) from the Piute Mountains, at the southern extremity of the Sierra Nevada, while from the San Diego region there are many specimens taken during December, Janu- ary, and February. These winter birds are all from points west of the divide that separates the Pacific slope from the desert, though there is no reason to doubt that the subspecies winters also on the desert slopes of these same mountains. The northernmost winter specimen at hand from the coast region is from Nordhoff, Ventura County, and there are many points of capture from that station south practically to the Mexican boundary. The subspecies has not as yet been reported from any of the islands off the coast of southern Cali- fornia, nor from any loeality north of Ventura County and west of the San Joaquin and Sacramento rivers. The earliest arrival in the fall is one taken at Blue Canon, Placer County, August 31, though others have been secured far to the south of this but a few days later—at Cottonwood Lakes, 11,000 feet altitude, Inyo County, September 3, and at Lytle Creek, San Bernardino County, September 2. In Kings River Canon, in 1916, slate-colored fox sparrows were migrating in small numbers the middle of Septem- ber, the first taken on September 12, though others, believed to be of the same subspecies, had been seen on the 10th. A few days later, at Horse Corral Meadow, a high ridge just south of the Canon, the 1920] Swarth: Revision of Avian Genus Passerella 157 migration was evidently in full swing, and specimens of schistacea were shot from time to time along with several other subspecies of Passerella. The latest taken specimen in the spring is one from Ojai HS | Gey ral oe | Ne iS \Z ~h P. iliaca schistacea, breeding P. iliaca schistacea, non-breeding P. iliaca canescens, breeding P. iliaca canescens, non-breeding DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA Fig. X. Map showing stations of occurrence of Passerella iliaca schistacea and P. i. canescens in Nevada and California, as established by specimens exam- ined by the author. The portion of southern California enclosed within the outline is believed to be the main winter habitat of schistacea in this state. Note the general avoidance of the coastal region. Valley, Ventura County, April 25. In the vicinity of Lone Pine, Inyo County, in 1912, slate-colored sparrows were seen daily about the middle of April, migrating northward, specimens being secured from April 10 to 16. 158 University of California Publications in Zoology [Vou 21 The capture of an individual of this subspecies in midsummer (July 8, 1908) in the San Jacinto Mountains, Riverside County, Cali- fornia, remains an anomalous occurrence, difficult to understand (Grinnell and Swarth, 1913, p. 281). There are four specimens of schistacea available from Arizona. The winter range of schistacea is generally given as including Arizona and New Mexico (ef. A. O. U. Check-List, 1910, p. 277; Ridgway, 1901, p. 396), but the published records of occurrence in the former state hardly suffice as a basis for the statement. I know of but four such locality records: forty miles south of Camp Apache, September 1, 1873, one specimen (Henshaw, 1875, p. 2938); San Francisco Moun- tain, September 29, 1889, one specimen (Merriam, 1890, p. 97); Huachuca Mountains, November 20, 1894, one specimen (Fisher, 1904, p. 81); Big Sandy Creek, near Signal, Mohave County, February 6 and 9, 1880, three specimens (Stephens, 1914, p. 259). Brewster’s (1882, p. 197) record from Tucson was a mistake, as pointed out by Stephens (loc. cit.). The additional specimens now at hand are two from the Sierra Ancha, Gila County, November 7 and 12, 1916 (nos. 1803, 1804, coll. G. Willett), one collected by A. Van Rossem in Bonita Canon, Chiri- cahua Mountains, January 24, 1915 (no. 3179, coll. J. E. Law), and one taken by Dr. E. A. Mearns at Payson, Yavapai |= Gila?] County, February 18, 1888 (no. 52996, Amer. Mus. Nat. Hist.). It may be only due to the failure to collect in the proper localities that the sub- species has not been found to be of regular and abundant winter occurrence in Arizona, but on the basis of the above mentioned speci- mens (all that are known to me as occurring in that state), schistacea can hardly be regarded as of much more than casual occurrence. The four Arizona specimens at hand are in appearance essentially like the breeding bird of northern Nevada, and are probably migrants from that general region. Passerella iliaca fulva Swarth Warner Mountains Fox Sparrow Original description.—Passerella iliaca fulva Swarth, 1918, p. 162. Type specomen.—No. 14795, Calif. Mus. Vert. Zool.; adult male; Sugar Hill (5000 feet), Warner Mountains, Modoe County, Califor- nia; May 19, 1910; collected by W. P. Taylor and H. C. Bryant; original number 2887. 1920] Swarth: Revision of Avian Genus Passerella 159 Range.—In summer the extreme northeastern corner of California, in the mountains of Modoe and Lassen counties; north into central Oregon; east of the Cascades. The northernmost specimen at hand is from Warmspring, Crook County, the easternmost from Burns, Harney County. Winter home unknown. Specimens examined.—s89 (see list, pp. 201-202). Distinguishing characters——Of the Schistacea group (see p. 89). Bill intermediate in size between schistacea and mariposae; of about the same size as in monoensis, but more slender and attenuated than the short and rather heavy bill of that subspecies (see fig. F). Color- ation more brownish than in mariposae and monoensis; about as in schistacea. Wing and tail measurements somewhat less than in mari- posae, about the same as in schistacea (see table 4). Remarks.—Size of bill will suffice to distinguish fulva from any of the other fox sparrows of the Schistacea group save monoensis. From the latter form fulva may be told by the somewhat differently shaped bill, and by difference in coloration. In worn summer plumage eolor differences are more or less obscured, but in freshly molted fall specimens they are readily apparent. The large series of breeding birds at hand from tthe Warner Mountains, California, is quite uniform in appearance throughout. Breeding birds from various points in central Oregon exhibit consider- able diversity in size and shape of bill and in relative grayness and brownness of color, though all, I believe, are best referred to fulva. This variation is in accordance with the geographical position of the several specimens, showing different degrees of intermediateness between fulva, schistacea, and mariposae. Birds from extreme southern Lake County are typical fulva. One specimen from Burns, Harney County, the easternmost point represented, in size of bill approaches schistacea. Examples from Fort Klamath and Sisters, the westernmost points of record, have larger bills, apparently tending toward mariposae. These latter specimens also are decidedly grayish in color, as compared with Warner Mountains birds, in this respect again showing an approach to mariposae. There is an abundance of breeding specimens of fulva at hand, and from enough different localities to give a fairly accurate idea of the summer habitat of this form, but there is practically no material that would serve to shed light on migration route and winter home. There is a specimen in the Mailliard collection (no. 4253), a typical example of fulva, collected at Eagle Lake, Lassen County, October 1, 1899; and one in the collection of Allan Brooks, collected by A. Van Rossem [ Vou. 21 160 University of California Publications in Zoology at Sierra City, Sierra County, October 6, 1911. These would indicate a rather late tarrying near the summer habitat. An immature male (no. E. 301, coll. D. R. Dickey) taken at Lawson (= Lassen) Creek, Modoe County, August 23, is so different from midsummer birds of the same age and stage of plumage from the near-by Warner Moun- tains, that, although I include it under the name fulva, I believe that it was a migrant at the point of capture, probably from somewhere in central Oregon. It has the gray coloration and the type of bill seen in birds from that region. There is one specimen in the Emerson collection taken in the Volean Mountains, San Diego County, February 9, 1884. This bird is unquestionably fulva. It can be very closely matched by selected specimens from the Warner Moun- tains (allowing for the different degrees of wear on the feathers), and it is absolutely the only speci- men of which this can be said, that has come to light among the hundreds of winter collected southern Californian fox spar- rows examined. There has been Fig. Y. Bills of two subspecies of fox sparrow, natural size. a. Passerella iliaca mariposae, adult female; no. 25691, Mus. Vert. Zool.; Yosemite ‘Point, Yosemite Park, Cali- fornia; June 4, 1915. b. Passerella iliaca megarhynchus, female; no. 12402, U. S. Nat. Mus., type specimen; Fort Tejon, Kern County, California. very little bird collecting done in the mountains of San Diego County in winter, not over a dozen Passerellas from this whole county having been examined by the present writer, and it may be that future work will disclose the winter home of fulva to lie in this unexplored part of the state. There is a skin in the collection of A. B. Howell (no. 6827), taken at Bluff Lake, San Bernardino Mountains, October 11, 1918, and one in the collection of W. L. Dawson, taken in Santa Barbara County, January 11, 19138, that also appear to be fulva, though not so un- The Bluff Lake specimen has the brown coloration of fulva, but not the attenuated bill characteristic of adults of this subspecies. The relatively stubby bill of this bird may be a result of immaturity, or it may, on the equivocally so as the Volean Mountains specimen. other hand, be an indication of intergradation, perhaps towards megarhynchus. Size and shape of bill are about intermediate between 1920] Swarth: Revision of Avian Genus Passerella 161 the two subspecies. The Santa Barbara specimen is close to typical fulva. Two specimens in the W. M. Pierce collection (nos. 1772, 1796), from Cucamonga and San Dimas eafions, southern California, I have also considered as fulva, but they are far from typical of the form. If these several points in San Bernardino and Los Angeles counties represent northern and western limits of the winter habitat of fulva, as seems possible, it would not be surprising to find at these outposts non-typical birds, visitors from the borderland of the summer home of the race. Passerella iliaca megarhynchus Baird Thick-billed Fox Sparrow Original description.—P [asserella]. megarhynchus Baird, 1858, p. 925. [Note that I use this name, megarhynchus, as it is written by the original deseriber, Baird. He, too, used it in this form in conjunction with the generic term Passe- rella. | Type specimen.—No. 12402, U. S. Nat. Mus.; female; Fort Tejon, Kern County, California; collected by J. Xantus de Vesey; original number 1397; date of capture not on label. Range.—Common winter visitant to the Pacific slope of southern California. Summer home unknown. Specimens examined.—104 (see list, pp. 202-204). Distinguishing characters.—Of the Schistacea group (see p. 89). From all other subspecies of this group except brevicauda, megar- hynchus may be distinguished by its heavy, stubby bill and relatively brownish coloration. In both these features it approaches brevicauda, but it has a less heavy bill, and it is not as brown colored as in the extreme of that form. In bill structure megarhynchus is about inter- mediate between the type of brevicauda on the one hand and fulva, of southern Oregon, on the other (see fig. Y). Remarks.—In assorting the confused series of ‘‘thick-billed spar- rows’’ contained in the several collections available from southern California, there proved to be in each lot a rather large percentage of birds, winter visitants only, with the prominent characteristics of broad, stubby bill, and relatively brownish coloration. Through the courtesy of the authorities of the United States National Museum I have been permitted to examine the type specimen of Passerella megarhynchus Baird, and it proves to be a bird of this character. The name megarhynchus is thus shown to be properly applied (as it has been in the past) to the most common form of ‘‘thick-billed spar- row’’ found in winter in southern California, but it can no longer 162 University of Califorma Publications in Zoology [Vou 21 be used for the breeding bird of the Sierra Nevada. Series of summer specimens from various parts of those mountains are available for comparison, and prove to be of a different type of bird (see page 173). It is not possible at present to indicate the breeding range of megar- hynchus, but the relatively thorough working-out of the summer homes of stephensi, mariposae, monoensis, fulva, and brevicauda, made pos- sible by the series of specimens here assembled, narrows down the territory in which this breeding area may be expected to be found. Megarhynchus as here restricted, is, it may be remarked, perfectly distinct from any of the above mentioned subspecies, and our ignor- ance of its summer habitat is not to be explained by a mistaking of seasonal or other variation of individual birds for the subspecifie variation of geographic races. Although there are only winter speci- mens of megarhynchus at hand, there is sufficient material representa- tive of the other closely related subspecies to demonstrate the reality of the indicated differences, regardless of season. Megarhynchus is most nearly like brevicauda, in fact the material at hand indicates perfect intergradation between the two. Both are brownish colored birds, compared with mariposae, monoensis, and stephensi, and both have the same shaped bill, in different sizes. Series of breeding birds at hand show that the subspecies of the Sierras are decidedly grayish. Stephensi, mariposae, and monoensis, extending from the San Jacinto Mountains to Mount Shasta, are all of this pale type, and all have similarly shaped, rather slenderly-pointed, bills. Obviously there is no room here for the insertion of a brownish- colored, stubby-billed subspecies; also there are enough breeding stations represented along these mountain chains to show that there are no fox sparrows there of the megarhynchus type. On the other hand, the available material shows the existence of a chain of brownish-colored subspecies of the Schistacea group, extend- ing from schistacea of northern Nevada, through fulva of southern Oregon, and the Warner Mountains, California, and ending with brevicauda of the Yolla Bolly Mountains. In its salient characters megarhynchus apparently fits in between fulva and brevicauda; it is intermediate between the two, though appreciably nearer the latter. According to this reasoning the breeding grounds of megarhynchus would he in the coastal mountains of extreme northwestern California, and, probably, southwestern Oregon. This section is not represented by a single specimen in the material at hand, and apparently there has been little or no bird collecting done there. As far as I know 1920] Swarth: Revision of Avian Genus Passerella 163 there are no published references to the occurrence of fox sparrows in that region, at any rate nothing to shed light on this problem. Among the winter birds from southern California here ascribed to megarhynchus, there are many that are difficult to distinguish by color from mariposae. It is worth noting that rather grayish-colored individuals are frequent in the Claremont series, while brown-colored ones predominate about Pasadena and Los Angeles. The gray-colored Claremont specimens, however, have the stubby bill characteristic of megarhynchus, and I consequently place them in that category. I do not claim to possess a perfect understanding of the exact relation- ships existing between megarhynchus, mariposae, and fulva, which, indeed, can not be attained without much more information than I possess regarding the summer habitats of all three; I can not do more than point out the features of the winter birds examined, as correlated with places of capture. The suggestion may be made, however, that here again is evidence of fairly close restriction of winter habitat ; furthermore, winter birds with the various characters above described should in the future be checked up with summer birds from points as yet unrepresented, in the more minute working out of relationships and routes of migration. Megarhynchus is a common winter visitant to the Pacific slope of southern California, but there are very few specimens from other points indicating the route traversed between this region and its breed- ing ground. There are three specimens at hand from Tower House, Shasta County, taken on March 2 and 7, and one from Sisson, Siskiyou County, September 5; whether these birds were winter visitants at the points of capture, or merely migrants, there is no way-of telling. At any rate they may be taken as evidence that the breeding ground of megarhynchus lies at least as far north as this. The subspecies certainly does not breed in the immediate vicinity of either Tower House or Sisson. One bird from Lakeport, Lake County, taken on February 27 (no. 11486, Calif. Acad. Sci.), is in large part albino, but is un- equivocally megarhynchus nevertheless. There is one specimen of megarhynchus from Nicasio, Marin County, taken January 30, the only one of this subspecies that has come to light among the many fox sparrows examined from this county. The dates of capture of the two birds last mentioned are indicative of their being winter visitants, though the subspecies is evidently extremely rare at the points where they were taken. There is a record of megarhynchus 164 University of California Publications in Zoology [Vou 21 as seen at Alton Junction (near Eureka), Humboldt County, in late October (Ferry, 1908, p. 43), but in the absence of specimens there is no certainty as to the oe actually encountered. ma oo Ps = Passerella iliaca mariposae Passerella iliaca megarhynchus Passerella iliaca fulva DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA Ss pecace Fig. Z. Map showing winter occurrence of Passerella iliaca ‘mariposae, P. 4. megarhynchus, and P. i. fulva in California, as established by ‘specimens examined by the author. The region enclosed within the dotted outline is believed to be the main winter habitat of megarhynchus. The locality stations of mariposae and fulva denote the capture of but a single specimen at each point, apparently strag- glers from the still unknown center of winter abundance. Along the line of the Sierra Nevada, among the scores of fox sparrows collected, there are but two examples of megarhynchus. One is from Blue Cafion, Placer County, taken on October 19, the other from El Portal, Mariposa County, on November 28. The sub- species is evidently rare in these mountains, but may be assumed 1920] Swarth: Revision of Avian Genus Passerella 165 to winter in small numbers as far north as the latter station. The Pacific slope of southern California is evidently the winter metropolis of megarhynchus. In the foothills of this section it is in winter per- haps the commonest of the fox sparrows, certainly the most abundant of the Schistacea group. Dates of capture, in the region about Pasa- dena, range from September 15 to April 17; in the Santa Monica Mountains, from October 26 to February 22; from the Ojai Valley, Ventura County, there is one as late as April 27. There is but one specimen (no. 30446, Mus. Vert. Zool., Seaside, Monterey County, December 30) from a coastal point between Santa Barbara and San Francisco, but it would not be surprising should the subspecies be found regularly, migrating at least, along the coast ranges of this section. On the whole there is a notable contrast in the abundance of this subspecies in its winter home in southern California, and its scarcity as a migrant farther north in the state. An explanation of this con- dition may be found in this bird again being one that leaves its breed- ing ground abruptly and departs by a long flight to its winter home, passing over much of the intervening country. Passerella iliaca brevicauda Mailliard Yolla Bolly Fox Sparrow Original description.—Passerella iliaca brevicauda Mailliard, 1918, p. 138. Type specimen.—No. 23924, Calif. Mus. Vert. Zool.; female adult ; one-half mile south of South Yolla Bolly Mountain, in Trinity County, California; August 7, 1913; collected by A. C. Shelton and George Stone (original number 385). Range.—There are breeding birds at hand from South Yolla Bolly Mountain, which lies at the junction of Trinity, Tehama and Mendo- cino counties, from Snow Mountain, in the northwestern corner of Colusa County, and also one juvenal from Mount Sanhedrin, in north- eastern Mendocino County. Winter birds have been examined from the near vicinity of the coast, from Marin County, central California, and from Santa Barbara, Ventura and Los Angeles counties, includ- ing Santa Catalina Island, southern California. Specimens examined.—73 (see list, pp. 204-205). Distinguishing characters——Of the Schistacea group (see p. 89). Of the subspecies within this group brevicauda is superficially most nearly like stephensi, the two being noticeably alike in the enormous development of the bill. Between stephensi and brevicauda there are differences of color and proportions. Brevicauda is brownish colored, stephensi grayish ; brevicauda has the tail shorter than stephensi (see table 4), has weaker claws (see fig. AA), and a somewhat differently shaped bill (see fig. F). 166 University of Califorma Publications in Zoology [Vow 21 Remarks.—Although brevicauda is one of the more recently differ- entiated local races of the species, it so happens that there is more information at hand regarding the seasonal distribution of this sub- species than there is regarding that of any of its near relatives. The available data include facts of decided interest, the locally restricted winter habitat being especially worthy of emphasis. In its summer home brevicauda occupies the westernmost point of any of the schistacea group, and in its winter home also it keeps just as far to the westward as possible. Other closely related subspecies occur quite as far west, it is true, but brevicauda in winter does not extend east- ward (inland) for any distance, as do the others. It hugs the near vicinity of the coast. The local restriction of the race in Marin Fig. AA. Feet of two subspecies of fox sparrow, natural size. a. Passerella iliaca stephensi, adult female; no. 2096, Mus. Vert. Zool.; San Jacinto Mountains, California; July 10, 1908. b. Passerella iliaca brevicauda, adult male; no. 23922, Mus. Vert. Zool.; South Yolla Bolly Mountain, Tehama County, California; August 1, 1913. County, just north of San Francisco Bay, has been commented upon by Mailliard (1912, p. 63). Here it oceurs in winter apparently with fair regularity upon certain rocky, brush-covered ridges near the coast, and nowhere else. East of San Francisco Bay, on the University of California campus, this subspecies has not yet been detected among the several kinds of fox sparrows that frequent these grounds in winter. Neither is it included in the extensive series of Passerella collected over a period of years at Hayward, a few miles to the southward, by W. Otto Emerson. South of San Francisco Bay there are no record stations for this subspecies until Santa Barbara is reached, though it can hardly be doubted that it oceurs in winter at suitable spots. In this region, as elsewhere, it is probably restricted to certain limited areas. In Los Angeles County the peculiarly local distribution of the race receives special emphasis from the large number of specimens of Passerella available from many points within the county, rendering it out of all question that the predominance of certain subspecies at certain spots 1920] Swarth: Revision of Avian Genus Passerella 167 should be purely a coincidence (see pp. 110-111). Some of the writer’s earliest experiences in bird collecting (in the vicinity of Los Angeles) were with fox sparrows, and he soon discovered that the place to hunt for ‘‘thick-billed sparrows’’ (quite satisfactorily thick-billed they were, too!) was in certain brushy canons between Los Angeles and the ocean, while for ‘‘Townsend’s sparrows’’ (any of the Unalascheensis group) it was usually necessary to make trips to the mountain tops and canons to the eastward above Pasadena. In its history thus far, brevicauda has been regularly confused with stephensi. Winter specimens of the former at one time were cited by Mailliard (1912, p. 63) as demonstrating the winter occurrence of the latter in Marin County. Oberholser (1900, p. 233) lists speci- mens of stephensi from Santa Catalina Island in April, and Bowles (1911, p. 175) from Santa Barbara in August. The specimens upon which the above records were based have all been examined by the present writer, and all prove to be examples of brevicauda. It will be noted that Oberholser (loc. cit.) comments upon the brown dorsal coloration of the Santa Catalina Island specimens, one of the dis- tinguishing features of brevicauda as compared with stephensi. While brevicauda has thus been confused with stephensi wherever the former has been collected, I nevertheless do not believe the rela- tionships of the two to be especially close. The large bill, common to both, is, of course, a most conspicuous feature, the presence of which would be apt to lead the observer astray, for without strongly con- firmatory evidence one would hardly assume the existence of two subspecies of the same species, each characterized by exceptional development of the bill. With this feature conspicuously in view, the differentiating characters of color and proportions are not such as to impress the observer so strongly, especially with but a limited number of specimens for comparison. With an adequate series, how- ever, it can be seen that the peculiar characters of brevicauda are further accentuations of the features, not. of stephensi, but of megar- hynchus. The latter is a rather brownish colored bird, compared with mariposae and stephensi, and with a broad, stubby bill. Brevi- cauda is still browner in color, and with the same shaped bill greatly cnlarged (see fig. F). In typical stephensi, at least in adult birds, the bill is rather longer and more slenderly pointed, very much the shape seen in mariposae, though, of course, of greater size (see fig. E). Individual variation in subspecific characters is manifested in the series of brevicauda to about the same extent as in the other races 168 University of California Publications in Zoology [Vou. 21 of Passerella. Such variation in most cases takes the form of inter- gradation toward megarhynchus, sometimes in slight degree, occasion- ally to an extent that makes the placing of the individual specimen a matter of arbitrary decision. It is noteworthy that the birds from Snow Mountain, the southernmost known breeding station of brevi- cauda, have the bill perceptibly larger than in the Yolla Bolly birds, to the northward. The latter thus appear to show in some degree intermediateness toward megarhynchus. In but a few cases, among winter birds, is there any occasion for confusion with stephensi. In faded and worn plumage there may sometimes be difficulty in deciding between these two subspecies, but there are no fresh-plumaged birds at hand that are at all equivocal. In addition to the normal variation just described, there is one specimen at hand (coll. W. L. Dawson, male, ‘‘Santa Barbara Co., Cal.,’’ November 22, 1913) that shows a noteworthy trace of albinism. It has a pure white area on the forehead, extending from eye to eye, from the base of the bill backward to a maximum width of six milli- meters. On the left side it includes the lores and extends below the eye over the cheek. This white marking is of interest through its appearance of being an unusual extension of the two whitish spots between eyes and nostrils, present to a greater or less extent in prac- tically all fox sparrows. The series of breeding birds at hand sheds little ight upon the dates of arrival and departure of this subspecies upon its nesting ground, for they are all midsummer specimens. The winter birds from Marin County were taken on dates ranging from September 23 to April 29. Specimens collected by Maillard on September 25 were believed by him to have been raised possibly at the spot where they were taken (Mailliard, 1912, p. 63), a supposition that has not as yet been verified. One specimen at hand from Nicasio, Marin County (Carnegie Museum, no. 29698), bears the date of ‘‘June’’ 28. The label now attached to this skin is obviously not that of the original eollector. In all probability a mistake was made in copying, so that this skin can not be admitted as evidence of the breeding of brevicauda at this place. From southern California by far the earliest fall migrant is Bowles’ specimen from Little Pine Mountain, Santa Barbara County, taken August 30. From the Santa Monica Mountains there are speci- mens from October 4 to January 19, but there is little doubt that the birds arrive earlier and depart later than is indicated by these dates. 1920] Swarth: Revision of Avian Genus Passerella 169 There is one skin at hand from the Ojai Valley, Ventura County, April 18. From Santa Catalina Island a specimen collected April 19 affords the latest date for southern California. Specimens collected on the same island in mid-December attest the wintering of the bird there. Passerella iliaca canescens Swarth White Mountains Fox Sparrow Original description.—Passerella iliaca canescens Swarth, 1918, p. 163. Type specimen.—No. 28439, Calif. Mus. Vert. Zool. ; immature male (in nearly complete first winter plumage) ; Wyman Creek at 8250 feet altitude, east slope of White Mountains, Inyo County, California; August 15, 1917; collected by A. C. Shelton; original number 3549. Range.—In summer apparently confined to the White Mountains, in Inyo and Mono counties, California. Migration route and winter habitat unknown, save as the species is represented by a few speci- mens from Los Angeles and Santa Barbara counties and the Colorado River, California. Specimens examined.—23 (see list, p. 205). Distinguishing characters——Of the Schistacea group (see p. 89). Distinguished from all other subspecies within this group, save schistacea, by its diminutive bill (see fig. D). From schistacea, canescens may be differentiated by its decidedly more grayish ecolor- ation. This is strikingly apparent when freshly molted birds of the two subspecies are compared, and it is also evident in the juvenal plumage. In worn midsummer birds these color differences naturally are obscured. Remarks.—The type series of canescens consists of twelve speci- mens from the Boreal Zone on the White Mountains, in Inyo and » Mono counties, California. The series includes three adults in rather worn summer plumage, two adults undergoing the annual molt, but mostly in the new winter plumage, and six in juvenal plumage, some of them showing a few feathers of the first winter plumage. There are a few winter birds at hand from southern California that appear to belong to this subspecies, all taken at points some distance trom the coast. One from Mount Wilson, November 4, 1894 (no. 34, Swarth eoll.), appears to be typical canescens, extremely small-billed, of decidedly slaty coloration, and the breast spotting rather sparse and with no trace of the reddish brown peculiar to schistacea. Others from Blythe, Riverside County (on the Colorado River), San Antonio Canon, San Bernardino County, San Dimas and Palmer’s canons, Los Angeles County, and Santa Barbara, though not so extreme in character, are apparently also to be referred to canescens. There is a notable difference between these birds and the brownish colored schistacea found migrating in the Sierra Nevada. 170 University of Califorma Publications in Zoology [Vou. 21 The southern California winter specimens do not suffice to indicate the winter habitat of this subspecies. Apparently, however, canescens, like the other Sierran and Great Basin forms, avoids the vicinity of the coast, and must be looked for farther inland. The points of winter capture here recorded may indicate the northern and western limits at that season. It is a noteworthy fact that in all the bird collecting carried on by the Museum of Vertebrate Zoology in Owens Valley and in the Death Valley region, not a single migrating example of canescens was secured. As these places are directly south of the White Moun- tains, they would certainly be traversed by migrating canescens, unless —as seems probable—the birds pass over these regions in long, direct flights toward and from the winter habitat. Accumulating data points more and more convincingly toward this latter mode of travel as prevalent among many of the subspecies of Passerella. Passerella iliaca monoensis Grinnell and Storer Mono Fox Sparrow Original description—Passerella iliaca monoensis Grinnell and Storer, 1917, p. 165. Type specimen.—No. 26930, Calif. Mus. Vert. Zool.; male adult; Mono Lake Post Office, altitude 6500 feet, Mono County, California ; May 21, 1916; collected by Joseph Dixon; original no. 4644. Range.—Breeds on the east slope of the Sierra Nevada in the vicinity of Mono Lake; possibly also in the Panamint Mountains. Occurs in winter in the western foothills of the Sierra Nevada and on the Pacific slope of southern California. Specimens examined.—38 (see list, p. 206). - Distinguishing characters.——Of the Schistacea group (see p. 89). A gray-colored fox sparrow, in bill character intermediate between canescens and mariposae (see fig. E). Of the pale coloration common to the canescens-monoensis-mariposae-stephensi series. In this line of subspecies monoensis occupies a position comparable to that of fulva in the schistacea-fulva-megarhynchus-brevicauda chain. Specimens of monoensis and fulva are sometimes very much alike in bill structure, but may be distinguished by coloration, fulva being a brownish colored bird, monoensis a grayish colored one. Remarks.—There is a fox sparrow at hand, an adult male in rather worn plumage, collected at Jackass Springs, Panamint Mountains, California, June 25, 1917 (no. 28430, Mus. Vert. Zool.) that I tenta- tively refer to monoensis. The capture of this specimen in midsum- mer, a bird which was in full song and in breeding condition, is fair 1920] ‘Swarth: Revision of Avian Genus Passerella eal . iliaca fulva, breeding . iliaca mariposae, breeding . iliaca stephensi, breeding . iliaca brevicauda, breeding . iliaca monoensis, breeding . iliaca monoensis, doubtful breeding record ey, Ze pees eee DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY Cob See Fig. BB. Map showing summer habitat of Passerella iliaca fulva, P. %. mari- posae, P. %. monoensis, P. i. stephensi, and P. i. brevicauda. The symbols indicate ae of breeding stations, as established by specimens examined by the author. 172 University of California Publications in Zoology [Vou. 21 evidence that the species was nesting in the mountains where it was taken, though no more of the birds were seen. If I am correct in ascribing this specimen to monoensis, it may indicate that the sub- species occupies a rather extensive summer habitat on the summits of certain of the desert mountains. However this may be, the only breeding specimens at hand that appear to be unequivocally of the race monoensis (aside from the one exception above noted) are all from a relatively limited region about Mono Lake. Specimens from Mammoth, immediately to the southward, appear to be mariposae, as are the birds still farther south on the east slope of the Sierras (see p. 175). Furthermore, between Mono Lake and the Panamint Mountains lie the White Mountains with the very different subspecies canescens. It is hardly reasonable to suppose that the latter race would be thus interposed between the known habitat of monoensis at Mono Lake, and an outlying colony such as seems to be indicated by the Panamint Mountains specimen. Altogether, the capture of the latter is an extremely puzzling occurrence. While engaged in this study of Passerella, I took occasion to request Mr. Donald D. McLean to collect for the Museum specimens of fox sparrows from the vicinity of his home, near Coulterville, Mariposa County, California. There were but few of any of the subspecies at hand from the Sierra Nevada foothills, and he had told me that there 2? were many ‘‘thick-billed sparrows’’ in that region during the winter months. To my surprise, of the nine birds of this type that he sent in, collected during December and January, eight proved to be mono- ensis. Coulterville is at the west base of the Sierra Nevada, almost due west of Mono Lake, though, of course, with the whole width of the mountain chain between the two points. There is a specimen of monoensis at hand taken in Yosemite Park, October 1, and four others from Horse Corral Meadow, in the mountains of Fresno County, secured between September 19 and 23, indicating that the subspecies spills over the crest of the Sierras on to the western slope in the migration from the summer home. If the main winter habitat should prove to be in the western foothills of the Sierra Nevada as is indi- cated by the Coulterville birds, it would imply a rather remarkable migration route—almost due east and west, rather than north and south. There are a few scattered winter birds at hand from points in Los Angeles County, including Santa Catalina and San Clemente islands, all taken west of the desert divides. Extreme dates of capture 1920] Swarth: Revision of Avian Genus Passerella 173 are September 24 (San Antonio Cafion) and March 28 (San Clemente Island) ; other specimens have been taken during October, December, January, and February. Passerella iliaca mariposae Swarth Yosemite Fox Sparrow Original description—Passerella iliaca mariposae Swarth, 1918, p. 161. Type specimen.—No. 25693, Calif. Mus. Vert. Zool.; adult male; ridge at 7000 feet, near Chinquapin, Yosemite Park, California; June 10, 1915; collected by J. Grinnell; original number 3284. Range.—Summer visitant to the Canadian and high Transition zones in the northern and central Sierra Nevada. Breeding stations represented by specimens in hand range from central Siskiyou County (head of Little Shasta River) at the north, south at least to the Yosemite region on the west slope of the Sierras, to Kearsarge Pass on the east slope. There are a few winter specimens at hand from scattered points in San Diego, Los Angeles and San Bernardino counties, but too few to indicate accurately the entire winter habitat or migration routes. Specimens examined.—206 (see list, pp. 206-209). Distinguishing characters.—Of the Schistacea group (see p. 89). With the gray coloration common to the canescens-monoensis-mariposae- stephensi series, and not to be distinguished in color and markings from the other members of this chain of subspecies. In bill structure mariposae lies between monoensts and stephensi (see fig. E). In gen- eral size and length of tail in this line there is progressive increase from canescens to stephensi, and in these particulars as in size of bill, mariposae occupies its appropriate intermediate position (see table 4). Remarks.—While there is at hand an abundance of summer speci- mens of mariposae, there is a notable dearth of winter collected birds. These facts, coupled with the reverse condition in megarhynchus, of which I have seen winter specimens only, may be used by some as arguments proving the two to be identical, and only suffered to masquerade under different names through a misunderstanding of seasonal differences. In contravention to this idea it may be pointed out that there are many specimens in the available series of mariposae taken at summer stations just prior to the birds’ departure therefrom in the fall, and after they had assumed the winter plumage, these specimens sufficing to show the color differences distinguishing mari- posae from megarhynchus. Shape of bill is an excellent differentiating character between adults at any season. There are a few examples of mariposae at hand taken in winter at various points in southern California, but they are so few in 174 University of California Publications in Zoology [Vou 21 number compared with the abundance of certain other subspecies associated with them that there seems no doubt that the places of capture of these scattered individuals do not represent the main winter home of the race. Then, too, some of these winter birds are not absolutely typical of mariposae, though most nearly resembling that form. Altogether, it is evident that the winter metropolis of mariposae is still to be discovered. In this connection it may be pointed out that we are similarly ignorant of the winter homes of the other Sierran fox sparrows, stephensi and monoensis, and of their neighboring relatives fulva and canescens. In each case there are scattering winter birds and migrants at hand, but nothing to show where the bulk of the populations winter. It is not to be supposed that all five of these subspecies necessarily depart to the same place, but it seems possible that their modes of departure and arrival are very similar, and that the explanation of the puzzle in each ease lies in the existence of a relatively restricted winter habitat, and one that has not been thoroughly scrutinized for these birds at that season. It may be, of course, that mariposae winters in some parts of the Sierran foothills, but there is little evidence of this either in the avail- able specimens, or in published literature. Belding (1890, p. 170), speaking of the Sierran bird (under the name megarhynchus), says: ‘‘Not at Alta and Colfax November 17-21, nor at Red Bluff in warm winter of 1884-85. I never see it in the lower foothills of Calaveras County in winter, though P. wnalaschcensis is common there at that time.’’ That the detailed mapping of summer ranges of the forms of Passerella breeding in the Sierra Nevada will yield interesting and valuable results is indicated by the general facts already gathered. The habitats of stephensi and mariposae are each known to be inter- rupted along certain lines where thorough exploration has been prose- cuted, and as there are other parts of the Sierras presenting similar conditions it is evident that at these places also there must be breaks in distribution, and that each of these subspecies in summer is scat- tered into a number of colonies of varying extent. The Kings River Cafion section of the Sierra Nevada may be cited in its bearing on the fox sparrow problem. This canon runs far back into the mountains, a broad, gently sloping valley, Transition zone clear to its uppermost confines and therefore not inhabited in summer by Passerella. The surrounding walls are high and steep, and drop from plateaus, zonally of high Transition, Canadian and higher. In 1920] Swarth: Revision of Avian Genus Passerella 175 this elevated region south of the Kings River Cafion the present writer found stephensi in abundance in the late summer, evidently on its breeding ground. This is assumed to be the northernmost point reached by this subspecies in the Sierra Nevada, though no collecting has as yet been done on the high country immediately north of the Kings River Canon. This gorge evidently acts as a barrier to the continuous distribution of Passerella in this direction. From the head of the Canon (Transition Zone) steep slopes arise, carrying an extremely limited belt of Canadian Zone (where we saw no Passerella) and leading abruptly to the extensive strip of Hudsonian and Alpine- Arctic along the crest of the Sierras. These latter belts are not occupied by Passerella, hence they act as a barrier between colonies on the east and west slopes of the mountains. On the east side, in Kearsarge Pass, mariposae is abundant in summer, the southernmost point at which the subspecies has been found breeding. This station on the east side of the mountains is directly opposite that point on the west side where stephensi occurs at its northern limit, the two being separated by the intervening strip of Alpine-Arectic and Hudsonian. Thus mariposae occurs consider- ably farther south on the east slope of the Sierra Nevada than on the west slope. There are one or two specimens of Passerella at hand from Mammoth, Mono County, on the east slope and close to the habitat of monoensis. These appear to be referable to mariposae, and indicate the probable extent of territory occupied by this subspecies on the east slope of the Sierras, a territory cut off from the western colony for its entire length by the Alpine-Arctic and Hudsonian zone of the summit. It is difficult to understand why this isolated division should not be of the monoensis type. There is no obvious barrier between it and that subspecies, and reasoning d@ priori one would expect the features of the latter race to be developed over this entire eastern exposure where Passerella occurs. Such is most emphatically not the ease at the southern limits of mariposae, however, Kearsarge Pass specimens being unmistakably similar to mariposae of the Yosemite region, and impossible to confuse with their nearer neighbors, stephensi on the one hand and monoensis on the other. Farther north, in Eldorado County, there is what appears to be intergradation toward monoensis. Birds from the western part of this county are typical mariposae, but in the series from the immediate vicinity of Lake Tahoe and Mt. Tallae there are some individuals quite as small-billed as selected examples of monoensis. So, too, as regards 176 University of Califorma Publications in Zoology [Vou. 21 three skins from Douglas County, Nevada, southeast of Lake Tahoe. Two of these are about as small-billed as monoensis, while the third is an average mariposae. Monoensis is not a strongly marked form, and its characters, such as they are, obviously show intermediateness between mariposae and canescens, between which races monoensis is situated geographically. This fact may have some bearing on the relative restriction of the region in which typical monoensis occurs. Doubtless also, a careful working out in greater detail than has yet been done of the breeding ranges of the several subspecies concerned will go far toward explain- ing facets the meaning of which now seems obscure. There is little data at hand to indicate precise dates of arrival and departure of the subspecies mariposae at the summer and winter homes. In Kearsarge Pass, on May 27, male fox sparrows were in full song, evidently settled at their nesting grounds, and on May 29 a set of three well incubated eggs was taken. There is a female mari- posae at hand (no. 29090, Mus. Vert. Zool.), which had laid part of its set, taken May 15 at the head of Little Shasta River, Siskiyou County, the northernmost point of record for the subspecies. Of fall dates in the Sierra Nevada, there is a large series of mariposae at hand taken at Cisco and Blue Canon, Placer County, from August to October, the latest from Blue Cafion, October 14. There is one bird from Tuolumne River, Yosemite Park, October 1. There is one speci- men at hand taken at Yermo, on the Mohave Desert, May 28. In view of the above nesting dates this last must be regarded as an extremely belated migrant. This bird is a typical example of mariposae; the point of capture, of course, is many miles from the nearest point in this subspecies’ summer habitat. Two specimens from Mt. Wilson, Los Angeles County, taken April 1 and 4, respectively, may indicate more nearly the usual limit of its stay in the spring in the south. Passerella iliaca stephensi Anthony Stephens Fox Sparrow Original description.—Passerella iliaca stephensi Anthony, 1895, p. 348. Type specimen.—No. 15387, Carnegie Institute Museum; male adult; San Jacinto Mountains, California, at 8000 feet; July 14, 1895 (see p. 181). Range.—Summer visitant to upper Transition and Canadian zones in the southern Sierra Nevada and the Sierras of southern California. There are several isolated tracts occupied by this subspecies, namely : in the Sierra Nevada, from Hume and Horse Corral Meadow, Fresno 1920] Swarth: Revision of Avian Genus Passerella 177 County, south through Tulare County; on the summit of Mount Piiios, Ventura County; in the San Gabriel Mountains, Los Angeles County ; the San Bernardino Mountains, San Bernardino County; and the San Jacinto Mountains, Riverside County. The winter home is unknown. Specimens examined.—186 (see list, pp. 209-211). ia ae wt ru fos a Passerella iliaca stephensi Passerella iliaca brevicauda Passerella iliaca monoensis DISTRIBUTION MAP MUSEUM OF YERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA Ss 7 rd ra We = cra ~ 7 Fig. CC. Map showing winter occurrence of Passerella iliaca stephensi, P. i. brevicauda, and P. %. monoensis in California, as established by specimens exam- ined by the author. Note the restriction of brevicauda to the near vicinity of the coast. The locality stations for stephensi denote the capture of but four specimens, apparently stragglers from the still unknown main winter habitat. Distinguishing characters——Of the Schistacea group (see p. 89). Most nearly hike mariposae, from which it is distinguished by slightly greater general size and notably large bill (see fig. E). The enor- mous bill distinguishes stephens: from all other fox sparrows except brevicauda. From the latter it is distinguished by rather longer, 178 University of Califorma Publications in Zoology [Vou. 21 more slenderly pointed bill (compared with the blunt, stubby bill of brevicauda) (see fig. F), by more grayish coloration, and by longer tail (see table 4). Remarks.—In all its characters stephensi represents the cul- mination of the lne of development beginning with canescens and continuing through monoensis and mariposae. Gray coloration is common to all these forms, as compared with the more brownish hue of the other subspecies of the Schistacea group, and there is no appar- ent difference in this respect between these several southern subspecies. There is, however, gradual increase in general size from canescens along the line indicated to stephensi, with notable increase in size of Fig. DD. Bills of two subspecies of the Schistacea group (races that are distinguished mainly by differences of size in this member), showing how variation of adult and immature may be mistaken for variation differentiating subspecies. a and b are, respectively, adult and undeveloped immature of stephensi, c and d, of mariposae; b is more nearly like ¢ than like a. a. Passerella iliaca stephensi, adult male; no. 20505, Mus. Vert. Zool.; Taylor Meadow, Sierra Nevada, Tulare County, California; July 19, 1911. b. Passerella iliaca stephensi, immature male; no. 27388, Mus. Vert. Zool.; Hume, Fresno County, California; August 18, 1916. c. Passerella iliaca mariposae, adult male; no. 25693. Mus. Vert. Zool.; Chin- quapin, Yosemite National Park, California; June 10, 1915. ad. Passerella iliaca mariposae, immature male; no. 23195, Mus. Vert. Zool.; Blue Cafion, Placer County, California; August 28, 1912. bill, and there is also, to a lesser extent, some increase in proportionate length of tail. Stephensi has a strikingly disconnected range, occurring in a number of widely separated high mountain localities. In the southern Sierra Nevada the subspecies is found in high Transition and Can- adian from southern Fresno County (at points just south of Kings River), south through Tulare County. The localities of occurrence are all on the west slope of the range. Fox sparrows at this latitude do not ascend into the Hudsonian and Alpine-Arctie zones, which constitute so large a part of the summit of the Sierra Nevada. This strip along the divide is an effective barrier to the general dispersal of these birds, and in the case of stephensi it has apparently restricted the subspecies to the western slope. 1920 | Swarth: Revision of Avian Genus Passerella 179 South of the Sierra Nevada the next breeding station of stephensi is a limited area on the summit of Mount Pinos, some eighty miles southwest of the southernmost station in the Sierras. Then, still farther south, there are widely disconnected areas on the San Gabriel Mountains, the San Bernardino Mountains, and the San Jacinto Mountains, separated from each other by many miles of lower zones. In the San Jacinto Mountains points of occurrence are mostly from 8000 to 9000 feet, though a few birds were seen in one part of the mountains as low as 6000 feet. In the San Bernardino Mountains the subspecies occurs at about the same elevation, mostly from 7400 to 9000 feet; in the San Gabriel Mountains it is known only from the higher slopes of Mount Waterman from 7000 feet upward to the summit (7752 feet altitude) ; and on Mount Pinos (altitude 8826 feet) from 8000 feet upward. In the Sierra Nevada the vertical range is from 7000 to 10,000 feet. It is a notable fact that, despite the isolation of these several colonies and the demonstrated plasticity of the species throughout its range, there are no discernible differences between specimens of stephenst from the several regions occupied. Considering the varia- tion existing between the many closely connected subspecies, and the way in which each represents a step from one extreme to another, it might be expected that such widely parted groups as those into which stephensi is divided would illustrate separate and distinguishable stages as they advanced farther and farther from the starting point, but such is not the ease. In the series at hand I am unable to appre- ciate any tangible features serving to distinguish birds even from the extremes of the subspecies’ range. The most that can be said regard- ing such differences is that in the series from Hume, Fresno County (the northernmost point at which stephensi has been collected), there are certain small-billed individuals that may be taken to illustrate intergradation with the nearby form mariposae, but in the same series there are other specimens with quite as large bills as the maximum from any other place. In other words, while stephens? is the culmin- ation of a series of steps from canescens and through monoensis and mariposae, the maximum of change is reached as soon as the habitat of the subspecies is entered at the north, where mariposae and stephensi come together. There is no further accentuation of character in the extensive but disconnected habitat of stephensi as it extends southward. Stephens fox sparrow is a summer visitant only upon its breeding grounds, but there are neither specimens nor notes available indicating 180 University of Califorma Publications in Zoology [Vou.21 dates of arrival in its summer home. All the breeding birds at hand were taken long after they had settled down to the duties of nesting. As regards the time of departure at the summer’s end, this can be fairly definitely fixed. In the San Jacinto Mountains I revisited in September, 1914, certain sections where the Stephens sparrow had been found in abundance in the summer of 1908. On September 9 one was secured near Strawberry Valley, somewhat below the breeding level in this range. On September 11, after most careful search, four were seen in Tahquitz Valley, in the heart of the nesting ground, and on September 15, one other was noted, the last observed, though I remained in the region some days longer. In the San Bernardino Mountains Stephens sparrows ‘‘were seen in small numbers at Bluff Lake as late as September 3’’ (Grinnell, 1908, p. 99). At Hume, Fresno County (altitude 5300 feet), Stephens sparrows were abundant during the middle of August, 1916, evidently on their nesting grounds from the number in juvenal plumage, and - numerous specimens were collected from August 17 to 23. On Sep- tember 25 and 26, this point was revisited, and while fox sparrows were again numerous they proved to be practically all examples of more northern subspecies on their southward migration. One stephensi was collected, on September 26. The main winter home of the Stephens fox sparrow is unknown, and it seems evident that when the birds leave their summer habitat it is an abrupt departure, and possibly to a relatively great distance. They do not descend the mountains to the foothills and adjacent valleys, which are occupied in winter by other subspecies, or they would surely have been detected in such places. Stephensi has been reported in winter from Santa Catalina Island (Oberholser, 1900, p. 233), but the specimens upon which the statement was based prove to be of another subspecies, brevicauda (see p. 167). In partial contradiction to my general statement regarding the winter habitat, I must admit to having before me four fox sparrows, taken in midwinter and to all appearances unequivocal examples of stephensi. The data for these skins are as follows: No. 8106, coll. F. 8S. Daggett, male, Franklin Canon, Santa Monica Mountains, Los Angeles County, California, November 24, 1915; no. 1424, coll. J. E. Law, female, Hollywood, Los Angeles County, California, February 22, 1912; no. 1762, coll. W. M. Pierce, male, Palmer’s Cafion, Los Angeles County, California, January 11, 1919; coll. A. Brooks, male, Santa Barbara, California, January 11, 1913. |) ciutticce Ch ane RAEN ceed SPA fae Carnegie Mus. 32666 Onn Mus. Hist. Sci. Art 21491 | oO im. PU! || 6H coscce PONE Noe acess 21621 QO im. 21541 Oo ee U.S. Nat. Mus. 81390 | oO ad. 81391 9 ad. 170231 O ad. Victoria Mem. Mus. 4080) hot a 4081 Coe. 4105 One D. R. Dickey A MIRO We ee J. H. Fleming 25392 (oe J. Grinnell 1673 ols ee 1820 | See 1821 Ol ae 2097 oft eas SVB |) asta 3280 Of ees A. B. Howell AQ33 1a een ces AQG 2a On nee 2139! OR. GEVBM |) Cf! cau 6828! Chow. 68251 Sue 68291 OR J. E. Law TIS | Aeon J. & J. W. Mailliard sera a ~ See ue me Cw Se art 2 aes ~ aoa e « Pi ee ere 7 cae, emerges ppp hs aS “= -_ = oe a UNIVERSITY OF CALIFORNIA PUBLICATIONS iN “ ZOOLOGY Vol. 21, No. 5, pp. 225-238, 1 text figure May 20, 1920 A STUDY OF THE CALIFORNIA JUMPING MICE OF THE GENUS. ZAPUS BY -. . A. BRAZIER HOWELL 6, aes Ys UNIVERSITY OF CALIFORNIA PRESS BERKELEY UNIVERSITY OF CALIFORNIA PUBLICATIONS : Note.—The University of California Publications are offered in exchange for the publi- | cations of learned societies and institutions, universities, and libraries. Complete lists of - all the publications ofthe University will be sent upon request. For sample copies, lists of publicaticnsor other information, address the MANAGER OF THE UNIVERSITY PRESS, BERKELEY, CALIFORNIA, U. 8..A. 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Ea “$5.00 Volume 15. 1915-1916, 360 pages, With 38 plates -..s.i coco cocicicccccensanncaencaccennegnenesenenadeencene== $5.00 ~~ Volume 16, 1915-1917, 522° pages, With 46 plates -...n. ccs cece penececceneeecenenstceceseepecnenennen SS $5.00 —— Volume 17, 1916-1918, 545 pages, with 24 plates -..2cc..22e--w--ciecctitecesceenetsssteeenectereneacensteanes DOO Volume 18, 1917-1919, 519 pages, with 20 plates. (Index im preparation) «...-..--...-.- $5.00 tC ye ¢ : ee witch yakan! 5 hawk Shee Ves ok UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Vol. 21, No. 5, pp. 225-238, 1 text figure May 20, 1920 A STUDY OF THE CALIFORNIA JUMPING MICE OF THE GENUS ZAPUS BY A. BRAZIER HOWELL (Contribution from the Museum of Vertebrate Zoology of the University of California) PREFATORY NOTE i Sep ed aS In attempting to identify certain specimens of Zapus from’ Caf: fornia it became necessary for me to study the whole genus in so far as it is represented within the state. The results of this study con- stitute the present paper. The material used first in making com- parisons consisted of adults selected at random from the series at hand, yet with due regard to locality, season, and condition of pelage. The conclusions reached from examination of this material were then checked and rechecked with the full series. The tables of measurements are of properly comparable specimens from as near the type localities of the several forms as possible. All measurements are in millimeters. The names of the colors employed are those used in Ridgway’s Color Standards and Color Nomenclature (1912). The specimens examined in the present revision are contained chiefly in the collection of the Museum of Vertebrate Zoology of the University of California. In addition, material was borrowed from Donald R. Dickey, D. E. Brown, and the Bureau of Biological Survey of the United States Department of Agriculture. For help rendered during the preparation of this paper, the writer is under obligation to Dr. Joseph Grinnell, Mr. Harry S. Swarth, and Mr. Tracy I. Storer, of the staff of the Museum of Vertebrate Zoology. 226 University of California Publications in Zoology (Vou. 21 Two hundred and eighty-six specimens of Zapus have been exam- ined from forty-eight localities within the state of California, as well as small series of some of the forms from north of this state. GENERAL DISCUSSION The cranial differences which obtain between the forms of Zapus are not great, and there is seldom any one diagnostic character which makes identification possible without further comparison. The amount of individual variation in the skulls, even of topotypes of well marked species, is often considerable, but this is taken fully into account in defining the species and subspecies. In spite of the minor differences in any one series, there are usually several constant characters which, taken together, make final determination entirely satisfactory. There is available no series consisting of specimens taken in a single locality in all months during which the animals are active. This is a misfortune, for seasonal variation between spring and fall is considerable in some forms. However, this difference seems to be entirely due to fading and wear. Although none of the skins are in new pelage, it is assumed that there is only one annual molt, and that the new coat is acquired some time after the middle of September. Too much stress should not be laid on the exact shade of coloration within a series unless the material at hand be strictly comparable as to age and stage of pelage. Nor, of course, should too much confidence be placed in the measurements in the flesh made by different collectors, because of difference in method of measuring. There is seemingly no one character which can be considered as more important than any other. All parts of the cranium must be carefully considered. Rostrum including nasals, brain-case, incisive foramina, bullae, and the pterygoid fossae, all present features of value. The proportion of black to lighter hairs on the dorsal area is variable, but the tone of coloration of the sides and underparts is of importance. IJmmatures are usually a trifle paler dorsally than are adults. Coloration seems to have been less easily affected by evolutionary influences than the cranial characters, and by following this more conservative criterion, of color, it is found that California species fall into three natural groups, inhabiting three main geographic areas. 1920] Howell: A Study of the California Jumping Mice 227 These groups comprise: (1) Zapus trinotatus eureka and Z. orarius, in the humid coast belt; (2) Z. pacificus alleni, in the Boreal Zone of the main Sierran mountain mass; and (3) Z. major, in the isolated Warner Mountains of extreme northeastern California. Carrying our enquiries to the northward, it is found that group 1 is replaced at some indeterminate point by Z%. trinotatus trinotatus, and that the latter continues up the coast of Washington and also towards the interior, into the Cascade Mountains of that state. But the Cascade Mountains of Washington and of Oregon are separated by the Colum- bia River gap, and in the latter state there is an evident barrier between the coast and the mountains, constituted by differences in relative humidity and in temperature. Hence, on the main mountain mass of Oregon occurs Zapus pacificus pacificus, which is apparently distinct from any form which has been found to the north of that state. The relationships of the two races of pacificus (Z. p. pacificus and its southern neighbor Z. p. alleni) are discussed elsewhere. Zapus major, of the third group, undoubtedly has its closest affinities with the jumping mice which occur in the isolated mountain ranges of the Great Basin region. With this group is placed Z. mon- tanus, for, although that species is found on the southeastern border of the Cascades, and hence adjacent to the range of Z. p. pacificus, its relationship clearly is not close to the latter form. It is nearer to group 3, although specifically distinct from any form now known to the writer. The type localities of pacificus and montanus are only about thirty miles apart, and are separated by a remarkably even slope, containing not the slightest obvious geographical barrier. It must be inferred, therefore; that the two forms arose separately, in different faunal areas. Of montanus, nine, and of pacificus, five, topotypic specimens are available. This is a larger number than has heretofore been assembled, and it seems desirable, therefore, although not strictly within the scope of this paper, to compare these two species on the improved basis. When Preble (1899) monographed the genus, but two typical examples of pacificus were known, and these were in faded, washed-out pelage. My specimens were taken during the latter part of July and the middle of August, and appear to be normal in every way. The underparts of both species are pure white; the pattern of coloration is as usual for Zapus, but the sides of pacificus are ochraceous buff, while those of montanus are much duller, and close to tawny olive. In the dorsal area of the former, the black hairs are very slightly in the majority ; 228 University of California Publications in Zoology [Vou. 21 that area of the latter is still darker. In short, pacificus is a light- colored, ‘‘yellow-sided’’ form, while montanus is a dark, ‘‘buff-sided’’ form. The skulls of the two species are equal in size and in length of rostrum and nasals, but the rostrum of pacificus is more tapering, the bullae are smaller, and the incisive foramina are slightly narrower and more constricted anteriorly. The brain-case is slightly less expanded, and the upper tooth rows converge posteriorly instead of being parallel as in montanus. I am unable, however, to distinguish even the slightest difference in the interpterygoid fossae (see Merriam, 1897, p. 104). 1. Zapus trinotatus ewreka 2. Zapus orarius 3. Zapus pacificus allent 4, Zapus major Locality from which one or Q more specimens have been examined ® Type locality DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA SCALE 25 $0 —— MILES ORAFT oF 1919 u Fig. 1. Distribution in California of the species and subspecies of Zapus. 1920] Howell: A Study of the Califorma Jumping Mice 229 List oF Forms oF ZAPUS OCCURRING IN CALIFORNIA, WITH TYPE LOCALITIES Zapus trinotatus eureka, new subspecies Fair Oaks, Humboldt County, California Zapus orarius Preble Point Reyes, Marin County, California Zapus pacificus alleni Elliot Pyramid Peak, Eldorado County, California Zapus major Preble Warner Mountains, Oregon Kry TO THE CALIFORNIA FORMS OF ZAPUS 1. Underparts strongly suffused with the color of the sides. 2. Foot averaging more than 32.5 mm.; tail sharply bicolor and 60 per cent of total length; skull usually more than 24 mm. long -...-........... eureka 2’. Foot averaging less than 32.5 mm.; tail not sharply bicolor and less than 58 per cent of total length; skull usually less than 24 mm. I oh a ipa ee eee Sos Pere a Pee SERS ER IM ene 22S eee A ee Dee eee Me orarius 1’. Underparts white. 3) Coloration of sides) bright ochraceous): WU seeascee cece serene ene alleni 37 Colorations of sid@es cle Genwimiya olive sees eee seer eee major Zapus trinotatus eureka, new subspecies Humboldt Jumping Mouse Type——Female adult, skin and skull; no. 11703, Mus. Vert. Zool.; Fair Oaks, Humboldt County, California; August 27, 1910; collected by J. Dixon; original no. 1743. Range.—Northwestern coast region of California, from Mendocino City, Mendocino County, north at least through Humboldt County. Number of specimens examined, 24, from the following localities in California. Specimens in Mus. Vert. Zool., unless otherwise indicated. Mendocino County: Mendocino City, 1. Humboldt County: Eureka, 11; Fair Oaks, 6; Arcata, 5; Loleta, 1 (coll. D. R. Dickey). Diagnosis—A rather large, ochraceous-bellied Zapus, with sharply bicolor tail, nearest to Z. t. trinotatus (5 skins from Seattle and 3 from Puyallup, Washington; D. E. Brown and A. B. Howell collec- tions; assumed to be typical). From that species it differs in having larger bullae and pterygoid fossae, narrower basioccipital, and longer rostrum and nasals, the latter being wider posteriorly. The brain- case is slightly less inflated and the bases of the anteorbital foramina are heavier. Underparts more suffused with the color of the sides. External characters——Collectors’ measurements of all available adults indicate that eureka has a longer foot than has trinotatus, but my own measurements of museum specimens fail to substantiate this. In eureka and trinotatus in early fall the general color effect is close to tawny olive, and there is little difference in the coloration of the 230 University of California Publications in Zoology [Vou 21 dorsal areas and of the sides, except that in the case of eureka the tone is somewhat brighter in mass effect, and the dorsal areas of the least worn specimens are often paler. Skins of the two may be dis- tinguished by a glance at the underparts, which in trinotatus are mainly pure white, with the central area and a small portion of the lower throat tinged with a lighter shade of the lateral color, while the underparts of ewreka are evenly and uniformly so tinged. In the single spring specimen that is at hand of eureka, the dorsal area is fairly well defined, but is a trifle paler anteriorly, with a grayish cast. The inguinal region and the underparts immediately contiguous to the forefeet are white, while the hairs of the remainder of the under- parts are strongly tipped with ochraceous buff, and the lateral tone is a more intense shade of the same color, very much as in orarius. Skull—tThis form and orarius have especially tapering rostra and large bullae. Eureka differs from trinotatus as remarked above. For comparison with orarius, see under that species. Measurements —Average, in the flesh, of 15 apparently adult specimens from Humboldt County, California: length, 228.3 mm. ; tail, 144; foot, 33.1. Average of 11 Z. t. trinotatus from northern Washington: length, 230.5; tail, 145.4; foot, 31.1. Remarks.—The California coast forms of Zapus are evidently con- fined to the narrow, humid strip between the dense forests of the Coast Range and the sea, though they may extend up the more open river valleys for varying distances. This part of the coast is chiefly within the Boreal Zone. It is extremely moist, and drenched with ~ heavy fogs during the drier season. In the main, its fauna is definitely separated from that of the Boreal caps on the mountains on the west of the upper Sacramento Valley by a wide stretch of semi-arid Transition throughout which it is practically certain that Zapus does not occur. Eureka probably grades into trinotatus to the northward, but at just what point it is now impossible to say. There are specimens in the Field Museum of Natural History from Requa and Crescent City, and it is most unfortunate that they cannot be examined at this time. They may be trinotatus, but for the present they are provisionally referred to eureka. Similar treatment is accorded the two specimens from California which were listed by Preble (1899, p. 27) under trinotatus. 1920] Howell: A Study of the California Jumping Mice 231 Zapus orarius Preble Point Reyes Jumping Mouse Zapus orarius Preble (1899, pp. 29-30). Type—Male adult; no. 250; coll. E. A. and O. Bangs; Point Reyes, Marin County, California; May 14, 1893; collected by C. A. Allen; original no. 618 (fide Preble, 1899, p. 29). Range—Bunch grass marshes on the uplands of the Point Reyes Peninsula, Marin County, California. Number of specimens exam- ined, 17 (2 in coll. A. B. Howell), all from the type locality. Diagnosis —An ochraceous-bellied species with rather short, dully bicolor tail. Distinct from, but perhaps nearest to, Z. tf. eureka; coloration brighter, foot and tail shorter. Skull shorter and more slender, the supraoccipital region hghter, rostrum shorter, lighter, and narrower at all points. The interorbital width is less, incisive fora- mina shorter and comparatively wider. The interpterygoid fossa is narrower and the antcorbital foramina and bullae slightly smaller. External characters—In this species, the tail averages less than 58 per cent of the total length, while in all other California forms, this member is 60 per cent. Only May and June skins are available, and in these, the dorsal areas are rather poorly defined, the black hairs outnumbering the lighter ones but slightly if at all. The head is uniform with the back and the general color tone is bright ochraceous buff. In most of the skins, the hairs of the underparts are uniformly tipped with the color of the sides, but a few specimens are somewhat lighter in this region. Skull.—Orarius has the smallest skull with narrowest interorbital width, shortest incisive foramina and shortest molar series of any form in the state. It and eurcka have especially tapering rostra and large bullae. Measurements.—Average, in the flesh, of 11 apparently adult topo- types: length, 224.3 mm.; tail, 129.4; foot, 31.6. Remarks.—Z. orarius is a well defined species and apparently intergrades with no other form. However, it is not impossible that specimens which are intermediate between it and eureka may be found along the coast of Sonoma County. Its closest relationship is certainly with the latter form. From alleni, it is separated by a wide stretch of Upper Sonoran territory. When Preble (1899, pp. 29-30) named the species, he had but two specimens from the type locality, and with these he listed an individual 232 University of California Publications in Zoology [Vou 21 from Eureka and another from Mad River, Humboldt County. Con- sidering the paucity of specimens from the coast then available, and the fact that the former skin was without a skull, it seems safe to assume that both of these examples should now be referred to eureka, especially when it is considered that, at certain times of the year, the two forms are quite similar in coloration. Zapus pacificus alleni Elliot Allen Jumping Mouse Zapus alleni Elliot (1898, pp. 212-213). Type.—Female; Pyramid Peak, Eldorado County, California; collected by W. W. Price (fide Elliot, 1898, p. 212). Range.—Boreal Zone of the Sierra Nevada from Kern Peak, Tulare County, California, north to Mount Shasta; thence west and south through the Trinity Mountains to northern Trinity County. Also South Yolla Bolly Mountain, Tehama County. Number of speci- mens examined, 215, from the following localities in California: Tulare County : Jordan Hot Springs, 10; Kern Peak, 1; Red Rock Meadow, 1; Sequoia National Park, 1. Fresno County: Hume, 1. Mono County: Mammoth, 17 (coll. A. B. Howell) ; Mono Lake P. O., 4; Walker Lake, 5; Leevining Creek, 2; Williams Butte, 2. Mariposa County : Wawona, 1 (coll. Biol. Surv.) ; Merced Grove Big Trees, 6; Indian Cafion, 9. Yosemite National Park: Chinquapin, 12; Mono Meadow, 4; Lyell Canon, 8; Poreupine Flat, 9; Mount Hoffman, 4; Merced Lake, 5; Yosemite Creek, 9; Yosemite Falls, 8. Eldorado County: Fyffe, 1; Phillips, 5; (coll, .D. Ri: Dickey) ; Emerald Bay, 6 (coll. Biol. Surv.). Plumas County: Bucks Ranch, 5 (coll. D. R. Dickey). Shasta County: Mount Lassen, 3 (coll. Biol. Surv.). Siskiyou County: Mount Shasta, 12 (6 in coll. Biol. Surv.) ; Sisson, 1 (coll. A. B. Howell) ; Salmon Creek, 2; Rush Creek, 6; Bear Creek, 1; Grizzly Creek, 1; Jackson Lake, 1; south fork Salmon River, 7. Trinity County: north fork Coffee Creek, 32. Tehama County: South Yolla Bolly Mountain, 15. Diagnosis —A bright-colored Zapus, usually with a white tip to the tail. Typical skulls differ from those of pacificus in having the brain-case more inflated and the whole cranium with a more robust appearance. The interorbital width is greater, rostrum heavier with wider nasals, incisive foramina wider and not so constricted posteriorly, interpterygoid fossae slightly wider, and bullae smaller. 1920] Howell: A Study of the Califorma Jumping Mice 233 External characters—lIn coloration, typical allenic differs but slightly from pacificus; the sides are a shade brighter, and the dorsal area is usually darker, but not invariably so. The average length of foot in five topotypes of pacificus measured in the flesh by the writer is 32.2 mm., so that this member is not shorter than in allen, as was indicated by the type and cotype (Preble, 1899, p. 31). The tail of this form is slightly more hairy than is usual in the genus. In the spring and summer pelage, the normal coloration is a bright shade of ochraceous buff, and the dorsal area is well defined, with the black hairs predominating on the middle of the lower back, but fewer in proportion anteriorly, so that the top of the head is but little darker than the cheeks. There are two extreme types of coloration, with every degree between. In one, the dorsal area is very dark and the lighter hairs are almost absent, while in the other extreme, the black hairs are decidedly in the minority, giving a poorly defined dorsal area. This difference seems to follow neither season nor age. Immatures are usually lighter on the back than are adults, but not invariably so. Most of the skins taken during the last of August, and all of the September specimens (from the southern Sierras), have the general color tone, buff, with the dusky bases of the hairs showing through and fewer black hairs in the dorsal area. There are no examples taken later in the fall by which the significance of this change may be judged, but the pelage is then usually shorter, and as there is no evidence of new hairs, the duller shade may be due to fading, wear, and general devitalization of the old hairs preceding the change to the winter coat. Skull.—Typical alleni has the smallest bullae of any race in the state. Measurements.—Average, in the flesh, of 10 apparently adult specimens from Eldorado County, California: length, 228.3 mm. ; tail, 134.5; foot, 31.5. Average of 4 apparently adult topotypes of Z. p. pacificus: length, 235.3; tail, 140.7; foot, 32.8. Remarks.—Although alleni has usually been considered a sub- species of trinotatus, it seems to be definitely separated from that species geographically, and its characters certainly place it closer to pacificus. Many more specimens from the mountains of Washington and Oregon are necessary before this point can be considered settled, but the inference is that pacificus and trinotatus are specifically dis- tinct, separated as are their ranges by the topographical barriers pre- viously mentioned. It is possible that alleni is specifically distinct as 234 University of California Publications in Zoology [Vou. 21 well, as its range is separated from that of pacificus by the Klamath River gap. Although intergradation between these two races has not been demonstrated, they are so similar in many ways, and alleni is subject to such geographical variation within its range, that it seems better, for the present, to consider one the subspecies of the other. The segregated Boreal areas in which allent is found, comprise (a) the small cap of South Yolla Bolly Mountain in Tehama County, apparently separated by twenty miles or so of Transition Zone from (b) the Siskiyou Mountains and Mount Shasta, between which there is uninterrupted distribution, to some extent at least, through the Sisson region. This area is separated from (c) Mount Lassen by the Pit River gap, which in turn, is divided from (d) the main Sierran mass to the south by the Feather River gap, but the latter may or may not be a barrier to the genus (see Merriam, 1899, pp. 69-70). In certain respects, the skulls of examples from Mount Shasta show a tendency to approach pacificus (as in rostrum and _ nasals), but all that I have seen are distinct from the latter. It is unfortunate that the specimen from Little Shasta, which Preble (1899, p. 31) pro- nounced ‘‘not typical’’ pacificus, is not available, but the animal was immature, and I strongly suspect that it may present an appearance similar to a very pale juvenile which I took at Sisson. Average meas- urements, in the flesh, of 10 apparently adult specimens from Mount Shasta: length, 230.5 mm.; tail, 139.8; foot, 33.1. Skulls from the Siskiyou Mountains in Trinity County are large, with incisive foramina even larger than in typical alleni, and rostrum varyingly intermediate between that form and pacificus. Average measurements, in the flesh, of twenty apparently adult specimens from Trinity County: length, 231.8 mm.; tail, 140.7; foot, 32.4. Skulls from South Yolla Bolly Mountain, in Tehama County, show a very shght leaning towards orarius (small size, small rostrum, and short incisive foramina), as might be expected on geographical erounds, but they are definitely distinct from that species and clearly belong to the pacificus-allent group. The rostrum is slightly shorter than is typical, brain-case proportionally more inflated, incisive fora- mina shorter, pterygoid fossae smaller, and bases of the anteorbital foramina considerably heavier. The dorsal area is shghtly paler than usual and there is a yellowish tinge to the underparts of the specimens at hand, which darkens to brownish mid-ventrally in three skins. However, all of these were taken at the same time, and it is safer to infer, at least until more material is at hand, that the unusual tinge 1920] Howell: A Study of the California Jumping Mice 235 to the underparts is merely a fortuitous stain. Average measure- ments, in the flesh, of 10 apparently adult individuals from South Yolla Bolly Mountain, Tehama County : length, 231.6 mm.; tail, 139.4; foot, 33.2. Although there is apparently not the slightest climatic or topo- eraphical barrier between the type locality in Eldorado County, of alleni, and the Sierras to the south of it, specimens from the Yosemite region are not at all typical; and, what proves disconcerting, they vary slightly towards pacificus in some respects, having a narrow brain-case and incisive foramina, large bullae, and small pterygoid fossae. Average measurements, in the flesh, of 18 apparently adult specimens from the Yosemite National Park: length, 227.4 mm.; tail, 136.5; foot, 32.4. Still another variant is found in the Sierras of Tulare County, which is closer to typical alleni as far as small bullae are concerned, but the incisive foramina are even narrower than in Yosemite examples. The pterygoid fossae are very small, the pterygoid plates short, and the rostrum rather short and tapering, with the nasals of equal width throughout, as in pacificus. Average measurements, 10 the flesh, of 5 apparently adult specimens from the mountains of Tulare County : length, 220 mm.; tail, 132.4; foot, 31.2. All the cranial differences of the specimens from these several regions are rather subtle, but unmistakable in series. It might be justifiable, to recognize subspecifically one or another of these variants, but that could hardly be done without naming at least one, and prob- ably two other divisions of alleni, which would then be subspecies of pacificus. Such a course of action would be eminently out of place until suites from a large number of intermediate points are available, and skins of Z. p. pacificus are more numerous. Zapus major Preble Warner Mountain Jumping Mouse Zapus major Preble (1899, pp. 24-25). Type.—Female adult; no. 79983, Bur. Biol. Sury.; Warner Mountains, Oregon; August 4, 1896; collected by C. H. Merriam and V. Bailey; original no. 5720 (fide Preble, 1899, p. 42). Range.—High Transition and Boreal zones in the Warner Moun- tains of Modoe County, California, from Lassen Creek south to War- ren Peak. Number of specimens examined, 30, from the following 236 University of California Publications in Zoology [Vou. 21 localities in California. Modoe County: East Warren Peak, 1; Sugar Hill, 1; Parker Creek, 19; Dry Creek, 3; Goose Lake Meadows, 4; Lassen Creek, 2 (coll. A. B. Howell). Diagnosis.—Major is the largest species of Zapus in the state, and skins may be distinguished by the combination of white underparts and dull-colored sides. The skull differs from that of typical alleni in being heavier and slightly longer, with proportionally robust ros- trum, smaller pterygoid fossaec, and larger bullae and incisive fora- mina, the latter averaging less constricted posteriorly. This species differs from montanus in having a heavier skull, rostrum less tapering, incisive foramina larger, molar rows longer, basioccipital slightly wider, and bullae smaller. External characters—lIn the skins of spring and early summer, the general tone of the sides is warm buff, but this is modified to a slightly greater extent than is usual with California Zapus by the admixture of black hairs. Normally, the latter greatly predominate in the dorsal area, the boundaries of which are poorly defined, but a few of the adults have paler backs. The ears are dull black, faintly marked with lighter hairs, but in none of the specimens before me are these members edged with white, as is said to be the case with the type (Preble, 1899, p. 25). In August the skins are slightly duller in tone but are otherwise similar. Skull—Major has the longest skull, with heaviest rostrum, largest incisive foramina, and longest molar row of any form in the state. Measurements—Average, in the flesh, of 14 apparently adult specimens from the Warner Mountains, California: length, 231.9 mm. ; tail, 139.4; foot, 30.6. Average of 7 apparently adult topotypes of Z. montanus: length, 226.7; tail, 1386.4; foot, 32.7. Remarks.—The difference in coloration between major and alleni is so great that the two need never be confused. Specimens taken in August can hardly be distinguished as to color from montanus, but the hind foot of major seems to be a trifle heavier. In California, major has been taken only in the Warner Mountains of the extreme northeastern corner of the state, but it may occur in the few other suitable spots that are situated in the Modoe region. So far as I am aware, but the one animal has been secured at the type locality. 1920] Howell: A Study of the California Jumping Mice 9 o_ MEASUREMENTS (IN MILLIMETERS) OF APPARENTLY ADULT SPECIMENS OF THE CALIFORNIA ForMS OF ZAPUS Zapus trinotatus eureka 37 a = = = : 3|2|8_ (2 els a zg < | g | 2/58] 32 |i] ge |. | 2) = |=] 2 | Se/83| s5| sel fe s> |2 Locality - H Pe Gee |e eae men CI led 11693 | | Eureka, Humboldt Co. 242 .0)152.0/35.0/24.9)13.1|4.5]11.7/9.5)3.8 11696 | 7| Eureka, Humboldt Co. 233 .0/148 .0/34.0/24.1)12.9/4.5)11.6/9.2/3.8 11699 | #| Fair Oaks, Humboldt Co. /218.0/142.0/34.0/24.2)12.3/4.2/10.8/9.3/3.9 11703 | 9| Fair Oaks, Humboldt Co. [237 .0/148 .0/32.0/24. 1/13 .0/4.2/11.0/9.3/3.7 11704 | 9} Arcata, Humboldt Co. 220 . 0/137 .0/31 .0/23 9/12 .3/4.2/10.9/9.3/4.0 11705 | 9} Arcata, Humboldt Co. 234 0/141 .0/32.0)24.8)12.9/4.6)11.3/9.7/4.2 Average 230. 7/144. 7/33 .0/24.3/12.7/4.4/10.2/9 413.9 Zapus orarius 20288 | 9 | 3 mi. w. Inverness, Marin Co.|222.0/131 .0/31 .0/23 .3]12.0/3.9/10.8/9.0/3.8 20293 | 2| 3 mi. w. Inverness, Marin Co./230 . 0/137 . 0/32 .0/23 .3/12.0/3.8/11.0/9.0|/3.9 20294 | 9| 3 mi. w. Inverness, Marin Co.|249 . 0/148 .0/34.0/25 . 5/12. 2/3 .8/10.9/9.3/3.8 20291 | "| 3 mi. w. Inverness, Marin Co./223 . 0/128 . 0/31. 0/23. 7/12 .6/3.8/10.9/8.9|3.7 20292 | | 3 mi. w. Inverness, Marin Co.]222.0/130.0/30.0/23 . 9/12 .2/3.8/11.0/9.3/3.8 20295 | o7| 3 mi. w. Inverness, Marin Co./225 0/121 .0/30.0/24.1/12.0/4.1/10.6|9.5|3.7 Average 228 . 5j137 . 5/31 . 3/23 . 8/12. 2/3 .9}10.9|9.2/3.8 Zapus pacificus alleni 88267!| | Emerald Bay, Eldorado Co. |222.0/132.0/31.0/24.0/12.3/4.8/10.6/9.5|3.7 88268!| "| Emerald Bay, Eldorado Co. |245 0/151. 0/33 .0/25.2/12.8/4.9/11.4/9.8/3.7 88276} o'| Emerald Bay, Eldorado Co. |236.0/148 .0/32.0/24.6]12.4/4.3/11.1/9.5/3.9 882781] «| Emerald Bay, Eldorado Co. |226.0/134.0/30.0/24.0/12.7/4.3/11.0/9.4/3.7 88281! «| Emerald Bay, Eldorado Co. |227.0/138.0)/34.0/24.1/12.4/4.5/11.1/9.6/3.8 Average 230. 1/139 6/32 .0/24.4/12.5/4.6/11.0/9.6|3.7 Zapus major 11270 | | Parker Creek, Modoc Co. /221.0/133.0/30.0/24. 7/12 .6/4.6/11.2/9.5)/4.5 11282 || Parker Creek, Modoc Co. |227.0/138.0/30.0)25.3/12.8/4.5/11.1/9.7/4.3 11273 | #| Parker Creek, Modoc Co. |226.0)137.0/31.0/24.9)12.6/4.5)11.0/9.5/4.2 11272 | 9) Parker Creek, Modoc Co. /|240.0/144.0/30.0/25.4/12.7/4.6]11.2/9.8/4.4 11288 | 9} Parker Creek, Modoc Co. = |250.0/150.0)32.0/24 9/12 .8/4.8/11.3/9.6/4.5 11271 | | Parker Creek, Modoc Co. (|248.0/146.0/33.0/25.4/12.5/4.6/11.1/9.6)/4.5 Average 235 3/141 .3/31.0/25.1/12.7/4.6]11.1/9.6/4.4 1From U. 8S. Biological Survey collection. 238 University of California Publications in Zoology [Vou. 21 LITERATURE CITED ELuiot, D. G. 1898. Lists of species of mammals, principally rodents, obtained by W. W. Price ...in... California .... Field Columb. Mus., zool. ser., 1, pp. 193-216. MERRIAM, C. H. 1897. Three new jumping mice (Zapus) from the northwest. Proe. Biol. Soc. Wash., 11, pp. 103-104. 1899. Results of a biological survey of Mount Shasta, California. U.S. Dept. Agric., Div. Biol. Surv., N. Amer. Fauna, 16, pp. 1-179, 5 pls., 46 text figs. PREBLE, H. A. 1899. Revision of the jumping mice of the genus Zapus. U. 8. Dept. Agric., Div. Biol. Surv., N. Amer. Fauna, 15, pp. 1-42, 1 pl., 4 text figs. Rimeway, R. 1912. Color standards and color nomenclature. (Washington, D. C., published by the author), pp. 8+ 44, 53 col. pls. Transmitted March 15, 1920. 2 9. 10, aie 12. 138. 14. x 15. 16. 17. 18 Vol. 18, 1. 2. 3. 4, 5. 14, 15, UNIVERSITY OF CALIFORNIA PUBLICATIONS — (Continued) Notes on the Systematic Status of the Toads and Frogs of California, by Charles Lewis Camp, Pp. 115-125, 3 text figures. February, i hk? BE pecs A Distributional List of the Amphibians and Reptiles of California, by tpg ees and Charles Lewis Camp, Pp. 127-208, 14 figures in text. pilin: peers WB RY RGU ais Son epi Sree POU esta Wim eS SOME a Cone a A Seley ion ety e, se A Study of the Races of the White-Fronted Goose (Anser albdifrons) ‘oceur- ring in California, by H. S. Swarth and Harold C. Bryant. EDs 209-222, 2-figures-in text, plate 13.”; October, -1917: 334 ee ee A Synopsis of the Bats of California, by Hilda Wood Grinnell, Pp. 293-404, plates 14-24, 24 text figures. .January 31, 1918 .22.2.20 ee The Pacific Coast Jays of the Genus Aphelocoma, by H. 8S. Swarth, Pp. 405-422, 1 fignre-in text.. February 23,°1918 <2..2.20 Se Six New Mammals from the Mohave Desert-and Inyo Regions of California, by Joseph Grinnell, Pp. 423-430; — - Notes on Some Bats from Alaska and British Columbia, by Hilda Wood Grinnell, Pp. 431-483. Nos. 14 and 15 in one cover. April, {91g pS RAAT SAE ES Se AR SS Fe Revision of the Rodent Genus Aplodontia, by Walter P. -Taylor. Pp. 435- 504, plates 25-29, 16 text -figures.. May, 1918 ........ SS SPSS dg SS. AA age The Subspecies of the Mountain Chickadee, by Joseph Grinnell. Pp. 505- GAGS PORE OUT ESS: TEA Ys O38 oe Te ee are es ok ee ee Excavations of Burrows of the Rodent Aplodontia, with Observations on the Habits of the Animal, by Charles Lewis Camp. Pp. 517-536, 6 figures te CORA DUNC S LOTR. ee es re Oe CS Be Index, pp..537-545. Mitosis in Giardia microti, by William ©. Boeck. Pp. 1-26, plate 1. Octo- ihe 1A YSZ ite cares Soe etd cede eee sine ae eee Sea ee ee: A a Pe aw RO An Unusual Extension of the Distribution of the Shipworm in San Fran- . cisco. Bay, California, by Albert L. Barrows. Pp. 27-43. December, 1917. Description of Some New Species of Polynoidae from the Coast of Cali-: fornia, by Christine Essenberg. Pp. 45-60, plates.2-3. October, 1917 .. New Species of Amphinomidae from the Pacific Coast, by Christine Essen: berg. Pp. 61-74, plates 4-5. October, 1947 2.0020 ese lie eked Crithidia euryophthalmi, sp. nov., from the Hemipteran Bug, Hijo ophibaleaie convivus. Stal, by Irene McCulloch. Pp. 75-88, 35 text figures. Decem- GR OT Tc oe ON re Sone Sees an ae eae On the Orientation of Erythropsis, by Charles Atwood Kofoid and Olive Swezy. Pp. 39-102,.12 figtires in text. December, 1917 __-.......08. cos : . The Transmission of Nervous Impulses in Relation to Locomotion in the Earthworm, by John F, Bovard. Pp. 103-134, 14 figures in text. J anaeys BB A: Sake mats Bc aN) Sec Nap YODER GOR BR a er ny Re STONERS SSO apes Pn Re en ee ROR a tee . The Function of the Giant Fibers in Barthworms, by John F. Bovard. Pp. 135-144, 1 HEUTE-In BERT. sd) AU ATY, ALG oe sooo adda acca nace aaicsoeetedendes A Rapid Method for the Detection of Protozoan Cysts in Mammalian Faeces, by. William C. Boeck. Pp. 145-149, December, 1917 .........22.... The Miusculature of Heptanchus maculatus, by Pirie Davidson... Pp..151-170, 27 AD -fiouresAttext.s Wiarchy, VOLS ees ios aa sites ce sacs ieee cp cece aaeaet dace The Factors controlling the Distribution of the Polynoidae of the Pacific Coast of North America, by Christine Essenberg. Pp. 171-238, plates 6-8, Qofivures a eS a ~ 10 -9. A Rapid Method for the Detection of Protozoan Cysts in Mammalian - -Faeces, by William C: Boeck. Pp. 145-149, December,:1917 0.02... .05 - 40; The Musculature of H eptanchus maculatus, by Pirie Davidson. Pp. 151-170, to Heyresin text, Mrareh: AOL rc ee ae aan at cose ce oa Machen amos ED: 11, The Factors Controlling the Distribution of the Polynoidae of the Pacific Coast of North America, by Christine Essenberg.. Pp. 171-238, plates 6-8, & figures in tekst, 2 March sy VOLS ec ee ec se sone target aoe Sested (3 Sheer ; 12. Differentials in. Behavior of the Two. Generations -of* Salpa democratica yates Rinks - Relative to the Temperature of the Sea, by Ellis ‘L. Michael. Pp. 239-298, Ras plates-9-11; 1 figtre in text=March,*1918 4422 so ee. .65 es, 13. A Quantitative Analysis of the Molluscan Fauna of San Francisco Bay, by E. L. Packard. ‘Pp. 299-336, plates 12-13, 6 figures in text. April, 1918 =. —. .40 S 14. The Neuromotor Apparatus of Luplotes patella, by Harry B. Yocom. Pp. 337-396. plates. 14-16: September, 1918 <2). oh eek 570 mes 15, The Significance of Skeletal Variations in the Genus Peridinium, by A. Ts, Noe Barrows. Pp. 397-478, plates 17-20, 19 figures in text. June, 1918 _...2...... .90 16. The Subclavian Vein and Its Relations in Elasmobranch Fishes, by J. Frank : Daniel. Pp. 479-484, 2 figures in text. August, 1918-022 W2282.--e 10 17. The Cercaria of the Japanese Blood Fluke, Schistosoma japonicum Katsu- rada, by William W. Cort. Pp. 485-507, 3 \figures in text. keen 18. Notes on the Eggs and Miracidia of the Human Sree ORCEeS, by William ieee : W, Cort. Pp. 509-519, 7 figures in text. Ber ots Nos, 17 and 18 in one cover. January, 1919 fie ast So ipee ice, tee 35 Se : Index, pp. 521-529, UNIVERSITY OF CALIFORNIA. PUBLICATIONS—(Continued) Vol. 19. A fs Reaction of Various Plankton Animals with Reference to Their Diurnal Ki ._ Migrations, by. Calvin O. Esterly. Pp. 1-83. “April, 1919 222. te 2. The Pteropod Desmopterus pacificus (sp. nov.), by. Christine Essenberg. “Pp. Pea tie 85-882 fignres:in text. “May, 1919-2 so er a ee oe ee ea es SOR a 3. Studies on Giardia microti, by William C,: Bosch. ‘Pp. $5- 136, plate 15° 19 F Ya Some eh ba figures In texts: Apri OE Sapa cca ate Reve asap nce cheecdeh uotaeaa teas GOS ey 4. A Comparison of the Life. Cycle of Crithidia with that ‘of fr "ypanosoma in. ena a. i the Invertebrate Host, by shone McCulloch. Pp 135- 190, plates:2-6, 3° ‘ fitures in text." October; D919 c.25 550 Po Ree i eee ee ee ae a OG er aes 5. A Muscid Larva of ‘the San Francisco Bay Region Which ‘Sucks the Blood Aa gt Owe - ef Nestling Birds, by 0. HE. Plath, Pp. 191-200. February, 1919 =... Sar AS ase? Fe 6. Binary Fission in Collodictyon. triciliatum Carter, by Robert Clinton Rhodes. Bip Pe tale ss Pp..201-274, plates 7-14; 4 figures in text.. December, 1919 (000.2228... “$100 ss See 7. The Excretory System of a Stylet Cercaria, by William W. Cort. Pp. 275- 7h, SNCs eee Si, 3 figure in text. Angust, 1919 = one ea oe a eet a TOs ier eae 8. A New Distome from Rana aurora, by William W. Cort. Pp. 283-298, bs artery tant oh figures in: text. November, 1919 28 oss ae ee Me ie RE aes Bere a eS) 9. The Occurrence of a Rock-boring Isopod along the Shore of San Francisco~ Bay, California, by Albert L. Barrows. Pp. 299-316, plates 15-17. De- SS ee tet GOIN DORs 3 LOT Re ey Ra NO See ak Te Shep a IE cha aE Ns cok ate teee ogee ae SDB: Aes ate 10. A New Morphological Interpretation of the Structure of N. ocliluca, and Its sR Ae ve ag ~ Bearing on the Status. ofthe Cystoflagellata | (Haeckel), by. Charles” A. Ta ieee ay | Kofoid. Pp. 317-334, plate 18, 2 figures in text. February, 19207 2:2... Sate ee ee 11. The Life Cycle of Echinostoma revoluium (Froelich), by John C, Johnson. _ ee Pp. 338-388, plates 19-25, 1 figure in text. May, OU Shh Sti eae eee Bea its 12. On Some New Myriopods Collected in India in 1916 by C. A. Kofoid, by ..= = =. Ralph V. Chaniberlin. Pp. 389-402, plates 26-28.° August, 1920 222.0... 5 Qe Scere 13.. Demonstration of the Function. of the Neuromotor Apparatus in Euplotes : by the Method of Microdissection,: by Charles V. Taylor. Pp. 403-470, plates 29-33, 2 figures in text. October, 1920 2 ee es 86. Se ze Index in preparation. * apes patie Vol. 20. 1. Studies on the Parasites of the Termites. I. On Streblomastia striz, a Poly-- ©. = gr mastigote Flagellate with a Linear Plasmodial Phase, by Charles Atwood ae ee Kofoid and Olive Swezy.- Pp. 1-20, plates 1-2, 1 figure in text. July, 1919. 25 | 2. Studies on the Parasites of the Termites. II: On Trichomitus termitidis, a Polymastigote Flagellate with a Highly Developed Neuromotor System, — by Charles Atwood Kofoid and Olive Swezy. Pp, 21-40, plates $-4, 2. « 0. figtres in text. cFUly; 1910. eee eR) a as eee BT eel posh y 3. Studies on the Parasites of the Termites. III. On Trichonympha Scanpanate iad sian tena sp. nov., by Charles Atwood Kofoid and Olive Swezy. Pp. 41-98, plates ree ee tee 5-12, 4 figures In PExt.. 2 SUV Ole hee Ae ey eS ee a ek St ee ee 4. Studies on the Parasites of the Termites. IV. On Leidyopsis sphaerica gen. Shen nov., sp. nov., by Charles Atwood Kofoid and Olive Swezy. Pp. 99-116,.55.. eee plates 18-14, 1 Hopre-4n-bexh.*.d tly; AGT: hs oe a OR oes ee eee ee 5.-On the Morphology and Mitosis of Chilomastiz mesnilt (Wenyon), a Comino: ok em ane Flagellate of the Human Intestine, by Charles A. Kofoid and Olive Swezy. ng aerials Pp. 117-144, plates 15-17, 2 figures.in text. April, 1920 2W oe ASO ee 6. A- Critical. Review of the Nomenclature of Human Intestinal Flagellates, Paani Sea Cercomonas, Chilomastiz, Trichomonas, and Giardia, by Charles A. Kofoid: = = © Pp. 145-168, Sfisures im text.— Tune, 1920) 2 ne eee Mepis Yorn y, os 3 7. On the Free, Encysted, and Budding Stages of Councilmania lafleurt, a Para--—- sitic Amoeba of the Human Intestine, by Charles Atwood Kofoid and Olive Swezy. Pp. 169-198, plates 18-22, 3 figures in text. . June, 1921... .60 1. A Revision of the Microtus califarnicus Group of Meadow Mice, by Reming. 22 pee os ton Kellogg. Pp. 1-42, 1 figure in text. December, 1918 _..:. SiR Ves es SS S3BO. Sitcoms 2. Five New Five-Toed. Kangaroo Rats from California, by J beeen Grinnell. ior tot Pp.-48-47 6 larch, DONO oi es a A cer ahaa a ee aR pecs ata ‘OBR Fagus 8. Notes on the Natural History of the Bushy-tailed Wood Rats of Californias... ee, 4 Vol. 21. by Joseph Dixon. Pp. 49-74, plates 1-8, 3 figures in text. December, 1919 25°. . Revision of the Avian Genus Passerella, ‘with Special Reference to the Dis- =< tribution and Migration’ of the Races in California, by H. §. Swarth. SPP + 75-224, plates 4-7, 30 fignres in text. September, 1920) 20. ccc A Study of the California Jumping Mice of the Genus Zapus, by A. ‘Brazier Howell. Pp. 225-238, 1 figure in text. May, DOA ie Sk Sn CEES ey : Two New Rodents (Genera Thovomys and Marmota), from the faster Border of California, by Joseph Grinnell. Pp, 239-244; 6 BENE in-text. APOVEMID EL, LODE kal Ps ra Ra la es gt n a ae a Vol. 22. 1. A Quantitative and Statistical Study of the Plankton of-the San Joaquin River and Its Tributaries in and near Stockton, California, in-1913, by Winfred Emory Allen. Pp. 1-292, plates 1-12,.1 figure in text. June, 1920.. $3.0 00. Vol. 23. 1. The Marine Decapod Crustacea of California, by Waldo L. ‘Schmitt. . Pp. . 1-470; plates 1-50, 165’figures in ‘text... 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Spelerpes platycephalus,.a New Alpine Salamander from the Yosemite National Park, California, by Charles Lewis Camp. Pp. 11-14. Reptens ber, 1916 22... a aBiagOe -t SIME ah ey ee a ed SY Lap SOMME SESS eR ae i SR 7B AES 05 4. A New Spermophile from the San Joaquin Valley, California, with Notes : on Ammospermophilus nelsoni nelsont Merriam, by Shoah as P. Taylor. Pp. 15-20, 1 figure in text... October, 1916 22.02 ee 05° : Habits and Food of the. Roadrunner in California, by Harold C.. Bryant. on . Description of Bufo canorus, a New Toad from the Yosemite ‘Wational Park, a o2) . The Subspecies of Sceloporus occidentalis, with Description of 2 New Form from the Sierra Nevada and Systematic Notes:on Other California Lizards, . .. by Charles Lewis Camp. Pp. 63-74. December, 1916 2.2.22... +10 8. Osteological Relationships of Three Species’ of Beavers, by F, Harvey : 9. Notes on the Systematic Status of the Toads and Frogs of California, by. Charles Lewis Camp. 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March, 1917 -....-....... 40... i UNIVERSITY OF CALIFORNIA PUBLICATIONS - IN ZOOLOGY Vol. 21, No. 7, pp. 245-274, 25 figures in text April 18, 1922 A STUDY OF THE CALIFORNIAN FORMS OF THE MICROTUS MONTANUS GROUP OF MEADOW MICE BY REMINC '¥ J KELLOGG (Contribution from the Museum of Verteorate Zoology of the University of California) INTRODUCTION The present paper is a further report upon the writer’s studies on the meadow mice of the genus Microtus in California. The first report consisted of ‘‘A revision of the Microtus californicus group of meadow mice’’ (Kellogg, 1918). The material upon which the paper now offered is based consists of three hundred and sixty-seven speci- mens, representing sixty-three localities within the state of California, and further series of specimens bearing upon the problem from outside this state. Microtus montanus, under the name Arvicola montana, was first made known to science by Titian R. Peale, its discovery being one of the results of his explorations through northern California. The specific name he applied to the mouse was doubtless in allusion to the nature of the type locality, which was near the headwaters of the Sacramento River close to Mount Shasta. Within the state of Cali- fornia the species ranges zonally from Transition to Hudsonian, each inclusively. It is apparently restricted to alpine meadows in the Sierra Nevada and the Trinity Mountains, and to the lowland marshes of the Modoe region, in the northeastern part of the state. Since the discovery of Microtus montanus, several closely related forms have been found, the group thus constituted ranging far beyond the con- fines of California. One race has recently been described from so far distant as Wyoming. Altogether, six forms of this group are recog- nized, of which three, the ones occurring in California, are treated in considerable detail in the present paper. 246 University of California Publications in. Zoology [Vou 21 DIAGNOSTIC CRITERIA The diagnostic criteria that appear to be best adapted for dis- tinguishing the several races of Microtus montanus have been deter- mined by assorting the various characters into certain categories, and ascertaining the mean manifestation with respect to each character. It is a conspicuous fact that the three races in question do overlap to a considerable extent in respect to many characters. Specimens of this group frequently cannot be assigned to any particular sub- species unless the locality at which they were collected be known. The diagnoses of the crania describe the average, or the mean, of all individuals examined of any particular race rather than any particular type or individual. Color, in the Microtus montanus group at least, is too unreliable a feature to be of much use in separating the different races in California. Therefore the color terms used in the following descriptions relate to extremes, not to averages; it 1s impossible to designate any particular shade as the average, because there is such wide variation in color in each race. Furthermore, many of the shades to be seen in any one subspecies can be matched in individuals of a different form. VARIATION There is evidently a wide range of individual variation within each subspecies with respect to color, external measurements (1e., total length, length of tail, length of hind foot, and height of ear above crown), enamel pattern of the molariform series, and configuration of the skull. The variation within a given subspecies is frequently as great as the differences between the means of two subspecies. In view of the emphasis laid on cranial characters as a means of differentiating closely allied subspecies, it is well to point out the sort of variation that takes place in the configuration of the skull and in the form of the separate bones. In general, the contour of the brain ease, the relative size and shape of the interparietal and frontals, and the development of the interorbital ridges are altered materially as the animal reaches maturity; and as old age approaches, the ridges and tuberosities, the angulation of the zygomata, and other peculiar- ities of the skull become more and more accentuated. In young individuals the occipito-nasal length is usually greater than the con- dylobasal length; as the animal reaches maturity and approaches old age, there is a gradual shifting forward of the brain case until finally the reverse is often true. In skulls of young individuals, the brain 1922] Kellogg: Californian Forms of Microtus Montanus 247 ease is disproportionately large in comparison with the rostral and palatal regions of the skull. Certain variations occur in skulls of the same sex and of approx- imately the same age among mature individuals of the same subspecies. There appear to be but few elements of the adult skull that are con- stant, either in form or in relative size as compared to other parts. Thus the interparietals are extremely variable in appearance. The posterior border may be nearly straight or may exhibit many degrees of convexity, from a type that is gently convex to another in which it is nearly acutely convex. The lateral terminations may be either straight, rounded, emarginate, or sloping, and the median projection is variable with respect to size and to length. The posterior termina- tions of the nasals may be attenuate, emarginate, truncate, or even rounded. The interorbital ridges vary from a type in which there is a distinct suleus between them to one where but a single interorbital erest is discernible. By selecting specimens near the same age, thus eliminating those variations which are the result of different stages of growth, a fairly reliable series was obtained for comparative study. Specimens of Microtus montanus montanus from Sisson, California, old and young adults, were chosen to illustrate in graphic form (see tables I, II, and III) the extent of variation that might be expected in any par- ticular locality, although, of course, some age difference is also repre- sented. In general, total length indicates the age very closely, as a eritical study of the skulls shows that the oldest individuals are, as a rule, the largest. Table I shows that there is little or no correlation between length of tail and length of hind foot. This table probably contains inacecu- racies. The specimens were obtained and the measurements taken and recorded by several collectors, employing different methods. The animals themselves were by no means all in the same condition, some being freshly killed, in some rigor mortis had set in, and some had been dead so long as to be altogether relaxed. The data for tables II and III, however, were prepared by one person, the writer, and he was carefully consistent in the methods employed. It will be observed from table II that the mastoid width increases very little with general growth. The condylobasal length, as pre- viously noted, increases during growth in correlation with changes in the cranium. However, no such correlation is apparent between the types of enamel folding exhibited by the second upper premolar and the fourth lower premolar (see table III). University of California Publications in Zoology [Vou. 21 248 GLT-OLT “WOTDEI[OD AVAING [RoIsOoIg oy} WOT 218 S,000'86 94} UL perequinu [][e pus :ADOTOOT ayvIGo}I9A JO WINES] IY} WOIZ 01 S,QOO'TS OY} UI peroquinu [we !uOTZdaT[Oo [JaMOF “g “VY EY} UIOIZ d1B 8.00% AY} UT periequinu susuttoads [Ty “BIusoyeD ‘AquNOD NOATYsIG ‘UOSsSIg UIOIJ oI SOTqe} BUIMOT[O} Pue Sty} UT pasn sueUTIOads Oy} [Ty “suOTydvo Aq payeorpuUT sv syUdUIEINsvoUT 9014} 07 SUIPI090B8 UStUIO9ds YOu JO UOT} BOOT[V YIM SJUIUIAINSVIUT [VUIO}X9 0} BUIPIOIV snUDJUOW SNUDJUOW sNjowLPY JO S[BNPIAIPUT UT UOTPBIUBA ZULMOYS oq, Foy peeoeaes| cea cere beeen [eect FG ice BG oer ecedes SopoSRaaBA| Ee RpaRbaee| Hepatonees oe ride pa een | oe cee cere | cee eeacree ere BE eso | meters saeteeapersennecnsnel| Spry =| ssteareene|eoaeenn ot naterean Seteane TZ [oe een e| ge petespeere node this leeecosee TUG eco pence pence ecco ance enact PEELE | Reenee Te [ooops Sa acnauono| FaBrddbee! Kadena ree Ge cera ocdace side Pao fee eeeancs tier oe echeen| ace Retbrnsdlneco sess TUCe Nees eg | eee | eed oe BSAECESH Beard0ce| eeenese| Seeeonbr GI oe eteete pores Spartan ses cuneel tareBeee (oe ee ae OG [ores eettefeetteefeeeeeeeeteeefeoeeeees 0¢ hareriBod| |seeebor: es) ee So nOOG ZG [oe fcesttefeeeetcetfeteretetceee eoeeeene ecensceas [eceaerenebeneennnen Reanebes Feige Poo ee coat aco ect cece portres tr aro rey aoa cea Ee CO CS 0g Seta Ae ee en eG meson rae lt oe ea aka Racin eae bets anaes Da og Og foe |ceeefeeeeee fesse eeeteeeeeeefeeeeeeeeeceneenee LP eect notte TG [oc fee feet feceeeseesfeseetees [eseeeeeenal cece teens €% | G3 | 16 | 02 | 6I LG-G¢ | PS:¢G | 1S:6h | 8h: 9F | SF EF yoo} purzy IBIQIJIIA [IC], GF: OF S61:98T S8T-O8T | 6ZT:9LT 69T:99T | SOT:O9T yyBuey] [e301 6ST: 0ST © 68986 5 8986 © 08986 5 19986 5 16986 5 98986 © FE0G © §60% 5 8986 5 SZL86 5 F6986 Y 1F0Z Y 61986 © ZE0Z © 06986 © F00Z © 960Z © G00Z 5 €00z © 180% ‘ON uinesnjl I A@TaAVL a 1922] Kellogg: Californian Forms of Microtus Montanus 249 TABLE II Height of Condylobasal Zygomatic cranium at Mastoid length width bullae width Museum No. Sex 26 27 28 14 15 16 9 10 11 12 98698 ot 49}-(0) |} cco ae ses ice fil U5) 274 eee ee Oe: See | P4624 2004 fof PROPS ||) se ane aa a (29) 0 a eae ea LONG 12.5 2032 rot PAB MN woe are Pega aillsereois eee Pee LOR Feet | alpzae! 2037 fot KD) st) || aoe es SA ei eee. Ren pee i al Opes al ea ta Oe as 2036 fof PRO) sth] see ae sae 6 9.6 WA see eee M Li Redbook teal vere tal Tf eceme aaL a 1271 2041 rot Px etS) |) ears ae pa nate ee co || LOs@ Ae piso 2028 of PAD 5 08) a] | dane mn sooo || dhe 10.0 soa 2033 rot DH ily LOR ee | hilpaes 2034 of DW oe 15 1 So AOr Ste ig Ee 98684 9 27.5 115)308) |), fence OF OR meet ey pallet 98697 Q Die Cnr net zee |) does en Ons Je Nelle aees) 98680 ot DEMO MW! eset soe | aang on) LOLS coe. |p ed!) 98651 je) MM eee won NG LORS eee le2e0 98686 fe) Pape MW ieee Lge Seo: OOO |e eee: bead gaa evah 98682 ie) BRSEY IY Gane sao || hOee} Oe: Pe Table showing allocation of the skulls according to the different cranial elements selected for measurement; all specimens from Sisson, Siskiyou County, California. DENTITION The writer’s attention was first directed to the wide variation in the enamel pattern of the molariform series in the genus Microtus by Miller’s (1896, p. 26) figures of the variation in enamel folding in Microtus pennsylvanicus. Miller estimates that 75 per cent of the specimens possess a normal enamel pattern, though he illustrates with diagrams eighteen types of variation in both the M2 and Pm; which were present in one hundred and seventy specimens of that species. This variation is considerably greater than has been currently assumed, and will be discussed somewhat fully. The author is indebted to Mr. Gerrit S. Miller, Jr., for advice as to the nomenclature to be employed for the enamel crown divisions of Microtus, but the writer himself is to be held responsible for any errors he may have made in interpreting Mr. Miller’s remarks as to the homologies of the elements of the enamel pattern. Numerous attempts have been made in the past to assign certain symbols or names to the divisions of the molar crowns of rodents, but there has been much disagreement as to the homologies of the divisions con- cerned. Among the first to adduce specific evidence in support of their findings were Winge (1883, pp. 67, 68-69) and Forsyth-Major 250 University of California Publications in Zoology |Vou-21 (1897, pp. 695-720). Winge advocated the use of numerals to desig- nate the various molar crown divisions. For the following translation of Winge’s views the writer is indebted to Dr. A. Boving, of the Bureau of Entomology, U. S. Department of Agriculture. The three primitive cusps, which are innermost on the teeth of the lower jaw, and outermost on the teeth of the upper jaw, are designated as 1, 2, and 3; the cusp marked 2 is the oldest, corresponding to the single cusp of the incisors and canines and to the middle cusp on the two anterior molars. The two more recently added cusps, the outermost on the teeth of the lower jaw, the innermost on the teeth of the upper jaw, are designated as 4 and 5. The original single heel of the teeth in the upper jaw is designated as 6, the more recently added enlarge- ment of the heel is designated as 7 (Winge, loc. cit., p. 67). On the two following pages Winge (loc. cit., pp. 68-69) gives a further explanation of his views on this subject : Synopsis of the changes of anterior molars in some mammals. The three original cusps, innermost in the lower jaw, outermost in the upper jaw, are designated as 1, 2, 3; the two more recently added cusps, outermost im the lower jaw, innermost in the upper jaw, are designated as 4~~5; the heel on the teeth of the upper jaw is designated as 6 whenever it is single; as Ga whenever it is doubled by the addition of a new cusp; as 6+7 when the two are fused or when one of them has disappeared. In his ‘‘Mammals of Denmark,’’ Winge apparently does not adhere to some of his early views. In this work (1908, p. 77), cusps 4 and 5 are shown as being outermost in the teeth of both upper and lower jaws of Microtus arvalis, and according to these figures his numbers for the cusps would have the following equivalents: In the upper molars, 4 and 5 are the paracone and metacone, respectively, and 6 and 7 the protocone and hypocone. In the first lower molar, 2 and 3 are equivalent to the metaconid and entoconid, and 4 and 5 are equivalent to the protoconid and hypoconid. Winge considered that his cusp ‘‘6’’ was a later addition and that his cusps ‘‘1, 2, and 3’? (=parastyle, mesostyle, and metastyle, according to Barrett- Hamilton, 1914, pl. 28) were the oldest. Barrett-Hamilton (loc. cit.) makes the statement that cusp ‘‘2’’ is present in some of the American Cricetinae, while in all the Microtinae this cusp is fused with ‘‘4’’ (== paracone), and thus has completely disappeared. He held, fur- thermore, that the lower molar series was the inverted image of the upper. The considerations which were advanced by Fleischmann (1891, p. 891) and Mahn (1890, p. 652) to show that the anterior end of the lower molars is homologous with the posterior end of the upper ones and vice versa, have been refuted by Scott (1893, pp. me ES pn ee 1922] = Kellogg: Californian Forms of Microtus Montanus 201 410-417). The evidence adduced by Mahn was derived from the resemblance in form of the second upper molar to the fourth lower premolar, and from the assumption that in Arvicola the position of the cement bands on the teeth indicates the homologous crown divi- sions. Forsyth-Major (loc. cit.) also used numerals in his studies on various groups of rodents. Most of the crown divisions in the molars of Oryzomys as figured by Goldman (1918, p. 11) are present in the hypsodont teeth of Microtus. pm, Bin m! Mm; 2 m mM ht Figs. A, B. Diagrams of enamel patterns of left lower and left upper molari- form series of Microtus montanus yosemite. X 6. Mus. Vert. Zool., no. 22365, 3g; Tuolumne Meadows, Yosemite Park, California. Explanation: Pm4, fourth premolar; M1, first molar; M2, second molar; end, entoconid; esd, entostylid; hy., hypocone; hyd, hypoconid; me., metacone; med, metaconid; mesd, mesostylid; ml., metaconule; mts., metastyle; pa., paracone; pad, paraconid; pl., pretoconule; pr., protocone; prd, protoconid; pred, protoconulid ; ps., parastyle; sec. ml., secondary metaconule; sec. pad, secondary paraconid; sec. pred, secondary protoconulid. It is not the writer’s intention to take up any theoretical discussion involving the origin of the various cusps, for that can be accomplished better by some one having more material, especially fossil material, at his disposal. However, there are one or two points that might well be mentioned in connection with the present diagrams (figs. A to D). The entire loop anterior to the protocone in fourth upper premolar, which was designated by Winge as ‘‘1’’ (= parastyle), cannot be designated as that cusp with any degree of certainty because of its direct connection with the protocone. The parastyle is a direct off- shoot of the paracone. It may be possible, however, that the parastyle loses its connection with the paracone and unites secondarily with the protocone. The data which are available relative to the sequence and 4 252 University of California Publications in Zoology [Vou 21 manner of the addition of cusps in other groups of mammals make such an interpretation seem improbable, though it must be admitted that at present we do not have definite evidence as to the formation of these loops. It is more probable that the internal portion, at least, of the anterior transverse loop of the fourth upper premolar repre- sents the protoconule, but even in that case its position is very unusual. pm* PM 4 m! m, m2 ie Figs. C, D. Diagrams of enamel patterns of left lower and left upper molari- form series of Microtus californicus californicus. 7. Mus. Vert. Zool., no. 3655, g; Palo Alto, Santa Clara County, California. Explanation: Pm4, fourth premolar; M1, first molar; M2, second molar; end, entoconid; esd, entostylid; hy., hypocone; hyd, hypoconid; me., metacone; med, metaconid; mesd, mesostylid; ml., metaconule; mts., metastyle; pa., paracone; pad, paraconid; pl., protoconule; pr., protocone; prd, protoconid; pred, protoconulid; ps., parastyle; sec. ml., secondary metaconule; sec. .pad, secondary paraconid; see. pred, secondary protoconulid. In many rodents having a hypsodont type of dentition, the anterior deciduous molar continues its function of slicing and shearing while the permanent molars are coming in. Evidence bearing on this is afforded by the beaver, Castor. The view that this anterior molari- form tooth is a deciduous molar is also held by Forsyth-Major and Martin A. C. Hinton, the latter basing his view upon the extraordinary complexity of the anterior cheek teeth and upon various theoretical considerations. Support is also lent to this view in an instance recorded by Winge of the occurrence of a small fourth posterior cheek tooth in Microtus agrestis. If this anterior molariform tooth is the deciduous molar, it would necessarily possess more primitive characters than the permanent 3 1922] Kellogg: Californian Forms of Microtus Montanus 25 molars, and for that reason the internal portion of the anterior trans- verse loop would be the protoconule instead of the parastyle. The protoconule is a cusp which is present in the early stages of develop- ment of the mammalian molar crown, whereas the parastyle is a later addition, or specialization. TABLE III Types of Types of second upper molar fourth lower premolar K L M N O P E F G H I J PAV ASNC RN Rik ers hale eeeh 1 || mabe eian Ill eoeeta Ga ree Pearce dl | eaeteies| Wee Fee (MASON Bey en ee en ies, coer tins Menara Pid Wsccls [Eine a eraceadl eet ene leak Gel eee aeepioael be eee acme US EON Re ne as arene anc sanctal eaeeenearie sCal( ee sec cnmeal aa ed C7) [et lence gees pp cel rey PAV SS oil Ua atin aap pins) Uber cee eens] [n geamae ieee bes ale al Uae a a Sea |Past | Ral oral PADD SS Gil saeetccare te lNeecestusbee Phoceestetiscentel tee reecccrsel seme neat Dal eel bine ane |e PAD Ye SIL GS see eae sh Ee ce al Va fl | ( S| | eres | eon ater [cereal Ee oka PLO OV Oalee emer cade lt isdes terra |PRELne cecal Cece eo ets ell keecee D daa Beet Stbe| eaekich a mee tel hg einen PAD SYSNG IN] Bie mea eet eeesee aI lar Leen QR Re ee Salle (ee le eae Se ec] We Seats (Meee ae PAU DOO} Bide ac SRNR ee ese MRA ar een ML | A ls al leccceetall aac ae este PAID Sie bpeameees ae [F-test cc tpn ke Manat Meese Lie Me lyre I oe ge Seige eat Weer pO sd OXON LOD ak ARES Eee | eee | arenes Sl eee Lele ey IE CON eae tenes Wp tae See ae a DAVES | eames taal tee Pa vedas. elas a ae ck reget Loe oral ale MeL ab Se iseescalinanetee EHSL NOLO) el ads eaters Ka sae || Mcrae lee ae aslo alle TNR LW SC seo [aneatoe. OSGSO isles eee seem Joke ccc tae ale a tee ane pa | Be on Weta ale eee ARO a OSGSAION| &, Sor Sean We DA. |b ieee elec tet eae bee eae [eons ke dP Be QSOS |S aereerceat Rees eaten | ienect aces eS eaeealll ees cae SCO Caisse [ee cars sees eee QR RO) | ee eee eter eee teeny tke | eer Sha Bo reese tee ee taal eve co OS Oe Bea eciars [tec es Loe OLE Bo see | ea SCAM Uae ROM Aeon wba [ie se ee OSG 9470"! Bei menral steer nace re-sale See ae Stale Peer [eceweradlaaeceaee eats CFV ON ES ill Ma ed ela en een eale el ALA SCI oreo esses teeta | ose Re PAOLO SCT IN, Saat ool Me ae en Sg ca bse ca (le A SA |i eel etoile ee PRE OS Bac ates acocel acamteteee sel eesadee eer eer ecacel erence Sho ee ce haart mel | Met QSOS2iO eee: Merce nest | eect ar see rae lNawe rial Ae | ga e 6 Medes lose OS erie Oa sen eee ter (ese oes fac es eae ccne [Each er ra Oe ae cr ae ee | Oe SON canna QSGSO oa | Rees sre ea eee a behace eet (ht cee Beamer lio. pall ees I Renee Sau eee OROSO MEd eke ly eaten cectawel mt eves aren eeaivertnl| ed I A al aoe 230, | ay one Dail | ence DSA cialllee cette ears ee ee ot amet eats eel Mm eee heat rT GN Ue LTD yd |e x OS GSS Only cmemee ales meme scale emcee te) [ei eerra | eoncty ee hare eee eee | eee x 2 OAR iy ere cr ected teeter ceed (eee eealberee SCAM Iss Alle reel vee ee pe Of 8 aed ee Ae eee FRR An Oe Sali S see ere tee Ve ee ee PANO} Yeni Beacen se nsanel uae scbse] eesanee Daal getaceed | Petar len ere | eee 2037 & Dil Le ae | aN IA Table showing correlation and allocation of the second upper molar and the fourth lower premolar in thirty-two skulls, each being referable to six types of variation in enamel pattern, illustrated in the text-figures as lettered. There are also additional or supernumerary loops present in the fourth lower premolar which are very difficult to homologize. The sequence of their succession is uncertain and they have therefore 254 University of California Publications in Zoology (Vou. 21 been designated in this paper as accessory cusps. The diagrams of the second upper molars and fourth lower premolars indicate the possible methods of the addition of these secondary cusps. A careful study of the enamel patterns of the molariform series in each of nearly two thousand specimens, belonging to three species of the genus Microtus, has conclusively shown that the closure or opening of any particular section enclosed by enamel is extremely uncertain. The amount of variation is so great that little reliance can be placed on the use of any especial feature of this sort as a diagnostic character. peeete Figs. E-J. Diagrams of enamel pattern of fourth lower premolars of Microtus montanus montanus, from Sisson, Siskiyou County, California. X 6. Fig. E, no. 2035, 9, coll. A. B. Howell; fig. F', no. 2033, g, coll. A. B. Howell; fig. G, no. 2038, 6, coll. A. B. Howell; fig. H, no. 2032, ¢, coll. A. B. Howell; fig. I, no. 98689, dg, coll. Biol. Surv.; fig. J, no. 98688, 9, coll. Biol. Surv. Hm Am Ute Figs. K—-P. Diagrams of enamel pattern of second upper molars of Microtus montanus montanus, from Sisson, Siskiyou County, California. X 6. Fig. K, no. 2028, 3, coll. A. B. Howell; fig. L, no. 2032, ¢, coll. A. B. Howell; fig. M, no. 2003, 9, coll. A. B. Howell; fig. N, no. 2004, g, coll. A. B. Howell; fig. O, no. 21098, 3, Mus. Vert. Zool.; fig. P, no. 2037, ¢, coll. A. B. Howell. The exact enamel pattern of any particular second upper molar is rarely if ever duplicated. The enamel patterns align themselves, however, in certain categories, that is, modifications of the normal enamel pattern of the tooth as is illustrated by the diagrams. It was observed in the course of this study that nearly all degrees of varia- tion in the enamel pattern were present to form a continuous series 1922] Kellogg: Californian Forms of Microtus Montanus 255 from one type to another in whatever direction one might wish to illustrate it, except in case of certain abnormal or aberrant variations such as is shown in figure P. Wr tm ti Me Figs. Q-T. Diagrams of enamel pattern of second upper molars of Microtus montanus yosemite. X 6. Fig. Q, no. 22361, 9, Mus. Vert. Zool., Tuolumne Meadows, Yosemite Park; fig. R, no. 24012, 9, Mus. Vert. Zool., Mono Lake (Salmon Ranch), Mono County; fig. S, no. 22363, 9, Mus. Vert. Zool., Tuolumne Meadows, Yosemite Park; fig. T, no. 23988, 4, Mus. Vert. Zool., Farrington’s Ranch, Mono Lake, Mono County. Hie We Oe Ot Figs. U-X. Diagrams of enamel pattern of second upper molars of Microtus montanus dutcheri. 6. Fig. U, no. 15601, 9, Mus. Vert. Zool., Whitney Meadows, Tulare County; fig. V, no. 15619, g, Mus. Vert. Zool., Whitney Creek, Tulare County; fig. W, no. 15622, 9, Mus. Vert. Zool., Whitney Creek, Tulare County; fig. X, no. 15595, 9, Mus. Vert. Zool., Whitney Meadows, Tulare County. Considerable amplitude is to be allowed in interpreting the eate- gories of variations as shown by table III. In some instances the assign- ment of certain specimens to any particular type was arbitrary, though in most cases the particular tooth in question was clearly a modifica- tion of some one of the regular types of variation. Of the second upper molars, figure L represents the mean, and of the fourth lower premolars, figure F is the type most often present. No two fourth lower premolars are exactly alike. However, there are fairly definite categories of these variations, which group themselves around certain types as illustrated by table III, with especial reference to figures E, HaG, Bel and J: Figures showing the’ molariform series of Microtus californicus with tritubereular nomenclature are included in the present paper in order that reference may be made to it in later studies, and that it may be available for use of other workers on that group. 256 University of California Publications in Zoology |Vou. 21 GENERAL CONSIDERATIONS Our present information relative to the Microtus montanus group points to the conclusion that ecological segregation is a factor as potent as any other in limiting the dispersal of the group as a whole and at the same time favoring its multiplication in particular areas. Microtus montanus is often absent from areas which apparently are favorable to its presence, and in certain localities, where conditions are suitable, it is present in rather large colonies. This, coupled with its wide distribution, would naturally result in slightly different, geographic variations becoming more or less characteristic of each colony. Any such variation that did arise would have a better chance of surviving, thus isolated, than if there was free intercrossing with the other colonies which comprise any given subspecies. Each marshy area becomes a center for differentiation. What one is really dealing with is a set of colonies, the individuals of each more or less variable, though usually with less individual variation in such a community than is observed among the individuals of the whole subspecies. The differences between the colonies are so shght that one cannot treat each aggregation as a separate subspecies. To recognize any one of these colonies by a subspecifie name would result in more confusion than convenience. On the other hand, the placing of any particular set of these colonies within one subspecies to which fairly definite limits of variability can be given does not really indicate their true importance, though for the sake of convenience it is necessary to make such a disposition of the material. METHODS USED IN STUDYING SPECIMENS The methods used for studying skins and measuring skulls (Thomas, 1905, pp. 191-196) are described in a previous paper of the writer on the Microtus californicus group (1918, p. 2). The color terms are taken from Ridgway (1912). ACKNOWLEDGMENTS All the specimens of the Microtus montanus group and the data pertaining thereto in the Museum of Vertebrate Zoology were placed at the writer’s disposal. For the loan of important, additional material the writer is indebted to the following: Dr. E. W. Nelson, chief of the Bureau of Biological Survey of the United States Department of Agriculture; Mr. Gerrit S. Miller, Jr., curator of the Division of 1922] Kellogg: Californian Forms of Microtus Montanus 257 Mammals, United States National Museum; Mr. A. Brazier Howell, of Pasadena, California; and Mr. Chase Littlejohn, of Redwood City, California. The writer also wishes to acknowledge his thanks for the help in many ways that he has received from Dr. Joseph Grinnell, director of the Museum of Vertebrate Zoology, and from Mr. Harry S. Swarth, Mr. Tracy I. Storer, and Mr. Joseph Dixon, of the staff of the same institution. LIST OF SUBSPECIES OF MICROTUS MONTANUS, WITH TYPE LOCALITIES Microtus montanus montanus (Peale). Headwaters of Sacramento River [prob- ably near Sisson], near Mount Shasta, California. Microtus montanus arizonensis Bailey. Springerville, Apache County, Arizona. Microtus montanus caryi Bailey. Milford, Fremont County, Wyoming. Microtus montanus dutcheri Bailey. Big Cottonwood Meadows, [southeast of | Mount Whitney, Inyo County, California. Microtus montanus rivularis Bailey. St. George, Washington County, Utah. Microtus montanus yosemite Grinnell. Yosemite Valley, Mariposa County, California. KEY TO SUBSPECIES OF MICROTUS MONTANUS 1. Size large, total length of old adults 177-190 mm.; skull large (condylobasal length about 28 mm.). 2. Bullae larger. 3. Skull with well developed longitudinal ridges; tail medium, 47-58 mm. 4. Colors dark, upperparts bister or brownish, strongly lined with black- ish overhairs; hind feet large, 20-23 mm.; lateral pits of palate OG PO 2a ccc care tates eet Re eee oe tee Ree Nae ct ce meen due cates beset cad ieee montanus 4’ Colors bright, upperparts yellowish or rusty brown, less heavily lined with blackish overhairs; hind feet smaller, 20-21 mm.; lateral puts (Of spallater shall OW Crna: cece cae os oa sec ceas serene = cesar eee arizonensis 3! Skull with longitudinal ridges poorly developed; tail short, 43-48 mm. Colors dull and pale, upperparts dull bister, lined with blackish over- Ives} loutayals sitereyne lbanefex es | AL} ea eas cece ge eee eee See rivularis 2! Bullae smaller. Skull angular, with well developed longitudinal ridges; tail medium, 45-58 mm. Colors dull, upperparts buffy or grayish, lined with blackish OVerhairs hind reeb lar OCs 20 — on Wnt lee eee ene eee esa ep eaeeeat yosemite U Size medium, total length of old adults seldom exceeding and usually less‘ than 190 mm.; skull small (condylobasal length less than 28 mm.) and with well developed longitudinal ridges. 5. Colors bright, upperparts reddish or brownish; underparts with buffy suf- fusion; tail medium, 47-56 mm.; hind foot large, 21-23 mm.; occiput oUt ys tir Cae te esac we Recetas ee ors a earn dutcheri 5! Colors dull, upperparts buffy or grayish; underparts without buffy suf- fusion; tail short, 41-45 mm.; hind foot smaller, 20-21 mm.; occiput SOKO (Od OF Ka KEY ire rab AK CEE ce ee che a aE ne caryt 258 University of California Publications in Zoology (Vou. 21 1. Microtus montanus montanus. 2. Microtus montanus yosemite. 3. Microtus montanus dutcheri. Specimens examined. Type locality. \ Ae J) a, S :: “. 4 y) iP “A Ne Se DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA SCALE 75 100 — MILES Fig. Y. Californian distribution of the Microtus montanus group of meadow . mice. enn ———————————_ Sr 1922] Kellogg: Californan Forms of Microtus Montanus 259 Microtus montanus montanus (Peale) Peale Meadow Mouse Arvicola montana Peale (1848, pp. 44-45). Arvicola montana, Baird (1857, pp. 528, 741); and of authors. Arvicola longirostris Baird (1857, p. 531); and of authors. Arvicola (Myonomes) riparius, Coues (1874, p. 189), part; and of some authors. Microtus longirostris, Trouessart (1897, p. 564). Microtus montanus, Merriam (1899, p. 95). Microtus montanus, Bailey (1900, p. 27); and of some authors, part. Nicrotus montanus montanus, Miller (1912, p. 216), part; and of some authors. Type specimen.—Sex unknown; U. 8. Nat. Mus. (no. unknown) ; headwaters of Sacramento River, near Mount Shasta [probably close to Sisson, 3600 feet altitude, Siskiyou County], California; October 4, 1841; collected by Titian R. Peale. (Type specimen cannot now be found. ) Range.—Siskiyou region of northern California, from Siskiyou Mountains east to Tule Lake, and south to Fort Crook and Dana, Shasta County. Vertical range from 2700 feet up to 6500 feet; zonal range Transition to Canadian. (See map, fig. Y.) Total number of specimens examined, 60, from the following local- ities in California. Siskiyou County: Stud Horse Canon, Siskiyou Mts. (6500 feet), 1 (coll. Biol. Surv.) ; Brownell (4800 feet), 5 (coll. Biol. Surv.) ; Mayten or Big Spring (2700 feet), 1 (Mus. Vert. Zool.) ; Sisson (3600 feet), 32 (coll. Biol. Surv., 16; coll. A. B. Howell, 13; Mus. Vert. Zool., 3). Shasta County: Dana, 2 (coll. Biol. Surv.) ; Fort Crook (3000 feet), 17. (coll. Biol. Surv.). Modoe County: Carr’s Ranch, Rhett [Tule] Lake, 2 (coll. Biol. Sury.). Diagnosis —Size large (hind foot 21 to 23 mm., condylobasal length of skull in largest individuals, 28.3 mm.) ; skull relatively large (ratio of zygomatic breadth to condylobasal length about 56 per cent), and ridged in old adults; incisive foramina narrow and constricted posteriorly ; bullae large and rounded; pelage long, usually dark; ears very hairy, relatively large, and not concealed by fur; hip glands conspicuous in old adults. Measurements —Average of 27 specimens from Sisson, California: total length, 165.5 mm. (142-192); tail vertebrae, 48.44 mm. (41- 55); hind foot, 21.55 mm. (20-23); condylobasal length, 26.86 mm. (24.9-28.3) ; occipito-nasal length, 26.29 mm. (25.2—-28) ; basilar length, Hensel, 24.31 mm. (22.5-26) ; nasal length, 7.55 mm. (6.8-8.2) ; zygo- matie width, 15.45 mm. (14.2-16.9) ; interorbital constriction, 3.6 mm. (3.4-4) ; interparietal width, 8.27 mm. (7.6—-8.8) ; shelf of bony palate, 14.21 mm. (12.8-14.9); height cranium at bullae, 10.21 mm. (9.6— 10.7) ; mastoid width, 12.25 mm. (11.6-12.9); length upper molar series, 6.61 mm. (6.2—7.3) ; length mandible, 17.34 mm. (16.4—-18.9). 260 University of Califorma Publications in Zoology [Vou.21 Color—Mass effect of dorsal area ranging from dull Prout’s brown to dark bister, with varying suffusion or admixture of long black overhairs, and heavily tipped with warm buff on cheeks, neck, and along the sides. In fresh pelage: hairs of upperparts slate color basally, with hght-tipped portion grading from cinnamon buff to snuff brown, the long overhairs varying from aniline black to blackish brown. Sides lighter than upperparts, decidedly more buffy, with light tipped portions of hairs approaching pinkish buff. Rump darker than rest of upperparts because of predominance of blackish over- hairs. Underparts varying from white to pallid neutral gray, irregu- larly darkened by the slate-colored basal underfur showing through. Inguinal region occasionally washed with buffy; anal area pure white: Terminal portion of whiskers whitish with blackish bases. Hairs of upper and lower lips whitish. Nose usually lighter than area in front of eyes. Ears large, not concealed by fur, the anterior border furred with hairs the same color as light-tipped hairs of sides. Tail distinctly bicolor, dark blackish brown above and pallid neutral gray below. Hands and feet covered with hairs varying from mouse gray to pallid neutral gray. In worn pelage: much duller, lighter, with grayish overeast in region of shoulders, and with a diminution of the long dark overhairs. In some specimens the long dark overhairs are so plentiful as to conceal the hghter hairs, giving the individual a grayish appearance. Skull—The skull of montanus is relatively large and massive, though slightly smaller than yosemite, becoming more ridged and angular in fully adult individuals. Dorsal profile, with exception of interorbital region, is uniformly convex. Brain case rather deep. Occiput rounded off behind, not obliquely truncate, concealing occipi- tal condyles for the most part when skull is viewed from above. Normally with considerable variation in contour of brain case due to differences in age. In sexually adult individuals, the skull is rather long, generally smooth, rounded at sides, convex superiorly, and slightly truneated anteriorly by postorbital tuberosities. In fully adult or very old individuals the brain case is subrectangular, flattened superiorly, but not abruptly truncated anteriorly by the postorbital tuberosities, which spread out laterally. Interorbital constriction nar- rower than anterior portion of rostrum. Interorbital ridges distinct, rather closely approaching each other, usually coalescing and extend- ing back to parietals, where in very old individuals they unite with longitudinal ridges. Interparietal strap-shaped, its greatest antero- posterior diameter, exclusive of median projection equaling one-third or more of greatest transverse diameter, and its lateral extremities abruptly truncated. Frontals variable posteriorly, either truncate or emarginate. Rostrum long, slender, gradually tapering anteriorly, the least depth behind incisors somewhat greater than width in same region. Posterior terminations of nasals variable, in some straight and in others attenuate, rounded or slightly emarginate, exceeded by nasal branch of premaxillae. Nasals at their widest region in front are conspicuously narrower than rostrum, abruptly constricted anterior to middle, and may extend as far as, but not beyond, plane of incisors. Anterior narial opening normally with vertical diameter greater than transverse. Anteorbital foramen wide and rather high, with superior i ea ae 1922] Kellogg: Californian Forms of Microtus Montanus 261 width usually not more than twice inferior width. The plate forming the outer wall, and continuous with the lacrimal internally, present, though incomplete superiorly. Occiput not depressed, the median depth averaging more than 59 per cent of greatest width across lambdoidal ridge of brain ease. Contour of foramen magnum variable, occasionally somewhat flattened superiorly, with vertical diameter usually less than, sometimes equal to, transverse. Paroccipital processes heavy, directed more downward than backward, with their extremities applied to bullae. Posterior margins of paroccipital processes continued upward as well defined ridges on supraoccipital though seldom reaching lambdoidal ridge superiorly. Basioccipital elevated and with well defined median ridge. Width of basioccipital at suture between it and basisphenoid usually less than one-third its median length. Auditory bullae well rounded, and rather large, nearly equaling those of Microtus californicus aestuarinus in old individuals. Posterior border of bulla rather oblique, very slightly indented by fenestra rotunda. Palate rather narrow, with palatine sulci shallow and indistinct. Median ridge variable, but lateral bridges well developed. Lateral pits of palate large and fenestrated. Pterygoids and hamular pro- cesses gradually diverging posteriorly. Interpterygoid fossa relatively narrow as compared with californicus, with posterior margin of palate variable, normally rather squarely truncate, occasionally rounded. Posterior palatine foramina small. Incisive foramina long, narrow and constricted posteriorly, equalling about 64 per cent of distance between base of incisors and anterior border of alveolus of first pre- molar. Zygomatic arches divergent anteriorly, relatively heavy, with median portion nearly parallel to main axis of skull and shightly indented at junction with premaxillae. Mandible similar to that of californicus. Ridge for masseteres laterales prominent. and well developed. Coronoid process large, its base broad, its extremity rising above level of condyle and curved stronely backward. Angular process rather wide, with its main axis curving outward and upward. Articular process over base of incisor marked by well developed protuberance. Base of mandibular foramen on a level with or slightly above cutting surface of last lower molar. Mental foramina on outer face below superior surface of diastema. Teeth.—Premolar and molars relatively light, their enamel pattern well defined, all the elements distinct, with deep salient and reentrant angles. Fourth upper premolar is not noticeably longer than first molar and is of approximately the same Jength as second molar. First upper molar with four closed sections. Fourth lower premolar normally with six closed triangles (fig. F). Fourth upper premolar normally with an anterior transverse loop and four alternating closed sections; two deep reéntrant angles are normally present on each side; these are sharply pointed, their points extending across long axis of tooth, those of outer side deeper than those of inner side; anterior loop with enamel enclosing protoconule normally flattened superiorly and shorter than same element of parastyle, its long axis at nearly right angles to that of tooth row. Triangles forming protocone and hypocone normally with sharper apex than triangles forming paracone and metacone. Loop forming metaconule with sharp internal angle. 262 University of California Publications in Zoology [Vou. 21 First upper molar with anterior transverse loop prolonged into narrow parastyle, its long axis in reverse direction to long axis of transverse loop of Pm*. The enamel enclosing the section designated as metaconule considerably constricted and prolonged. Second upper molar with an anterior transverse loop, its long axis nearly parallel to long axis of anterior transverse loop of Pm4, but with considerable variation in outline; closed sections are smaller than those of M+; enamel section enclosing metacone may or may not be closed (figs. L, M); just posterior to metacone, the long terminal loop is subtended externally by a slight notch, and internally by one or two reéntrant angles; this loop in its simplest form (fig. L) is indented at middle by a deep reéntrant angle which imparts to it a strongly curved erescentic outline, the anterior metaconule longer than secondary metaconule; the posterior limb of crescent enclosing meta- style may form an additional internal reéntrant angle (fig. O). Triangle enclosing hypocone may fuse with that of paracone (fig. P). Fourth lower premolar with posterior transverse loop, normally with three inner and three outer closed triangles, and a large anterior loop usually indented by deeper reéntrant angle on inner side than on outer. Inner side of tooth with four or five well developed re- entrant angles; outer side of tooth with three deep reentrant angles and a shallower fourth one. The large anterior loop designated as secondary paraconid of variable outline, often forming an external prolongation (fig. E). Reéntrant angles separating enamel sections enclosing secondary protoconulid, protoconulid, and protoconid, deeper at base than reéntrant angles that separate enamel sections enclos- ing metaconid, mesostylid, and entoconid. Posterior loop narrowly erescentic, with long axis slightly oblique with reference to that of tooth row, the internal portion enclosing entostylid longer than that designated as hypoconid. First lower molar with transverse posterior loop, two inner and two outer closed sections, and with two inner and two outer deep reéntrant angles. The closed triangles designated as paraconid, proto- conid, and metaconid of approximately the same size (in Microtus mordax, paraconid and metaconid are larger than protoconid). Second lower molar with three transverse loops, each side with two reentrant angles, the antero-external angle normally present; posterior loop larger than either of the other sections, the internal portion enclosing the entostylid curved forward at tip. Internal angle of metaconid not deflected forward as in Microtus californicus. Remarks.—The type of Arvicola montana was collected at the headwaters of the Sacramento River and described by Titian R. Peale (1848, pp. 4445). Vernon Bailey (1900, p. 28) after careful com- parison of various specimens from that general region decided that the type specimen agreed in all essential details with specimens from Sisson. Dr. T. 8S. Palmer in a conversation informed the present writer that various statements in Peale’s journal indicate that Bailey’s selection of Sisson as the type locality is correct. 1922] Kellogg: Californian Forms of Microtus Montanus 263 As remarked in the introduction, Microtus montanus has an inter- rupted distribution, being restricted for the most part to colonies in wet meadows and apparently limited to the vicinity of Mount Shasta and the Siskiyou Mountains of California. The subspecies montanus shows about the same range of individual variation as is exhibited by yosemite and dutcheri. Specimens from Brownell and Tule Lake are apparently closer to montanus than to yosemite. Specimens from Dana and Fort Crook are apparently best disposed of by referring them to montanus, while certain specimens from Cassel sted under yosemite show an approximation to montanus. On the other hand specimens from Bieber, Adin, and Pine Creek exhibit characters within the limits of variation of yosemite. During the year 1855, Dr. J. S. Newberry collected the type of Baird’s Arvicola longirostris at some point between Fort Reading and Fort Vancouver. He was then attached as geologist and botanist to Lieutenant R. S. Williamson’s party, which was making a survey for a railroad route from the Sacramento River Valley to the Colum- 8 : ——~, now 1 aS: ] 9990? NOW in the U. S. Nationa Museum, is labeled as having been collected during September, 1855, on the upper Pit River. Various circumstances indicate that the bia River. The type specimen, no. allocation of the name will be difficult. If the specimen was actually taken on the upper Pit River, it was probably collected on the west bank, somewhere north of the upper end of the canon (possibly near Happy Camp). It seems hardly possible, however, that the type could have been collected on the upper Pit River, for the explorers were in Oregon by August 13, and arrived at Fort Vancouver on October 9, 1855. He Cite tt Figs. F-P. Diagrams of enamel pattern of second upper molars of Microtus mordaxz sierrae. X 6. Fig. F, no. 95667, ¢, Biol. Surv. coll., Lassen Peak, Shasta County; fig. G, no. 106169, 2, Biol. Surv. coll., Grizzly Mountains, Plumas County ; fig. H, no. 17448, 9, Mus. Vert. Zool., Little Onion Valley, Sierra Nevada, Inyo County; fig. I, no. 23160, ¢, Mus. Vert. Zool., Fletcher Creek, Yosemite Park, Tuolumne County; fig. J, no. 17457, 9, Mus. Vert. Zool., Little Onion Valley, Sierra Nevada, Inyo County; fig. K, no. 95665, ¢, Biol. Surv. coll., Lassen Peak, Shasta County; fig. L, no. 15750, 9, Mus. Vert. Zool., Little Brush Meadow, Olancha Peak, Tulare County; fig. M, no. 25024, ¢, Mus. Vert. Zool., Bullfrog Lake, Fresno County; fig. N, no. 25035, 9, Mus. Vert. Zool., Kings River Canon, Fresno County; fig. O, no. 25031, 9, Mus. Vert. Zool., Kings River Cafion, Fresno County; fig. P, no. 13455, g, Mus. Vert Zool., Head Rush Creek, Siskiyou County. 1922 | Kellogg: Synopsis of Microtus Mordax 279 Figs. Q-T. Diagrams of enamel pattern of fourth lower premolars of Microtus mordax sierrae. X 6. Fig. Q, no. 22383, J, Mus. Vert. Zool., Merced Grove Big Trees, Mariposa County; fig. R, no. 22384, 9, Mus. Vert. Zool., Merced Grove Big Trees, Mariposa County; fig. S, no, 22382, 3, Mus. Vert. Zool., Merced Grove Big Trees, Mariposa County; fig. T, no. 22446, 9, Mus. Vert. Zool., head of Lyell Caiion, Yosemite Park, Tuolumne County. ti (tt > Figs. U-Y. Diagrams of enamel pattern of second upper molars of Microtus mordax angusticeps. X 6. Fig. U, no. 11528, g, Mus. Vert. Zool., 3 miles west of Arcata, Humboldt County; fig. V, no. 11526, 9, Mus. Vert. Zool., 3 miles west of Arcata, Humboldt County; fig. W, no. 20103, 9, Mus. Vert. Zool., Mendocino City, Mendocino County; fig. X, no. 1725, 9, coll. A. B. Howell, Crescent City, Del Norte County; fig. Y, no. 25588, 9, Mus. Vert. Zool., Fort Bragg, Mendocino pe CHR Cte Z AA BB Figs. Z-BB. Diagrams of enamel pattern of second upper molars of Microtus mordaz bernardinus. X 6. Fig. Z, no. 6264, Jf, Mus. Vert. Zool., Bluff Lake, San Bernardino Mountains; fig. AA, no. 6299, 2, Mus. Vert. Zool., San Bernardino Mountains; fig. BB, no. 6290, 3, Mus. Vert. Zool., San Bernardino Mountains. As a further illustration of the amount of variation likely to be encountered, attention should be ealled to the incisive foramina exhibited by the skulls on plate 8. The expansion or contraction of these foramina is subject to considerable variation. In fig. 7b, the incisive foramina are narrowly constricted posteriorly as in Microtus montanus, while in fig. 3b they are wide open. Other individual differ- ences are normally present in an ordinary series. For instance, the sides of the foramina are nearly parallel in fig. 5b, and a further type of modification with both ends narrowed is shown in fig. 8D. 280 University of Califorma Publications in Zoology TABLE I ILLUSTRATING THE REMARKABLE UNIFORMITY IN ZYGOMATIC WIDTH OF SKULL AMONG INDIVIDUALS SHOWING GREAT DIvERSITY IN ToTaL LENGTH oF Bopy Microtus mordax sierrae [ Vou. 21 CONDYLOBASAL LENGTH AND 491 Oy Oy, Oy Ss QQ Ss “Sei 0-0 A™A AAA AAA AAAAABRAWDAA A Total length 195 mm 195 mm 196 mm 196 mm 196 mm 197 mm 197 mm 26.8 MEASUREMENTS Condylobasal length 27.2 27.6 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. 28. noe ee nor RWE OC 29.1 29.3 14.8 Zygomatic width 15. 15. 15. 15. 15. 15. 15. 15 15. 15. 15. 15. SINT EH 00 DOOR 16.3 16.4 16.4 16.4 16.6 16.4 1922] Kellogg: Synopsis of Microtus Mordax 281 TABLE I—(Continued) MEASUREMENTS Museum No. Sex | Total length Condylobasal length Zygomatic width 15665 | 9 198 mm ae Ald 28.4 ay Bie “o 16.0 26412 | Q 198 mm ee Rae ese ee we ve 16.3 30054 | o 198 mm . Le — 28.7 ae ae 115)_.05) 22414 | o 199 mm ee ae ae 29.1_ os 15.8 25060 | 9° 200 mm is cae 28.0 OnE sock 15.6 13469 | o« | 200 mm aie ask 28.5 ae re 15.3 ar 22409 | & | 200 mm “ch mae ae 29.0 fos ae 16.4 23182 | Q 201 mm Ase sta 28.0 a coe 15.9 i ee 25052 | & | 201 mm oe ae ees 29.5 ae 53 ae 17.0 26422 | «| 204mm bee Es 28.6 aN nal 1/5) -45) Jes ‘oF 25043 | o& | 204 mm aif ect Bee 29.1 Hees ar sre il7/ ell 22420 | & | 207 mm ba sake 28.8 sais ae 15.8 19316 | &@ | 208 mm ss ae ae 29.5 st ee elOn 13456 | ow | 210 mm me ae 28.4 fas aie 50 13452 | @ | 221 mm Lee 27.9 as save a, 15.6 AGE AND SEX VARIATION It would be difficult to determine the sex of any individual from a study of the cranium alone. In practically all instances the range of individual variation is considerably greater than that of sex varl- ation. Thus, in the study of Microtus mordax, sex variation may be in most cases disregarded ; but it is by no means possible to eliminate, altogether, age variation. In most cases, when making comparisons of typical series of adults, it is very difficult to determine what con- stitutes maturity. In a series of fifty specimens of Microtus mordax sierrae, ranging in total length from 190 mm. to 221 mm., it was found on tabulation that there was but slight variation in the condylo- basal lengths of the skulls of these same individuals (see table I). There are two alternative interpretations which can be drawn from these data: either the total lengths as noted by the collector are unreliable, or the crania increase in size very slowly after the animals reach sexual maturity. Of the first interpretation it may be pointed out that errors would be on the average just as much below the true 282 Umwversity of Califorma Publications in Zoology [Vou 21 measurements as above and would make little or no difference in the conclusions derived from the data. The second interpretation seems to be justified. LIST OF CALIFORNIA SUBSPECIES OF MICROTUS MORDAX, WITH TYPE LOCALITIES Microtus mordax mordaz (Merriam). Sawtooth Lake, east base of Saw- tooth Mountains, Blaine County, Idaho. Microtus mordaz angusticeps Bailey. Crescent City, Del Norte County, California. Microtus mordax bernardinus Merriam. Dry Lake, at north base of San Gorgonio Peak, San Bernardino Mountains, San Bernardino County, California. Microtus mordaz sierrae, new subspecies. Tuolumne Meadows, Yosemite National Park, Tuolumne County, California. KEY TO SUBSPECIES OF MICROTUS MORDAX IN CALIFORNIA 1. Size medium, total length of old adults seldom greater than 180 mm., and usually less; longitudinal ridges of skull normally poorly developed. Skull medium (condylobasal length about 27 mm.) and narrow; colors bright, PEC Gish Gru DrO WISH. vc seen est. eee ae URE hee eee Ee ee angusticeps 1! Size large, total length of old adults at least 190 mm., sometimes over 200 mm.; longitudinal ridges of skull well developed. 2. Skull large (condylobasal length about 29 mm.) ; longitudinal ridges promi- MENG PAGO! OTs Gets lese OTe O Ryans eeees meena eeeeer succes ae seen ee ey sierrae 2° Skull smaller (condylobasal length about 28 mm.); longitudinal ridges less prominent. 3. Incisive foramina constricted posteriorly; colors light, sides grayish; dorsal tail stripe sharply contrasted with ventral surface....... mordax 3: Incisive foramina not constricted posteriorly; colors dark, sides dusky; dorsal tail stripe not sharply contrasted with ventral surface... bernardinus 1922 | Kellogg: Synopsis of Microtus Mordax 283 Microtus mordax mordax. Microtus mordax angusticeps 4) Microtus mordax bernardinus. Microtus mordax sierrae. bo em ww Specimens examined. Important published records. Type locality. DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA SCALE 25 75 400 MILES DRAFT Of 1919 Fig. CC. Californian distribution of the Microtus mordax group of meadow 284 University of Califorma Publications in Zoology [Vou. 21 Microtus mordax mordax (Merriam) Cantankerous Meadow Mouse Arvicola (Mynomes) mordax Merriam (1891, p. 61, pl. 2, figs. 3, 4). (Microtus) mordax, Trouessart (1899, p. 564). Microtus mordax, Bailey (1900, p. 48), part [Goose Lake]. Microtus mordaz mordax, Miller (1912, p. 221), part. 24231 31635” coll. Biol. Surv., U. 8. Nat. Mus.; Sawtooth Lake (or Alturas Lake), east base of Sawtooth Mountains, 7200 feet altitude, Blaine County, Idaho; September 29, 1890; collected by Dr. C. Hart Merriam and Vernon Bailey; original no. 1903. Range.—Modoe or Great Basin region of northeastern California. Vertical range from 4800 feet up to 8700 feet ; zonal range Transition to Hudsonian. (See map, fig. CC.) Total number of specimens examined, 39, from the following loceali- ties in California. Modoe County: east face of Warren Peak, Warner Mountains (8700 feet), 11; head of N. Fork of Parker Creek (7300 feet), 11; Parker Creek (5000-5500 feet), 9; Goose Lake Meadows near Sugar Hill (4800 feet), 2; Goose Lake Meadows near Davis Creek (4800 feet), 4 (1 Biol. Surv.) ; Lassen Creek (5600 feet), 1 (Biol. Surv.) ; Goose Lake (5150 feet), 1 (Biol. Surv.). Diagnosis —Size medium (hind foot, 21 to 22 mm., condylobasal length of skull in largest individuals, 28.1 mm.) ; second upper molar without posterior internal loop or triangle; first lower molar with four reentrant angles on outer side; skull, medium, slender and lightly built (ratio of zygomatic breadth to condylobasal length about 54 per cent) ; interorbital region not developing a crest with age. Pelage grayish and normally light colored; underparts nearly pure white; ears moderate in size (10 to 138 mm.). Measurements.—Average of 9 specimens, including 2 topotypes: Total length, 177.55 mm. (168-185) ; tail vertebrae, 63.88 mm. (61— 70) ; hind foot, 21.27 mm. (20-22.5) ; condylobasal length, 26.84 mm. (26.2-28.1) ; occipito-nasal length, 26.24 mm. (25.3-27.8); basilar length, Hensel, 23.9 mm. (23.3-24.8) ; nasal length, 7.82 mm. (7-8.8) ; zygomatie width, 14.96 mm. (13.9-16.1); interorbital constriction, 3.78 mm. (3.1-4.1) ; interparietal width, 8.56 mm. (8.1-8.9); shelf of bony palate, 13.46 mm. (12.9-14.3) ; height cranium at bullae, 9.77 mm. (9.5-10.1) ; mastoid width, 12.22 mm. (11.3-12.8) ; length upper molar series, 6.34 mm. (5.9-6.7) ; length mandible, 15.94 mm. (15.3- TGS). Color.—Mass effect of dorsal area ranging from tawny olive to light cinnamon brown, with varying admixture of long dark overhairs. Dorsal stripe strongly contrasted with rest of upperparts. In fresh pelage: hairs of upperparts blackish plumbeous basally, with light tipped portion grading from cinnamon buff to dark clay color; the long overhairs grading from bone brown to aniline black. Color of Type specimen.—Male adult; no. 1922 | Kellogg: Synopsis of Microtus Mordax 285 upperparts shades into light gray on face, sides and rump, the indi- vidual hairs sometimes tipped with pinkish buff; forehead of same color as dorsal stripe. Underparts whitish, dulled by the plumbeous bases of hairs; anal area pure white. Whiskers pallid neutral gray except short basal portions which are aniline black. Ears moderate in size, with anterior border concealed by grayish hairs of face. Tail distinctly bicolor, dusky drab or dark blackish brown above and pallid neutral gray below, though when tail is scantily haired the yel- lowish scales show through. Hands and feet covered with pallid mouse gray hairs. In worn pelage: general coloration as a whole lighter. Skull.Separable as a rule from that of sierrae by lighter and less spreading zygomatic arches, the relatively shorter rostrum and smaller bullae. Dorsal profile essentially the same. Occiput rounded, con- cealing for the most part the condyles when skull is viewed from above. All skulls examined, characterized by narrower brain case than in sterrae, with sides rounded and rarely with postorbital tubero- sities sufficiently developed to make any change in contour of the skull. Longitudinal ridges poorly developed, continuous posteriorly with lambdoidal ridge and anteriorly with interorbital ridges. Inter- orbital ridges variable in appearance, seldom prominent enough to show a median suleus. Interparietal with essentially same variations aS In sierrae. (See plate 8.) In superior view the slender rostral region tapers gradually toward anterior end. Rostrum narrow, rarely wider at narrowest point behind incisors than interorbital width, the least depth behind incisors greater than width in same region. Nasals long, spatulate, widening at a point anterior to middle, their posterior terminations emarginate. Ascending arms of premaxillae exceeding the nasals and extending to or beyond plane of lacrimals. Vertical diameter of opening for anterior nares about equal to transverse. Anteorbital foramen narrow and high, with superior portion considerably wider than inferior. Occiput not depressed or flattened, the median depth averaging about 58 per cent of greatest width. across lambdoidal ridge of brain ease. Outline of foramen magnum essentially as in sierrae. Par- occipital processes usually directed more downward than backward and applied to bullae at their extremities. Posterior margins of these processes continued upwards on supra-occipital as sharp crests, termin- ating below lambdoidal ridge. Basioccipital with well defined median ridge. Auditory bullae smaller than in sierrae, flattened below meatus, with their posterior margins indented by foramina. Palate slightly narrower than in sierrae, with rather deep palatine sulci and elevated median ridge. Lateral bridges light but well developed. Lateral pits of palate exhibiting a variable amount of fenestration. Interpterygoid fossa relatively narrow, the pterygoids and their hamular processes gradually diverging posteriorly. Incisive foramina narrow, expanded anterior to middle and equaling about 61 per cent of distance between base of incisors and anterior borders of alveoli of first premolars. Zygomatic arches diverging rapidly anteriorly, with region of greatest convexity at junction of jugal with maxillary root of zygoma. Jugal normally expanded, its head shallowly mortised into maxillary root of zygoma. 286 University of California Publications in Zoology [Vou. 21 Mandible light and slender, similar to that of sierrae. Ridge for masseteres laterales well developed. Coronoid process long, its base narrow, its extremity rising above level of condyle and curved back- ward at tip. Angular process relatively stout, curved outward, but with posterior border not extending as far as plane of condyle. Peri- pheral swelling over base of incisor, lying for the most part within masseterie ridge. Base of mandibular foramen above or on a level with cutting surface of last lower molar. Mental foramen on lateral face situated below superior surface of diastema. Teceth.—Both upper and lower molariform series differ from those of Microtus montanus in a general tendency toward wider reéntrant angles, with apices of both salient and reentrant angles more rounded, and with enamel section designated as protoconid smaller than either paraconid or metaconid. Compared with those of Microtus californi- cus the molariform series are slightly smaller; otherwise only certain details of the enamel folding constitute the diagnostic differences. Fourth upper premolar is longer than either the first or second molars (fig. B). First upper molar with four closed sections. Fourth lower premolar with five and occasionally six closed triangles. Fourth upper premolar normally with anterior transverse loop and four alternating closed sections; the external reentrant angles deeper than the internal, their points extending across long axis of the tooth; anterior portion of transverse loop designated as proto- econule somewhat flattened superiorly, and long axis of loop forms an oblique angle with long axis of tooth. Triangles designated as protocone and hypocone with apices more rounded than either para- cone or metacone. Loop forming metaconule with no internal pro- longation. First upper molar with anterior border of transverse loop irregular in appearance, its long axis in reverse direction to long axis of transverse loop of Pm4. Metaconule is not conspicuously lengthened posteriorly. Second upper molar with long axis of anterior transverse loop nearly parallel with long axis of anterior transverse loop of Pm, but normally with more convex anterior profile. The closed sections are smaller than those of M+; enamel section enclosing metacone normally closed posteriorly (fig. C) though sometimes open superiorly (fig. E) ; the terminal loop is subtended externally by notch at base of enamel section designated as metacone and internally by one reentrant angle ; this loop in its simplest form (fig. D) is indented at middle by deep reéntrant angle, though external outline is quite variable (fig. E), seldom erescentic ; an incipient internal reéntrant angle is occasionally formed on posterior limb of loop designated as metastyle (fig. D). Triangle enclosing hypocone may fuse with that of metacone (fig. E). Fourth lower premolar with posterior transverse loop, normally witb three inner and two outer closed triangles, and a large anterior loop, usually with deeper internal reéntrant angles than external ones. Inner side of tooth with four or five well developed, reéntrant angles; outer side of tooth with three deep reentrant angles and a shallower fourth one. Contour of large anterior loop designated as secondary paraconid variable, rarely forming an external prolonga- tion. Posterior loop narrowly erescentic, with oblique long axis. 1922 | Kellogg: Synopsis of Microtus Mordax 287 First lower molar with posterior border of transverse loop unevenly convex, two inner and two outer closed sections and with internal reéntrant angles much deeper than external ones. The closed triangles designated as paraconid and metaconid, larger than protoconid. Second lower molar with three transverse sections or loops, the internal reéntrant angles much deeper and narrower than external; posterior loop but shghtly if at all larger than adjoining loop, the enamel border enclosing the paraconid considerably produced so that tip lies in same plane as tips of metaconid and entostylid. Remarks.—The writer has not examined the type specimen of Microtus mordax, though a topotype series of eleven specimens, loaned for study by the Biological Survey, was critically studied. These specimens agree in all essential details with specimens of this subspecies from the Modoe region of California, and from the Pine Forest Mountains of northern Nevada. As originally described, Microtus mordax was a wide ranging form, with frequent interruptions or gaps in its distribution, but with sup- posedly little variation throughout its range. The accumulation of a considerable series of specimens in the Museum of Vertebrate Zoology from many points within California has enabled the writer to better elucidate the relationships of the various local varieties which make up this composite. Microtus mordax mordaz, as now restricted, occupies within the state only the Modoe or Great Basin region. Unfortunately no specimens have been collected in the Pit River region east of Bear Creek, and as the specimens from the latter locality have not been examined, the area of intergradation between mordax and sierrae remains to be determined. A few specimens from Parker Creek exhibit certain characters which indicate that this entire Modoc region may be one of intergradation, though the series as a whole average smaller both with respect to skulls and to external measure- ments. There is often considerable uncertainty as to the exact dis- position of individual specimens from intermediate areas, and this is certainly true of various specimens from the Modoe Region. The number of specimens of Microtus mordax mordax available for study is much too small to permit accurate comparison with the material of sierrae, though it does indicate that the latter form is a much more robust and heavily proportioned meadow mouse than typical mordax. 288 University of California Publications in Zoology [Vou. 21 Microtus mordax sierrae, new subspecies Sierra Nevada Meadow Mouse Microtus mordax, Bailey (1900, p. 48), part. Microtus mordax, Elliot (1907, p. 296), part. Microtus mordax mordax, Miller (1912, p. 221), part. Type speciomen.—Male adult ; no. 22437, Mus. Vert. Zool. ; Tuolumne Meadows, Yosemite National Park, 8600 feet altitude, Tuolumne County, California; July 7, 1915; collected by Charles L. Camp; original no. 2159. Range.—The Sierra Nevada, White and Trinity mountains, of eastern and northern California, from Taylor Meadow, Tulare County, north to Lassen Peak and Goose Nest Mountains, Siskiyou County, and adjoining parts of Oregon. (Extralimital range unknown.) Vertical range from 2000 feet up to 11,500 feet; zonal range low Transition to Arctic-Alpine. (See map, fig. CC.) Total number of specimens examined, 568, from the following localities in California. TRINITY cCouNTY: head Grizzly Creek (6000 feet), 18; head Bear Creek (6400 feet), 10. siskryou county: South Fork Salmon River (5000 feet), 18; Saloon Creek Divide (6850 feet), 1; Jackson Lake (5900 feet), 10; Sisson (3600 feet), 4; Mount Shasta (7000 feet), 9; head Rush Creek (6400 feet), 10; Castle Lake (5434 feet), 2; Goose Nest Mountains, 1 (Biol. Surv.). SHASTA COUNTY: Warner Creek, south slope Lassen Peak (5500 feet), 1 (Biol. Surv.) ; Lassen Peak (Hudsonian east side), 15 (Biol. Surv.). PLUMAS county: Twenty miles s.w. Quiney (5000 feet), 2 (Biol. Surv.) ; Grizzly Mountains (4100 feet), 2 (Biol. Surv.) ; Sierra Valley (5100 feet), 2 (Biol. Surv.). NEVADA county: Independence Lake (7000 feet), 3. PLACER couNTY: Blue Canon (4700-5000 feet), 2; Cisco (6000 feet), 6. ALPINE couNTYy: below Hope Valley (6800 feet), 2 (Biol. Surv.). Mono county: Farrington’s Ranch near Mono Lake (6800 feet), 1; Walker Lake (8000 feet), 1; one mile south Williams Butte (6900 feet), 1; Warren Fork of Leevining Creek (9200 feet), 8; McCloud Camp, Cottonwood Creek, White Mountains (9200 feet), 1; McAfee Meadow, White Mountains (11,500 feet), 4; Big Prospector Meadow, White Mountains (10,200 feet), 4; Poison Creek, White Mountains (9000 feet), 2. TUoLUMNE couNTY: Tuolumne Meadows (8600 feet), 6; Lyell Canon (10,000 feet), 6; head Lyell Canon (9300— 10,700 feet), 10; Glen Aulin, Tuolumne River (7700 feet), 5; Ten Lakes, Yosemite Park (9200 feet), 1; Aspen Valley (6400 feet), 1. MARIPOSA COUNTY: Gentry’s, Big Oak Flat Road (5800 feet), 3; Yosemite Valley floor (4000 feet), 4; El Portal (2000 feet), 1; Yosemite, Yosemite Valley (4000 feet), 3; Power House Meadows (4000 feet), 3; Yosemite Creek (4000 feet), 3; Mount Hoffmann (10,000 feet), 5; Poreupine Flat (8100 feet), 14; Mereed Grove Big Trees (5400-5500 feet), 15; Chinquapin (6200-6400 feet), 9; Crane Flat. (6300 feet), 2; Mono Meadow (7300 feet), 8; East Fork Indian 1922] Kellogg: Synopsis of Microtus Mordax 289 Canon (7300 feet), 3; 1 mile east Merced Lake (7500 feet), 5; Vogel- sang Lake (10,300 feet), 7; Fletcher Creek near Vogelsang Lake (10,100 feet), 4. mvvo county: Carroll Creek (5500 feet), 1; Little Onion Valley, Sierra Nevada Mountains (7500 feet), 28; Onion Valley, Sierra Nevada Mountains (8500 feet), 45; Kearsarge Pass (10,500 feet), 25; Roberts Ranch, Wyman Creek, White Mountains (8250 feet), 6; Silver Canon, seven miles east of Laws (7000 feet), 20; Little Cottonwood Creek, Sierra Nevada Mountains (9500 feet), 5. FRESNO county: Hume (5300 feet), 8; Bullfrog Lake (10,600 feet), 13; Bubbs Creek (9500 feet), 4; King’s River Cafion (5000 feet), 29; Horse Corral Meadow (7600 feet), 10. TULARE couNTY: Taylor Meadow (7000 feet), 8; two miles east Kern Peak (9300 feet), 1; Jordan Hot Springs (6700 feet), 7; Cannell Meadow (7500 feet), 24; Siretta Meadows (9000 feet), 18; Jackass Meadow (7750 feet), 17; Monache Meadow (8000 feet), 12; Whitney Meadow (9800 feet), 18; Little Brush Meadow, Olancha Peak (9750 feet), 9; Twin Lakes, Sequoia National Park (10,238 feet), 6; Sherman Creek, Sequoia National Park, £ Diagnosis —Size medium (hind foot, 20 to 22 mm.; condylobasal length of skull in largest individuals, 29.5 mm.) ; second upper molar without postero-internal triangle ; first lower molar with four reéntrant angles on outer side; skull long, of moderate width, angular and much ridged in old adults (ratio of zygomatic breadth to condylobasal length about 55 per cent), the intertemporal region rarely developing a ridge with age; auditory bullae large and inflated; pelage darker than in mordax, more brownish; ears moderate in size (10 to 14 mm.). Measurements.—Average of 51 specimens: total length, 195.97 mm. (190-221) ; tail vertebrae, 68.39 mm. (55-77); hind foot, 22.2 mm. (21-24); condylobasal length of skull, 28.21 mm. (26.8—29.5) ; occipito-nasal length, 27.68 mm. (25.6-29.9) ; basilar length, Hensel, 25.0 mm. (24.1—26.3) ; nasal length, 8.4 mm. (7.38-9) ; zygomatie width, 15.8 mm. (14.8-16.9) ; interorbital constriction, 4.05 mm. (3.7-4.5) ; interparietal width, 8.7 mm. (7.9—9.6) ; shelf of bony palate, 14.15 mm. (13.5-14.8) ; height cranium at bullae, 10.01 mm. (9.5-10.8) ; mastoid width, 12.77 mm. (11.9-138.3) ; length upper molar series, 6.78 mm. (6.1-7.4) ; length mandible, 16.94 mm. (15.6-18.3). Color.—Upperparts and sides as a rule darker than mordar. Mass effect of dorsal area more strongly mixed with brownish and blackish overhairs. Dorsal stripe less sharply contrasted with coloration of sides. In fresh pelage: hairs of upperparts blackish slate basally, with light tipped portions normally clay color, but may vary from cinna- mon buff to tawny olive; the long overhairs grading from bone brown to aniline black. Sides lighter than upperparts, more grayish, with long overhairs not materially darkening the ground color; rump erayer than rest of upperparts. The plumbeous hair bases are usually visible in pelage of sides and rump, and affect general coloration. Underparts pale gray, with blackish slate hair bases showing through at surface to a varying degree in different individuals; anal area pure white. Whiskers mixed with white and aniline black. Upper and lower lips whitish. Cheeks grayish with slight infusion of cinnamon 290 University of California Publications in Zoology (Vou. 21 buff tipped hairs. Nose and forehead strongly mixed with dark over- hairs. Ears small, almost concealed by fur, with anterior border con- cealed by blackish slate hair bases, while posterior border is covered with hairs of same shade as light tipped hairs of upperparts. Tail distinetly bicolor, aniline black above and pallid neutral gray below, thongh when tail is scantily haired the yellowish seales show through. Hands and feet covered with dull white hairs. In worn pelage: gen- eral coloration as a whole much duller or more grayish. The blackish slate hair bases enter into general coloration of upperparts to a larger extent. Skull—tIn fully adult individuals the skull attains a length equal to that of Microtus califormcus mariposae, but width of zygomata rarely exceeds this dimension in the latter, so that the general form is somewhat less robust and less angular. Dorsal and ventral profiles essentially the same as in mariposae except that nasals are usually less abruptly depressed anteriorly and depth of brain ease is slightly shallower. Occiput rather obliquely truncate, concealing part of occipital condyles, when skull is viewed from above. (See plate 8.) The majority of skulls have a rather long brain case, as compared with anterior portion of skull, subovate in outline, but with moderately developed postorbital tuberosities that exhibit a tendency to spread out laterally along posterior border of orbit, though they are suf- ficiently prominent to give the anterior border of brain case a squarish appearance. Surface of brain case, in mature individuals, with well indicated longitudinal ridges along outer edges of parietals. The inter- orbital region is very broad, more so than in other races of this group. Interorbital ridges usually poorly defined, but normally with distinct suleus between the two. Interparietal variable in outline, usually sub- rectangular, its posterior border decidedly convex, its lateral extrem- ities rather obliquely truncate, and with median projection of normal size. Frontals large, truncate, or emarginate posteriorly. Preorbital region long and slender, with gradually tapering ros- trum, slightly wider than greatest breadth of interorbital constriction and rather shallow proximally; least depth behind incisors slightly greater than width in same region. Nasals not projecting beyond incisors, usually terminating posteriorly on a level with nasal branches of premaxillaries, somewhat constricted a little anterior to middle, attenuate posteriorly but rarely as noticeably as in angusticeps. An- terior narial opening with transverse diameter about equal to vertical. Anteorbital foramen rather shallow and wide, with superior breadth considerably greater and in most cases at least twice inferior breadth. Depth of occiput, when viewed from behind, shallower than in Microtus c. mariposae, and with flattened occipital region, the median depth in majority of skulls examined about 57 per cent of greatest width across lambdoidal ridge of brain case. Foramen magnum variable, with vertical diameter equal to or less than transverse. Paroccipital processes light, directed more downward than backward and rather closely applied to bullae; their posterior margins con- tinued upward as sharp crests, rarely touching the lambdoidal ridge superiorly. 1922] Kellogg: Synopsis of Microtus Mordax 291 Basioccipital in most of skulls examined possesses a poorly defined median ridge. Width of basioccipital at suture between it and basi- sphenoid equal to or less than one-third its median length. Auditory bullae moderate in size, smaller than in mariposae, and considerably inflated; differs from the latter in that posterior margin of bulla is not so strongly indented by foramen, and superior rim of meatus is shorter and not so flaring. Palate narrower, more depressed, and with deeper palatine sulci than in mariposae. Lateral bridges and posterior palatine foramina present but not conspicuous. Lateral pits of palate seldom noticeably fenestrated. Pterygoids and hamular processes normally very nearly parallel. Interpterygoid fossa wider than in Microtus californicus group, with posterior margin of palate concave. Incisive foramina equal to or less than two-thirds distance between base of incisors and anterior border of alveoli of molariform series, wide open and narrow- ing at both ends, but with widest portion usually slightly anterior to middle. Zygomatie arches not widely or abruptly spreading, with jugal portion parallel to main axis of skull. Jugal slender, about equal in length to that in mariposae, but shallower. Zygomatie arches may or may not be notched at anterior junction with premaxillaries. Mandible very lght, long and slender. Ridge for masseteres laterales well developed. Coronoid process slender, its base narrow, its extremity rising to or above level of condyle, but with posterior curvature at tip variable. Angular process well developed though rather slender, shorter than in mariposae, not curved as strongly outward. Articular tuberosity over base of incisor, conspicuous. Base of mandibular foramen on a level with cutting surface of last lower molar. Mental foramen on outer face considerably below superior surface of diastema. Teeth.—In structure the teeth do not differ appreciably from those of Microtus mordax mordax, though the enamel pattern in general is somewhat variable, especially in M2. The elements of the enamel pattern of the remainder of the molariform series are very similar to those in mordax ; M2 occasionally lacks the notch posterior to meta- cone (fig. F'); terminal loop seldom erescentie (fig. H), often with incipient external angle (figs. J, K, M, and N); an incipient internal reentrant angle is occasionally formed on posterior limb of loop designated as metastyle (fig. O) ; hypocone may fuse with paracone (fig. I) ; metacone may (figs. F, G, I, L, and P) or may not (figs. H, J, K, M, N, and O) be open inferiorly; internal reéntrant angle on terminal loop may be almost wanting (fig. L.) or may be deep (fig. J). Large anterior loop of Pm; designated as secondary paraconid, rarely forms an external prolongation (fig. R); section designated as secondary protoconulid often considerably reduced in size (fig. T) ; internal reéntrant angles (figs. Q-T) relatively deeper than those of mordax. 292 University of California Publications in Zoology [Vou. 21 Remarks.—This race of the Microtus mordax group exhibits con- siderable geographic variation throughout its range in California. Specimens from the Trinity Mountain region and from the southern part of its range in the Sierra Nevada are much darker than those from intermediate localities. Similarly, specimens from the White Mountains are much lighter than any from the nearby Sierra Nevada. There can be little doubt, however, that there is an opportunity for continuous dispersal along streams which empty into Owens River from the Sierra Nevada and thenee up similar streams which drain the White Mountains. Sierrae is common throughout its range in suitable places such as high mountain marshes and wet woods and, more especially, along the banks of cold mountain streams. Indi- viduals often descend these streams for a considerable distance below the normal zonal position of the species. So far as is known sierrae is separated from bernardinus by a wide interval. Also no specimens are known from the considerable width of territory between the head of Rush Creek, Siskiyou County, whence sierraé has been obtained, and the northwest coastal strip occupied by angusticeps (see map, fig. CC). There is no reason known to the writer which would prevent the dispersal of sierrae down Rush Creek to the Klamath River and thence to the seacoast ; future collect- ing may supply the necessary material to establish the fact of inter- gradation between sierrae and angusticeps. In fact large specimens of mordax have been collected near Crescent City and Eureka which are very perplexing and can only be explained on this assumption. Elliot (1907, pp. 295-296) referred these large individuals to Microtus townsendi, but they certainly belong to the mordax group. In some respects these specimens are intermediate between typical angusticeps and sierrae. 1922 | Kellogg: Synopsis of Microtus Mordax 293 Microtus mordax angusticeps Bailey Northwest Coast Meadow Mouse Microtus angusticeps Bailey (1898, p. 86). Microtus angusticeps, Bailey (1900, p. 51). Microtus angusticeps, Elliot (1903, p. 177). Requa, Del Norte County. Microtus townsendi, Elliot (1907, pp. 295-296). Eureka, Humboldt County. Microtus mordax angusticeps, Grinnell (1913, p. 320). GOST nak A - 34008” Biol. Surv. coll., U. 8. Nat. Mus.; Crescent City, Del Norte County, California; June 16, 1889; collected by Theodore S. Palmer; original.no. 151. Range.—Northern humid coast region of California from Mendo- eino, Mendocino County, north to Crescent City, Del Norte County. Vertical range, so far as known, not over 100 feet above sea level; zonal range Transition. (See map, fig. CC.) Total number of specimens examined, 44, from the following localities in California. Mendocino County: Fort Bragg, 16; Mendo- cino, 1. Humboldt County: Trinidad, 3; three miles west of Arcata, 16; Eureka, 4. Del Norte County: Crescent City, 4. Diagnosis—Size medium (hind foot, 20 to 23 mm.; condylobasal length of skull in largest individuals, 28.4 mm.). Skull normally small, narrow and lightly built (ratio of zygomatic breadth to con- dylobasal length about 55 per cent), and not conspicuously ridged in adults; auditory bullae small and flattened below meatus; pelage in mass effect approaching russet; ears relatively large (14 to 15 mm.). Measurements —Average of 10 specimens: Total length, 174.3 mm. (162-194) ; tail vertebrae, 61.2 mm. (56-74); hind foot, 22.0 mm. (21-23) ; condylobasal length, 26.59 mm. (25-28.4) ; occipito-nasal length, 26.6 mm. (25.5-28.2) ; basilar length, Hensel, 23.32 mm. (22- 24.9) ; nasal length, 7.77 mm. (7.2-8.6) ; zygomatic width, 14.89 mm. (14.1-15.7) ; interorbital constriction, 3.71 mm. (3.5-4) ; interparietal width, 8.43 mm. (8.1-8.9) ; shelf of bony palate, 13.06 mm. (12.4-13.9) ; height cranium at bullae, 9.92 mm. (9.5-10.4) ; mastoid width, 11.88 mm. (11.3-12.5) ; length upper molar series, 6.5 mm. (6.1—7) ; length mandible, 15.98 mm. (15.2-16.8). Color.—Upperparts and sides as a rule brighter than in any other race of the mordax group in California. Dorsal area not so strongly darkened by presence of long overhairs. In fresh pelage: ground eolor of upperparts varying from argus brown to auburn. Light terminal portions of hairs grading from ochraceous buff to warm buff ; basal portions of hairs blackish plumbeous. Long overhairs ranging in shade from mars brown to seal brown. Color of upperparts changes rather abruptly to lighter color of sides. The hight or warm buff hairs of sides are more conspicuous because of absence of long overhairs, and sides are not grayish in mass effect. Underparts with pale Type specumen.—Male adult; no 294 University of California Publications in Zoology (Vou. 21 ochraceous buff suffusion irregularly darkened by plumbeous hair bases showing through; anal area pure white. Terminal half of whiskers pallid neutral gray, basal half blackish. Upper and lower lips white. Ears relatively large, not concealed by fur, with anterior border concealed by plumbeous bases of lght tipped hairs. Tail bicolor, dark mars brown above and scarcely covered with light grayish hairs below, with the yellowish scales showing through. Hands and feet covered with light mouse gray hairs. Young: color of upperparts duller and darker than in adults, with larger proportion of long dark overhairs, though in some specimens there is a conspicuous brightening due to presence of buffy hairs. Skull—Although the skull of this form is lighter and shorter than in typical sierrae, it is similar in general outline. The depth of brain case is relatively greater, though dorsal profile is essentially the same. Occiput rather obliquely truncate, permitting posterior portion of condyles to be visible when skull is viewed from above. (See plate 8.) Compared with skulls of sierrae the brain ease is rather long and narrow, slightly rounded at sides, but with postorbital tuberosities not sufficiently developed to produce any break in contour of skull. The brain case is slightly inflated superiorly, with fairly well indicated longitudinal ridges along outer edges of parietals. Interorbital region very narrow, more so than in other races of this group. Interorbital ridges usually coalesced in adults, forming a shallow median erest. Interparietal variable in outline, its area decidedly less than that of parietal. Width of interparietal at extremities contained at least three and occasionally four times in its greatest length. Frontals convex or emarginate posteriorly, but more elevated anteriorly than in other races in this group. In comparison with Microtus m. mordax, the rostrum is essentially the same, though ratio of length to proximal depth is more noticeable as contrasted with shallow depth of former. Least depth behind incisors usually greater than width in same region. Nasals at widest point but shghtly narrower than rostrum, and projecting as far as or beyond plane of incisors, their posterior terminations usually exceeded by posterior margins of premaxillaries. Opening for an- terior nares with transverse diameter about equal to vertical. Ante- orbital foramen rather short and narrow with superior portion at least twice width of inferior. Depth of occiput when viewed from behind more than half width of brain case; the median depth may equal as much as 61 per cent of greatest width across lambdoidal ridge of brain case. Dorsal profile more convex, and ratio of depth to length greater, than in mordaz. Foramen magnum roughly triangular, not flattened superiorly as in mordax, with vertical diameter slightly exceeding the transverse. Paroccipital processes light, directed more backward than downward, not applied to bullae at their extremities, and their posterior margins continued upward, as in mordax. The median ridge of basioccipital poorly defined. Auditory bullae small and constricted, their posterior boundaries more obliquely truncated than in mordax, with posterior margins slightly indented by foramina. Palate about the same as in bernardinus, but with shallower palatine sulci, broader median ridge, and more strongly developed ia 1922 | Kellogg: Synopsis of Microtus Mordax 295 lateral bridges. Lateral pits of palate variable, usually with distinet fenestration. Pterygoids and hamular processes slightly diverging posteriorly. Posterior margin of palate usually concave, sometimes notched. Interpterygoid fossa averaging a little narrower than in typical mordax. Posterior palatine foramina open. Incisive fora- mina narrow, somewhat constricted at ends, and equal to about 62 per cent of distance between base of incisors and anterior border of alveoli of molariform series. Zygomatie arches hght, broadly spreading posteriorly, expanded at the middle and narrowing anteriorly. Zygomatie arches deeply notched at anterior junction with premaxillaries. Jugal heavier, deeper and more firmly mortised into zygoma than in mordaz. Mandible small, relatively short and deep. Inferior border of ramus nearly straight, forming a more abrupt angle with chin than in californmicus. Ridge for masseteres laterales well developed and but seldom as sharp as that exhibited by mordax. Coronoid process very slender, its extremity rising to or slightly above level of condyle, and curved strongly backward at tip. Condylar process slender and long, much more so than in typical mordax or californicus. Angular process well developed but not narrowing abruptly near extremity and curved but little outward. Surface of mandible over base of incisor not marked by such an obvious protuberance as in mordaz. Mandibular and mental foramina essentially as in mordaz. Teeth.—While in no way essentially different from the average of those of Microtus mordax mordax, the teeth of M. m. angusticeps appear to be more uniform in the enamel folding of the crown. The metacone of M2? may (figs. U. V, and X) or may not (figs. W and Y) be open inferiorly ; the internal reéntrant angle on terminal loop may be rudimentary (fig. X), or may be conspicuously developed (fig. V) ; an incipient secondary internal reentrant angle may be present on terminal loop (fig. W). Remarks.—Microtus mordax angusticeps is a depauperate form of the mordax group, with a notably restricted range, being known at present only from a narrow coastal strip of northern California lying between Point St. George and Cape Mendocino. It is most com- monly met with in marshes or in damp redwood forests a few feet above sea level. It is an apparently isolated offshoot, probably pre- vented from free interbreeding with sierrae, with consequent little or no echanee for geographic intergradation, by some disadvantageous intervening condition of territory. Angusticeps resembles both sierrae and mordax in many respects, though certain external characters make it difficult to decide which should be considered the immediate parent stock. In the vicinity of Eureka and Crescent City, individuals of large size have been col- leeted which are unquestionably referable to the mordax group. As regards cranial characters, it will be noted that the skulls are larger and more angular, yet the outlines of the incisive foramina resemble 296 University of California Publications in Zoology [Vou. 21 very closely the typical condition in angusticeps. In comparison with sierrae, the molariform teeth are lighter and the bullae smaller. The angularity of the skull and the shape of the zygomata are in most respects intermediate in character between angusticeps and sverrae. Externally the specimens in question are more richly colored than in sterrae, though the tail is not noticeably bicolor. In size they are as large as old adults of sierrae. According to current usage it is not permissible to recognize two subspecies of the same species in the same region. To regard these large individuals as either sierrae or mordax would lead to the anom- alous situation of two subspecies of the mordax group occurring in the same area and living in the same ecological association, which is hardly in keeping with our present ideas regarding speciation. The most satisfactory disposition of these large specimens would seem to be to refer them to angusticeps and to recognize the fact that they are individual extremes intermediate in character between the latter and sverrae. Microtus mordax bernardinus Merriam San Bernardino Meadow Mouse Microtus mordax bernardinus Merriam (1908, p. 145). Type specimen.—Male adult; no. 150632, Biol. Surv. coll., U. 8. Nat. Mus.; Dry Lake, 9050 feet altitude, San Bernardino Mountains, San Bernardino County, California; August 21, 1907; collected by Vernon Bailey ; original no. 8749. Range.—San Bernardino Mountains of southern California. Ver- tical range from 7500 feet to 9050 feet ; zonal range Transition through Canadian. (See map, fig. CC.) Total number of specimens examined, 47, from the following locali- ties in California. San Bernardino County: South Fork Santa Ana River (8500 feet), 6; Bluff Lake (7500 feet), 41. Diagnosis—Size medium (hind foot, 21 to 23 mm.; condylobasal length of skull in largest individuals, 28.2 mm.). As compared with Microtus mordazx sierrae, skull smaller, zygomata narrower and shorter, brain ease less ridged, and with relatively longer rostrum (ratio of zygomatic breadth to condylobasal length about 57 per cent) and not conspicuously ridged in adults; incisive foramina wide open, not con- stricted posteriorly ; auditory bullae slightly smaller than in sverrae; pelage dark and usually dull; ears smaller than in sverrae (10 to 12 mm)’. Measurements —Average of 10 specimens: Total length, 183.8 mm. (170-198) ; tail vertebrae, 62.6 mm. (53-68); hind foot, 22.4 mm. (21-23) ; condylobasal length, 27.07 mm. (26.2—28.1) ; occipito-nasal 1922] Kellogg: Synopsis of Microtus Mordax 297 length, 27.19 mm. (26.5—28) ; basilar length, Hensel, 23.82 mm. (22.8- 24.8) ; nasal length, 8.31 mm. (7.9-8.8) ; zygomatic width, 15.46 mm. (14.9-16.1) ; interorbital constriction, 3.94 mm. (3.84.1) ; interparie- tal width, 8.65 mm. (8.3-9); shelf of bony palate, 13.5 mm. (12.9- 14.6) ; height cranium at bullae, 9.85 mm. (9.7-10) ; mastoid width, 12.15 mm. (11.8—12.7) ; length upper molar series, 6.6 mm. (6.3-6.9) ; length mandible, 16.48 mm. (16.1—17.3). Color—General hue of dorsal area much darker than in sierrae, the exact shade near snuff brown, but with an evident darkening; middorsum usually overlaid with long aniline black overhairs. In fresh pelage: ground color of upperparts approaching bister but light tipped hairs often as hight as pinkish buff; basal portions of hairs blackish plumbeous. Long overhairs ranging from mars brown to aniline black. Color of upperparts shading into buffy on sides, and this in turn giving way to pallid mouse gray. Underparts pallid neutral gray, with varying admixture of white tipped hairs and with faint suffusion of buff in the inguinal region; anal area nearly pure white. Whiskers for the most part pallid neutral gray, with blackish slate bases. Upper and lower lips white. Ears large, not concealed by fur, with anterior border concealed by hairs of same color as those of upperparts. Tail distinetly bicolor, dull mars brown above and pallid mouse gray below, though tail is often so scantily haired that the yellowish scales show through. Hands and feet covered with pale drab gray hairs, occasionally with hairs as dark as light drab. In worn pelage: much duller, lighter, and with an apparent reduction in amount of long overhairs on upperparts. Young: coloration of upperparts of young individuals very dark as compared with adults because of preponderance of aniline black overhairs, though there is a sheht suffusion of buffy on sides and on‘head. Immature individuals are characterized by the increase in amount of ght tipped hairs, and by the increased size of the grayish patch’on each side of the body. Skull.—tIn all general features the skull agrees with that of sierrae, but brain case is relatively smaller, narrower, and not so convex dorsally. Occiput not abruptly truncate, though occipital condyles are barely visible when skull is viewed from above. (See plate 8.) This race has a rather long and narrow brain ease, slightly rounded at the sides. Postorbital tuberosities usually small, producing no con- spicuous change in contour of skull. Longitudinal ridges along outer edges of parietals well indicated anteriorly, but tending to disappear posteriorly. Interorbital region resembling that of mordax; inter- orbital ridges variable, normally indistinct in interorbital region and with median suleus very shallow if present. Interparietal similar to that of mordax, but with posterior margin biconvex and lateral extrem- ities less obliquely truncate; its anteroposterior width, exclusive of median projection, equal to one-third or more of its greatest trans- verse diameter. Frontals normally broadly coneave posteriorly. The proximal width of rostrum slightly greater than in sverrae. Nasals extending forward to or beyond front plane of incisors and posteriorly not quite so far as ends of ascending branches of pre- maxillaries. Lateral borders of nasals, when: viewed from above, essentially straight with slight constriction anterior to middle. An- terior narial opening, with vertical diameter about equal to transverse. 298 University of California Publications in Zoology (Vou. 21 Anteorbital foramen essentially as in sierrac, though in some cases slightly broader, with inferior termination more angular and _ less rounded. Outhne and depth of occiput when ‘viewed from behind, similar to that of sterrae, the median depth equalling about 56 per cent of greatest width across lambdoidal ridge of brain ease. Foramen magnum essentially the same as in sierrae. Paroccipital processes applied to bullae, their posterior margins continued upward as crests which usually terminate below lambdoidal ridge. Median ridge of basioecipital poorly defined. Auditory bullae usually smaller than those of sverrae, their posterior boundaries slightly indented by fora- mina. Palate similar in many respects to that of sterrae. Palatine sulci well indicated, but with narrow median ridge. Lateral bridges light, walls of lateral pits somewhat fenestrated and pterygoids and hamular processes nearly parallel. Posterior margin of palate usually concave. Interpterygoid fossa shghtly narrower than in sierrae. Posterior pala- tine foramina wide open. Incisive foramina wide open, much more so than in other members of this group, with widest portion usually anterior to middle. Zygomatie arches divergent anteriorly, the maxillary root strongly eurved backward. Jugal relatively heavy, essentially as in sierrae, though usually broader. Zygomatic arches slightly notched at anterior junction with premaxillaries. Mandible light, with relatively deeper horizontal ramus than in slerrae. Ridge for masseteres laterales very well developed. Coronoid process small, short, its extremity not rising above level of condyle, and curved but little backward. Angular process relatively narrow, attenuate, but not curved strongly outward. The articular tuberosity over base of incisor prominent, but included in masseteric ridge, and not lying for the most part above as in other races of this group. Mandibular and mental foramina the same as in sierrae. Teeth Except for their greater size, a character noticeable in both the incisors and molariform series, the teeth do not. differ appre- ciably from those of Microtus mordax sierrae. The metacone may (figs. Z and AA) or may not (fig. BB) be closed inferiorly ; hypocone may fuse with metacone (fig. Z). No incipient internal reéntrant angle was found on terminal loop of M2? in any of the specimens examined. Remarks.—It is of peculiar interest to note the existence of this race in the remote San Bernardino Mountain area. It is probable that the mordax group at one time had a much wider distribution within the state than at present, and there is little reason to doubt that it formerly had continuous distribution from the southern Sierra Nevada to the mountains of southern California, and that geological changes have resulted in the isolation of this form within its present limited habitat. Another point of interest is offered in the bearing of the distribution of Microtus californicus neglectus, which extends 1922] Kellogg: Synopsis of Microtus Mordax 299 in the San Diegan faunal district uninterruptedly from the tidal marshes of the seacoast to the boreal zone of adjacent mountains, upon that of bernardinus. This meadow mouse extends as high up in the San Bernardino Mountains as Bluff Lake, 7500 feet altitude, where it lives within the domain of MW. m. bernardinus. It has been pointed out by Grinnell and Swarth (1918, pp. 348-349) that Microtus cali- formcus (neglectus) occupies, to the exclusion of any other Microtus whatsoever, the ecological niche of Microtus mordax in the more southern San Jacinto Mountains. Possibly mordax once lived in the San Jacinto Mountains, as it does now in the San Bernardino Moun- tains, but it has been replaced in the former by californicus through a process of upward invasion by that species from its adjacent lowland habitat of a much larger extent. 300 University of California Publications in Zoology [Vow. 21 LITERATURE CITED BaiueEy, V. 1898. Descriptions of eleven new species and subspecies of voles. Proc. Biol. Soc. Wash., 12, 85-90. 1900. Revision of American voles of the Genus Microtus. U. 8. Dept. Agric., Div. Biol. Surv., N. Am. Fauna, 17, 88 pp., 5 pls., 17 figs. in text. ELLIOT, D. G. 1903. A list of mammals obtained by Edmund Heller, collector for the Museum, from the coast region of northern California and Oregon. Field Columb. Mus., Zool. ser., 3, pp. 175-197. 1907. A catalogue of the collection of mammals in the Field Columbian Museum. Ibid., 8, pp. viiit 694, 92 figs. in text. GRINNELL, J. 1913. A distributional list of the mammals of California. Proe. Calif. Acad. Sci., ser. 4, 3, pp. 265-390, pls. 15, 16. y GRINNELL, J., and Swart, H. 8. 1913. An account of the birds and mammals of the San Jacinto area of south- ern California, with remarks upon the behavior of geographic races on the margins of their habitats. Univ. Calif. Publ. Zool., 10, pp. 197-406, pls. 6-10, 3 figs. in text. KELLoGe, R. 1922. A study of the Californian forms of the Microtus montanus group of meadow mice. Univ. Calif. Publ. Zool., 21,, pp. 245-274, 25 figs. in text. MeERrRIAM, C. H. 1891. Results of a biological reconnoissance of south-central Idaho. U. S. Dept. Agric., Div. Ornith. & Mamm., N. Am. Fauna, 5, 132 pp., 4 pls., 4 figs. in text. 1908. Four new rodents from California. Proc. Biol. Soc. Wash., 21, pp. 145-148. Miuuer, G. S., Jr. 1912. List of North American land mammals in the United States National Museum, 1911. U. 8. Nat. Mus. Bull., 79, pp. xiv + 455. TROUESSART, HE. L. 1897-1904. Catalogus mammalium tam viventium quam fossilium (Berlin, Friedlinder), fase. 3, pp. 453-664, 1897; Quinquennale Suppl., fase. 2, pp. 289-546, 1904. Transmitted August 5, 1920. ———— nro ee ULE) th ae gry)! ay Okie a EXPLANATION OF PLATE 8 Dorsal and ventral views of crania of four subspecies of Microtus mordaz; X 2. Figs. la, 1b. No. County. Figs. 2a, 2b. No. County. Figs. 3a, 3b. No. tains. Figs. 4a, 4b. No. nardino Mountains. Figs. 5a, 5b. No. Park. Figs. 6a, 6b. No. County. Figs. 7a, 7b. No. Mountains. Figs. 8a, 8b. No. tains. Microtus mordax angusticeps. 11522, J, Mus. Vert. Zool., 3 miles west of Arcata, Humboldt 11530, 3, Mus. Vert. Zool., 3 miles west of Arcata, Humboldt Microtus mordax bernardinus. 6279, J, Mus. Vert. Zool., Bluff Lake, San Bernardino Moun- 6285, J, Mus. Vert. Zool., South Fork Santa Ana, San Ber- Microtus mordax sierrae. 22437, J, Mus. Vert. Zool., Tuolumne Meadows, Yosemite 25052, g, Mus. Vert. Zool., Kings River Cafion, Fresno Microtus mordax mordaa. 11193, J, Mus. Vert. Zool., east face Warren Peak, Warner 11166, g, Mus. Vert. Zool., Parker Creek, Warner Moun- [302] UNIV. CALIF. PUBL, ZOOL, VOL. 21 [KELLOGG] PLATE 8 Seas hy * UNIV: ERSITY | oF CALIFORNIA . PUBLICATIONS (Continnea) “4a, Six. Ne ew Mammals from the: Mohave Desert and Inyo Regions of California, Ries oes DY eséph Grinnell. Pp, 423-430. Por dss Notes on Some Bats from Alaska and British Columbia, by. Hilda bf ; Grinnell.” Pp- 431-433." . Nos. 14 and 15 in one cover. April, OTS eee Sy aera > 604, plates 25-29; -46- figures’ in text: >: May; 1918 a ee ‘17. The Subspecies of the Mountain Chickadee, by Joseph Grinnell. Blb23 figures in text>: Mag< 1918 ok es ee 18. Excavations of Burrows of the Rodent Aplodontia, with Observations on the Habits of the Animal, by Charles Lewis Camp. Pp. 517- 536, 6 figures pe ear hese UG, EGP es os a Ee a oe ee Index, pp, 537-545, ‘ : ‘Vol. 18. 1, Mitosis in Giardia microti, by William Cc. Boeck. Pp. 1-26, plate 1. October, 1917 HELPS ree hy a ge ee ee ee hr re eee ee ee et ee ee ee ee ea ren Bay; California, by Albert L. Barrows. Pp. 27-43. December, 1917 3.. Description of Some New Species of Polynoidae from the Coast of: Cali- fornia, by Christine Essenberg. Pp. 45-60, plates 2-3. October, 1917... New Species of Amphinomidae from the Pacific Coast, by. Christine Essen- berg. Pp, 61-74; plates 4-5..° October, 1917 0... Ss Crithidia euryophthalmi, sp. nov., from the Hemipteran Bug, Euryophthalmus convivus Stal, by Irene ‘McCulloch. Pp. 75-88, 35 figures in text. ABE, DOPE oe oc Viet eee uA ee Bee oN dT BE NS On the: Orientation of Erythropsis, by Charles Atwood Kofoid. ahd Olive Swezy. Pp.-89-102, 12 figures in text. December; 1927 1..2...0000 The Yransmission of Nervous Impulses in Relation to Locomotion in the Earthworm, by John F. Bovard. Pp. 103- 134, 14 figures in text. J anuary, 1918 4, 5. 6 A Bee i eee ee ete eer err 135-144, 1 ficure in text. January, 1918 “ Faéces; by William ©. Boeck.. Pp. 145-149. December, 1917 10. The Musculature of Heptanchus maculatus, by Pirie Davidson. IZhpures Uitext.:- Marchy 1018. se Res ee Oe eee 11. The Factors Controlling the Distribution of the Polyneidae of the Pacific ~ Coast. of North America, by Christine Essenberg. Pp. 171-238, plates 6-8, & fisutes in text. °- March; TOUS 23s so Ae a a ee oe 12. Differentials in Behavior of the Two Generations of Salpa democratica Relative to the Temperature of the Sea, by Ellis li. Michael. Pp. 239-298, plates (0-11. 1 fenre in-texts- arch): 1918" 0 ho ee es a A. Quantitative Analysis of the Molluscan Fauna of San Francisco Bay, by E. L. Packard. | Pp. 299-386, plates 12-13, 6 figures in text. April, 1918 _ 4. The Neuromotor Apparatus of. Huplotes: patella, by Boy B. Yocom. Pp. 837-396, plates 14-16. -“September, 1918° oe The Significance of Skeletal Variations in the-Genus Peridinium, ‘by Aw i. ' Barrows. Pp. 397-478, plates 17-20, 19 figures in text. June; 1918 The Subclavian Vein and Its Relations in Elasmobranch Fishes, by J. Frank te Daniel, Pp. 479-484, 2 figures in text. Amgust, 1918 2200000) c 17. The Cercaria of the Japanese Blood Fluke, Schistosoma. japonicum Katsu- , Z rada, by William W. Cort. Pp. 485-507, 3 figures in text: Fat ns 18. Notes on the Eggs and Miracidia of the Human. Schistosomes, ees WW. Cort. Pp.'509-519,-7 figures in text. Nos. 17 and 18 in ore cover. January, 1919 Index, pp. 521-529. “¥Yol. 19. 1. Reaction of Various Plankton Animals with ee picanti to Their- Diurnal peepee err terre 13. 15. 16. Retrr: tay Migrations, by Calvin O. Esterly. Pp. 1-83. April, 1919 0220... i 2. The Ptéeropod Desmoptcrus pacificus (sp. nov. ), by Christine Essenbers. Pp. Bre THA 85-88 ,2 figures in toe b.) Miay, LOLS cio ee ea ea Aa ge ata tie GEA eee 3. Studies on Giardia microti, by William C. Boeck. Pp. 85-136, plate 4, 19 figures in text. April, 1919 _0.....:... oa te x BR aE ted Rene CaP SNONNR el INCCS gc le ERE . 4, A Comparison of the Life Cycle of Crithidia with that.of Trypanosoma in “the Invertebrate Host, by irene McCulloch. Pp. 135-190, plates 2-8, 3 ‘figures in text.. October, 1919 A Muscid Larva of the San Francisco Bay Region Which Sucks the Blood .of Nestling Birds, by C. BE. Plath. Pp..191-200. February, 1919 22.0... Ss Binary Fission in Collodictyon triciliatum Carter, by Robert Clinton Rhodes. Pp. 201-274, plates 7-14, 4 figures in text. December, 1919. ..22.2.00. 2. The Excretory System of a Stylet Cercaria, by William W. Gort. Pp. 275- 281, 1 figure in text. August, 1919 LA New Distome from Rana aurora, by William W. Cort. Pp, 283-298, 5 neaeen in text. ONE OEES 1019 Cee ee eee ee ern en ees 5. [=>) 7. waceehesedesandustee sebapsrdduegeddecsws ue Jud poncouwswlanccbatadsecasce o eee eee nwa scan tenth een Rv oman dap eaeeb ar haces ae ane enens cesawecs nen eeneeeee® Pp. 151-170, - _ 16. Revision of the Rodent Genus 4 plodontia, by Walter ie: Taylor: Pp, 435-- Pp. 505-:. 20 2, An Unusual Extension of the Distribution of the Shipworm in-San Francisco ‘i Decem- © The Function of the Giant Fibers in Earthworms, by John F. Bovard. Pp.- 9. A Rapid Method for the Detection of Protozoan Cysts in Mammalian © by William: — ape UNIVERSITY OF CALIFORNIA. . PUBLICATIONS —(Continued) 9. The Occurrence of a Rock-boring Isopod along the Shore of ‘San ‘Francisco. 3 a + Bay, California, by Albert L. Barrows. LP. 299-316, ‘plates 15- = Wy Oss Pe. COMPS TOL Oo aoc Ke ie ea a a oe ee ene ered a 10. A-New Morphological Interpretation of the Structure of. Nodtiluca, and its Bearing on the Status of the Cystoflagellata (Haeckel), by Charles A. Kofoid. Pp. 317-334, plate 18, 2 figures in text. February, 1920 02:2... nah 11. The Life Cycle of Echinostoma revolutum (Froelich), by ib ta Cc. Johnson. Pp. 338-888, plates 19-25,-1 figure intext. May, 1920 2.22.20 12. On Some New Myriopods Collected in India in 19i6- by C A. Kofoid, by ss Ralph V. Chamberlin. Pp. 389-402, plates 26-28... August, 1920 2.2...02...-. 13. Demonstration of the Function of the Neuromotor Apparatus in Luplotes — by the Method of Microdissection, by Charles V. RANTS Pp. bgt ty plates 29-33, 2 figures in text. Octoner, 1920... PRE SEES aaNet h UES AE ae Index in preparation. — sy Vol. 20, 1, Studies on the Parasites of the Termites. ° I. On Streblomastiz striz, a Poly- as mastigote Flagellate with a Linear Plasmodial Phase, by Charles Atwood Kofoid and Olive Swezy. . Pp. 1-20,-plates 1-2; 1 figure in text. July, 1919 ~ 2. Studies on the Parasites of the Termites. -T1.,On: Trichomitus termitidis, a -Polymastigote Flagellate with a Highly Developed Neuromotor System, ~ - by Charies Atwood Kofoid and Olive Swezy. Pp. G1-40, plates 3-4, 2° < APUTES ID PEXt. EY A OIO Ge ee BN A Ree a ae tee ae 8. Studies on the Parasites of the Terinitos. TIt. On Trichonympha campanulg sp. nov., by Charles Atwood Kofoid and Olive Swezy. Pp. 4i- 98, plates 5-12, 4 figures in-text, July: 19108 ee ee a ee ee acs 4 Studies on the Parasites of the Termites. IV. On Leidyopsis sphaerica gen. plates 13-14, 1 figure.in téxt.codaly; 191s oo ee ee a ee 6. On the Morphology and Mitosis of Chilomastix mesnili. (Wenyon), a Common -Flagellate of the Human Intestine, by Charles A. Kofoid and Olive SWEET: Pp. 117-144, plates 15-17, 2 figures-in text. April, 1920 <0... ne ne 6. A Critical Review of the Noimenclature of Human Intestinal Flagellates, Cercomonas, Chilomastia, T'richomonas, and Giardia, by Charles A. Kofoid. Pp.445-168,"9 figures in text.< June, LO20. es eee oc ea sence ater 7. On the Free, Encysted, and Budding Stages of i RO ae lafleurt,-a Para. sitic Amoeba of the Human Intestine, by Charles Atwood Kofoid and Clive Swezy. Pp. 169-198, plates 18-22, 3 figures in text.: June, 1921 pss 8. Mitosis and Fission in the Active and Encysted Phases of Giardia enterica (Grassi) of Man, with a Discussion of the Method of Origin of Bilateral ‘Symmetry in the Polymastigote Flagellates, by Charles A. Kofoid and- Olive Swezy. Pp. 199-234, plates 23-26, 11 figures. in text. March, VQ ODS Fa ae gr ee a in, eB ge tae een De OPE. Rane ee Se 9. The Micro-Injection of TEER: by Chas. Wm. Rees. ae 239- 242, PV, VOD Ci Roh ae en cake eS hin Biniea ad Ng ceed ence ae aR cs = Vol. 21. 1. A Revision of the Microtus ealifornicus Group of Meadow Mice, by Reming- ton Kellogg.’ Pp. 1-42, 1 figure in text. December,’ 1918... cok 2. Five New Five-Toed Kangaroo Rats from California, by J oseph Grinnell. reo PHL aS Tey Maren; AOL ce ee ee eR a ae oe es ees 3. Notes on the Natural History of the Bushy-tailed Wood Rats of. California, by Joseph Dixon. - Pp. 49-74, plates 1-3, 3 figures in text. -December, 1919. 4, Revision of the Avian Genus Passerella, ‘with Special Reference to the Dis- tribution and Migration of the Races in California, by H. S. Swarth. Pp. 75-224, plates 4-7,.30 figures in text.” September, 1920 22.00 5. A Study of the California Jumping Mice of the Genus Zaps, by A. Brazier Howell. Pp. 225-238, 1 figure in text... May, 1920. 407 nov., sp. nov., by Charles Atwood Kofoid and Olive Swezy. FP. 99-116, _ pte 6. Two New: Rodents (Genera. Thomomys and Marmota)y from the Bastern Border of California, by Joseph Grinnell. - Pp, 239-244, 6 figures in text. NOVEM HOR 1B 5 ea a ae Be Sata clea niga agac tae apnsees 7. A Study of the Californian Forms of the Microtus montanus Group of Meadow. Mice, by Remington Kelloge: «Pp, 245-274, 25 figures in text. 8..A Synopsis of the Microtus mordax Group of Meadow Mice in’ California, by Remington Kelloge. - Pp. 275-302, plats 8,29 figures in text. Nos; ‘7and:-8:in’ ene cover: April 192R ic oo Se a is aor as Vol. 22.1. A Quantitative and: Statistical. Study of the Plankton of the San Joaquin _-River and Its Tributarles.in and near Stockton, California, in 1913, by Winfred Emory Allen. Pp, 1-292, plates:1-12, 1 figure in text. June, 1920. Vol. 23..-The Marine Decapod Crustacea of California, by Waldo Ll. Schmitt. Pp, 1-470, plates 1-50, 165 figures in text. May, 1920 2.2. es Vol, 2¢. 1..A Geographical Study of the Kangaroo Rats of California, by Joseph ~ Grinnell. Pp. 1-124, plates 1-7, 24 figs. in text. bickctay OO 2) Se ae tae eS one as»: UNIVERSITY OF CALIFORNIA PUBLICATIONS g IN ZOOLOGY Vol. 21, No. 9, pp. 303-312 ; '- February 1, 1923 BEHAVIOR OF THE LEAF-NOSED SNAKE, PHYLLORHYNCHUS DECURTATUS BY SARAH ROGERS ATSATT UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA 1923 UNIVERSITY OF CALIFORNIA PUBLICATIONS - Note.—The University of California Publications are offered in exchange for the publi- cations of learned societies and institutions, universities, and libraries. Compiete lists of all the publications of the University will be sent upon request. For sample copies, lists ~~ of publications or other information, address the MANAGER OF THE UNIVERSITY PRESS, BERKELEY, CALIFORNIA, U.S, A. All matter sent in exchange should be addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY, CALIFORNIA, U.S.A. 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As there is no published account of the habits of this leaf-nosed snake, the writer took the opportunity of making observations on this individual. Ultimately the snake was preserved (no. 8174) in the collection of the Museum of Vertebrate Zoology of the University of California. ‘The sealation of the snake is essentially as given by Cope (1900, p. 823). The large rostral plate is the basis for both the generic name, Phyllorhynchus, and the common name, leaf-nosed. The short tail, one-tenth of the total length of the animal, is another diagnostic character and makes the specific name of decurtatus appropriate. While the snake was alive the pinkish cast due to the blood showing through the translucent skin was noticeable. The effect was similar to that commonly seen in the banded gecko and other animals that live in the dark. After the snake was preserved, the ground color became dead white and made the pattern of brown spots on the back more prominent than in life. Whenever the snake became warm and active a pink spot on the head between the occipitals grew more warmly pink. In the dead snake this spot was translucent, not dead white. 304 University of California Publications in Zoology [V0u. 21 LOCOMOTION If the snake was left in a box of sand open to light it spent most of the time buried in the sand. When dug out and placed on top of the loose sand it soon reburied itself. In getting into the sand the large rostral plate served as a resisting surface while the head was being pushed in. At the same time the head was moved from side to side. The are of this swing was usually not so long as the head itself. The ‘‘neck’’ or portion just back of the head was held a little arched above the ground. The course of the head when once completely under the sand rarely continued in one direction; it might even be doubled back under the rest of the body which was following through the original entrance to the sand. The direction pursued seemed to be entirely at random except as the snake was turned aside by resisting objects or by harder areas of earth. Many times; by random movements, the head appeared above the sand before the body was entirely in or while part of the body was still going under the sand. At various times this emergence was due to the shallowness of the sand, or to the compactness of the sand, or to the fact that the snake did not make a sufficiently sharp angle down- ward when entering the sand. The snake took only a brief time to get under the sand. When especially observed in this regard on April 13, the animal had been active for twenty minutes in response to warmth and light from an electric bulb and had made nine attempts to bury its head and one attempt to dig a hole. At 3:39 p.m. the head was started under in the loose earth. A minute later, with its head partly under, the snake rested. Then the head was covered and the snake rested again. The head was then worked in deeper. By 3:42 the last part of the hody had been pulled under by intermittent slight motions but at the same time the back part of the head and the neck became exposed. Though the head was thrust in deeper it soon reappeared. In a few minutes continued activity resulted in the snake working wholly out of the sand. Later that afternoon, after various periods of activity and rest, the snake was again exposed to the electric light. In three minutes it hid itself in the sand except for a small section near the tail. This time it remained quiet under the surface. On another occasion, on April 16, the snake was transferred to faint 1923 | Atsatt: Behavior of the Leaf-nosed Snake 305 sunlight after forty-five minutes of activity during exposure to lght. The snake crept into the sand in three and a half minutes, resting three times during that period. Ten minutes later the snake took only two minutes to get entirely into the sand except for an inch of the tail. Often, attempts to hide in the sand were not completed and the head only was covered. And often, successful burying of itself in the sand was not followed by any period of quiet and so the animal became uncovered again. A few times the snake was observed to use another method of eetting into the sand, namely, by digging a hole. When the snake was originally found on the desert floor, it was under a rock in a small depression with a burrow leading to it from the ground surface. On April 18, while the snake was being observed with relation to heat and light, it made four attempts to creep into the fairly compact shallow sand. Each time, the head was covered but was raised again to the surface. Then the snake scooped a hole by turning its head to the right and piling up the sand. This action was not repeated there, but in another spot the animal worked its head in and then up again to the light. When the earth was loosened by the observer, the snake successfully worked its way into the sand within the next few minutes. This method of digging a hole was used again on April 15 after the snake had been exposed to light for about forty minutes, part of the time to direct sunlight and part of the time to sunlight through a blue filter. The snake made attempts to bury itself in the sand, but each time the head reappeared in the light. A small stone was put in the shadow on one side of the box. Twice the snake started to creep under the stone but did not get its head covered. Then it dug a hole by loosening the sand and turning its head to one side and pulling back the anterior half of its body so as to form loops. The sand held between the head and first loop of the retracting body was dropped when the snake straightened out again. The snake used either right or left side; for instance, right, left, right, right, left, right. With its head at the bottom of the hole thus started, it finally erept into the sand. On April 16 the snake was stimulated by blue light. After seven- teen minutes of rather constant motion, including attempts to get out over the top of the box and three attempts to cover its head, the snake started at 4:03 p.m. to dig in the southwest corner in a shadow east by the side of the box. Either the posterior third or the middle portion of the body was held tight on the ground. The snake con- 306 University of California Publications in Zoology (Vou. 21 tinued to dig until 4:09 and then it moved slowly along the west side of the box. It continued to move around trying the earth and the glass cover until, at 4:13, it again started to dig in the southwest corner. Digging lasted two minutes. After more traveling around the box it dug two more minutes in the shadow. Meanwhile the sun had moved so that only a small patch of sunshine was left in the southeast corner. At 4:23 the snake moved slowly down into the southwest corner; when two minutes later the blue filter was lifted the snake was found two-thirds coiled in the depression it had made. On April 21, under blue light, the snake made several attempts to cover its head, then dug once to the right in the northwest corner. Twelve minutes later, at the middle of the side of the box, the snake dug to the right and to the left. It tried’ twice to cover its head in the sand, then dug again to the right and moved off. Six minutes later in the southwest corner it dug to the left, moved a trifle, and dug to the right. A minute later it dug in the northwest corner once to the left and tried to bury its head. This was the last time the snake was seen to dig. This cireum- stance may have been due to the fact that the extreme stimulation of blue light was not used again. The first attempt, however, on April 13, was made under electric light. More likely the loss of the habit was due to the growing weakness of the snake, for the digging required more effort than merely creeping into the sand. If a larger rock had been put in the box soon after the snake was eaptured and the sand left entirely undisturbed the snake might have made a hollow under the rock. In loose sand the snake could bend its body freely and back up rapidly. On April 11 when an attempt was being made to measure the length accurately the snake was persuaded to creep into a glass tube which was shghtly greater in diameter than the animal’s body. The tube was put over the head and the snake poked until it moved in to over two-thirds of its length. Then it suddenly began to back. The tube was turned vertically with the snake’s head downward; still the snake backed out. When two-thirds of the body was out a finger of the observer was placed under the tail. With that leverage the snake finished the backing out process rapidly. When the animal was on a smooth table it moved only with difficulty. The snake coiled rarely. Only once was it observed in that posi- tion and then it was in a loose coil. Usually it was not even looped unless it was in a small ean in which it did not have room for stretch- ing out more or less at full length. 1923 | Atsatt: Behavior of the Leaf-nosed Snake 307 RESPONSE TO STIMULI The snake’s movements seemed little controlled by a positive reaction to contact. To be sure, it traveled along the edge of the box rather than across the open surface of the sand but occasionally it did go across. When the thermometer was in the box it glided over this object or between it and the nearby side of the box. The snake crawled under a rock, under a piece of palm leaf and out again, but it crawled over smali strips of glass so fixed that they offered easy opportunity to crawl under them. Light here seemed to play a more important part than contact in determining the course of progress. One problem was to determine whether the activities of the snake were controlled by response to light, or to heat, or both. The negative response to direct sunlight was very marked. When the reptile was first found, shortly before noon, it was in a depression under a rock and hence in the dark. When it was transferred to a tin box with holes punched in the top it hid itself in the sand. On April 4, on a warm afternoon when the open box of sand was half in sunshine and half in shadow, the snake always turned away when it found itself in the direct sunshine. It would pull back its head sharply when only half an inch of the head extended into the area of sunlight. Occasionally it would get more into the light but it always went back promptly into the shade. Finally it buried itself in the sand. On the next day when no direct sunlight fell on the sand a flat mirror was used to throw a beam of light on the snake’s head. The snake responded immediately by turning away to get out of the spot of light. It did this repeatedly and finally buried itself to get rid of the annoyance. On April 18 record was kept of the snake turning back sharply from bright sunshine twenty-five times in as many minutes. The response was as consistent at the end of that period as at the beginning. Furthermore, when the snake was kept in a ean. with only a few holes in it and placed in a cool dimly lighted room, the animal was quiet on the surface of the sand. It was invari- ably found on the surface when the cover was lifted either in the daytime or at night. An experiment on combined temperature and hght control was undertaken on April 13 in a room with dim daylight. An electric bulb with reflector was placed seven inches above the can containing 308 University of California Publications in Zoology [Vou 21 the snake. In eight minutes the temperature of the air rose from 28° C to 34°. The animal made frantic attempts to bury itself in the sand in the usual manner and it also attempted to dig. When the hght was snapped off the snake remained quiet on the surface of the sand. After the temperature had dropped several degrees the hght was turned on again. The temperature rose half a degree in the quarter of a minute that elapsed before the snake began to move. Again the light was turned off and the animal immediately became quiet. When the temperature had dropped lower than at the start the light was turned on. The snake became active and continued active until it seemed to be becoming exhausted. Again the light was turned off. When next the light was turned on, the snake buried itself in the sand. The following day an effort was made to test the effect of tem- perature alone by using a dark room with red light and a box with warmed sand at one end and cool sand at the other. The sand at the warm end varied because of cooling from 47.8° C to 41° and at the cool end from 27.5° to 25.3°. The snake moved around actively and showed no differences in reaction on warm and cool sand. When the red glass was withdrawn so that electric light came on the snake ° from above and to the right the snake generally withdrew from the light into the shadow east by the right end of the box. It did not always do so; occasionally it crawled into the light. When it was transferred to cooler sand, with the warm end 37.6° to 34.2° and the cool end 27.5° to 26.4°, the snake showed no reaction to temperature. As long as the white light was on, the animal spent more time in the shadow than in the light. If it got its head into the light it withdrew it. It made few attempts to get into the ground at either the warm or the cool end but tried to get out over the low wall of the box. On April 18 in a box of sand partly in direct sunshine and partly in shadow the snake consistently drew back from the sunshine. The can was turned at intervals so that the warm sand was in the shadow and the cool sand in the sun. The snake moved into the shade and thereby into much warmer sand. On April 15 the effect of light was studied while the factor of temperature was eliminated as far as proved possible. Over the box of sand was placed a erystallizing dish containing a blue solution (alkaline copper sulphate) three or four inches deep, to cut out the red heat rays. The temperature of the sand in sunshine was 37° C; under the blue light it was 33.5° at the start and 31° twenty minutes ee 1923 | Atsatt: Behavior of the Leaf-nosed Snake 309 later. The temperature of the air under the blue light varied from 30° to 28°. The snake crawled around actively, attempting to get into the sand and trying the glass at the corners several times. Then it was allowed to get out of the box and into strong sunlight on the table. It moved into the shadow east by the blue filter and traveled along in the neighboring shadow east by a book. Next, the snake was put into a box of sand in which the temperature ranged from 26° to 27° under the blue filter. For the next fourteen minutes the animal moved around trying to get out. When the blue filter was taken off, the temperature of the sand rose eight or nine degrees in twenty-five minutes, while within one minute of taking off the blue light the snake tried to burrow in the sand in the shaded portion. It kept for the most part in the shade; if it got into the sunshine, in a second it quickened its rate of locomotion. It dug a hole by the side of a small rock and crawled under and beyond it into the sand, where it remained quiet. ; On April 16 this experiment was repeated, but with the tempera- tures lower. The temperature of the sand under the blue filter varied from 18° to 19.5° and of the air from 19° to 21°. During the time of experimentation, from 3:46 to 4:35 p.m., the snake was in constant activity. It wandered around trying to get out, especially at the corners, or trying to bury its head in the sand. In general it kept at the edge of the box either near the thermometer at the east end or more often in the shadow in the west third of the box. Oceasion- ally it went diagonally across the sunny area. After two attempts at digging a hole it partially coiled itself in the hole it had dug in the southwest corner in the shade. On April 21 the blue filter was placed over a box of sand warmed at one end to 42° and at the other to 24°. The snake itself had been in sand. at 36°, in the dark. Under the blue light it displayed great activity and moved into the shaded end which happened to be the cool end. After about six minutes of comparative quiet spent in the shade, it began more rapid activity, and continued for twenty-five minutes. It made attempts to burrow, to dig, and to get out over the corner of the box. There was no apparent difference in the amount of time spent in the cool and warm ends of the box, but most of the time was spent in the shadow along the south and west sides of the box. When the snake was put in cool sand of 24° in direct sunshine, the same type of movement continued. When it was allowed to creep over the edge of the box to the table it crawled into the shade east 310 University of California Publications in Zoology [Vou 21 by the box, under some paper, back to the box, and so on. It was frantie when it got from the shade directly into sunshine. The tem- perature in the sun was 36° and in the shade about 28°. The reaction was so quick that the snake itself could not have become warmer. The snake was rather exhausted when put back into sand of about 30° and the cover put on the can. There it remained quiet on top of the ground for over an hour. Although these experiments do not completely eliminate heat as a controlling factor, it is evident that light is a much stronger control. The action under the blue filter, when the heat rays were partially cut out, was the same as in direct sunshine. The temperature under the blue filter made no difference in reaction, for on April 16 when the temperature was rather low, between 18° and 21°, the snake acted exactly as it did every time under the blue filter. It showed signs of extreme stimulation and made frantic endeavors to get out of the box or into the sand; it even made attempts to dig a hole, and event- ually tried to coil in one of the depressions scooped out. Moreover, the responses to direct sunlight were so rapid as to make it improb- able that they were due alone to the quickening of muscular and nervous activity as a result of a rise in temperature of the snake. Again, the snake showed no objection to sand artificially heated con- siderably above the temperature of the sunlight to which it was exposed. FOOD AND WATER Various kinds of food were offered the snake, but all solid food was consistently rejected. Since the snake must be out on the surface of the ground in dim light only, and since it is small in size, the most probable food would seem to be insects. Ants were tried in vain. When an ant crawled on the snake, the snake gave evidence of panic and thrust its head into the sand. When ants carrying pupae and therefore unable to bite were placed in the can the snake tried to get away by burying its head. Green aphids, gray aphids, woolly aphids, mealy bugs, house flies, fruit flies, larvae and adults of mealworm beetles, ladybird beetles, caterpillars, small moths, doodle-bugs, sow- bugs, and spiders were tried. They brought no response except appar- ent fear and efforts to get out of the way. Raw meat, cooked meat, old odoriferous meat, and cheese offered no inducements. Vegetation aroused no interest. The snake was too small to be fed forcibly with 1923 | Atsatt: Behavior of the Leaf-nosed Snake oalut solid or liquid food. Ditmars (1912, p. 219) says, ‘‘The feeding habits of Phyllorhynchus are practically unknown, but judging from structure, it apparently feeds upon small lzards, other snakes and possibly the young of the smaller rodents.’’ Unfortunately this indi- vidual was too small for such food. eh On April 16 the snake refused water when a small drop was put on its snout. That drop was wiped off by movements of the head in the sand. Another drop caused excited burrowing of the head in the corner of the box. On April 21 when the snake was exhausted from exposure to blue lght it paid no attention to warm water dropped on its snout. On April 22 the snake was moved from dim light where it had been over-night to a shaded spot out of doors. A small drop of water of about 30° was placed on the rostral plate. At first there was no response; then the drop was wiped off by the snake against the side of the dish. Another drop was placed on the snout. The snake stuck out its tongue slightly, then withdrew it. Then with no movement of the tongue outside of the mouth the snake drew that drop into its mouth. Next it took seven more small drops, an amount equal to three large drops from an ordinary pipette. With the succeeding drop hanging from its mouth the snake moved away and finally the drop touched the sand and was absorbed. The animal refused more water but ran its tongue out freely as it crawled along. The snake started to burrow and covered its head several times slowly and easily. If drops of warm milk or of beaten egg were put on the rostral plate of the snake they were rubbed off if possible. If the animal were placed on the observer’s hand so that its head and neck hung below, it could not reach anything on which to rub its snout. Twice in this position it got a few drops of beaten egg by licking them off of its rostral plate with the tongue in an effort to get rid of the offend- ing substance. On June 6 the snake took a little water from a pipette. On June 16 it drank water from the surface of a hand. On June 21 it drank with its head held high in the air; there were two periods of wiping off its snout and of swallowing rapidly. On June 24 more water was taken. On June 30 the snake was observed to open its mouth twice. When given water it drank some quickly and followed this by frequent swallowing and a few deep breaths. In the preparation of this paper the writer acknowledges her indebtedness to Dr. Loye Miller and to Dr. Joseph Grinnell. 312 University of California Publications in Zoology [Vou 21 SUMMARY The first specimen of Phyllorhynchus decurtatus reported from California lived in captivity from March 25 to July 20, 1921. The snake showed a marked tendency to hide itself in the sand. It could creep in by side movements of the head and body. Complete burial by this process required from two to three and a half minutes. It made several attempts to dig a hole by scooping out a hollow in the sand. Apparently it ean dig a burrow for itself under a rock. Light rather than temperature seemed to be the controlling factor leading it to hide in the sand. Under electric hght and in direct sunlight the snake drew back into shadow. Under red light the snake showed no reaction to differences in temperature. When the heating effects of sunlight were cut out to a large extent by using a blue filter the snake showed marked stimulation, regardless of high or low tem- perature or of contrasted temperatures. Insects crawling on the body and liquid placed on the snout also brought about the reflex of hiding in the sand. Food was persistently rejected except for some beaten egg, and that was licked off the snout and swallowed only when the snake could not reach a surface against which to rub it off. Several times a few drops of water were drunk from the end of a pipette or from a surface. LITERATURE CITED ArsatT, S. R. 1921. A snake new to California. Copeia, no. 96, 38-39. Cork, HE. D. 1900. The crocodilians, lizards and snakes of North America. Ann. Rep. U.S. Nat. Mus., 1898, pt. 2, pp. 151-1294, 36 pls., 347 figs. in text. Dirmars, R. L. 1912. The feeding habits of serpents. Zoologica, 1, 197-238, 13 figs. in text. Transmitted November 17, 1922. UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) 41. The Life Cycle of Echinostoma revolutum (Froelich), by John C. Johnson. Pp. 338-388, plates 19-25, 1 figure in text. May, 1920 ...-. wes 12. On Some New Myriopods Collected in India in 1916 by C. A. Kofoid, by _ Ralph V. Chamberlin. Pp. 389-402, plates 26-28. August, 1920 us $a, 18. Demonstration of the Function of the Neuromotor Apparatus in Euplotes by the Method of Microdissection, by Charles V. Taylor. Pp. 403-470, plates 29-33, 2 figures in text. October, 1920" Pear ee EE See eS Index, pp. 471-480. - Vol. 20. 1. Studies on the Parasites of the Termites. I. On Streblomastiaz ate a Poly- mastigote Flagellate with a Linear Plasmodial Phase, by Charles Atwood Kofoid and Olive Swezy. -Pp. 1-20, plates 1-2, 1 figure in text. July, 1919 2. Studies on the Parasites of the Termites. II. On Trichomitus termitidis,-a Polymastigote Flagellate with a Highly Developed Neuromotor System, by Charles Atwood Kofoid and Olive Swezy. Pp. 21-40, plates 3-4, 2 © HPUTOR ING TEX Gis PLY A LO ee EO Ee ae a mane 8. Studies on the Parasites of the Termites. ITI. Ox Trichonympha campanula sp. nov., by Charles. Atwood Kofoid and Olive Swezy. Pp. 41-98, plates B22 4 fiosiresiny text Anda ys VOLO eT ral ee ee 4, Studies on the Parasites of the Termites. IV. On Leidyopsis sphaerica gen, nov., sp. nov., by Charles Atwood Kofoid and Olive Swezy. Pp. 99-116, plates 13-14, 1 figure in text. July, 1919 ..._.. ON et eR RGN dle Aa ee th 5. On the Morphology and Mitosis.of Chilomastiz mesniit (Wenyon), a Common Flagellate of the Human Intestine, by Charles A. Kofoid and Olive Swezy. Pp. 117-144, plates 15-17, 2 figures in text. April, 1920 sve 6. A Critical Review-of the Nomenclature of Human Intestinal Flagellates, Cercomonas, Chilomastiz, Trichomonas, and Giardia, by Charles A. Kofoid. Pp.2445-168;-9:fisures:in texts, “June 1920 ose sks Sinn posed iwntnniseon sce fee 7, On the Free, Encysted, and Budding Stages of Councilmania lafleuri, a Para- sitie Amoeba of the Human Intestine, by Charles. Atwood Kofoid and Olive Swezy. Pp. 169-198, plates 18-22, 3 figures in text. June, 1921 __. 8. Mitosis and Fission in the Active and Encysted Phases of Giardia enterica (Grassi) of Man, with a Discussion of the Method of Origin of Bilateral Symmetry in the Polymastigote Flagellates, by Charles A. Kofoid and Olive Swezy. Pp. 199-234, plates 23-26, ll figures in text. “March, gL 0 12 17 RC ris Si ide igs oi eek bas as tint caine OL RK tab: Syste aah OBER Oia Mean os Clee PN t ee Nit 9, The Micro-Injection of Paramaecium, by Chas. Wm. Rees. Pp. 235-242, April,.1922) 20... PESCSS RENN LEN AR I Ser Sree Paget eee hg IETS Ue aes OU, aha acmenead de ee 10. On Balantidiwm coli (Mailmsten) and .Balantidium suis (sp. nov.), with an account of their neuromotor apparatus, by J. Daley McDonald. Pp. 243-300, plates 27, 28, 15 figures in text. May, 1922 2. 11. Mitosis in Endamoeba dysenteriae in the Bone- Marrow in Arthritis de- formans, by Charles Atwood Kofoid and Olive Swezy, Pp. 301-307, 7 figures in text. 12. Endamoeba. dysenteriae in the Lymph Glands of Man in Hodgkin’s Disease, by Charles A. Kofoid, Luther M. Boyers, M.D., and Olive Swezy. Pp. 309- 312, 4 figures in text. ~ Nos. 11 and. 12.in-one eever.: (April; 1922 oo ee 18. Mitosis in the Encysted Stages of Hndamoeba coli (Loesch), by Olive Swezy... Pp. 313-332, plates 29-31/\. May, 1922 ooo canna cece senses cheep tse aee 14, The Neuromotor Apparatus of Paramaecium, by Charles William Rees. Pp. 333-364; plates 32-36, 5 figures in text, November, 1922... 15. A Comparison of the Cysts of Hndamocha coli and Councilmania lafleurt in Congo Red, by Nathaniel Bercovitz, M.D, Pp. 365-371. February, 1923. Vol.21. 1. A Revision of the Microtus californicus Group of Meadow Mice, by Reming- ton Kellogg. Pp. 1-42, 1 figure in text. December, 1918 ....2. 2. 2. Five New Five-toed Kangaroo Rats soe California, by Joseph Grinnell. GRE OSE By NL «ll IS Racha ee em stp Re ne TS ee ec dere ee eA chien A 3. Notes on the Natural History of the Bushy-tailed Wood Rats of California, by Joseph Dixon. Pp. 49-74, plates 1-3, 3 figures in text, December, 1919 4, Revision of the Avian Genus Passerella, with Special Reference to the Dis- tribution and Migration of the Races in California, by H. 8S. Swarth. Pp. 75-224, plates 4-7, 30 figures in text. September, 1920 ..............-...---..-..--.-. 85 Py {a UNIVERSITY OF CALIFORNIA PUBLICATIONS—(Continued) . A Study of the California Jumping Mice.of the Genus Zapus, by A. Brazier Howell. Pp. 225-238, 1 figure in text. May, 1920 ~..2222 2000. o eee ee 6. Two New Rodents (Genera Thomomys and Marmota) from the Eastern Border of California, by Joseph Grinnell. Pp. 239-244, 6 figures in text. INGO WOM Dears LO Dare oe TS A cee edna Biprac ean pacastgn tte Saeit lesssthecsonnaed 7. A Study of the Californian Forms of the Microtus montanus Group of . Meadow Mice, by Remington Kellogg. Pp. 245-274, 25 figures in text. 8. A Synopsis of the Microtus mordax Group of Meadow Mice in reams. by Remington Kellogg. Pp. 275-302, plate 8, 29 figures in text. Nos. 7 and’ 8th: one*cover. ~ April, 1922 as a a 9. Behavior of the Leaf-nosed Snake, Phyllorhynchus decurtatus, by Sarah Rogers Atsatt.. Pp. 303-312. February,: 1923 ©... oe. ence ence cee eenecccentenee ao 10. A Systematic List of the Mammals of California, by Joseph Grinnell. Pe SIS-SL44; J ANUARY 5 LOGS eo aa ats canons dadinn we bbe apna cotingst te : Sa) fers = ; i _ . + A ae rae - . pga | . ws * 7 /* . ' University or CALIFORNIA PUBLICATIONS, In ZooLocy Vol. 21, No. 11, pp, 325-332, plates 9-10 Issued April 7, 1923 THE SYSTEMATIC STATUS OF THE MOUNTAIN LION OF CALIFORNIA BY JOSEPH GRINNELL anp JOSEPH DIXON (Contribution from the Museum of Vertebrate Zoology of the University of California) As far as we can learn from the literature no one has hitherto made any special effort to determine the systematic status of the mountain lions inhabiting the rather distinct biotic division of western North America known as the Californian fauna. C. Hart Merriam, in his important revisionary paper (1901), refers to California only incidentally (p. 590) as included in the geographic range of his Felis hippolestes olympus. Satisfactory material was evidently not then available for determining the exact relationships of the Californian animal. The collection gathered during recent years in the California Museum of Vertebrate Zoology now warrants attention to the subject, and as a result of some considerable study the present writers find that there are good grounds for recognizing a new race, as follows: Felis oregonensis californica May, revived name California Mountain Lion Felis concolor (Richardson, 1839, p. 6), and of many subsequent authors. Felis Californica (May, 1896, p. 22): first use of this name. Felis oregonensis (Merriam, 1899, p. 104), and of some other authors. Felis hippolestes olympus (Merriam, 1901, p. 590), part, as far as California is concerned. Felis oregonensis oregonensis (Grinnell, 1913, p. 297), and of some other authors. Basis of present description——Male, mature adult, skin and com- plete skeleton; no. 31252, Mus. Vert. Zool.; ‘‘Lynchburg’’ [more exactly, 4300 feet altitude at head of Brushy Creek, a branch of the Middle Fork of the American River, 814 miles east-southeast of Michigan Bluff], Placer County, California; July 21, 1920; collected by Jay C. Bruce and Joseph Dixon; orig. no. 7924, J. Dixon. Diagnosis.—A eat of the Felis concolor group, nearest related to Felis oregonensis oregonensis Rafinesque (see Stone, 1899, pp. 34-35). Differs from the latter form in slightly paler tone of coloration and in some restriction of dark markings, but most especially in cranial 326 University of California Publications in Zoology [ Vol. 21 features, as follows: Frontals not bulging dorsally but flattened, and with no marked median sulcus (see Merriam, 1901, p. 590); nasals longer and nasal profile sloping gently, not ‘‘humped’’; jaw less massive and less undershot; skull throughout, smaller than in oregon- ensis: dentition weaker ; canines notably slenderer. (See pls. 9, 10.) Measurements.—Of male, no. 31252, data as above: Total length, 6 feet, 614 inches; tail, 2 feet, 6144 inches; hind feot (from heel), 11 inches; ear, height from crown, 334 inches; height of animal at shoulder, 2814 inches. Of female, no. 31251, probably mate of preced- ing, same place, July 16, 1920 (orig. no. 7923, J. D.): Total length, 6 feet, 414 inches; tail, 2 feet, 6 inches; hind foot (from heel), 11 inches; ear, height from crown, 314 inches; height of animal at shoulder, 2714 inches. For cranial measurements, see accompanying table. Comparisons should be made with figures given for Felis oregonensis hippolestes by Roosevelt, 1901, p. 435, and Merriam, 1901, p. 587; for F. o. oregonensis by Swarth, 1912, pp. 98-99; and for F. 0. browni by Grin- nell, 1914, p. 253. Distribution—The metropolis of this subspecies lies within the state boundaries of California. Its range appears to be limited on the east by the sage plains of the Great Basin and by the Mohave and Colorado deserts. To the south it occurs as far at least as the Cuyamaca Mountains of San Diego County and it is possibly this form that extends south into the San Pedro Martir Mountains. To the northward, we do not know what happens beyond the Oregon line; but we think it quite likely that mountain lions formerly occupied the territory continuously across Oregon into Washington and British Columbia and that complete intergradation between californica and oregonensis took place. We have no material at all from Oregon. Life zone, chiefly Transition. Comments.—Thirty skulls of mountain lions from California have been before us at one time for study. The first impression gained from a survey of this series is that a very great amount of variation exists. But persistent study brings to the fore the facts that (1) there are great differences between the sexes, and that (2) aging is accompanied by conspicuous changes in proportions as well as by general massiveness of skull. Taking fully into account these two "factors, sex and age, we come to the precise conclusion that Merriam (1901, p. 579) did, namely, that ‘‘the limits of variation for adults of each sex fall within surprisingly narrow bounds.’’ We have read closely the criticism by Elliot (1904, pp. 454-455) of Merriam’s ‘‘Revision.’’ It is curious that anyone with the oppor- tunities and wide experience that Elliot may be supposed to have enjoyed could have so failed to apprehend the facts. We take this occasion to defend Merriam’s paper as an admirable piece of system- atic work, one which involved a very careful and correct distinguish- 1923 | Grinnell-Dixon: The California Mountain Lion 327 ment of phylogenetic characters (those of real systematic value) from the products of age and sex variation so generally confused with the phylogenetic characters by some other writers on the Felidae. It was Mr. Gerrit S. Miller who discovered that there is an old name available for the California Mountain Lion. In the ‘‘ Addenda’’ to his List of North American Land Mammals in the United States National Museum, 1911 (1912, p. 400) he writes: Page 117, in synonymy of Felis oregonensis oregonensis Rafinesque, insert: 1896. ? Felis californica May, California Game ‘‘marked down,’’ p. 22. (Kern County, California.) We have had opportunity of examining a copy of the publication in question, which was issued for advertising purposes by the Southern Pacific Company. In response to an inquiry, Mr. Chas. S. Fee, Passenger Traffic Manager of the Southern Pacific Railroad, wrote us under date December 15, 1919, that he believed the author to be Dr. Wm. B. May, who in 1896 was employed in the advertising de- partment of the Southern Pacifie Company. Someone is likely to question the propriety of using a name printed so casually, and unaccompanied by any characterization such as would link it up definitely with the subspecies in question. In answer, we would point out the following facts. On page 22 of the pamphlet cited there appears a half-tone photo- graph (58 X 91 mm.) of a mounted specimen of mountain lion. Beneath this half-tone is the caption: California Lion. Felis Californica. This photograph and caption fall within a section (pp. 20-25) headed ‘‘Kern County.’’ On page 21 we read: ‘‘The mountain district furnishes ideal haunts for bear—grizzly and black, brown and cinnamon, with California lions and other feline beasts of prey, not to mention wolves and coyotes.’’ The specimen photographed was probably destroyed in the San Francisco fire of 1906. Thus, a binomial name for the mountain lion which inhabits the Sierra Nevada of California has been published (in the nomenclatural sense) ; and our present view is that this name must be employed for the subspecies we here characterize. [ Vol. 21 University of California Publications in Zoology [ 328 a[Apuod 07 sisaydurdAs ‘Mee OI ocoONn OG WG IES Giz WENGE WAWA = Nee i GZ | Olen | Stews zz leogt poe 9 FF GS KOaAL | 4) te US HG ME Tel 4, Ooi? 7G 19 €1 | €8 10 6S (Ge NEIL WT 4 AS; Go |Z VI | c& |Z 0S (OM 'AN) i' Ss g mi 5 25 ob 2. bs me | ee | Se | = Bool sel 4 3 a al & OO & Yon} LD SH LO Yes) Yen) wd QS 0 oo =) uw ud (IPE. zUINI4SO1 a © mon Wd — *A[LO1I9}S0d019},UB UOISUBUIIp 4S94veIF ‘AUO YS OUT, + “UIN[NSUID 4B A[IOTIe4ySOdo1a4uB paiNsva ‘9UO YYSI IY], ¢ ee —— ——————eaererere | “LE |T'L9 |9°ZL |0'6Z1\¢°89 |L SFTlg Zor “98 1€°89 1€°6Z |G OZTIE' SZ |O ESS TLT ‘28 1Z'69 10°64 |L' E211 OL |9 LEIP FOT GSE VE Ode lees as 9 6ZII8 EZ |9 SSTI¢ SLT ‘Gp | FL 19°S8 O'SFIIZ SZ |Z SOTIe Sst IP |2°12L |9°28 |% VPLS" 22 |0°E9T|0 ZEST PY |8°SL 10°88 lO 'SSTie Sz [629712 98T GP |Z LL |¢'16 [O'SFLIL ZS 8 ZLOTIE 681 OF |8°S9 18°88 | TFIIE 08 0° SZTI8 O61 rr IL TL 12°98 |[€ PHI TS |€ EZ1/9 TEL ge |e | oe | ce | ce | ce | oe es | 2ea/ §s | Ss ial ee | ae So ee eee ei hobo: Benlmerea rem heros | : ae |) B3 : ye z g LLY 681 est Z81 806 £06 I1G S16 606 80G [gneei Y}Su9] 4SeqRIITDH ‘OUTUED JO OSB YL Fulyjams Jo youq ysnez ‘G61 “4 ‘CGT ‘seo Aq pouyep sy 1 09 ourpreusag ug “sj OuIpieusog ueg ‘youRy s,xo9 Shee eect e eee e eee e en eeeeeseeereee 09 BsOdeyy “BUOM® AA Henne eee nee ta wetness aee onsen (oye) GSOdIIv Ay “BUOMB \A eee e cece ccc etesessneteceenesssssece 09 I9d¥[ J ‘SINQyOUATT 09 ourpasuieg ueg “Sq OUIpPIBUIEg UBg ‘UOAUBD YoeID TIN “OO BIBGIBG BIUBG ‘OYB'T BOT YSBO “TUT ae i O- esoduvyy ‘BUOME AA JSOM “TUT %4 J pee eee eee ects eeessarscsssecenssne re) Ide q ‘SINQUYOUATT Wee eee eee ener ener nena enter e nee ane , Ayunog vyseyg,, sicreeee og RysRYyg “BIg pNo[HoT YSve “TUT ZT ALITVOOT Or OF OF OF = ct Ho} Roo oye Xo) x99 SPIL PS9EST ES9ES ISZIé VESTS SZESE GSPLES GSGIE 1Z6P 8988 ‘ZAW ‘ON VOINYOMITVO SISNHYNODAYO SITAY LINGY AO STTAYG AO (SUALUNITMYY NI) SLNGWAXASVA] 1923 | Grinnell-Dixon: The California Mountain Lion 329 LITERATURE CITED ANONYMOUS [=May, W. B.]. 1896. California game ‘‘marked down.’’ (San Francisco, Southern Pacifie Co.), 64 pp.,"58 illus. (half-tone photographs); 12mo (173 X 129 mm.). Ewuiot, D. G. 1904. The land and sea mammals of Middle America and the West Indies. Field Columb. Mus., zool. ser., 4, part 2, xiii+441-850 pp., pls. XULII-LXVIII, text-figs. 83-167, LIX—CXLII. GRINNELL, J. 1913. A distributional list of the mammals of California. Proc. Calif. Acad. Sei., ser. 4, 3, 265-390, pls. 15, 16. 1914. An account of the mammals and birds of the lower Colorado Valley, with especial reference to the distributional problems presented. Univ. Calif. Publ. Zool., 12, 51-294, pls. 3-13, 9 figs. in text. MERRIAM, C. H. 1899. Results of a biological survey of Mount Shasta, California. U. S. Dept. Agric., Div. Biol. Surv., N. Am. Fauna, 16, 1-179, pls. 1-5, 46 figs. in text. 1901. Preliminary revision of the pumas (Felis concolor group). Proce. Wash. Acad. Sei., 3, 577-600 (December 11, 1901). MILLER, G. 8., Jr. 1912. List of North American land mammals in the United States Na- tional Museum, 1911. U.S. Nat. Mus., Bull., 79, xiv + 455. RICHARDSON, J. 1839. Mammalia [pp. 1-8] in The zoology of Captain Beechey’s Voyage . . . performed in his majesty’s ship Blossom. (London, Henry G. Bohn), 180 pp., 47 pls. ROOSEVELT, T. 1901. With the cougar hounds. Scribner’s Magazine, 30, pp. 417-435 (Octo- ber, 1901), idem, pp. 545-564 (November, 1901) ; many photographic illus. STONE, W. 1899. The pumas of the western United States. Science, n.s., 9, 34-35 (January 6, 1899). SwarTH, H. S. 1912. Report on a collection of birds and mammals from Vancouver Island. Univ. Calif. Publ. Zool., 10, 1-124, pls. 1-4. THOMAS, O. 1905. Suggestions for the nomenclature of the cranial length measurements and of the cheek-teeth of mammals. Proce. Biol. Soe. Washington, ile}, IS also. Transmitted January 13, 1923. PLATE 9 Fig. 1.—Lateral view of skull of Felis oregonensis oregonensis, adult male; no. 12453, Mus. Vert. Zool.; Nootka Sound, Vancouver Island, British Columbia; August 11, 1910. X .55.. Fig. 2.—Lateral view of skull of Felis oregonensis californica, adult male; no. 31252, Mus. Vert. Zool.; Lynchburg, Placer County, California; July 21, 1920. Note in particular, as compared with above, the straighter frontal and nasal outlines, the less massive jaw, and the slenderer canines. X .55. [330] ——— VINIVenGAL ES PUB, ZOOL VOR 21 [ GRINNELL—DIXON ] PLATE 9 Ld al A) 2 CENTIMETERS Fs] cy ra % PLATE 10 Fig. 3.—Anterior view of skull of Felis oregonensis oregonensis, adult male; no. 12453, Mus. Vert. Zool.; Nootka Sound, Vancouver Island, British Columbia; August 11, 1910. X .60. Fig. 4.—Anterior view of skull of Felis oregonensis californica, adult male; no. 31252, Mus. Vert. Zool.; Lynchburg, Placer County, California; July 21, 1920. Note, as compared with above, the flatter top of the skull, the lesser mass of bone at the base of each postorbital process, lesser width of rostrum, the less massive jaw, and the slenderer canines. X .60. [332] ee UNIV CALE RUBE, ZOOL VOLE. 24 [ GRINNELL—DIXON } PLATE 10 2) 4 LJ | pn) ad = Be Zz ermemnnen (NY bad v UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 11. The Life Cycle of Echinostoma revolutum (Froelich), by John Cc. Johnson. Pp. 338-388, plates 19-25, 1 figure in text. May, 1920 00.000 12. On Some New Myriopods Collected in India in 1916 by C. A. Kofoid, by : Ralph V. Chamberlin. Pp. 389-402, plates 26-28. August, 1920 ........0... 18. Demonstration of the Function of the Neuromotor Apparatus in Hiplotes . by the Method of Microdissection, by Charles V. Taylor. Pp. 403-470, plates 29-33, 2 figures in text. October, 1920 ...o.cc..cccecccl cc ceceecceeccececcesdeccees ze Index, pp... 471-480. ~ Vol. 20, 1, Studies on the Parasites of the Termites. I. On Streblomastix striz, a Poly- ‘ mastigote Flagellate with a. Linear Plasmodial Phase, by Charles Atwood Kofoid and Olive Swezy. . Pp. 1-20, plates 1-2, 1 figure in text. July, 1919 _2. Studies on the Parasites of the Termites. II..On Trichomitus termitidis, a Polymastigote Flagellate with a Highly Developed Neuromotor System, by Charles Atwood Kofoid and Olive Swezy. Pp. 21-40, plates 3-4, 2. fisuressin: fext.<' duly, 2010~ -October, 1999 jes NE en eee ee .60 5. A Muscid Larva of the San Francisco Bay Region Which Sucks the Blood of Nestling Birds, by O. E. Plath. Pp. 191-200. February, 1919 ..._......... Bxaesmy |] 6. Binary Fission in Collodictyon triciliatum Carter, by Robert Clinton Rhodes. Pp. 201-274, plates 7-14, 4 figures in text. December, 1919 ...-....2...2.00.. 4.00 7. The Excretory System of a Stylet Cercaria, by William W. Cort. Pp. 275- 2e1; 1 fivure in:text. “Ausust,'1919 22s Se ee a ee 10 8. A New Distome from Rana aurora, by William W. Cort. Pp. 283-298, 5 figures in:-text.: November, 1919 on a ee 20 9. The Occurrence of a Rock-boring Isopod along the Shore of San Francisco Bay, California, by Albert L. ati ca Pp. 299-316, plates 15-17. De- Comber, =1919 i ee eee oe tee Le A SN ae ASS Ee 25 10. A New Morphological Interpretation of the Structure of Noctéluca, and Its Bearing on the Status of the Cystoflagellata (Haeckel), by Charles A. Kofoid. Pp. 317-834, plate 18, 2 figures in text. February, 1920 ........ SS NEW SUBSPECIES OF BIRDS FROM PATAGONIA BY ALEXANDER WETMORE UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Vol. 21, No. 12, pp. 333-337 Issued June 16, 1923 NEW SUBSPECIES OF BIRDS FROM PATAGONIA BY ALEXANDER WETMORE Comparison of birds from an excellent collection secured in northern Patagonia by Mr. J. R. Pemberton with specimens available in the U.S. National Museum has revealed four geographic forms not hitherto recognized. These forms are described in the present paper. The Pemberton collection, made during the course of a geologic survey of parts of the territories of Rio Negro, Chubut, and Santa Cruz, has been deposited in the Museum of Vertebrate Zoology of the University of California. The type of one of the subspecies here characterized comes from the series in the institution just mentioned. The other three are based on skins in the U.S. National Museum taken in the Straits of Magellan by naturalists on the U.S. Fish Commission steamer ‘‘ Albatross.”’ Pteroptochos rubecula nemorivaga subsp. nov. Characters.—Similar to Pteroptochos rubecula rubecula Kittlitz (1831, p. 179), but decidedly darker ; upper surface, flanks, and under tail coverts darker brown; rufous brown of chest and head deeper. Description—tType, U.S. Nat. Mus. Cat. No. 116272, male, from Port Otway, Straits of Magellan, Chile, collected in February, 1888, by naturalists of the U.S. Fish Commission steamer ‘‘ Albatross.’’ Crown slightly darker than mummy brown; hind neck, back, secapulars, and rump mummy brown; upper tail coverts Prout’s brown; remiges fuscous; wing coverts and outer margins of flight feathers Prout’s brown; rectrices fuscous, margined externally with a wash of Prout’s brown; superciliary streak, extending from base of nasal fossa, cover- ing space before eye and then extending back along sides of crown, hazel; space beneath eye to ear coverts sooty black, the feathers below eye cinnamon-buff basally and on ear coverts streaked narrowly along shafts with whitish; sides of head behind and above ears deep mouse gray; chin cinnamon-buff; throat and upper breast between tawny and russet, becoming hazel on sides of neck; feathers of lower breast and abdomen deep mouse gray basally, barred narrowly with dull black and dull white, the latter with a wash and intermixture of cinnamon-buff ; sides deep mouse gray, barred sparingly like breast ; 334 University of California Publications in Zoology [Vou. 21 flanks Prout’s brown, toward abdomen barred indistinetly with blackish ; under tail coverts Prout’s brown; lesser under wing coverts deep mouse gray; greater under wing coverts mikado brown; edge of wing mixed mikado brown and mouse gray. Bill dull black, washed obscurely with brownish at base of mandible; tarsus and toes verona brown (from dried skin). Measwrements.—(In millimeters): Male (one specimen, type), wing 73.2, tail 64.0, exposed culmen 14.9, tarsus 38.4. Female (one specimen), wing 69.0, tail 57.5, exposed culmen 14.5, tarsus 37.6. Two specimens with sex not indicated, wing 66.6-72.2, tail 56.6— 67.2, exposed culmen 14.7-15.5, tarsus 37.0-39.3. Range.—Straits of Magellan, Chile (known from four specimens from Port Otway). Remarks.—The present form is based on four skins, two of which bear date of February 10, 1888, secured by naturalists of the. “* Alba- tross’’ at Port Otway, in the Straits of Magellan. Though the date on two of these specimens is not indicated it is certain that all four were collected at the same time. Kittlitz described Pteroptochos rubecula from a male obtained at Concepeién, Chile. In making com- parisons in the present connection I have considered a series of four skins from Valdivia as representative of the typical form. Tenioptera pyrope ignea subsp. nov. Characters.—Similar to Tenioptera pyrope pyrope (Kittlitz [1831, p. 191]), described from Tomé, Concepcién, Chile, but decidedly darker gray above, with little or no olivaceous wash; breast duller gray; under wing coverts darker. Description.—Type, U. S. Nat. Mus. Cat. No. 116246, collected at Laredo Bay, Straits of Magellan, January 20-22, 1888, by naturalists of the U.S. Fish Commission steamer ‘‘Albatross.’? Crown, hind neck, back, and wing coverts deep mouse gray; rump and upper tail coverts mouse gray; feathers of crown with narrow, indistinct, shaft streaks of black; lores and forehead grizzled indistinetly with grayish white; primaries, secondaries, and secondary coverts dull black; secondaries margined faintly with pale olive-gray to whitish; tail mouse gray with shaft stripes of black and an outer edging of whitish to pale olive-gray ; auricular region deep mouse gray mixed with storm gray ; throat white with shafts of feathers black ; breast dull pale smoke gray, with a faint wash of drab; abdomen and under tail coverts white ; sides grayish olive; tibiae deep mouse gray ; under wing coverts mouse gray, washed with drab. Bill, tarsi, and toes black (from dried skin). Named with reference to the fiery hue of the eye. Measwrements.—(In millimeters): Males (11 specimens), wing 109.7-117.6 (112.2), tail 85.0-93.6 (89.9), culmen from base 17.0-18.8 (18.1) [average of 10 specimens], tarsus 26.7-29.6 (28.0). 1923 | Wetmore: New Subspecies of Birds from Patagonia 335 Female (one specimen), wing 111.5, tail 89.0, eulmen from base 17.3, tarsus 28.2. Type (male), wing 110.0, tail 87.3, culmen from base 17.0, tarsus Pee Range.—Straits of Magellan (Port Otway and Laredo Bay) north to Lake Nahuel Huapi, Rio Negro. (Pacific Slope opposite head of Rio Chico, Santa Cruz; Rio Fetaleufu, and Puesto Burro, Maiten, Chubut; Bariloche, Rio Negro.) Remarks.—The present form is described from thirteen skins, from the localities listed under the geographic range, which have been com- pared with nine skins of typical pyrope from Santiago and Concon, Chile. Birds in the Pemberton collection from western Chubut and Rio Negro are paler than those from farther south, but are nearer ignea than the typical form. Pyi ope Kitthtz, Cabanis (1859, p. 45), a name that has been applied to this flycatcher, is a synonym of Muscicapa pyrope Kittlitz. Turdus magellanicus pembertoni subsp. nov. Characters.—Similar to Turdus magellanicus magellanicus King (1851, p. 14), described from the Straits of Magellan, but much paler both above and below; under surface grayish. Description—tType, Museum of Vertebrate Zoology, Cat. No. 43065, adult female, from Cerro Anecon Grande, Rio Negro, Argen- tina, collected January 7, 1912, by J. R. Pemberton (orig. No. 126), in somewhat worn breeding plumage. Crown, lores, and nape between fuscous and fuscous-black ; forehead bleached to brownish drab; sides of head, hind neck, back, and wing coverts between drab and hair brown, shading to mouse gray on rump; upper tail coverts hair brown; remiges, primary coverts, alula, and rectrices dull black; outer margin of secondaries mouse gray; throat whitish streaked with fuscous-black, save for a small area on lower throat which is immaculate; sides of neck, upper breast, sides, and flanks paler than drab; lower breast and abdomen washed with pinkish buff; under tail coverts white, bordered with mouse gray; tibia light drab washed with buff on inner side; under wing coverts light drab, bordered indistinctly with a wash of pinkish buff; inner webs of primaries bordered faintly with light drab. Bill cream-buff; tarsus and toes Isabella color (from dried skin). Measurements.—(In millimeters): Males (two specimens), wing 128.2-129, tail 88.6—92.6, culmen from base 23.8, tarsus 38.4-39.0. Females (two specimens), wing 126.0 (type)—127.5, tail 95.7-96.7 (type), culmen from base 21.5-22.4 (type), tarsus 35.4-36.7 (type). Range.—Gobernacion de Rio Negro, Argentina, except in moun- tainous western portion (Cerro Anecon Grande, and General Roca), perhaps north into the province of Mendoza. 336 University of California Publications in Zoology (Vou. 21 Remarks.—This robin is known at present from the Rio Negro at Roea, and from a tributary of the Rio Limay at Anecon Grande. Two skins in spotted juvenal plumage when compared with typical magel- lanicus of a similar age offer the same difference of pallid color that distinguishes pembertoni in adults of the two forms. An old specimen in the U.S. National Museum from the Gilliss Expedition marked ‘‘Chili’’? may have come from Mendoza, since many of Lieutenant Gilliss’ birds were taken in that province. The bleached form of the Magellanic robin here described seems to inhabit willow groves along streams in the more arid regions to the eastward and northward of the forested section of the southern Andes. True Turdus m. magellanicus is found in the more humid region at Lake Nahuel Huapi, and extends from there southward through the Andes to the Straits of Magellan. Hellmayr (1921, p. 238) considers 7. magellanicus a form of 7’. falcklandu from the Falkland Islands, but to me the two species seem to be sufficiently different to merit their recognition as distinct species. Spizitornis parulus lippus subsp. nov. Characters.—Similar to Spizitornis parulus parulus (Kittlitz [1831, p. 190]), described from Conecepeién and Valparaiso, Chile, but decidedly darker, less olivaceous, on back and rump; black of head duller; breast somewhat more abundantly streaked. Description—Type, U. S. National Museum, Cat. No. 116157, adult (sex not indicated), collected at Mayne Harbor, Evans Island, Owens Islands, Chile, lat. 51° 19’ S, long. 74° 7’ W (approximate), February 5, 1888, by the naturalists on the U.S. Fish Commission steamer ‘‘Albatross.’’ Crown, sides of head, and crest black, the erown streaked rather indistinctly with white; nasal tufts partly white; a white spot at upper anterior corner of eye; back, rump, and scapulars between hair brown and chaetura drab; wing coverts and alula dull black, the greater and middle coverts with indistinct lighter tips of slight extent; primaries and secondaries chaetura black; primaries edged with pale olive-buff to whitish; secondaries with proximal part of outer webs black; posterior two-thirds and tip bordered lightly with olive-buff; rectrices chaetura black, outer webs margined lightly with pale olive-buff ; throat and extreme upper breast whitish, streaked finely and indistinctly with black; lower breast, abdomen, and sides washed with pale yellow; center of abdomen immaculate; sides and breast streaked indistinctly with blackish ; under tail coverts whitish, with small concealed dusky spots; under wing coverts pale yellowish. Bill, tarsi, and feet black (from dried skin). Named with reference to the peculiar coloration of the eye in members of this species, where a purplish segment occupies the upper third of the otherwise pale iris. 1923] Wetmore: New Subspecies of Birds from Patagonia oa Measurements.—(In millimeters): Males (two specimens), wing 47.2-48.2, tail imperfect, culmen from base 8.4-9.0, tarsus 18.5-19.5. Type (sex not indicated), wing 49.7, tail 50.2, culmen from base 9.2, tarsus 18.4. Range.—Straits of Magellan (Punta Arenas and Mayne Harbor). Remarks.—The dull coloration of the dorsal surface distinguishes this form from other described subspecies of Spizitornis parulus as it is darker even than S. p. wquatorialis (Berlepsch and Taeza- nowski [1884, p. 296]), from Eeuador and Peru. It agrees with parulus and differs from patagonicus Hellmayr (1920, p. 51) from Argentina in lack of well-marked wing bars. Two specimens of lippus from Punta Arenas, Chile, taken January 27, 1888, with tails imperfect through molt, are similar to the type from Mayne Harbor. In the type specimen part of the crest is missing. The three mentioned from the Straits of Magellan have been compared with ten skins of typical parulus from Concon, Valparaiso, and Coneepecién Bay, Chile. It is supposed that S. p. lippus ranges through the humid region of extreme southern Chile. LITERATURE CITED BERLEPSCH, H. DE, ET TACZANOWSKI, L. 1884. Deuxiéme liste des Oiseaux recueillis dans 1’Ecuadeur occidental par MM. Stolzmann et Siemiradski. Proc. Zool. Soc. London, 1884, 281-313, pl. 24. CABANIS, J. 1859. In Cabanis und Heine, Museum Heineanum, 2, 2 + 176. (Published September, 1859.) HELLMAYR, C. E. 1920. Kin Beitrag zur Ornithologie von Siidost-Peru. Arch. f. Naturg., 85, 1919 (published November, 1920), 1-131. 1921. Review of the birds collected by Alecide D’Orbigny, Part IT. Novitates Zoologicae, 28, 230-276. (Published September, 1921.) GING) Pas: 1831. Characters of new genera and species of birds from the Straits of Magellan. Proc. Zool. Soe. London, pt. 1, 1830-31, 14-16. (Pub- lished January 6, 1831.) KirrTuitz, F. H. von 1831. Ueber einige Vogel von Chili, beobachtet im Marz und Anfang April 1827. Mém. Acad. Imp. Sci. St. Petersbourg, 1, 174-194. Transmitted March 28, 1928 UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued) 11. The Life Cycle of Echinostoma revolutum (Froelich), by John C. Johnson. Pp. 338-388, plates 19-25, 1 figure in text. May, 1920 0000 12. On Some New Myriopods Collected in India in 1916 by C. A. Kofoid, by .. Ralph V. Chamberlin. Pp. 389-402, plates 26-28. August, 1920 2.0... =e 18. Demonstration of the Function of the Neuromotor Apparatus in Huplotes by the Method of Microdissection, by Charles V. Taylor. Pp. 403-470, ~ plates 29-83, 2 figures in text. October, 1920-..2.0.220 = Index, pp. 471-480, Vol 20. 1. Studies on the Parasites of the Termites. I. On Streblomastiz striz, a Poly- mastigote Flagellate with a Linear Plasmodial Phase, by Charles. Atwood Kofoid and Olive Swezy. Pp. 1-20, plates 1-2, 1 figure in text. July, 1919 2, Studies on the Parasites of the Termites. II. On Trichomitus termitidis, a Polymastigote FlageHate with a Highly Developed Neuromotor System, by Charles Atwood Kofoid and Olive Swezy. Pp. 21-40, plates 3-4, 2 Agures. in text, iwiulys TOL Ae a a ee ee ee ae oH a eS eT oh 8. Studies on the Parasites of the Termites. III. On Trichonympha campanula sp. hov., by Charles Atwood Kofoid and Olive Swezy. Pp. 41-98, plates ; S-i2< 4 foures-in ‘tex. cd tly Ol Onc Coe) SN ee 4. Studies on the Parasites of the Termites. IV. On Leidyopsis sphaerica gen. nov., sp. nov., by Charles Atwood Kofoid and Olive Swezy. Pp. 99-116, plates 13-14,-1 figure in. text: July, 1019 yk a ee Se ee 5. On the Morphology and Mitosis of Chilomastic mesnili (Wenyon), a Common Flagellate of the Human Intestine, by Charles A. Kofoid and Olive Swezy. Pp. 117-144, plates 15-17, 2 figures in text.” April, 1920 2.0... -22. tee 6. A Critical Review of the Nomenclature of Human Intestinal Flagellates, Cercomonas, Chilomastiz, Trichomonas, and Giardia, by Charles A. Kofoid. Pp; 145-168, O-figures in-text. - June; 192040 ee Oe es - J. On the Free, Encysted, and Budding Stages of Couwncilmania lafleuri, a Para- sitic Amoeba of the Human Intestine, by Charles Atwood Kofoid and — Olive Swezy.. Pp. 169-198, plates 18-22, 3 figures in text. June, 1921 _. 8. Mitosis and Fission in the Active and Encysted Phases of Giardia enterica (Grassi) of Man, with a Discussion of the Method of Origin of Bilateral Symmetry in the Polymastigote Flagellates, by Charles A. Kofoid and Olive Swezy. Pp. 199-234, plates 23-26, 11 figures in text. March, BS AAT Bip Ren Olan gr en De cee a 4 UATE HCl RI PA Nea rr Mrs Sal De CORE, 9, The Micro-Injection of Paramaecium, by Chas. Wm. Rees. Pp. 235-242. April LOD 2 a a ee EG NE SP Rt green IS IE eS 10..On Balantidium coli (Malmsten) and Balantidiwm suis (sp. nov.), with an account of their neuromotor apparatus, by J. Daley McDonald. Pp. 243-300, plates 27, 28, 15 figures in text. May, 1922 02.000 11. Mitosis in Endamoeba dysenteriae in the Bone Marrow in Arthritis de- formans, by Charles Atwood Kofoid and Olive Swezy. Pp. 301-307, 7 figures in text. 12. Endamoeba dysenteriae in the Lymph Glands of Man in Hodgkin’s Disease, by Charles A. Kofoid, Luther M. Boyers, M.D., and Olive Swezy. Pp. 309- 312, 4 figures in text. INos.11-and:-12 in one-cover,.~ April 1922: 62. et 13. Mitosis in the Encysted Stages of Hndamoeba coli (Loesch), by Olive Swezy. Pp.-313-332, plates 29-31..-.May, 1922 coon. ecccc eect leceeepaccee 14, The Neuromotor Apparatus of Paramaecium, by Charles William Rees. Pp. 333-364; plates 32-36, 5 figures in text. November, 1922..........0002... 15. 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May, 1920. 2.2... une 6. Two New Rodents (Genera Thomomys and Marmota) from the Bastar Border of California, by Joseph Grinnell. Pp. 239-244, 6 figures in text. TNO VERIO 21 2 hin eee n 5 ewe ec aoc erro nace > een eae cb ocak cundeens bodte ne se=naperoneunah=saeeicnsocen 7. A Study of the Californian Forms of the Microtus montanus Group of Meadow Mice, by Remington Kellogs. Pp. 245-274, 25 figures in text. 8. A Synopsis of the Microtus mordaz Group of Meadow Mice in California, by Remington Kellogg. Pp. 275-302, plate 8, 29 figures in text. Nos.°7 and 8 in one cover. — April, 1922 1910-1912, 446 pages, with 12 is Saute acca Sucncsameeanere 1911, 357 pages, with 25 plates 2200263 ~ Cited as Uniy. Calif. Publ. Zool. RSA 5 Sel eee a ae 1911-1912, 865 pages, with 24 plates... SS Pa eae Rede Jan SOD ene me: 1912-1913, 1912-1914, Volume 12, 1913-1916; _¥Yolume 13,. 1914-1916, Volume’ 14, 1914-1918, Volume 15, Volume. 16, . 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Notes on the Natural History of the Bushy-tailed Wood Rats of California, "x by Joseph Dixon, Pp. 49-74, plates 1-3, 3 figures in text. December, 1919. “2B. eek ae Revision of the Avian Genus Passerella, with Special Reference to the Distribution and Migration of the Races in-California, by. HS. Swarth. ea ‘Pp. 75-224, plates 4-7, 30 figures in text. September, 1920 -...0<....c--10: 1.75 estes . A Study of the California Jumping Mice.of the Genus Zapus, by A. Brazier oo) 5 56o> Two New Rodents (Genera Thomomys and Marmota) from the Eastern - Border of California, by’ Joseph Grinnell. “Pp. 239-244, 6 figures.in text Howell. Pp, 225-238, 1 figure im text. May, 1920 oc ecnteencece | “36 . Noveintien JOR SU eee a a a cee Ae CY 7. A Study of the Californian Forms of the Microtus montanus. ‘Group. of Fa Meadow Mice, by Remington Kellogs. Pp. 245-274, 25 figures in text. 8, A Synopsis of the Microtus mordax Group of Meadow Mice in California, by Remington Kellogg, Pp. 275-302, plate 8, 29 figures in text, Nos. 7 and 8 in one cover. 9. Behavior of the Leaf-nosed Snake, Phyllorhynchus decurtaius, by. Sarah . 2 Rogers Atsatt. 10, A’ Systematic List of the Mammals of eadizornis,. by Joseph “Grinuell. Pp. 313-324. 11, The. Systematic Status.of the Mountain Lion of California, by Joseph (April: $002 Seco Ben a oo a oe Pp. $03-Si2; February 21028 soe ee ace 125 - : Warinary.AQ2S oe ee OU ee ee Re no Maa »~ Grinnell and Joseph Dixon. Pp. 325-332, plates 9-10. April, 1923 ws 25 ee 12. New Subspecies of Birds from Patagonia, by, Alexander ‘Wetmore: SEB: Shae eet SBS—337.~ PuUMe, BOBS soa sa erawee e an tevathet ce Sacer connegtn Astana Shere bannegs cs ebetonsed a esa 13. Revision of the Genus Lynx in California, by Joseph Grinnell and Joseph Dixon. Pp. 339-354, plate 11, 1 figure in text. January, 1924.00.) 25 14, Changes during Growth in the Skull of the Rodent Otospermophilus gram- . murmus beecheyi, by E. Raymond Hall. Pp. 355-404, AB. figures in text. ‘ ited Men Agee ee NOR OR MTA ene ne ae. aBecmeeer nc Heo BO. US 15. A-New Race of the White-breasted Nuthatch from. Lower. California, by Joseph Grinnell, Pp. 405-410. March, 1926. UAT so Sa eo ah Nae SS “2B. CHANGES DURING GROWTH IN THE SKULL OF THE RODENT OTOSPERMOPHILUS GRAMMURUS BEECHEYI BY E. RAYMOND HALL UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Volume 21, No. 14, pp. 855-404, 43 figures in text Issued March 9, 1926 UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA CAMBRIDGE UNIVERSITY PRESS Lonpon, ENGLAND »s CHANGES DURING GROWTH IN THE SKULL OF THE RODENT OTOSPERMOPHILUS GRAMMURUS BEECHEYI BY E. RAYMOND HALL CONTENTS PAGE NRE TG LUIGI eee eee ete te he ot ae. ons Bele tml bo aly cued VEAL A te adr oe 356 Nev Cl aYGn cr FESO aa NIST ger aes eee nA, S16 he eT a eR AON IO. = CEA De Rec ee,© 356 WAT eRIDI Seer Ree ee eer settee Gee eit te eee ryt de Bd eae LENE be mY, 1 | 357 Individual skulllelements.:-2-4.-.....ne-eeiee ee + Lsrhostes, No lata mre Us kane) tems ei 358 INAS) Seeeeen es ee er eee fe eR cet Ne ROM ST is pt te NN a 358 Premaxillae........... RR ee ees NI af oc Le ns oe Ee ae OM OMe SiR Te Oe ed 358 Viet eve ieee secrete ee PRN ceca ce ctigs eye tee eee) A ERA ie One: Senne Me LS 359 IIsfaV eFC) IC a 2a eee. Le eee a, OL: Reece SO ony ALE eC PeR ERR 360 Ont alse ewer reek ee eee. Disk bee oe dosh c Ea ER RO eee Ne DEL 361 PATI Lal Sea eet core Mea pre ele ANI MT old Hoy oh ecole 2 ee me oral ve CR get 367 SSC LUE LATO SLSR Bet Reena ere eee et a eae EE tds eke 8, Ra ee Seas 370 “UCI IS Rate a elle een a eR RRR PRM RRO. tlle ot aR Re | 371 (Oye orig fe} 01 AVS TON (0 ESL a gate ea a eR nen ERAN Daisy, 2 Seo feature 9 371 Alisphenotds = .2....0..5: Fe PUA ena oe eee Pes eRe ave Le PERO Ee ct SI) alates eee ee Ee eke de ahh Ba Dl so ee MeN neha WS etar elm ds 87/8) J EAR EU ETE CH (OSV cl a ee ee ee eee RN Se, Cee NE DS ee PP Do 374 Wome nrancl te tim Old Siesteet vie serge crs ene ee aa ete ae me See Can 375 PRE SPACING Clemente ee CA Sat Ske, See VS ute sae ARETE Ans Sc ee 376 | BSEVEVLS}C) OEY CVC)ING Sia whee na sta ne a eer ean pr RT er EROS URL RRE RIN Noh 377 Basioceipitals 0. 4204./..400h0. A Dae Se Mae in seen ere Ue oe oe 377 EE Sip R UU ee ee Pye ean Src a aie the atarge ia vo Auceenn | ees be nora ne eRe 378 SSI PDILEL@ CG APSE het 8 cee ios Seid Se Weel vant eget ees taa ol eae Rt A 379 Jey in Sigf OPH SDI SE Me seme Le i ae oe aa eae a eed Re eet | Maw, CLE 379 CESTATI CSN UT GSEs ott, ARN GR RS ORE Meteo oN Petr Saar 6 Uap eee ME A Eee Sete CORNEA racials 381 TERE ERODE 2 0 lta OO cel SUS OO alee MEI Se OY See EES) eee ee ee 384 PMMTANTA My 1OS SUCLE Sian t Ne an tote tee che ca e120 tore eS Sock nn SR Ue ee et 385 AV UGiri Chill | @Weeeereeeerk She Meena tel lek saw Ook a Aaa Loae oad n pee Lek aie ae Se aR 386 A DEYSIH al gates bs OEE noe oe Oe eRe a OT PR ARTES NE NT Rae OLE RED! ted eat Aa) lls 390 (Sramiummgsrs enews Ol Cas researc ee ee TM eee ee Sede ne ee 391 TORT HER PATENT eee Bae ol ae Rone a ee NRE oc ye Se aA RRR ay Mere aed ammeter ek i 400 Siuclvaoimanl ane eGRseriess.tea ce kee ere oe eae Reeder Ey Ee Ree Soe 401 TIGA GUN HCIILE Cen een aN ec od SE Dene eer enemy a neas Uk. Watt Ae Cte IL cee cd 404 356 University of California Publications in Zoology [ Vol. 21 INTRODUCTION Changes taking place during postnatal growth in the bones of the rodent skull seem to have received little attention either from the systematist, who is primarily concerned with adult specimens, or from the embryologist, who has seldom continued his observation on the development of the skull beyond embryonic stages. In examining various rodents it has become evident to the writer that certain bones in the skulls of different genera differ widely as regards the relative amount of growth attained and the age at which this growth and the changes dependent thereon take place. It is the main purpose of the present paper to describe the postnatal changes that occur with growth in the bones of the skull of one particular rodent, the California Ground Squirrel, Otospermophilus grammurus beecheyi, of which satis- factory material happens to be available. Descriptions and figures of some embryonic conditions are also given. Upon comparison of these data with similar data for other forms it is hoped that generalizations of some phylogenetic as well as of ontogenetic significance may be arrived at. ACKNOWLEDGMENTS The author is indebted to Mr. C. D. Bunker, of the Museum of Natural History of the University of Kansas, and Mr. Donald R. Dickey, of Pasadena, California, for the loan of specimens to supple- ment those in the California Museum of Vertebrate Zoology; to Dr. C. A. Kofoid, of the Department of Zoology, University of California, for the use of a series of embryos; to Dr. Joseph Grinnell, Director of the Museum of Vertebrate Zoology, for the use of the collections in his charge and for guidance and detailed assistance while the work was under way. Dr. C. L. Camp has read the final manuscript critically. Figures 1 to 27 were made by my wife, Mary H. Hall, to whom acknowledgments are due as well as for much other assistance rendered in the preparation of this paper. 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 35 ~ MATERIALS Approximately 600 separate skulls, 175 skulls-with-skins, and 25 litters of embryos of the California Ground Squirrel have been avail- able for study ; nearly all of this material is contained in the California Museum of Vertebrate Zoology. In dealing with changes that occur with growth, only males have, as a rule, been used. The exceptions were specimens nos. 34289 and 34292, Mus. Vert. Zool., which are females of the exact ages of which no males were available. The measurements given of these two young skulls would not, it is believed, differ from the same measurements taken of male skulls of the same age, Since comparisons of male and female skulls slightly older showed no differences due to sex. Considerable pains were taken in selecting the male skulls believed to be typical of different ages. The entire series was arranged in probable order of age from youngest to oldest, using as criteria the degree of development of the various skull elements. Measurements taken to 0.1 millimeter were freely used in selecting specimens most typical in all respects of the different ages. Table 2 gives certain cranial measurements, ratios of these measurements to oecipito-nasal length, and amounts of increase and per cents of increase for each measurement, of 14 males selected in the above manner as well as for 2 females and for the average of 8 very old males. For convenience of text reference, letters A to Q are used for these skulls in place of the Museum catalogue numbers. All catalogue numbers given in this table and elsewhere in the text refer to specimens in the collection of the Museum of Vertebrate Zoology. A represents the skull of a 40 mm. embryo and B of a44 mm. embryo. C represents the skull of an animal found dead 6 days after birth, in the nest of a litter of young born in captivity. It was badly decomposed and represents a stage at birth or within a very few days thereof. D stands for the skull of an animal 7 days old, and F of an animal 15 days old. WN, and those designated by preceding letters, represent skulls of animals less than one year old. O, P and the series of skulls of 8 very old males designated as Q refer to skulls of animals more than one year old. The ratios for the different measurements are the ratios to the occipito-nasal lengths. For example, to obtain the ratio of the palatal leneth of A the following equation is made and solved for x, which in this ease is found to be 56.3. 11.2 (occipito-nasal length) : 100 :: 6.3 (palatal length) : x. oo Or 1e) University of California Publications in Zoology [ Vol. 21 INDIVIDUAL SKULL ELEMENTS Nasals—In A each nasal is represented by an ossification that is in contact laterally with the ossified premaxilla and medially with the other nasal. Posteriorly the nasals are separated from the frontals by a fontanelle that is bridged in B. In C the nasals are slightly nar- rower posteriorly than anteriorly and are still narrower in D. The posterior constriction in C and D results from the lateral border of each nasal bending slightly but sharply mesially and continuing in a straight line until it reaches the truneated posterior border. In EF the posterior half of the lateral margin becomes bent mesially near its middle. This median convexity of the posterior portion of the lateral border of each nasal and the constriction of the whole posterior four- fifths of the nasals become gradually accentuated from F# through the successively older specimens. As will be seen from table 2, the nasals become relatively (to their width) longer in the skulls successively older than E. In most of the skulls of Q the nasals are narrowest not at their posterior end but at a point 1 to 3 millimeters anterior to the posterior end. The transverse depression of the nasals, which is situated about one-third their total length from the anterior end, appears first in Z, and in F is as well developed as in any of the later stages. During postnatal development the nasals increase more than 314 times in length and they double in width (see table 2). Premaxillae—The premaxillae in A are thick ossifications in con- tact dorsolaterally and laterally with the nasals, and posteriorly both ventrally and laterally with the maxillae. No ossification exists between the anterior palatine foramina. The region of each posterior wing of the premaxillae, that later extends posteriorly between the maxilla and nasal and overlaps the frontal, is not yet ossified. In B ossification is complete between the anterior palatine foramina but an unossified area still exists between the dorsal wing of the premaxilla and the frontal. In C the inner part of the wing that hes beside the nasal is ossified and is in contact with the frontal. A fontanelle bounded by the frontal, laerymal, maxilla, and premaxilla persists on each side in specimens as old as HZ. The same fontanelles, but with the lacrymals excluded from their borders, persist as late as @ (see fig. 10). Of the several fontanelles these are the last ones to disappear. Figure 20 shows the incisor alveoli occupying a large area on the ventral surfaces of the premaxillae in A. Here the alveoli lie partly posterior and mostly lateral to the anterior palatine foramina. ‘The 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 359 posterior margins of the alveoli of the incisors in B (fig. 19) lie slightly anterior to the anterior palatine foramina and less lateral to them than in A. The distance gradually becomes greater between these foramina and alveoli. In A each premaxilla consists of two ossified portions (fig. 20). An unossified area extends diagonally back from the posterior border of the alveolus of the incisor to the maxillary border of the pre- maxilla. Traces of this division between the two parts of each pre- maxilla are also present in B. On the ventral side of the rostrum the premaxillae increase 314 times in length during postnatal develop- ment. The lateral wings of the premaxillae that meet the frontals vary greatly in the degree of posterior extension in different specimens of the same age. These wings sometimes extend farther posteriorly on the frontals, than do the nasals, and sometimes not so far. The increase in length of these two wings of the premaxillae is equal to the increase in length of the nasals. The postnatal increase in width of the two wings, 469 per cent, is more than the increase in width of the nasals, which is only 200 per cent. In E two slight depressions appear, one just behind each incisor tooth, on the ventral face of the pre- maxillae. These gradually deepen until the condition shown in Q is developed. Mazxillae—tIn A the maxillae are ossified and are in contact with ossified portions of the frontals, jugals, and premaxillae. In B the maxillae come inte contact with the lacrymals and palatines. The infraorbital canal remains constant in position, as does the process for muscular attachment immediately below it. The palatal surface is nearly flat in A to C, concave in D to K, and develops a median ridge in L which becomes most pronounced in Y. The distance between the first molars (measured in the young skulls from the median edges of the dental troughs) increases only 90 per cent from C to Y. From F to Q there is an actual decrease of 5 per cent. From H, where this distance reaches its maximum, to @ there is a decrease of nearly 16 per cent. The coneave palatal surface of the maxillae is present at a time, D to K, when the teeth are pushing down the regions of the maxillae that they occupy; thus the concave palatal surface of the maxillae seems to be due, in part at least, to the lateral borders being pushed down rather than to the center of the palatal region being pushed up. This coneavity is greatest in HL, just before the teeth begin to break through the gums, and disappears in K, the stage in which the last molar comes into use. These parts of the maxillae in which the teeth develop are bounded above by the eye and below by the buceal cavity. The increase in thickness of that portion of the maxillae 360 University of California Publications in Zoology [ Vol. 21 in which the teeth develop occurs below the level of the palate instead of above it. A purely mechanical explanation might be that the grow- ing eye constitutes a stronger barrier to growth of the maxilla upward than the buceal cavity constitutes to its growth downward. Many other factors much more complex in their effect probably help to _ determine the direction of growth. In A the zygomatic portion of the maxilla is weak and does not extend laterally beyond the outer margin of the infraorbital canal. In B a slight swelling marks the strong crest that, in adults, develops along the anterior portion of the zygomatie root. This is only slightly better developed in D than in A. The plate that is developed below this ridge and that faces ventrally in adults is part of the region where the masseter muscle has its origin. This plate is present in G as an oblique surface inclined outward and upward and at this time projects only 1 mm. laterally beyond the external border of the infraorbital foramen. The plate develops rapidly from H on and becomes modified in progressively older skulls until the condition in Q is reached. Here the sharp overhanging crest with a distinct lobe has a deep fossa beneath it. Coincident with these changes the zygomatic root of the maxilla becomes inclined outward at a wider angle from the longitudinal axis of the skull to permit of the zygomatic root taking part in the bowing outward of the whole zygomatic arch. The maxillary plate that faces ventrally and _ is developed beneath the zygomatic ridge develops very lttle until a iy eo) 2euetee 12 09 12 ee make possible the mastication of solid food. Apparently, the develop- sufficient number of the teeth, erupt from the gums to ment of this plate depends upon the use and growth of the masseter musele which has its origin partly on this plate. The zygomatic ridge extends to, but not beyond, the premaxillo-maxillary suture. Just behind the last molar and between the maxilla and the palatine, there is a notch for the transmission of a nerve. In EF the maxilla sends back, lateral to the notch, a thick process which meets the palatine bone and converts the notch into a canal. The process of the maxilla does not at once fuse with the palatine but grows thinner in suec- cessively older stages until O is reached. In O and in later stages this process of the maxilla is sometimes fused with the palatine, sometimes in contact with but not fused with the palatine, and sometimes it remains wide open as a mere notch. Measurements taken along the intermaxillary suture show a postnatal increase in length of 219 per cent. Lacrymals.—Each lacrymal is ossified in B except for the postero- ventral portion, which has become completely ossified in F. In Ba 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 361 mere edge of the lacrymal extends up between the jugal and frontal on the rim of the orbit, and gradually develops into the tubercle-like pars facialis of adults. It measures 2 to 3 mm. anteroposteriorly in Q. Within the orbit and immediately below the rim is the opening of the lacrymal canal. The opening of this canal does not change its position but the orbital ridge of the lacrymal overhangs it somewhat in old specimens; thus in Q only a small part of the opening is visible when the skull is viewed laterally. On the turbinal surface of this bone a wing is sent out which finally encloses the lacrymal duct. The lacrymal duet, then, runs not between the lacrymal and turbinal bones but through the lacrymal bone, at least in specimens older than J. The lacrymal les on the surface of and overlaps the jugal, maxilla, frontal, and turbinals. This superficial position together with the small size of the lacrymals and the fact that strong contacts are formed between the jugals and maxillae, and between the maxillae, premaxillae, and frontals, indicate that the lacrymals do not play an important part in transmitting the strains of the zygomatic arches to the other bones of the skull. Frontals—In A the anterior regions of the frontals, both dorsally and on the lateral wings that lie within the orbits, are the only portions ossified. In B the two frontals are largely ossified, but a large fon- tanelle exists between them and the two parietals, and there is some unossified tissue along the entire fronto-parietal suture. A supra- orbital ridge and slight notch are present in B, but not until C are there shght indications of postorbital processes. In D these processes are sharp and distinct. They are located about one-fourth of the total length of the frontal anterior to the frontal border of the parietal. From D to Q several changes gradually take place. These are: an elongation that results in the processes becoming more slender in pro- portion to their thickness, a movement that causes them to be inelined forward and downward, and an incorporation of a part of a parietal bone in the posterior border of each process. It is evident from its situation in D (noted above), that some change has taken place in the position of the postorbital process, relative to the position of the parietal. This change is accomplished by each parietal overlapping the frontal at the dorsolateral angle of the skull. When the parietal reaches the postorbital process, a portion of the parietal is carried out with the postorbital process to form a part of its posterior border. On the dorsolateral face of skull no. 29640, which represents the L stage of development, the parietal has overlapped the frontal for a distance of 5.7 mm., which is more than one-fourth of 22.1 mm., the total length of 362 University of California Publications in Zoology [ Vol. 21 Figs. 1-6. Dorsal view of ecrania. Fig. 1,@Q, X 1. Fig. 2, N, X 1. Fig. 3,- i, xX 1.- Fig. 4, BP, <0. Bip. 5) Ho to Big Oa, xe. 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 363 the frontals in this ease. Near the median line of the skull, however, the relation of these two bones is reversed, and the frontal overlaps the parietal to a distance of 1.3 mm. It is interesting to note that the place where the parietal overlaps the frontal is the place where the parietal and frontal first come into contact and that the place where the frontal overlaps the parietal is the place where the parietal and frontal last come into contact. This portion of the parietal that over- laps the frontal reaches the postorbital process in G. The supraorbital ridges gradually rise and in M project above the rest of the dorsal surface of the frontals. The ridges also increase in thickness and become very heavy in Q. Externally, between the orbits, the frontals are convex from A to H. Externally, they are also convex longitudinally from A to F. This longitudinal convexity is most pro- nounced in A and B where the frontals dip sharply downward to meet the nasals. This is perhaps partly the result of the relatively greater development, in these stages, of the brain than of the structures housed by the preorbital portion of the skull. The entire frontal suture is present in Z and the anterior part of this suture in M to P, but in Q it can not be made out, the frontals having fused throughout their entire length. The frontals are in contact with the following bones: nasals, pre- maxillae, maxillae, palatines, presphenoid, lacrymals, orbitosphenoids, alisphenoids, squamosals, parietals, and ethmoids. It has already been pointed out that the nasals come into contact with the frontals in B and later overlap the frontals, also that the premaxillae come into contact with the frontals in C and that the maxillae reach across the fontanelles that persist immediately in front of the lacrymal regions in H and at this time come in contact with the frontals at this point. Within the orbits, however, ossified parts of the frontals and maxillae reach one another in B. Within the orbit the frontal reaches the maxilla along most of the anterior half of their contact but the maxilla overlaps the frontal along the posterior half of their contact. The two wings of the premaxillae that lie alongside the nasals, like the nasals also overlap the frontals, but between each premaxillary wing and each nasal a process of the frontal, 2 or 3 mm. long, les at the dorsal surface of the skull and slightly overlaps the lateral border of the premaxilla. The contact of the frontals with the palatines is made in B. On each side in B and in C there is present what appears to be a separate center of ossification, namely, the region of the part of the frontal that lies immediately anterior to the region later occupied a ee Figs. 7-14, Lateral view of crania. Fig, 7, Q, X 1. Fig. Shoe Fig. 9, I,X 1. Fig. 10; G; 1a! Pig. 11, Fy & 1 Pig? 12> eax Lig: Rig a: X 2. Fig. “7%. Figs. 12 and 13 are of dried skulls which, due to Shrinkage 0 ions, show some distortion in the preorbital region, 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 365 by the orbitosphenoid. It is this part of the frontal that meets the palatine bone. The contact is never much greater than it is in B where it is first present. In fact, on the average, it becomes less, externally, in progressively older skulls. On the right side of L a process of the maxilla extends backward and meets the orbitosphenoid; thus exclud- ing contact of the frontal and palatine externally. This, however, is the exception rather than the rule. Ventrally the two frontals are separated only by the presphenoid and the thin wings of the orbito- sphenoids and palatines that extend forward with the presphenoid. The orbitosphenoids are first in contact with the frontals in EZ. In C anterior portions of the alisphenoids become ossified and are in contact with the frontals. The frontals are in contact with the lacrymals from the time the latter first appear, which is in B. The development and relation of the turbinals to the frontals and to the other elements con- cerned chiefly with the olfactory sense, were not made out in the youngest skulls since to study these parts it would have been necessary to disarticulate other elements which it was desirable to keep in place for comparative study. In adults, however, the frontals’ relation to the turbinals was noted. Here the cribriform plate is supported by the frontals along its entire border except for a narrow space ventrally where it comes in contact with the presphenoid, and perhaps the vomer. In three skulls examined the cribriform plate was situated on a trans- verse plane of the skull that passed in front of the supraorbital notches and through the last upper molars. Internally, the frontals bear ridges and coneavities that mark the position of the turbinal scrolls. Behind the cribriform plate the olfactory lobes leave two depressions. Other- wise, within the cranial cavity, the frontals bear but few impressions left by blood vessels or other structures of the soft anatomy. It is seen from table 2 that the least interorbital breadth of the frontals increases 212 per cent. The ratios given for this measurement of the series of skulls express the relations that exist in proportion between this width and the occipito-nasal length. The largest ratios are in Band D. The ratio grows less upward through the series until the adults are reached at which place the ratio increases again. This last increase represents the greater development of the orbital ridges in old individuals. Another change due to very great age, which is present in most skulls of Q, is the closure of the supraorbital notch that carries branches of the anterior facial vein, internal (?) carotid artery, and fifth (?) eranial nerve. In Q this notch has become changed into a canal (see figs. 1-6). 366 University of California Publications in Zoology [ Vol. 21 Figs. 15-20. Ventral view of crania. Fig. 15, @, X 1. Fig. 16, J, X 1. Fig. 17, E, X 1%. Fig. 18, D, X 2. Fig. 19, B, X 2%. Fig. 20, 4d, X 2%. 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 367 The least postorbital breadth shows an even greater difference in ratios in the different skulls than does the least interorbital breadth. It will be noted from table 2 that the total increase is 8.9 mm. or 1.7 mm. less than the increase in the least interorbital breadth. This is an increase of 127 per cent. Table 2 also shows that the ratio of this measurement decreases from the beginning ‘to the end of the series. In skulls H, K, L and N, however, the least postorbital breadth is greater than in Q. This proves to be a rather constant difference between adult and immature specimens. A series of fifty males and another series of forty-five females, both ineluding animals that ranged in age from those just shedding their milk premolars to very old indi- viduals, were arranged in order of youngest to oldest and then divided into four groups on a relative age basis. Table 1 gives the average, maximum, and minimum measurements of each group of these two series. Group 1 is the youngest and group 4 is the oldest. The female series 1s believed to average slightly younger than the male series. Both show an actual decrease in the least postorbital breadth with age. It is generally known that decreases with age occur in several parts of the skull of different mammals. Taylor (1918, p. 489), for instance, has pointed out an actual decrease with age in the interorbital con- striction of Aplodontia. TABLE 1 ACTUAL DECREASE OF LEAST POSTORBITAL BREADTH WITH INCREASE IN AGE Group 1 Group 2 Group 3 Group 4 Number of specimens.............. 8 7 14 16 OPTIMA CLAG Cy de cncc- ec Setccres eee: 15.4 16.4 15.5 15.4 Maximum and minimum........ 14.4-16.8 | 16.2-16.8 | 15.0-16.2 | 14.5-16.0 Number of specimens.............. 13 13 16 8 Chas | AN CRAG Cec Noee. 3. ce At, eee 16.3 5, a 15.4 15.4 Maximum and minimum........ 14.7-18.0 | 14.9-16.4 | 14.0-17.0 | 13.8-16.0 Parietals—tIn adults each parietal is in contact with a frontal, squamosal, mastoid, the supraoccipital, and the interparietal. The relations of the frontal contact have been pointed out. The squamosal overlaps the parietal for a considerable distance at all points where the two are in contact. This is reflected in the measurements given in table 2 for the width of the parietal bones dorsally. 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The interparietal overlaps the parietal. A firm contact is established between the supraoccipital and parietals. The supra- occipital gradually extends both under and up behind the parietals in successively older skulls. Contact of the parietals with the mastoid is gained by the parietal sending a process down beneath the squamosal to meet the mastoid. This process is relatively heavy and forms a strong support for the periotie and tympanic. The mastoids and parietals first come into contact in G. The parietal and parieto-inter- parietal sutures both disappear in / and the fronto-parietal suture in M. Figure 21 illustrates the contact relations of the bones along the OS \ S a 8 SS eo . ROPES S Om ww xe ° we ss Cy < oe ——$———— x Be XN < Fig. 21. A diagrammatic representation of the contact relations of the bones on the dorsal surface of the skull along the sagittal plane in skulls older than J. median line of the skull. The parietal ridges begin to develop in MW at which time two diverging lines extend from the postorbital processes back along the sides of the skull, bend inward to meet the short sagittal erest, and gradually develop into ridges. The ridges converge farther anteriorly in progressively older skulls. This causes the sagittal crest to become longer. The ridges also become straighter. Figures 1—2 rep- resent two stages in the development of these crests. The lambdoidal erest is also partly composed of the parietals. As the supraoccipital grows upward, it causes the posterior borders of the parietals to be reflected upward. Internally, the parietals are smooth except for a depression that extends along the region of the median suture. Impressions of a few blood vessels are also present. Squamosals—The zygomatic process of each squamosal and the small region from which this process arises are ossified in A. A gradual extension of this ossification carries the squamosal out over the other bones with which it has contact. Each squamosal comes into contact with a jugal and parietal in A, with an alisphenoid and tympanic in B, with a mastoid in F, and with the supraoecipital in G. The projection outward of the zygomatic process of the squamosal and the overlapping by this bone of the other bones with which it comes into contact are the main features to be noted in its development. Its relation to the supraoccipital is much like that of the parietal’s relation to the supra- 1926 } Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 371 occipital ; both the parietal and squamosal have their posterior margins reflexed upward to participate in the formation of the lambdoidal erest. Up to and ineluding J the outward projection of the zygomatic portion of the squamosal proceeds slowly, but after this time the curved portion of the squamosal is rapidly straightened. Jugals—In A each jugal is ossified and reaches from the zygomatic process of the squamosal to the lacrymal region. It extends along the inner side of the maxilla. In C the inner lip of the zygomatie portion of the maxilla that bounds the anterior part of the jugal has grown halfway up the side of the jugal. In # this part of the maxilla is pro- portionally as far extended over the mesial surface of the jugal as it is in Q. In D the zygomatic arch is bowed inward at a point near the middle of the Jugal bone and remains so up to and including G. In H and / it is straight. From this time on it gradually becomes bowed outward to a greater extent. The jugal bone increases in length from 6.5 mm. in C to 26.7 mm. in Q. In width it inereases from 0.6 mm. in C to 4.7 mm. in Q—an increase of 311 per cent in length and 683 per cent in width. Although the bone becomes very wide, in Q it is only 0.5 mm. in thickness, or 0.2 mm. more than in C. The main changes are an increase in width and a bowing outward. The jugal is also modified by muscles that attach to it. In Q a distinct lip, and also a transverse ridge, are developed on the dorsal margin of the jugal just back of its union with the outer slip of the zygomatic process of the maxilla. The position of the ridge relative to the length of the skull is modified by the central and posterior portions of the bone being moved relatively farther from the walls of the cranial cavity. This is partly accomplished by the bowing outward of the jugal but is more directly due to the straightening of the basal portion of the zygomatic process of the squamosal and to the development of a progressively wider angle between the longitudinal axis of the skull and the zygo- matic process of the maxilla. This causes the skull to be relatively wider in adults than in young specimens. For successively older skulls this increase is shown in table 2 by the increasing ratio of the zygomatic breadth. It is also readily perceived without the aid of measurements from figures 1—6 and 15-20. Orbitosphenoids—Of the several bones represented within the orbit the orbitosphenoids are the last to ossify. The first ossifications appear in D on the anterior, ventral, and posterior sides of the optic foramina and are in contact only with the palatines. In £ ossifications extend entirely around the optic foramina, and most of the posterior 372 University of California Publications in Zoology [ Vol. 21 parts of the orbitosphenoids are present in an ossified state. In F ossification is completed. For some time, however, these bones remain very thin as compared with the other bones of the skull. The sutures separating the orbitosphenoids from the other bones within the orbits ean be made out in very old specimens. The orbitosphenoids are in contact with the alisphenoids, frontals, palatines, presphenoid, and occasionally the maxillae. The orbitosphenoids extend down over the sides of the presphenoid; thus they are overlapped by the palatines. The alisphenoids and frontals also overlap the orbitosphenoids though only slightly. Alisphenoids—In A a slight ossification is present only in that portion of each alisphenoid that forms part of the pterygo-alisphenoid ridge. In B ossification has extended up along the anterior margin of the squamosal. In C the ossification has extended on dorsally, bring- ing the frontal into contact with the alisphenoid. At this time it is ossified little if any farther anteriorly than it is in B. After B ossification proceeds gradually to completion in F’. Each alisphenoid is in contact with the basisphenoid, a tympanic, a pterygoid, a palatine, an orbito-sphenoid, a frontal, and a squamosal. The orbito- sphenoids, frontals, basisphenoid, and sides of the pterygoids are overlapped by the alisphenoids. The palatines and tympanics meet the alisphenoids rather squarely. The basisphenoid and alisphenoids become firmly united in old individuals. The palatines also become firmly attached to the alisphenoids. Although the unions between the palatines and alisphenoids are firm, sutures can often be made out between the elements in adults. In the base of the alisphenoid two canals extend forward from the foramen ovale. The larger and lower of these two canals extends forward to the foramen lacerum anterius. The smaller of these two canals, posteriorly, begins just above the larger, only a thin partition of bone separating the two, and, anteriorly, opens into the orbit just behind and laterally to the foramen lacerum anterius. Only in the anterior part of its course is it a true canal for, above, it opens into the brain cavity for a short distance after leaving the foramen ovale. In Marmota, where the arrangement of the foramina and their relation to these two canals is essentially the same as in Otospermophilus, both of these canals carry branches of the fifth nerve. Neither carries a blood vessel. Under the heading of ‘‘General Morphology of the Rodent Skull,’’ Gregory (1910, p. 329) states that: ‘‘The alisphenoid canal (occasionally absent) is appar- ently not the large tunnel leading forward to the foramen lacerum 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 373 anterius but a small canal external to it.”’ This statement certainly does not apply to Marmota, and, although Gregory (1910, p. 329) lists Arctomys [== Marmota] as one of the genera in whieh the cranial foramina, at least, were examined by him, he makes no mention of its being an exception to his general statement regarding the alisphenoid eanal. Palatines—In A the palatines are separated from each other and from the maxillae anteriorly by tracts of unossified cartilage. Each pterygo-alisphenoid ridge is represented largely by a palatine at this stage. The term pterygo-alisphenoid ridge is non-descriptive in the ease of Otospermophilus because the elements comprising it are almost entirely the palatines and alisphenoids. Also, in Otospermo- philus, this ridge is more plate-like than ridge-like. Viewed laterally it is square in young skulls, but this shape changes with age until in @ it becomes oblong, being narrower anteriorly than posteriorly. It is also modified in older stages by the development on it of the pterygoid fossa. A median ridge that is continuous with, and con- temporaneous in time of development with, the median ridge men- tioned in connection with the maxillae, develops on the palatines. One of the most striking changes in the palatines is in the shape and extent of the palatal spine (see figs. 15-20). In the embryo skulls, A and B, this posterior margin of the palatines lies flat on the basi- sphenoid, but in C the posterior margin of the palatines is raised slightly above the basisphenoid. This margin continues to rise throughout the series and in Q lies about 3 mm. above the basisphenoid. In addition to the maxillae, orbitosphenoids, and alisphenoids with which the palatines are in contact, they also support the pterygoids, basisphenoid, and presphenoid. The palatines extend anteriorly to the forward end of the presphenoid. However, except on the posterior half of the basisphenoid, a narrow slit on either side of the pre- sphenoid and basisphenoid separates them from the palatines. These slits are never wide and in Q, at most points, become entirely closed. The support given the basisphenoid by the palatines is accomplished by their connection with the posterior half of each lateral border of the basisphenoid and by a slight contact of the palatines with the orbitosphenoids which in turn are in contact with the presphenoid. In measuring the palatines (exclusive of the palatal spine) it becomes evident that they are extremely variable in length in skulls of the same age; far more so than are the maxillae or premaxillae. In Otospermophilus grammurus beecheyi, at least, a less variable meas- 374 University of California Publications in Zoology [ Vol. 21 urement than the palatilar length or the palatal length, which are so often used in diagnosing various species of mammals, would be from the anterior tip of the premaxillae to the maxillo-palatine suture. Pterygoids—The pterygoids are not extensive at any stage of development and contribute to the skull only the processes bearing that name. In A, as shown in figure 20, each pterygoid is ossified and is in contact with a palatine. Its changes in shape and in relative size are shown in ventral view in figures 15-20. It is attached to the posterior end of the pterygo-alisphenoid ridge and in A, as may be seen from figure 20, only to the extreme posterior end of this ridge. In A to D the pterygoid is closely appressed to the underlying bones. In E£ it is slightly raised. In @ the pterygoid process les on a plane passing across the ventral surfaces of the tympanic bullae. In Q each process projects 1 mm. or more below this plane. At this time the anterior border of the pterygoid process projects downward from the floor of the palate at an angle of 35 degrees. In A to H the shape is profoundly modified by the tympanic, which crowds in upon the pterygoid and excavates its lateral border giving it the gibbous shape shown in figures 17-20. In the pig, according to Parker (1874, p. 322), the pterygoid dents the tympanic, instead of the tympanic denting the pterygoid, as is the case in Otospermophilus. In I and in later stages, a tendency to straighten this curved border is manifest but even in @ the process is curved outward. It would seem, how- ever, that the retention of this curve is due to causes other than the early influence of the tympanic. This opinion is supported by the fact that a rotation of 90 degrees occurs during the time that the process is lifted from its horizontal position to that of an inclination of 35 degrees whereby the border that was lateral in A becomes ventral (skull with palate facing downward) in Q; thus any effects that might remain in Q from pressure exerted by the tympanic in earlier stages would be present on the ventral border and not on the lateral face of the process. That this process does rotate is indicated by the fact that the palatal face of the pterygoid remains smooth whereas the lateral face is modified at its base by the formation of considerable osseous tissue, at precisely the proper point to cause rotation. The deposition of additional bony matter at this point and the ultimate shape of the hamular process are probably determined to a greater degree by the effects of muscular development than by the influence of the surrounding bones. The pterygoid processes fuse with the basisphenoid in G. On the median surfaces of the pterygoids the 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi Bw és) sutures separating them from the palatines can usually be made out in Y. At this time the pterygoids are very thin and have actually decreased in bulk. Their lengths in millimeters in several skulls are: C, 3; F, 8.6; 7,9; Q, 10.4. Instead of ending in a single point, the hamular process of each pterygoid has two divisions, as shown in figure 22. Fig. 22. External view Fig. 23. Ventral view of of right pterygoid process vomer in a skull from the OLION< 122 same litteras B. X 12. Vomer and ethmoids.—F igure 23 shows the condition of the vomer in a skull from the same litter as B. The two lateral wings are very thin and are turned up at the lateral margins, which are in contact with the maxillae. A considerable area of cartilage separates the posterior border of the vomer from the ossified portion of the pre- sphenoid. On the anterior portion of the vomer a round column of bone rises from the horizontal surface and establishes contact with the maxillae at the posterior borders of the anterior palatine foramina. In a skull from the same litter as #’, and approximately of the same age, the vomer is modified from the condition found in the skull from the same litter as B. In the older of these two skulls the vomer and rostrum are more elongated than in the younger. The column of bone that rises from the horizontal surface of the vomer and meets the maxillae is relatively shorter, but wider, in the older of the two stages mentioned ahove. The thickness of the vomer seems to be not much greater than in the younger skull. Posteriorly, in the skull from the same litter as F’, the vomer is not in contact with the pre- sphenoid. Other skulls of greater age show that the vomer is always a very thin element and that it is somewhat modified by the develop- ment of the turbinal and ethmoid elements. In the embryo skull of the same age as B these elements are little developed. In the skull of the same age as F, the maxilloturbinals, nasoturbinals, and four primary divisions of the endoturbinals are present in a well developed state, but all lack the complexity of the adult condition. The por- ve ~I Oo University of California Publications in Zoology [ Vol. 21 tion of this ethmo-turbinal complex that seems to affect the vomer most profoundly is the mesethmoid, which is composed of two sym- metrical halves that lie side by side and are separated by a mem- brane. The mesethmoid is not present in the skull of B age. In the skull of F age it is present as two unossified septa, and in a skull slightly older than LZ it is well ossified. Vertically the mesethmoid extends from the vomer to the frontals, and dorsally, it extends longitudinally from the cribriform plate to the naso-frontal suture. It is rounded on the anteroventral border, hence, ventrally, does not project so far anteriorly as it does dorsally. The vomer becomes intimately associated with the ventral border of the mesethmoid and, in adults, practically loses its identity. The cribriform plate is also supported in part by the vomer and perhaps influences its develop- ment. In one specimen, slightly older than L, the cribriform plate contains 187 foramina. These are arranged in three main tracts; one central and one on each lateral portion of the plate. Each lateral tract is divided into four primary regions on each side, but here the symmetry of the two sides ceases. Presphenoid.—The presphenoid is cartilaginous in A. In B it is represented by a small ossification which is slightly pointed anteriorly and is in contact only with the palatines. The small size and position of this ossification indicates that the presphenoid develops from a single center, a manner of development that Parker (1885, p. 114) has noted as perhaps a characteristic feature in rodents or at least one that occurs in some forms. He says: ‘‘The presphenoid is very rarely an independent ossification ; it attains to that special condition 9? most perfectly in the Marsupials and Rodents; He also says (1885, p. 204): ‘‘. . . It is also present in some of the lesser rodents 9? (at any rate in Arvicola, Mus, ete.).’’ Ina skull from the same litter as F the presphenoid is 7.3 mm. long, and bears the same relations to the surrounding bones as it does in older skulls. However, the presphenoid in this particular skull, throughout most of its length, is transverse in cross-section. One side of this triangle les hori- zontally and forms part of the floor of the brain case. At the anterior end the sides describe a nearly equilateral triangle, but this shape eradually changes toward the posterior end of the presphenoid. Here the ventral angle becomes more obtuse and the extreme posterior portion of the bone is flat. This shape is decidedly different from that in a skull slightly older than M, in which the presphenoid is quad- -angular in cross-section throughout its length. As in the specimen 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi S30 Of of the same age as F’, the presphenoid is widest posteriorly. In the specimen slightly older than M the presphenoid is 10 mm. long, 0.9 mm. wide anteriorly, and 2.6 mm. wide posteriorly. On its dorsal surface, between the two optic foramina, a fossa is present that partly accommodates the swollen region of the brain in which the optic chiasma lies. Basisphenoid.—A small transverse elliptical ossification represents this bone in A. At this stage the ossification is in contact with no other bone, but in B it has reached the alisphenoids, basioceipital, palatines, and pterygoids. The presphenoid and basisphenoid meet in F. In Q the basisphenoid is fused with all the bones mentioned above with which it was in contact before #. It also establishes con- tact with the fused margins of the petrosals and tympanies by a process that is sent out on either side. On either side one of these processes separates the foramen ovale from another foramen just posterior to it. Posteriorly, in all stages from C on, the basisphenoid abuts squarely against the anterior end of the basioccipital. No examples in which the two bones were fused were seen. The hypo- physeal fossa on the superior face of the basisphenoid is slightly anterior to the center. The fossa is about 2 mm. across and 1 mm. deep. No clinoid processes are present. In C the basisphenoid is 2 mm. long. A small canal that carries a blood vessel extends trans- versely through this bone slightly back of the center. In a skull from the same litter as F, the basisphenoid is 3.2 mm. wide anteriorly, 3.9 mm. wide posteriorly, and 5.4 mm. long. In a skull slightly older than M, the measurements in millimeters are: width anteriorly, 3.4; width posteriorly, 6.0; length, 8.8. In Q this bone averages 10.5 mm. in length. Basioccipital—Figures 15-20 show the changes in shape. One difference in the manner of development of the basioccipital and of the basisphenoid may be seen in figure 20. The ossification in the basisphenoid is elliptical, and the long axis les transversely with respect to the long axis of the skull. The elliptical oval area of ossification in the basioccipital has its long axis extending longitudin- ally with respect to the long axis of the skull. At this time, A, the area of ossification of the basioccipital is more than twice that of the basisphenoid, while the length of the basioecipital is always 1 to 2 mm. greater than that of the basisphenoid. A median ridge is present on the inferior surface of the basioccipital in adults. Traces of this appear first in G. Two large fossae develop at the same time that this ) 378 University of California Publications in Zoology [ Vol. 21 0 ridge, which separates them, develops. In adults, near the anterior end of the basioceipital, two processes, one on each side, extend ventrally as far as do the tympanic bullae. These are closely appressed to the sides of the tympanic bullae. This relation gives a broad support to the basioccipital and shields the bullae from the disturbing effects of muscular action. While these processes eventually grow as far ventrally as do the bullae the two structures by no means keep pace with each other in development. The bullae are well developed in F’, but only the first traces of these processes of the basioccipital are present at this time. In A the processes are one half the height of the bullae. In Q the bullae and lateral. basioccipital processes are equal in height. The posterior margin of the basioccipital is. thiek- ened. Part of this thickened border goes to form the ventro-mesial parts of the occipital condyles. The superior surface of the basi- occipital is concave but smooth. The two exoccipitals fuse with the basioccipital in L. During post-natal development the basioecipital increases from 3.0 to 12.2 mm., or 307 per cent in length, and from 2.7 to 9.3 mm., or 244 per cent in width. Exoccipitals—In A these bones are present as two small ossified areas entirely surrounded by cartilage. The portions bordering on the foramen magnum that will later form the larger part of the exoccipital condyles are slightly thicker than the rest of the bones. In C contact is made with the basi- and supraoccipital. Contact is made with the tympanic in C. In F, when the mastoids are ossified., these bones and the exoceipitals grow downward and form the par- occipital processes. At this time the paroccipital processes are com- posed almost wholly of the exoccipitals, but with inerease’in age the mastoids gradually contribute a larger share. In F the processes taper to blunt points and project straight downward. They are not appressed to the bullae. In G@, H, and J, however, the processes are closely applied to the bullae. Beginning with K continued growth carries the processes away from the bullae. During that period of growth from A onward, when the processes are growing downward, they also curve forward and inward. In Q they project nearly as far ventrally as a plane passing across the lower surfaces of the tympanic bullae. On the external surface of each exoccipital two condylar foramina are present. In several cases three foramina are present on the internal surface, but two of these three unite within the body of the bone; thus only two appear on the external surface. This is a slight advance over the condition not infrequently present in Mar- 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 379 mota, where three condylar foramina appear on the external face of each exoccipital. The exoccipitals fuse with the basi- and supra- occipital between M and Q. Supraoccipital——The large fissura bisupraoeeipitalis suggests that this bone may have come from two centers of ossification. Whether or not this is the ease figure 24 shows that in A ossification is certainly more extensive in the two lateral portions than in the central region. The interparietal is closely associated with the supraoceipital as early as A. In F ossified portions of the parietals, squamosals and mastoids come into contact with the supra- Fig. 24. Posterior view of supraoccipital in A showing fissura bisupra- the vertical axis, but between EH and F it oceipitalis extending dors- ; ; : ays ; ally from foramen mag- &@SSumes a more vertical position. In G and occipital. In D it leans shghtly forward from num. x 7. in H it again leans backward from a line drawn perpendicularly to the plane of the palate. From H on, its forward inclination increases. The change that occurs after H is very probably the direct result of muscular strains which, in fos- sorial and aquatic forms, very often tend to decrease the basicranial angle, that is, the occiput becomes inclined forward. Spalax and Balaenoptera represent extreme cases of development in this respect. Two lateral swellings mark the positions of the lateral lobes of the cerebellum. However, in some skulls of Q the external face becomes extremely rugose and the bone itself much thickened, pre- sumably from the effect of muscular pull rather than from brain pressure; thus muscles as well as the brain appear to have consider- able to do with the final form of the supraoccipital as well as with its inclination with respect to the plane of the palate. Externally a median vertical keel is present from J onward. An increase in height from 2.5 mm. in C to 10.0 mm. in Q, or 300 per cent, is made. TInterparietal—tIn A and B a striking feature of the interparietal is the relatively large area it covers in the roof of the skull. This is in marked contrast to the conditions shown by Parker (1885) of Centetes (pl. 32, fig. 1) of Rhynchocyon (pl. 36, fig. 1) and of Bos by Wilhelm (1924, text figs. 1-7). In these eases the parietals appear to contribute a relatively greater share of the skull’s roof in the embryonic stages than they do in Otospermophilus. In A and B ossification has hardly begun. In C the interparietal is well ossified. While no distinct suture is present that separates the bone into two parts, as is the ease in the skull of a young Erethizon that lies before me, there are indications 380 University of California Publications in Zoology [ Vol. 21 that the interparietal in C develops from two and possibly three centers of ossification. Tracts less densely ossified than the main body of the bone divide it into three parts. It will be seen from figure 25 that a considerable area of cartilage extends back- ward from the anterior border of the interparietal in D. This area of cartilage is seen to be somewhat smaller in H# (see fig. 5). As in the ease of the Fig. 25. Showing Supraoccipital these conditions indicate that pos- pa Fee CCE: sibly more than one center of ossification is in- into interparietal volved. However, careful search among specimens sag gee ce younger than D fails to reveal more than one center. Ossification appears to proceed from one point and to result at first in a crescent-shaped interparietal. The parietals and supraocecipital sur- round the interparietal. It meets the supraoccipital squarely and with the formation of the lambdoidal crest comes to have its posterior margin recurved. The parietals are overlapped by the interparietal. No trace of a suture between the parietals and interparietal can be made out in skulls older than K. In L a sagittal crest is present. The skulls of varying ages show that this crest develops first on the inter- parietal. In that part of the interparietal on which the sagittal crest develops sinuses also develop. Table 3 is significant in two respects. TABLE 3 SHOWING LENGTH AND WIDTH, IN MILLIMETERS, OF INTERPARIETAL BONE IN SKULLS OF DIFFERENT AGES Length of interparietal............ 2.4 Width of interparietal.............. 3.8 or bo ie) oO w When the measurements of the interparietal are compared with the measurements given in table 2 of the other elements comprising the dorsal surface of the skull it will be seen that the interparietal is rela- tively much larger in the first than in the last of the series. The peak of its development with respect to actual size is reached in F. After this there is an actual decrease both in width and in leneth. A, it will at once be noted, forms the single exception to this statement in that here the interparietal is 12 mm. wide. Comparison with other skulls of similar age shows that in A the interparietal is a wide departure from average conditions. In A two narrow lateral extensions have pushed outward between the supraoccipital and parietals and increase the 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 381 width 3 mm. over that of the normally shaped interparietal of this age. The general shape changes from the oval pattern of A and B (see fig. 6) to the square one of J shown in figure 3. Tympanics.—Only a very narrow half-ring of ossification is present in A. This ring hes in a horizontal plane. Membrane extends both laterally over the otic region to the external auditory meatus and postero-ventrally to the junction of the petrosal, basisphenoid and basioceipital. Ossification proceeds in this membrane as shown in figures 17-20. From figure 26 it will be seen that the tympanic lies Fig. 26. Showing relations of auditory ossicles and related structures in an embryo of 39 mm. from the same litter as B. Greatly enlarged. Explanation: ang., angle of mandible; cer., coronoid process; inc., incus; mal., malleus; me., Meckel’s cartilage; pet., petrous; sq., squamosal (overlying the end of Meckel’s cartilage and a part of the incus); st., stapes; tym., tympanic ring (anterisrly spreading out over Meckel’s cartilage and a portion of the malleus, posteriorly covering the stapedial process of the incus). against Meckel’s cartilage and that a part of the tympanic is extended posteriorly on the inside of the cartilaginous malleus and another smaller part anteriorly along Meckel’s cartilage. The presence of a small element slightly in front of the anterior end of the tympanic ring in C, but not noted in any other stage, suggests the possibility that all, or at least part of this portion of the tympanic that lies spread over Meckel’s cartilage represents the goniale. Careful scrutiny, however, fails to reveal any separation between the major portion of the tympanic ring and the part spread over Meckel’s cartilage. It is possible that a slightly later stage would reveal a separation of these two parts. However, at this time, in C, the malleus and incus, although still adhering to Meckel’s cartilage, are clearly defined from it and from each other. No separate element that could be interpreted as a goniale, as was doubtfully done with the element in C, was found in the stage shown in figure 26. Except for the spreading of the tympanic over Meckel’s cartilage, this relation of the tympanic to Meckel’s cartilage does not differ greatly from the relations in some other forms, judging from the figures given by Parker (1885) of Dasypus (pl. 7, 382 University of California Publications in Zoology [ Vol. 21 fig. 1), and of Sorex (pl. 30, fig. 16). Rather comprehensive changes in either of these types are required to accomplish the adult condition. As Doran (1878, p. 444) pointed out, the shrews, in common with other insectivores and marsupials, have the processus gracilis firmly united to the tympanic. Where the tympanic hes on the outside of the pro- cessus gracilis in the adult, but on the inside of the processus gracilis in the embryo, the condition seen in adults of the above mentioned genera would seemingly have to be arrived at: by an absorption of some of the parts of the elements that intervene between the processus gracilis portion of the malleus and the area encircled by the tympanic, by a rotation of the elements, or by the formation of the processus gracilis by direct outgrowth from the part of the malleus that is not covered by the portion of the tympanic that spreads over Meckel’s cartilage. These same three possible ways in which the adult condition might be brought about present themselves in the case of Otospermo- philus. Here the differences in the relative position of the processus gracilis and of the tympanic at embryo and at adult stages are, in the ease of Otospermophilus, fully as great as, if not greater than, those in the case of Sorex. In adults of Otospermophilus the processus gracilis is of course free, and directed downward. The material examined does not show which one of the above three general ways of change of posi- tion by the tympanie and processus gracilis was employed, but it does indicate, very strongly, that it was not the last, that is, not outgrowth of the processus gracilis directly from the portion of the malleus that is not covered by a part of the tympanic. From figure 26 it will also be noted that the tympanic lies not entirely around the petrous but partly upon the petrous. At this time the tympanic extends over the stapedial process of the ineus but terminates at the malleus. The tympanic is much wider posteriorly than anteriorly. This disparity in size between the anterior and posterior parts of the tympanic is rendered more apparent by reason of the fact that the anterior portion, beginning a short distance medially from where it meets Meckel’s cartilage, is inclined vertically whereas the posterior portion that extends over the stapedial process of the incus is nearly horizontal. Ossification of the bulla appears to proceed from this ring, no second center of ossification having been noted. In figure 17 of E it will be noted that a constriction is present in the tympanie which divides it into two parts. The basal or inner ring is the smaller of the two. Whether this division into two parts, seen only in £, is a fortuitous one is not known. At any rate it should not be taken to indicate that both a tympanic and an ento- tympanic are present, for earlier stages show no separation into two 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 383 parts. At the constriction between the two parts of the tympanic seen in #, a small inner third ring of bone, the tympanic annulus, to which the tympanic membrane attaches, is present and appears to be separated by a suture from the tympanic bone. Viewed laterally, there is nothing to indicate the place of attachment of the tympanie annulus from F to P. In Q, however, a clearly defined line (see fig. 15) mark- ing the position of the tympanic annulus is to be seen. Lateral to this line the tympanie is transparent. Mesial to this line the tympanic is often more opaque. In Q the tympanic annulus is firmly fused with, and is perpendicular to, the inner face of the bulla. The line of attach- ment of the tympanic annulus to the inner face of the bulla marks the limit of lateral extension of three bony lamellae which, when developed, divide the tympanie cavity into compartments. In a skull of N age, two lamellae extend from the petrous to the mesial and dorsal walls of the bulla. A third lamella extends in front of the petrous and parallel to the other two. The portion that extends lateral to the petrous bounds the petrotympanic fissure anteriorly. A fourth lamella, thicker than the others and shielding the ear bones, extends from the lateral border of the petrous to the lateral wall of the bulla and bounds the petrotympanic fissure posteriorly. The three septa that extend mesially from the petrous appear first in G. At this time the tympanic has practically completed its growth over the petrosal, the external auditory meatus being but little larger than in older skulls. The existence of an element concerning which I have not been able to come to any definite conclusion may be mentioned here. While, apparently, not intimately related to the tympanic, its posterior end hes near the anterior end of the tympanic ring. ‘This element in C is ossified and les between the squamosal and alisphenoid. It is much longer than wide and, longitudinally, extends from the posterior union of the alisphenoid and squamosal across the glenoid cavity and to a slight distance in front of the glenoid cavity. I am tempted to believe that this may be a constant element because indications of a differen- tiated region of bone at this same place can be made out in D. D and C, however, are from the same litter and the presence of this element might be expected in both animals even if it were merely a Wormian bone. That is, it might be so expected, if we may judge from analogous cases in the genus Marmota. The skulls of a few litters of Warmota monax monax that I have examined show, not only the presence of Wormian bones in the same position in skulls of the same litter but a striking correspondence in size as well. These elements were inter- 384 University of California Publications in Zoology [ Vol. 21 posed between the parietals immediately in front of the interparietal. Unhke Erethizon, Castor, and Marmota, Otospermophilus grammurus beecheyi has few Wormian or fontanelle bones. Aside from the possible occurrence in C only one small Wormian bone was noted in the approximately 800 skulls of this form that I examined. This series of 800 skulls represents animals ranging in age from small embryos to very old individuals. The Wormian bone noted occurs in J. Its shape, size and position are shown in figure 3. Petrosals—The petrous portion is well ossified in E. The mastoid portion, in F’, is represented only by a small ossification less than 1 sq. mm. in area. It is fully ossified in G. The lateral wall of the parafloceular fossa is the last part of the mastoid to ossify. Externally, in H, the mastoid appears as a square bone surrounded by the squa- mosal, temporal, parietal, supraoccipital, exoceipital and tympanic. Gradually, however, the squamosal encroaches upon the mastoid and in Q leaves only a small roughly triangular portion of it exposed. The base of the triangle bears the mastoid process. The mastoid fuses with the petrous at an early age. The two are not clearly separated from each other by suture in F. It will be recalled that at this time the mastoid portion is not fully ossified. Internally the mastoid is per- forated by the large aperture of the parafloccular fossa. Below the anterior half of this aperture, on the petrous, there is a depression into which the internal auditory meatus and a second canal, probably the facial, open. In a stage slightly younger than A three clearly defined parts of the petrosal can be recognized. The anterodorsal, or squamosal portion, soon fuses with the underlying petrous portion proper. At the stage slightly earlier than A, referred to above, the squamosal portion bears a downward facing shelf against which the posterior end of Meckel’s cartilage abuts. Later, a cup-shaped depression or socket develops at this place to receive the posterior end of Meckel’s cartilage. TABLE 4 MEASUREMENTS OF MANDIBLE—DIFFERENT AGES SELECTED ON BASIS oF EQUAL AMOUNTS OF INCREASE IN OCCIPITO-NASAL LENGTH Distance Greatest Length between Greatest Depth at | Height of length anterior glenoid thickness Pmyz coronoid to Pm; fossae below Pm; process Cee a ee Fa 14.0 5.0 9.0 1.0 ae 4.0 CEA ree: ee ee 19.0 6.4 15.0 240 4.2 6.7 1 ERS ASS ee RR. eo 24.0 8.2 AO 2.9 5.0 9.5 pee ee ace en ee aie 30.5 10.0 20.0 3.8 a0 14.1 (Oi eederest y eae GaMe br Aak s 40.0 12.0 27.0 4.6 9.3 213 Amount of increase...... 26.0 7.0 18.0 3.6 6.8 17.3 Per cent of increase....| 186 140 200 360 272 432 34 | | 35 i e / Bi; \ 8 ee Figs. 28-39. Views of left mandibles. Figs. 28-33, medial view of mandi- bles; Wig. 28)'0, >< Is Pig. 29.1 xX 1 Big, 30, FF, X ie Pig. 3k, xX 16: Fig. 32, C, X 1%. Fig. 33, B, X 3. Figs. 34-37, posterior view of mandibles. Innes ae, (5 Se le dei. Bas J OK IB, Bie ely Jao ISAS Lies Git Wn DS OY 7ic Figs. 38 and 39, dorsal view of mandibles. Fig. 38, Q@, X %. Fig. 39, EH, XA 388 University of California Publications in Zoology [ Vol. 21 D and E (see figs. 30-33) the condyle projects far behind the angle, but in older skulls the greater posterior projection of the condyle is much reduced. The inferior dental foramen comes to be located nearly half way between the condyle and M;. In £ it is immediately behind the pocket of Mz. The dental foramen likewise moves farther forward on the outside of the mandible. Viewed laterally the chief changes seen in the mandible are those due to the relatively greater increase in depth than in length. As will be seen from figures 28-37 and tables 2 and 4 these increases vary greatly in amount. In depth the relative amount of change is greatest in the post-alveolar portion of the mandible. Measurements of the height of the coronoid process, which also reflect the ventral growth of the angular portion of the mandible, show that the increase is 432 per cent whereas the increase in depth of the horizontal ramus is only 272 per cent. From E to Q the portion of the mandible anterior to Pm; increases in length 87 per cent, the tooth row 49 per cent, and the portion posterior to the tooth row 320 per cent. By far the greatest part of the increase is made behind the tooth row. This is in contrast to the relative amounts of increase in length of the corresponding parts of the cranium. That portion of the cranium anterior tg the tooth row increases 94 per cent from E to Q, which is less than the increase of 77 per cent made during the same time in that portion of the mandible posterior to the tooth row. In both the cranium and the mandible, however, there is a pro- portionately smaller amount of inerease in the length of the maxillary tooth rows than there is in the portions either anterior or posterior to them. Comparisons of amounts of increase in depth in these three corresponding regions of the cranium and mandible show that the greatest relative amounts of increase occur in the post-alveolar portion of the mandible and in the pre-alveolar portion of the cranium. The least relative amounts occur in the pre-alveolar portion of the mandible and in the post-alveolar portion of the skull. In depth the relative amount of increase in the alveolar regions of both the cranium and the mandible is intermediate in amount. The depth measurements of the skull are the three depth measurements given in table 2. The percentages of increase from posterior to anterior are: 90, 96 and 188. The post-alveolar and alveolar depth measurements of the mandible are given in table 4. The pre-alveolar depth measurement was taken at the point of least depth in the diastema of the mandible. The per- centage of increase in depth of the mandible at these three points from posterior to anterior are: 432, 272 and 64. From figures 38-39 it will be seen that the mandible straightens with inereased age. The 1926 ] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 389 proportionately greater increase in length of both the post- and pre- alveolar portions of the mandible than of the alveolar portion appears to be a factor involved in the straightening of the mandible. Another factor is the assumption by the whole post-alveolar portion of the mandible of a position lateral to the tooth row. The changes involved in the assumption of this lateral position are better shown by figures 34-37 than by any amount of description. Points that will be noted in these same figures in addition to those already indicated are: the reflexing of the angle, the change in position of the condyle, and the development of a more vertical posterior margin of the mandible. It may be noted that the assumption of a position lateral to the mandi- bular tooth row is reflected in the increase of 200 per cent in distance between the glenoid fossae from C to Q while the mandible is increasing only 186 per cent in length. Meckel’s cartilage was noted in D in the course of preparation of the skull but unfortunately its relations to adjacent structures were not recorded. In E the dentary has grown around Meckel’s cartilage, converting the Meckelian sulcus into atube. Within the lower portion of the posterior half of the horizontal ramus this tube is revealed by several openings over which ossification has not yet occurred. In this region of the mandible, Meckel’s cartilage appears to be absorbed. In stage B, represented by figure 33, there is a rather abrupt truncation of the cartilage proper a short distance behind the lower end of the ineisor pocket. Between the end of this pocket and the anterior end of Meckel’s cartilage there is a cylindrical area of tissue which repre- sents the anterior end of Meckel’s cartilage but which is, at this time, of an entirely different nature than the posterior part, having the appearance of being chondrified. Examination of this area in D shows a cavity ; thus it appears that the part of Meckel’s cartilage that hes within or along the median side of the mandible is largely if not wholly resorbed. In B Meckel’s cartilage is not straight but is bent at three places; it lies along the extreme ventral margin of the mandible and at this time actually projects shg¢htly below the ossified horizontal ramus (see fig. 33). The posterior end of Meckel’s cartilage fits in a socket of the petrosal, beneath the squamosal and slightly above the ventral margin of the latter. If Kingsley’s general statement (1917, p. 82) that: ‘‘In development the malleus is the posterior end [italics mine] of Meckel’s cartilage’’ apples in this case—as it prob- ably does—a considerable change has taken place in the relative posi- tions of Meckel’s cartilage, the malleus, and the ineus. As shown in 390 University of California Publications in Zoology [ Viol: 21 figure 26 the incus is much nearer the posterior end of Meckel’s cartilage than is the malleus. PaO) 2a ns 1207 ya ade The incisors both above and below appear in D. The fourth premolars Teeth—The dental formula of Otospermophilus is: both above and below appear in F. Pm appears in G. The first molars appear in G but M,; is farther advanced than M+. The second molars are present in H, M; again being better developed than M2. M,; appears in J. M2® appears later but is well developed in J. The deciduous upper premolars are both replaced between AK and L. The lower permanent premolar does not appear until N. It is seen, then, that the upper and lower incisors and milk premolars appear at Figs. 40-43. Buccal and crown views of unworn right upper premolars, X 5. Fig. 40, L, permanent Pms*4. Fig. 41, H, milk Pm‘, Fig. 42, L, permanent Pms?4, Fig. 43, H, milk Pms®4, Explanation: ac., anterior cingulum; me., metacone; mec., metaconule; pa., paracone; pr., protocone. Terms used are for convenience of reference and are not intended to imply homologies. approximately the same time; the lower molars and the upper per- manent premolars appear earlier than their respective opposites. The crown surface of permanent Pm, presents no striking differences from the crown surface of the milk tooth either in size or in number and arrangement of the conids. Pm in both the milk and permanent dentition is spike-like and of little or no functional value; the milk tooth is only two-thirds the diameter of the permanent tooth. The roots of milk Pm* are more widely spread than are the roots of the corresponding tooth in the permanent dentition. Pm‘ of the milk dentition is at first set squarely in the maxilla but, as the permanent tooth develops, the preorbital portion of the skull lengthens and the anterior root of Pm‘ is carried forward. This appears to be a mechan- ical adaptation for forcing the anterior root of Pm‘ out of its socket, thus raising the tooth. The following differences are to be noted between unworn Pm‘ of the milk and permanent dentitions: The milk. tooth is 0.1 mm. shorter, 0.3 mm. narrower, and more triangular in 1926 | Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 391 crown view than the permanent tooth. In the permanent tooth the protocone is relatively higher and wider, and on the lingual side slopes less abruptly downward than in the milk tooth. One of the most striking differences lies in the anterior cingulum, which in the per- manent tooth is lower than the paracone or metacone but in the milk tooth is fully as high as either the paracone or metacone. In the per- manent tooth the paracone is higher than the metacone; in the milk tooth the two are of about equal height. The metaconule is fully as well, if not better, developed in the milk tooth than in the permanent tooth. In the latter the ridge connecting the metaconule with the metacone is better developed but the ridge connecting the metaconule with the protocone is better developed in the milk tooth. The posterior cingulum extends almost to the lateral margin of the milk tooth, but in the permanent tooth terminates only a short distance beyond the valley which separates the metacone and metaconule. The permanent tooth is more molariform than the milk tooth. However, paradoxical as it may seem, the milk tooth, when all things are considered, is more complex than the permanent tooth. Other changes in the teeth that oceur with advancing age are due to wear. In very old individuals the metaloph, paraloph, and loph of the anterior cingulum are very broad and connect with the larger protoloph. CRANIUM AS A WHOLE The foregoing discussion has dealt largely with individual bones or components of the skull. Changes that occur in the skull as a whole remain to be considered. It is obvious that any adequate comprehen- sion of these changes involves constantly keeping in mind the changes that occur in the component parts. Portions of the skull which the measurements given in table 2 show to have increased three or more times between the two points from which measurements were taken are: length of nasals, 370 per cent; postpalatal length, 324; zygomatic breadth, 288; depth of skull at center of palate between first premolars, 279; mastoid breadth, 270; fronto-parietal length, 265; depth of skull at anterior border of basi- occipital disregarding crest, 264; frontal length, 263; occipito-nasal leneth, 250; condylo-basal length, 244; parieto-interparietal length (not given in table 2), 244; diastema, 227; depth of skull at center of palate near posterior margin of palatine bone, 214; least interorbital breadth, 212; greatest width of nasals, 200. Measurements that show an increase of less than three times, with the percentages of increase, are: palatal length, 197 per cent; maximum breadth of braincase at base [Vol. 21 University of California Publications in Zoology 392 UCee RGEC VON GtoenG Gielen sa| yo (70 SE) 6 Soy OFS 9) ORT. 8 FE. .-9 be 9996 Org rs [Byidro.01seq jO Jop1oq 10110} UB 4B [[NYs Jo yydoq IGS Pen RO SGe Osean eee GOP Se Paz ob OP bP Ol ws Oke, | OG 108 |s05G = Pe ourzeyed Jo uls1eur Iotieysod «ivou [[nys jo yydaq Cie aU Ore eOsces inert OOK |e Vee) OL Gil | Glee 9 —(se Teoh OG Qe cae: pautour -o1d 4s UseMyoq [[NyYs jo yydaq Cee Reece peer ioler Gailit. a |SelGe! OW i OlcG | F Zhe 2°]. |-O°8) Po Tk | S39 |eSt 88 oo sassoo0id o1yBUIOSAZ Jo aseq qB osvo uleiq JO YIpPIM WUMNUIIXEpY Po ees cet en OL es Guc le OSs 1G Oe |G. P| Ores leliGe (=F PI] OF6GS So. 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Ton | OnGte legis |OlG weet | 6 ery 2S. | TD pgs [eee res [Byarred -19}UI pue sjejolied jo y4suey Pie nC Ole ON Ota | ees 20 Gon leGr Zany |v Gla Gco.| 0-8 | 0 St FV OPee| 97g) PE | ee ere tre es syequody JO YySuaT OLE eee SPONGES ceatGsVGaleOrOvs el Sealer S Ol OT Oxce lee Olal Oo PL | OF G Wek | Lape aje ee ee s[Bseu JO Y}sue'T OlGee Pee sra soon st Ose OnOUs sora |e ME TGS Ole) eS see Le O9G: Seth £86, Pp Lhe eo qysuay Pesvu-ojdr199() eee eet ee lees eee (ooo lee lige it 72) esol ap 2 | ? BS | p= | 88 | Se 3 | Be 83 | 82 £3 | Be = 5 ge oe os 65 oe oe oe oe at Bee eel] sai= a 2] 2 a hee gel 8 ieee ile | a f | 3 YOO SUSVAYONT UWSHHL HOIHM LV AW, IHL ANV SEINANAYNSVAY TVINVUD IVYHAUG NI AUSVAYONT WO SINAD YA ANV SLNNOWY PNIMOHS GS WIdVL 1926] Hall: Skull of the Rodent Otospermophilus grammurus beecheyi 393 of zygomatic processes, 176; breadth of rostrum, 174; least postorbital breadth, 127; Pm. m. tooth row, 123; width of parietals (dorsal), 110; palatal width at alveoli of first molars, 90. Of the regions measured the percentage of increase is least for the palatal width at the alveoli of the first molars; thus it represents the most stable portion of the skull, of which measurements were taken. In fact, as will be seen from table 2, this measurement is greater in F and in a number of succeeding skulls than it is in Q. The measurement of the length of the nasals shows the greatest amount of increase, 370 per cent. This, then, appears to be an unstable region of the skull. Pebruary, AQe6 po ee ek “60 11. Mitochondria in Huglena gracilis Klebs, by Dayid Causey, Pp. 217-224, Ee a se ‘plates 19-20, Cae Sa EE 12. Mitochondria, th Noctiluea seintillans ‘Macartney, 1810), by. David Causey. ees fae Pp. 225-230, plate 21. “Mitochondria in Ciliates with Especial ‘Reference to Paramecium » Ebr A - by: David Causey. Pp. 231-250, plates 22-24. ; Nos.-11, 12; and 13 in one cover. February, 1926 .2....-20cc20 ee 14, Studies ‘on the Ingestion of Leucocytes, and on Mitosis, in Endamoeba: ~ - gingivalis, by Horace J. Child. Pp. 251-284, plates. 25-29, 9 figs. in text, ae PODTUBIY, = LO26s i eS Neg eee TS aa a na, cos > 415. On Ozymonas,’ a Flagellate with. an Extensile and Retractile Proboscis, from Kalotermes from British Guiana, by Charles A. Kofoid and -Olive oH ARG Swezy. Pp. 285-300, plate 30,5 figures in text) 0c Ae : 16. On Proboscidiella. multinudleata gen. nov., sp. nov., from Planocryptotermes he ae nocens From the Philippine: Islands, by Charles. A. Kofoid and Olive Swezy. Pp. 301-316, plates 31, 32, 4 figures in text 2.1.2 Nos. 15 and 16 in one cover, February, BOZG 28 Sie oe ae Ne eee ; 17. On the Cestode Genus Dipylidiwm from Cats and Dogs, by Theresa Marie : Millzner, Pp; 317-356, plates 33-39. February, 1926 220. 18.\A Useful. Modification of a. Clearing Fluid Formulated by, Spalteholz, by. . Franklin P. Reagan. Pp. 357-359. 19.-The Earliest Blood. Vessels of the Manimialian Embryo, Studied by Means of the Injection Method; by Franklin P. Reagan. Pp. 861-364. a =Nos. 18 and’ 19 in one cover. February, 1926 «2.0.0.0. ee we 20, The Biological Relationships of Leishmania and Certain Herpetomonads,. by. Hdna Hannibal Wagener and Doronty. Ann Koch. Pp. 365-388, < eet AQ—AS. 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Grinnell, Editors. > ans This series contains the contributions from the ‘Department ‘of - Zoology, 1 stem. ee a “Marine Laboratory of the Scripps Institution of Oceanography, “at La. he Calt- ee fornia, ann trom Sie California Museum of Vertebrate POOR Y in: “Rorkeley.. Bien Cited as ‘Univ. Calif. Publ. Zool.” ees Fie ie eae Volume 1, 1902-1905, 317 pages, with 28 plates 20.02... ostceinepternencaetense Die oe $3.50 Volume 2, 1904-1906, xvii +382 pages, with 19 plates - eg eS eee Volume 3, 1906-1907, 383pages, with 23 plates. Ee ee ai - Volume 4, 1907-1908, 400 pages, with 24 plates -.; Pia ae aE Es Volume 6, 1908-1910, 440- pages, with 34 plates -_.... Volume’6, 1908-1911, 478 pages, ‘with 48 plates ...... 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Behavior of the: Leaf-nosed: Snake, Phylloriynchus deourtatns, by Sarah” re ok Rogers Atsatt.. Pp. 303-312, . February, 1923 22.2... cceeic cece apntonn depen ABR cs tad ae. 10. A Systematic List of ‘the Mammals of California, by Joseph Grinnell. = eee ae Bh. S16 8245 < Fanary,” WOes ce Red ae el hic eaten rece bie wath ee - “41, The Systematic Status of the. Mountain Lion of California, by Toseph’ & Grinnell and Joseph Dixon. Pp. 325-332, plates 9-10... April, 1928 = 4,25 12. New. Subspecies. of Birds Jeece Patagonia, by: Alexander. Wetmore. VER, = Sir ie oy Jee prt ct een ks)? Gynt cde Gi font ocular’ Sermo tate coe eh moc ERT ET Se 2B 18. Revision of the Genus Lynz re California, by Joseph Grinnell and Joseph sek Se Dixon. Pp, 339-354, plate 11, 1 figure’in text. January, 1924... 26 < 14. Changes during Growth in the Skull of the Rodent Otospermophilus gram- ; murus beéecheyi,; by E: Raymond Hall. Pp. 355-404, 48 figures In text. gingivalis, by Horace J. Child. Pp, 251-284, plates 25-29, 9 Ags. in text, Sy : x) pipe Oe ie ESR So cece eins ey in MASP RCN AAS eS ee a BS oe from Kalotermes’ from British Guiana, by. Charles A. Kofoid and Olive “nocens From the Philippine: Islands, by Charles A. ‘Kofoid and Olive Swezy.. Pp. 301-316, plates 31, 32, 4 figures in text 2.05.00. k lie “*. . Nos. 15 and-16.in-one cover.- February, 1926 Ses lhe see ae 17. On-the Cestode Genus Dipylidiwm from Cats and Dogs, by Theresa Marie Ee -Millzner, Pp. 317-356, plates 33-39: February, 1926 —=.- ae ae aetna 18. A Useful Modification of a, Clearing Fluid Formulated by Spalteholz, by. ' Franklin P. Reagan... Pp..357—-359. 19, ‘The Earliest Blood Vessels of the Mammalian Embryo, Studied by Means ‘of the Injection Method, by Franklin P; Reagan. Pp, 361-364. se -+ Nos, 18 and. 19 in:ene cover, February, 1926 <2... on 22k octane cntine ‘20. The Biological: Relationships of Leishmania and: Certain ee oe ‘by Edna. Hannibal- Wagener and Dorothy Ann Se Pp. 365-388, : me cele 40-48. March, 1926 a ia apace ee eg Saacceeniee see acer tamed eee ee - 15, On. Oxymonas, a Flagellate. with an Extensile and Retractile Proboscis, ken, Swezy. Pp. 285-300, plate 30, 5 figures in. text: 222 2..s sci it cece ence poet d 2 or, On Proboseidiella multinuéleata gen. Nov., sp.-nov,, from Planocr purateraies = , 60” 10, On Oxyphysis. omytoxoides gen nov., sp, nov., A Dinophysoid DinoHagellate a, 25 40 ~ 40 ‘ny a | | UNIVERSITY OF CALIFORNIA PUBLICATIONS - eee ee - Note.—The University. of California Publications are offered in es for the pani © et ~ eations of learned societies and institutions, universities and libraries,. Complete lists of” Salk the publications of ‘the University ,will he sent upon request. For sample copies, lists _ of publications or ‘other information address the MANAGER. OF THE UNIVERSITY — PRESS, BERKELEY, CALIFORNIA; U.8,A.. All matter sent in exchange should ‘be. addressed to- EXCHANGE. 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December, 1919.25 4, Revision of the Avian Genus Passerella, with Special Reference to the i oe hoe Distribution and Migration of the Races in California, by H. 8. Swarth. Tk pa ES Pp. 75-224, plates 4-7, 30 figures in text... September, *1920 -...--_.--_. ae 1.75. , i 6. A Study of the California Jumping Mice’ of the. Genus Zapus, by A. Brazier if Warne Howell. Pp. 225-238, 1 figure in ests ERS, 4980) 2 a ee ere ti tana 6. Two New. Rodents (Genera Thomomys and- Marmota): from the ‘astern Border of California, by Joseph Grinnell. Pp. 239-244, 6 figures in'text A November, L920 onesies cec ete cetenennettncnerenenatntnnentencmesengnnentnntens st ieeenae See L Sa eae a 7. A-Study of the Californian Forms of the Microtus montanus Group of Batty, ae. ‘Meadow Mice, by. Remington Kellogg. Pp..245-274, 25 figures in text. “ 8. A Synopsis of the Microtus mordaz Group of Meadow Mice in California, Bor: by Remington Kellogg. Pp. 275-302, plate 8, 29 figures a: Geb Fh AMES ae ie Nos. .7 and 8 in: one cover. April, ODO gies a Ry at Racey {iets 9. Behavior of the Leaf-nosed Snake, Pliyllorhynchus decurtatus, by Sarali 0).7% % ~ Rogers Atsatt.. Pp: 303-312, February, 1923 1... snen snes snienenn nase ennennesene et SOD oe SO a 10, A Systematic List of the Mammals of. California, by. J oseph Grinnel Bay i Pp, 313-324. January, 1923 nlc eee at eaensac eens aup denn maensenasneeeeccmanenwewehbieresndacnncaeshnesaensne 3 .25 2 t =e Tl, The ‘Systematic Status of the Mointain Lion of California, by ouapr Koh ae, Grinnell and qesene Dixon, * re: oe plates’ 9-10. alae M923 oS OB gam ass St eee edhe? og = os TWO NEW RACES OF THE PINE MARTEN PROM Ee PACTEIC: COAST OF NORTH AMERICA BY JOSEPH GRINNELL anp JOSEPH DIXON UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Volume 21, No. 16, pp. 411-417, 9 figures in text Issued March 17, 1926 UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA CAMBRIDGE UNIVERSITY PRESS Lonpon, ENGLAND TWO NEW RACKS OF THE PINE MARTEN FROM THE PACIFIC COAST OF NORTH AMERICA BY JOSEPH GRINNELL anp JOSEPH DIXON (Contribution from the Museum of Vertebrate Zoology, University of California ) In working out the natural history of the fur-bearing mammals of California it has seemed desirable to establish the systematic status of all our species and races in so far as materials and opportunity might permit. Coming to the Pine Martens, our enquiry discloses that there are two well-defined subspecies within the state, one of which appears to be new; and comparisons of this subspecies with other Pacific Coast forms show that there is yet another subspecies that is unnamed, on Vancouver Island. The new races are now characterized, as follows. Martes caurina humboldtensis, new subspecies Humboldt Pine Marten Type locality—Ridge about five miles northeast of Cuddeback {[— Carlotta], Humboldt County, California. Type, male subadult, skull-only ; no. 19158, Mus. Vert. Zool.; obtained from local trapper by H. E. Wilder (orig. no. 1368) ; February, 1913. Material—Four skulls-only from the type locality; two skins with skulls (one with complete skeleton) from head of ‘‘Terwak’’ [= Terwer on U.S. G. S. Preston Peak quadrangle] Creek, 10 miles ‘northeast of Requa, Del Norte County, California. Diagnosis——In general external features, including dark tone of coloration, resembles Martes caurina caurina (C. H. Merriam [1890, p. 27]); decidedly darker, of richer golden brown tone, than M. c. sierrae Grinnell and Storer (1916, p. 2). Skull smaller than in either of these races, with rostrum slender, distinctly constricted behind roots of canines; frontal region and postorbital constriction narrow ; teeth small; molariform series not so crowded as in sierrae, So as to skew the premolars; last upper molar with heel greatly expanded anteroposteriorly, as much so, relatively to transverse diameter of this tooth, as in sierrae, and hence more than in caurina. (See figs. 1-8.) Fig. 1 Fig. 2 Fig. 1. Mustela caurina caurina, [gf] adult; no. 1063, coll. S. G. Jewett; Skagit Valley, 4 miles north of International Boundary, British Columbia; Sep- tember, 1920. X 1. Fig. 2. Mustela caurina vancouverensis, § adult; type, no. 12474, Mus. Vert. Zool.; Alberni, Vancouver Island, British Columbia; July 20, 1910. X 1. Fig. 3 Fig. 4 Fig. 3. Mustela caurina humboldtensis, § subadult; type, no. 19158, Mus. Vert. Zool.; Carlotta, Humboldt County, California; February, 1913. X 1. Fig. 4. Mustela caurina sierrae, & subadult; no. 24822, Mus. Vert. Zool. ; Bullfrog Lake, Fresno County, California; August 29, 1916. X 1. 1926] = Grinnell-Diron: Two New Races of the Pine Marten 413 Geographic range-—Known definitely only from within 50 miles of the seacoast in Humboldt and Del Norte counties, California. Evidence indicates probable occurrence, at least within historic times, north through the humid coast belt from the vicinity of old Fort Ross, Sonoma County, into Oregon; but material from the latter State is almost wanting, and nothing can now be said on any basis of fact as to what happens subspecifically between the California-Oregon line and the type-loeality of M. c. caurina (vicinity of Grays Harbor, Washington). See accompanying map (fig. 9). Measurements—The only reliable external dimensions available for this subspecies are of an adult male (no. 35372, Mus. Vert. Zool., taken near Requa, Del Norte County, California, January 14, 1926), as follows: Total length, 603 mm.; tail vertebrae, 210 mm.; hind foot, 84 mm.; height of ear above crown, 35 mm.; weight (lean), 755 grams. The weight indicates an animal decidedly lighter than sierrae (nine males of which average 988 grams) ; but the external dimensions fall well within the range of those of sierrae. Cranial measurements are given on the accompanying table. The last upper molar of the type of humboldtensis measures : trans- verse diameter 7.8 mm.; anteroposterior diameter of heel 5.2. The same tooth in the type of sterrae measures: transverse diameter 9.2 mm.; anteroposterior diameter of heel 5.9. The same tooth in caurina (no. 1063, eoll. S. G. Jewett) measures: transverse diameter 8.6 mm.; anteroposterior diameter of heel 5.5. Remarks.—Martes caurina humboldtensis is sharply distinet from M. c. sierrae. A series of seven martens at hand collected by Annie M. Alexander and Louise Kellogg in the Trinity Mountain region of northwestern California (see L. Kellogg, 1916, p. 355), while not typical of sierrae are both externally and ecranially very.close to that race and not, as might be expected, fairly intermediate toward humboldtensis. Indeed, no specimens yet available to us indicate intergradation between humboldtensis and sierrae. At the same time, normal geographic intergradation probably takes place to the north- ward between each of these races and caurinda. The only full specimen, that is, skin with skull and external measurements, available is unfortunately abnormal as to cranial characters (due apparently to early injury). And since the chief characters of the new race lie in the skull, it has thus been necessary to choose a skull-only for the type. Acknowledgment should here be made of the vigorous efforts in our behalf, by Mr. Harry E. Wilder, of Carlotta, to get specimens of this rapidly disappearing fur-bearer. 414 University of California Publications in Zoology (Vou. 21 Martes caurina vancouverensis, new subspecies Vaneouver Island Pine Marten Type locality—Golden Eagle Mine, 20 miles south of Alberni, Vaneouver Island, British Columbia. Type, male adult, skin with skull; no. 12474, Mus. Vert. Zool.; collected by E. Despard (orig. no. 41); July 20, 1910. Material.—Six specimens, skins with skulls and three of them with complete skeletons, from the type locality and from Errington, Vancouver Island, B. C. ; Fig. 5 Fig. 6 Fig. 7 Fig. 8 Fig. 5. Mustela caurina caurina, [gf] adult; no. 1063, coll. 8S. G. Jewett; Skagit Valley, 4 miles north of International Boundary, British Columbia; Sep- tember, 1920. X 1. Fig. 6. Mustela caurina vancouverensis, J adult; type, no. 12474, Mus. Vert. Zool.; Alberni, Vancouver Island, British Columbia; July 20, 1910. Xx 1. Fig. 7. Mustela caurina humboldtensis, g subadult; type, no. 19158, Mus. Vert. Zool.; Carlotta, Humboldt County, California; February, 19138. X 1. Fig. 8. Mustela caurina sierrae, 3 subadult; no. 24822, Mus. Vert. Zool.; Bullfrog Lake, Fresno County, California; August 29, 1916. X 1. Diagnosis —Externally like Martes caurina caurina (C. H. Mer- riam [1890, p. 27]), but skull markedly different: rostrum shorter ; frontal region and, indeed, whole cranium broader; zygomatic spread greater; molariform teeth more crowded and more skewed out of alignment; last upper molar decidedly smaller and its heel far less expanded anteroposteriorly. (See figs. 1-8.) Skull in general pro- portions more nearly like that of Mustela nesophila Osgood (1901, p. 338, pl. 5), but smaller (see Swarth, 1912, pp. 104, 105) and with weaker dentition; last upper molar much smaller, and heel even less expanded, thus nearly as in Mustela americana. Geographic range—So far as known, only Vancouver Island, British Columbia. 1926] Grinnell-Dixon: Two New Races of the Pine Marten 415 Measurements——The skull of the type presents the following dimensions: greatest length of skull, 78.8 mm.; condylobasal length, 77.2; basilar length of Hensel, 69.5; palatilar length, 37.2; zygomatic breadth, 49.4; mastoid breadth, 36.6; breadth across postorbital processes, 24.2; interorbital width, 19.3; width of rostrum at bases DISTRIBUTION MAP MUSEUM OF VERTEBRATE ZOOLOGY UNIVERSITY OF CALIFORNIA SCALE J ° 25 so MILES ORAFT of 1919 ub Fig. 9. Map showing distribution of Pine Martens in California: a, Mustela caurina sierrae; b, Mustela caurina humboldtensis. Based on eateh of licensed trappers of California for the five-year period 1919-1924, as reported to the State Fish and Game Commission, each dot being the location of a trapper who caught one or more martens. Presumed general range of each subspecies within history, outlined. of canines, 17.6; height of brain-case at bullae, 31.0; anteroposterior diameter of heel of last upper molar, 4.8; transverse diameter of same tooth, 8.3. For dimensions of other skulls of vancouverensis, as also of nesophila, and for body measurements of both these races, see Swarth (1912, p. 105); and for dimensions of skulls and teeth of caurina see C. H. Merrian (1890, pp. 28-29). [ Vou. 21 alifornia Publications in Zoology 1/ J University of € 416 € L6G 0 66 -UIBIG JO JYSIOFT avy[nq 4B aso VINUOAITV( ‘SAILNOOD ALHYON Taq ANV oO oD — SFr! oN 1d +H 19 a) ‘$I {II ud OD a4 UINISOI JO YBPIAL Ven) ~ MDH HO ‘OT ‘$I {VII na! ZL ‘BM (9)! yapea [B3tqs1010} UT jor) oS N sassao0id yeq1q103s0d SSO.108 YIPBeig ioe) N 6 Ui LLY 9 €& PSE 9 6 4°) Ws GIL 0°62 VINUOAIIVD ‘VOVAAN VaadIgQ AHL pion) oD APB PIowsE]y G88 € 8€ qypBesq OIVBULOSAT 8 08 PGE OSE v 9€ 8 9E yysug] epee N yyoug] Iepiseg 0°29 G89 [eseqo[Apuod Re ae ck ot ae So[BUldj JNpeqns puev J[Npe F jo ISBIIAYV sojvul J[Npeqns puB 4[NpwB OT Jo esvicAy WOUd ADIWAIS VULINDI $9) LD J [mys jo Y}sug] 4So}VII4) GZ6I “EZ “99 €16l “ue EI6l “UBL S16, “uBl S16l “49a a1eq ‘peqns ‘pe ‘peqns “‘peqns “‘peqns a5V 69ECE POT6I LST6I SOI6l SST6I 7° AW ‘ON LUIOMWAPT WOU SIsUa;pjoquny DULINDI sa)ID JY AO STIAMY JO (SAOJOUIT][LUT Ul) SLNAWAMASVATY 1926] = Grinnell-Dixon: Two New Races of the Pine Marten 417 Remarks.—Although vancouverensis and nesophila are both de- scribed from islands, they obviously belong to the same series of geographic races with caurina of the mainland. Furthermore, indi- vidual variation in the several diagnostic features brings very nearly complete bridging between them. For these reasons, the trinomial seems best to employ; and Osgood’s M. nesophila becomes Mustela caurina nesophila. The material as yet available for study indicates the existence of two distinct species of marten in northwestern America—Mustela caurina, with short bullae and other minor echar- acters, on the Pacific Coast from Admiralty Island south to east- central California; and Mustela americana, with long bullae, ete., in the interior of the continent and west at the north clear to the Alaska Peninsula. Each of these species contains several well-marked geographic races. The authors are indebted to Mr. Stanley G. Jewett, of Portland, Oregon, for the loan of a series of specimens of Martes caurina caurina from his private collection of mammals. LITERATURE CITED ’ GRINNELL, J., and Svrorer, T. I. 1916. Diagnosis of seven new mammals from east-central California. Univ. Calif. Publ. Zool., 17, pp. 1-8, 1 fig. in text. KELLOGG, L. 1916. Report upon mammals and birds found in portions of Trinity, Siskiyou, and Shasta counties, California, with description of a new Dipo- domys. Uniy. Calif. Publ. Zool., 12, pp. 335-398, pls. 15-18. MerrIAM, C. H. 1890. Descriptions of twenty-six new species of North American mammals. Deseription of a new marten (Mustela caurina) from the northwest coast region of the United States. U. 8. Dept. Agric., Div. Orn. and Mam., N. Am. Fauna, 4, pp. 27-29. Oscoop, W. H. 1901. Natural history of the Queen Charlotte Islands, British Columbia. U. S. Dept. Agric., Div. Biol. Surv., N. Am. Fauna, 21, pp. 7--50, pls. 1-5. SwartTH, H. S. 1912. Report on a collection of birds and mammals from Vancouver Island. Univ. Calif. Publ. Zool., 10, pp. 1-124, pls. 1-4. Transmitted February 1, 1926. ats cast Bin ee + Ae ae =e “UNIVERSITY oF CALIFORNIA PUBLICATIONS—(Continuea)_ * a8 ‘New Subspecies of Birds: from ‘Agee aee by Alexander ‘Wetmore. Pp. & 833-337. Jine, 1923 SRC SS AND RANMA Rat RTI LEED pt Myc BoP cal See NES RIG SEIN rn 18, Revision of the Genus Lyng in. California, by Joseph Grinnell and. Joseph Be Dixon. Pp. 839-364, plate 11,1 figure in text. January, BILE ie ae Maes a ‘Changes during Growth in the Skull of the Rodent Otospermophilus gram- ‘murs beecheyt, by. “a codes crachgt # iat, PP: 855-404) a8 figures. in text. : SERCH SALE To SS a ee aie eet ear aS Ls aged a 15; os New Race of. the White-breasted Nuthatch from Lower. ‘California, a by Joseph Grinnell. Pp. 405-410. ‘March, 1926... Ce Ne a. Sa ae p 16. Two New Races of ‘the Pine Marten from: the. Pacific. Coast of North.” America, by Joseph Grinnell and Joseph” pone Pp. 411-417, 9 figures ya Bers a Mad 1 GS a) | ENN ed «bie gue rN A eS Ue MMS US Ya ae aan CN NA aD SR 17. ‘The Trout of the Sierra San Pedro Martir, Lower California, ay John BAe hip Otterbein. ‘Snyder, Pp, 419-426. March, 1926" opts ae hee ES oe ee E Uhutene ses ‘Volume 22. 1920-1923, 485 pages, with 24 plates pn eeneemenneeeneen ns oe 3 Vol. 23, ‘The Marine Decapod Crustacea of California,. by. Waldo Tie Schmitt. ‘Pp. 1-470, plates: 1-50, 165 figures in text.® May, 1921 csc secieeesnceenetone se Vol. 24, 2s A ‘Geographical Study of the Kangaroo Rats of California, he Joseph ~ a : Grinnell. Pp..1-124, plates 1-7, 24 figures in: text. June, “1922 20s 2, Birds and Mammals of ‘the: Stikine River Region of Northern’ British’ eee Columbia and Southeastern Alaska, by H. 8. Swarth. Pp, 125-314, plate AB SA OUPOS AM TER. td WMO, 1 ORR a a a es i ~ Be Birds and Mammals. of the. Skeena ‘River. Region of Northern British © Columbia, by Harry S. Swarth. Fp. 315-394, plates oi) 1 sigure in text. January, - 7 Gg RUE SOARES Se ETS LDS Seer PT ERE, HRN AE a ge ale sae Fe Sa : x Vol 25. A Bibliography of Eugenics, by. ‘Samuel J. Holmes. Pp. 1-514. Bi anuary, 1924 aes Volume 26, 1923-1925, 453 pages, 34 plates esteem hr eta ae Sg BML PSIG Teel ARR SE : Wok. 27. A Synopsis of the Amphibia of California, by Tracy I. Storer, Pp. 1-342, ~~ < plates. 4-18,;-42 figures-in texts- June; 1920". “Vol, 28, 1, Mitochondria and Golgi Bodies in Endamoeba gingvoalie (Gros) ‘Brumpt, by David Causey. Pp. 1-18; ‘plates 1-3. 2, Mitochondria in. Leishmania brasilionste, by David Causey. Pp. ee plate 4, Nos, 1 and 2 in oné cover. November, MAQG kt pe ee en, Be 3. Mitosis in Ceratium hirundinella:O. F. M., with Notes on ‘Nuclear’ ‘Phenom- ena in Enucysted Forms and the Question of: Sexual Reproduction, by - Richard P. Hall. Pp. 29-64; plates 5-9, 5 figures in text. November, 1925. 4. The Cultivation of Endamoeba gingivalis (Gros), by Beatrice Fay Howitt. Pp. 65-126, plates 10-12, 4 figures in text. . November, 1925 2020000120. 6. The Behavior of Endamoeba dysenteriae in Mixed Cultures with: Bacteria, by. Charles A. Kofoid and Sue Hannibal ashes ecto Pp, 127-154, plates 13-15. 6, Studies of the Effects of ‘Certain Drugs upon ‘Endamoeba dysenteriae at) wares by Charles A. Kofoid and Edna Hannibal Wagener. Pp. 155-166. i “Nos. 5 and 6 in one cover. November, 1925.) ste eeteen sted cen ee aye Some New and Some ‘Previously Unreported Hydroids, Mainly from the > Californian Coast, by -C, McLean Eraser, atk 167-172, 7 figures in text, December, 1925 Bins CR eens een a Pate Sd DN ORE Or Ages BEATS CES Te Me NBER EIN aN RGIS xe 8, The Effect of Certain: Drugs and ‘Dyes ‘upon the Growth ‘of Endamoeba oe gingwatis (Gros) im vitro, by Beatrice Fay Howitt, Pp. 173-182, _ 9, Experiments with Endamoeba gingivalis (Gros) in Mixed Bacterial Cul- tures; in Filtered Saliva; on a Solid Base;-and. with Peritoneal Celis; 25 “and with Digestive Secretions, by. Beatrice Fay Howitt. eye 183-202, plates 46 and.17. - : _ Nos.-8 and 9 in one cover. February, L926 ene sannsnnnsiendepesnnncenenssngaton 10. On Oxy phasis ouytoxoides Zen nov,; sp. nov., A Dinophysoid Dinoflageliate Convergent toward the Peridinioid Type, by teal A. Kofoid.- Pp. Sita aie 19. Pisa 1926 a sieakes erontendnenteoneneassnusedisnenennsenaaatarnaanse == Bo “plates 19-20, - ie Mitochondria in Nostituea s scintilans. s (Macartney, 1810), b PD, sae fips 21 S f “AL. Mitochondria Jn fer oee oracitie Kies, by David Cousey. Pp. 2 21-204, 2 ssh: “Nos. i, 12, dia. a. in ¢ one “cover. een 1926. ere Studies ‘on -the™ Ingestion of ‘Leucocytes, ge on. ‘Mitosis, as Endamocba . +. gingivalis, by” Horace: J. Child. ‘Pp. 251-28. : _ February, 1926- aie aeons - Swery. Pp. 801-816, ~ Nos, 15 and 16 in‘one cover. February; 4926-22 * = on the Cestode Genus Dipylidiwm from Cats® and. Dogs, by 1 161 > 3 Milizner. Pp. 317-356, plates’ 33-39, February, 1926. : ; 18. A Useful Modification: of a Clearing Fluid Formulated oy, Senegels, y Franklin P, Reagan. ni Agaeeses eek : lates 31, 32, 4. [ggerea in text of the Injection Method, by. Franklin P. stages ie oe _ Nos. 18: and 19 in one cover, February, 1926. 2 20. The Biological Relationships of ‘Leishmania and: Certain. ‘Herpetomonads -- Spy Edna Hannibal Wagener: and Dorothy “Ann Koch, . Pp. 365-388 ae 40-43. fost. A926: = ‘UNIVERSITY OF CALIFORNIA PUBLICATIONS Boge ie ‘Note.—The University of California Publications are offered: in’ exchange: for the tonbtis - cations of learned societies and institutions, Universities and libraries. Complete lists of — all the publications of the University will be sent upon request. For sample copies, lists of publications or other information address the MANAGER OF THE. UNIVERSITY “PRESS, BERKELEY, CALIFORNIA, U.S. A. All matter sent in exchange should be — addressed to EXCHANGE - DEPARTMENT, sesh aogesers ‘auripcanmat BERKELEY, 2 CALIFORNIA, U.8.A. 0 ; = “Publications of the University of California’ ‘Press may be “wbtaltied from’ THE CAMBRIDGE” UNIVERSITY PRESS; FETTER. LANE, LONDON, E.C. 4, ENGLAND, a *40 which orders originating in’ Great Britain and Ireland san be sent. ©: : ZOOLOGY.—C. A; Kofoid and-J, Grinnell, ‘Editors. os = eae Z ‘This series contains the contributions. from the Depactiient of eS from ‘the: pn Marine Laboratory of the Scripps Institution of Oceanography, at La Jolla, Cali- aioe fornia, and from the California Museum. of Vertebrate: Zoology ia ek pa Re ‘ Cited as Univ. Calif. Publ. 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December, 1919.25. 4, Revision: of the- Avian Genus d = S SS N = Sak OPE OLDS S21 OST, S81 iteeraike SoC Tes Oc Ole O0G ee rOG SCAT aL BGI Se eae S19} OUT] [TWA UT yy Sua] = o¢ SF 6¢ ze GP 19 19 Lt e) ec zg rele) 89 ze 09 Spree nate sla arbtniatnpsvalat eA ia'e(ats/a:-ininte-atetebstalbin\s/a(etn|otelsinior atte ie(araMsa(afaiaiitays eiavalulsiare ool-laiaia sibiMem ais(s'efale uy pepneg ‘= zg 9¢ o¢ 1g SF OF 6g 1g 1g 6g Ze 6h Ig CF is Sweet hiece Son caries aie Saas ag cere a gant uy [esiog 3 e ¢ 8 6 0 62 Po 1z 9Z 0Z iG Ig 62 GZ I¢ Terre r ere re eee eee eee rere eee eee eee eee eee eee reer eT TT! 1ayaenb jerqueA IOII94SOg = re SITS Sch 28 68 doe Oo muGl aaluh ccele UG StObeeta sre (Oi Ss Se ee Jaqienb [esiop 10119}s0g e Z II Mike 0 ra 29 liz 9Z GE OL al 0g 0g ZG Ch Fem et eee eee t eee eee e meee tenes erate reese sues esenessoene iayr1enb [vaquoA IOLI0JUY A Co Vela Srl Pe; SP: Cr ere OGlemOSss Ohta ob CVs CO NOT acON Wet Be ie salon tees Oe Jay1enb [esiop 1olezUVy ‘= ee QZ OF OF Wee €Z 0g IZ 12 91 12 ee 0g Ze 6E dete eee n econ ee rece tance ttre pees enenseenessnssesecsaressensssscenesseseeasssessssessessesses peoyy a “JBOD ‘Ary oreleg ‘snap ourpeg TuOs[aU OWES = - S syods jo oquinyy ~~, S) = 3 WIaAVL ne s 422 1926 | Snyder: The Trout of the Sierra San Pedro Martir 423 In color the species may be considered as a profusely spotted form. In addition to the parr marks, which seem to become indistinct or to disappear entirely with increasing age, the whole upper portion of the body from the snout to the tip of the tail is pretty well covered with black spots, especially in the darkest individuals. The spots of the head are very distinct and round. On the opercles they are in some cases a little less distinct in outline, and always larger. The spots of the body are not regular in outline, as they are made up of closely associated pigment specks. In some regions they are inclined to be irregularly stellate, somewhat reticulate, or broadly linear. From about the middle of the body to the base of the caudal fin they grad- ually increase in size, an occasional one approaching the pupil in diameter. Not many spots occur below the lateral line, and most of these are small, indistinct, and somewhat linear in outline. ‘These are usually assembled in the region of the pectoral fin and on the caudal peduncle. An occasional specimen is well covered with spots on the entire lateral region excepting the belly. The spots on the dorsal fin are oblong and arranged in series, and those of the basal and anterior part of the fin are more nearly circular. The adipose fin is more or less completely edged with black and usually supports one or more round spots. The dorsal edge of the caudal is invariably supplied with from seven to ten well defined spots, while over the entire fin are scattered round or elongate spots, some near the posterior edge being markedly linear. About eleven broad parr marks appear along the lateral line. Alternating with these are smaller ones above and below. The anterior upper edge of the dorsal and the anterior lower edge of the caudal are broadly bordered with light color, as is also the anterior edge of the ventral. The pectorals, ventrals, and anal are without spots, although in some eases they are well diffused with black pigment. In order to express the numerical and regional variation in the number of spots with some degree of accuracy, they are tabulated (table 2) for ten individuals. The spots of five individuals of S. irideus are also arranged in the table. The body, conveniently divided by the lateral line into an upper and a lower half, is quartered by means of a vertical line through the anterior end of the dorsal fin. It would appear from this table that S. nelsoni might be sharply dis- tinguished among rainbow trouts as a color form; but when the field is widened and individuals from different localities are introduced, it does not stand out so sharply. The figure of a rainbow trout pub- lished in California Fish and Game (5, 1919, p. 114) furnishes a good 424 University of California Publications in Zoology [Vou. 21 example of an excessively spotted individual. The more intensely pigmented examples of S. nelsont have the dorsal, anal, and ventrals broadly edged with white, while in lighter-colored individuals the light area is often greatly restricted and much less evident. The species appears to exhibit a certain amount of reduction in the number of vertebrae, but until data from carefully prepared material representing a sufficiently extended geographic area are available for comparison the character cannot be relied upon as distinctive. An enumeration of twenty examples of S. nelsoni was made. In so doing the vertebral column was entirely denuded of tissue on one side so as completely to expose even the modified tail vertebrae, all of which are included in the count, which is as follows: TABLE 3 Number of specimens:: 2.3 oe ses ee vit eee cae tance iene 13 INtimMbentolavernte DEACw eee ere eeee ra ae 61 6 Non A tabulation (table 4) of the number of vertebrae of a series of specimens of Salmo irideus mostly from the coastal streams of north- ern California will serve for comparison with the above. It will be noted that, while an insertion of the counts of S. nelsoni might be TABLE 4 INtimiber role vierbe Dae anes: eee eet ee te 60 61 62 63 64 65 Noyo River, Northspur..............:.... Ba ktih 8 9 2 Tp! Tears TONED oat eheo sentence ctennceckonse 1 2 4 4 3 lata aytilanes Guia Type eee teeters eee eee 1 6 11 10 11 1 Klamath River, Hornbrook......................... 3 10 17 13 2 1 Camp:'Creek (Klamath)! aie. o iene ee 3 5 3 Malik @ reeks Chlammaitiin) peeeene eee meen: : 4 6 3 Bogus Creek (Mlamathy i ccc: 1 4 4 1 IMMUNE Creeka (Sacramento) mcrcsevecer tee: 2 1 1 5 26 57 47 20 2 made in this table without seriously changing its general appearance, yet the southern trout would have a decided preference for the left side. When compared directly with examples of Salmo irideus which are at hand—specimens from Coyote Creek and the Pajaro River, to 1926] Snyder: The Trout of the Sierra San Pedro Martir 425 be more explicit—it appears that the representatives of S. nelsoni are somewhat more deep, heavy and less trim, the caudal peduncle is deeper, the caudal fin perhaps less deeply notched, the head longer and somewhat blunter, the eye much larger, and the fins somewhat stronger. These very characters are subject to much variation in S. trideus, however. TABLE 5 SALMO NELSONI Ist year 2d year 3d year 4th year 87 135 107 134 79 125 86 135 99 145 100 124 95 138 162 106 163 200 93 166 185 94 164 175 61 146 202 92 164 190 63 145 195 94 165 185 fal 148 165 75 137 160 63 134 175 67 144 195 83 132 189 200 101 172 199 210 In Salmo nelsoni it is found that an inerease in the size of the head becomes apparent during the third year of age, as determined by a microscopical examination of the scales. In this connection it will be of interest to mention also that growth seems to be more rapid here than in the trout found farther north. For illustration, two tables are presented, the first (table 5) including the calculated stature of twenty representatives of nelsont at given ages, and for comparison, a second table (6) including the same data for ten examples of tridews from the Klamath River. The oldest specimens of nelsoni in the col- lection are in the fourth year of age. Most of the others are in the third year, a few are in the second, while two individuals measuring 69 and 95 millimeters respectively are clearly fishes of the year. They 426 University of California Publications in Zoology [Vou. 21 were all collected April 24 to 27, 1925. It will be noted then, that in each case the growth of the last year was interrupted in the latter part of April. The specimens of S. wideus had practically completed the second year. The lengths are recorded in millimeters. TABLE 6 SALMO IRIDEUS Ist year 2d year 41 95 82 152 Tl 145 79 137 55 139 56 105 48 114 59 117 44 104 48 122 NaturAL History MuSEuUM, LELAND STANFORD JUNIOR UNIVERSITY. Transmitted January 29, 1926. "UNIVERSITY. OF CALIFORNIA PUBLICATIONS — (Continued). - Dixon. Pp, 339-354, plate 11, 1 figure in text. January, 1924 20.0... i tranixes, during Growth in the ‘Skull of the Rodent. Otospermophilus ae. Ra Bed deh eke beecheyi, by. EB. "ci hema Hall. Pp. 355-404, 43 figures in HERR 2s ped arch, ASZ6. AC a Saati ee IAP ew Ba iL lr ee es a Se ata Nie as fi 5 ee Le A New -Race of. the ‘White-breasted. Nuthatch from Lower California, by Joseph Grinnell, Pp: 405-410. March, CA Re oie Re MASK ae REECE REDS ie - Otterbein Snyder. Pp. 419-426, March, 1926 Las Ge ae SC a Se Se se Volume 22, 1920-1923, 485 pages, with 24 plates. Se ie Vol, 23.” “fhe Marine Decapod Crustacea of California, by Waldo L. ‘Schmitt. Pp. Coa 4-470, plates 1-50, 165 figures in text. May, 1921.00 _ Vol 24, LA Geographical Study of the Kangaroo Rats of California, by ‘Joseph Grinnell. Pp. 1-124, plates 1-7, 24 figures in text, June, 1922 1.200002 2, Birds and Mammals of..the: Stikine River Region of Northern “Biitish ~~... Columbia and Southeastern Alaska, by H. 8. Swarth. Pp. 125-314, plate = GiiSt feares ii text... Sune. 19Ue ee a es a : 3 Birds and Mammals of the Skeena River Region of. Northern British . 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Grinnell and J oseph Dixon, Pp. 411-417, °9 figures a PN GORG.: Mare R SS TORG sn Leon aia UN eek ear care é 17, The Trout of the Sierra San Pedro Martir, Lower ‘California, By I ohn ~ os om Vol. 28. 1, Mitochondria and Golgi Bodies in Endamoeba gingivalis (Gros) Brumpt, a Mitochondria in” re gg brasiliensis, by David Causey. Pp. 19-28, - AO 7. Some New and Some Previously Unreported Hydroids, Mainly to the ‘Californian Coast, by ©. McLean Fraser. Pp. 167-172, 7 Benres in Met BISCO DOR, FO 20 es hisses ase eee aa Sa Sect dela ge Ml a wiclang Circus Sap oun apie gingivalis. (Gr08)- in vitro, by Beatrice Fay Howitt. Pp. 173-182," 9. Experiments with Hndamoeba gingivalis (Gros). in Mixed Bacterial Cul- --tures} in Filtered Saliva; ‘on a Solid Base; and with Peritoneal Cells; - and with Digestive Seid by Beatrice Fay Howitt. Pp. 183-202, plates 16 and. 17 : Nos, 8 and. 9 in one cover. Pernaty, 198023 Sr 10. On Oxyphysis ‘omytoxoides fen nov., sp. nov., A Dinophysoid Dinoflagellate Convergent toward. the Peridinioid Type, by Charles A, YOR: Pp, 208-216, plate 195," February, 1926 cs an icine eaten teas ecount ob costetntnentnnstuee - TL. Mitochondria in Huatene gracilis Kiebs, by David Cansey. Pp: 217-224, 12. - Mitochondria in Noetiluca scintillans. (Macartney, 1810), has David Causey. ; . Pp. 225-2380, plate 21. © 8. The Effect. of Certain Drugs and Dyes upom the Growth of Endamocba_ 25 -25 -- 16. On Proboscidiella multinucleata gen, nov.,. ‘sp. nOv., from Planocryptotermes - ees. Mitochondria in Gritavos with ‘Eopectal 3 agendind: to > Pordmecium Eh, - by. David Causey. - Pp. 231-250, plates 22-24. eh pact ‘Nos. 11, 12, and 18 in one cover, February, 1926 .. 14. Studies. on the ‘Ingestion of. ‘Leucocytes, ‘and. on ‘Mitosis, in “gingivalis, by Horace J. Child. Pp.:251-284, plates 26-29, ha irs, in. ‘text. = February, 1926 coset tae ie ens boneee ip Naneoantgmcneen see REESE ee ae ets “16. ‘On Onymonas, @ Plagellate with an Extensile and ‘Retractile ope a Swezy. Pp. 285-300, ‘plate 30, 5: figures in: text Selig teen age becene poetics is » 'nocens-From. the Philippine Islands, by Charles A. ore and eh -\ Swezy.. Pp. 301-316, plates $1, 32, 4 figures in. text . ~ Nos. 16 and 16 in one cover. February, 1926 .. ‘ 17. On the Cestode Genus Dipylidium from Cats and Dogs, by Theresa ‘Marie y : Milizner. --Pp. 317-356, plates 83-89. February, FODG iss ete ee -18. A Useful Modification of a Olearing Fluid Formulated by Spalteholz, ge = -.. Pranklin P. Reagan... Pp. 357-359. = = Bes 19. The Harliest Blood Vessels of the Mammalian Embryo,. Studied iby Means _ 1) =>. of the Injection, Method, by Franklin P, Reagan. Pp. 361-36 ss ~~ Nos. 18 and 19-in one cover.. February; 1926 es Re ep ere ‘ies 20. The Biological. Relationships of Leishmania and. Certain pcan an ~~. by. Edna Hannibal Wagener : and Dorothy” Ann Koch. - PRE 365-888, st n rian as Kereta 1926 ee sorte soriesconasttenargetotaceeuronecsenseocatiercasernentes ee 4 Lie oye AR Sy Pe Tests eS a “UNIVERSITY OF. CALIFORNIA ‘PUBLICATIONS. ee - Note,—The University of California Publications are offered in exchange for the Satie cations of learned societies and institutions, universities and libraries. Complete lists of all the publications of. the University will be sent upon request. For sample copies, lists... of publications or other information address the MANAGER OF THE UNIVERSITY _ PRESS, BERKELEY, CALIFORNIA, U.S.A. All ‘matter sent in exchatige should be —— addressed to EXCHANGE DEPARTMENT, UNV EReES, TAEEA RY, See TA: CALIFORNIA, U.S. A. 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Revision of the Avian Genus Passerella, with Special Refererice to the iv Distribution: and Migration of the Races in California, by H. S.. Swarth. Fe Peart ‘Pp. 75-224, plates 4-7, 30 figures in text. September, 1920 ........-2.-..-. ‘1.76. 5. A Study of the California Jumping Mice of the Genus Zapus, by A. Brazier tas Howell. Pp. 225-238, 1 figure in text. May, 1920-0 : #15 6. Two New. Rodents (Genera Thomomys and Marmota) from the Eastern — Border of California, by Joseph Grinnell Pp. 239-244, 6 figures in text | coo Woven ben, Ao br cy a LE ment gt aay WAR oma 7. A Study of the Californian Forms of the Microtus montanus Group of ; ‘ Meadow: Mice, by Remington Kellogg. Pp, 245-274, 25 figures in text. 8. A’ Synopsis of the Microtus mordaz Group of Meadow Mice in California, py Remington Kellogg. .Pp. 275-302, plate 8, 29 figures in text. ee Gee ee Nos. 7 and 8 in one cover. April, 1922 2. cn. ecennenneneenncnettemenes PY (7 Pah 9. Behavior of the Leaf-nosed Snake, Phyllorhynchus ° deourtatus, by Sarah BAR Rogers Atsatt. Pp. 803-312... February, 1928 2. ineciceseeee tects BO 10..A Systematic List of the Mammals of California, by. Joseph Grinnelh “at Pp, 313-324, “Tamuary, LOLS nea annensnencecnerarnccecaernesnnnnneannnhsnensnat netmrsetcees er Te AL. The Systematic Status of the, Mountain Lion of California, by Joseph Le ae Grinnell and Joseph Dixon. Pp. 325-332, plates 4 9-10. Pati: ecu Bs Bey * +, > Sole MAIC REVIEW OF THE -PACIFIC ‘COAST BROWN TOWHEES BY JOSEPH GRINNELL ann HARRY S. SWARTH UNIVERSITY OF CALIFORNIA PUBLICATIONS IN ZOOLOGY Volume 21, No. 18, pp. 427-433, 2 figures in text Issued April 6, 1926 UNIVERSITY OF CALIFORNIA PRESS BERKELEY, CALIFORNIA CAMBRIDGE UNIVERSITY PRESS LONDON, ENGLAND SYSTEMATIC REVIEW OF THE PACIFIC COAST BROWN TOWHEES BY JOSEPH GRINNELL ann HARRY S. SWARTH (Contribution from the University of California Museum of Vertebrate Zoology) Pipilo fuscus sencula Anthony was described from San Fernando, north-central Lower California. The acquisition of a series of brown towhees from that same general region (between latitudes 30° and 31°), and notably among these birds some taken in September and October, hence in fresh fall plumage, enables us to make comparison of birds from various parts of upper California with typical senicula, such as we, ourselves, have not before been able to do. Heretofore the assumption has been that the brown towhee of southern California has been of the subspecies senicula; but comparison of the material now available leads to a different view. Incidentally, examination of the Lower California birds has led to comparison with an exten- sive series from the near vicinity of Monterey, topotypes of Fringilla crissalis Vigors, which compels a further readjustment of our ideas regarding some of the California subspecies of this bird. In the deseription of Pipilo fuscus sencula, Anthony (1895, p. 111) aseribes southern California birds to that subspecies though at the same time noting slight differences in measurements. His Lower California specimens (at least those mentioned) were collected at seasons when fading and wear would to some extent have reduced color differences that are outstanding in fresh plumage. Compared with specimens from San Diego County our fresh-plumaged Lower Cali- fornia birds are slightly smaller throughout, a difference that at first glance is most apparent in size of bill. In body color they are darker and generally more slaty, as compared with the brownish-tinged upper California skins, and the ecrissum is a darker tone of reddish. The dark type of coloration extends to the remiges and rectrices. In life, the black tail of a flying senicula proved to be a character that impresses an observer who is used to seeing the more northern birds. 428 Unversity of Califorma Publications in Zoology (Vou. 21 The generally dark color of these birds extends even to the ‘‘soft parts,’’ for the feet of nearly all prepared specimens of senicula are appreciably blackish rather than the translucent light brown seen in specimens from San Diego and northward. In any discussion of the brown towhees of California it is neces- sary first to determine the proper application of the name Fringilla crissalis, This name was applied by Vigors (1839, p. 19) to a speci- men from Monterey. In the American Ornithologists’ Union Check- list (1910, p. 281) the type locality is given as ‘‘probably San Fran- Fig. 1. Bills of subspecies of Pipilo fuscus; natural size. a. Pipilo fuscus senicula, § ad.; no. 46393, Mus. Vert. Zool.; San José, Lat. 31°, Lower California, Mexico; September 27, 1925. b. Pipilo fuscus crissalis, § ad.; no. 27122, Mus. Vert. Zool.; Lytle Creek Wash, 2 mi. NE. Grapelands, San Bernardino County, Calif.; November 6, 1916. c. Pipilo fuscus albigula, g; no. 37433, coll. L. B. Bishop; La Paz, Lower California, Mexico; October 16, 1924. d. Pipilo fuscus petulans, 3; no. 36439, Mus. Vert. Zool.; Palo Alto, Santa Clara County, Calif.; January 26, 1901. cisco or Monterey’’; but inasmuch as Sharpe (1888, p. 754) lists the type specimen as in the collection of the British Museum, and as from Monterey, that locality may be accepted as fixed. In a previous study of the group by the junior author of the present paper (Swarth, 1918, pp. 117-121) the name crissalis was assumed to belong to the reddish colored form peculiar to the central coast region, an assumption in which he followed all previous writers on the subject. The specimens then available from the general region of the type locality of crissalis (Monterey) prove, however, not to have adequately represented the 1926] Grinnell_Swarth: Pacific Coast Brown Towhees Fig. 2. Distribution on the Pacific coast of the subspecies of Pipilo fuscus (see p. 481). 429 430 University of California Publications in Zoology [Vou 21 race of that section. They were few in number and were in worn plumage. Since that time the Museum of Vertebrate Zoology has acquired an excellent series of brown towhees from the coast of Monterey County, including fifteen fresh-plumaged skins from Seaside, immediately north of the town of Monterey. The fact now perfectly plain is that these Monterey specimens are practically indistinguish- able from southern California birds; they are not of the reddish type of coloration that distinguishes the brown towhee of the San Fran- cisco Bay region. The name Pipilo fuscus crissalis therefore applies to the brown towhee of southern California, from Monterey south at least to the Mexican boundary. Thus the brown towhees of the imme- diate coastal region are divided as follows: Senicula in north-central Lower California, not extending north of the international boundary ; crissalis ranging from about the boundary line north to Monterey ; and from Monterey northwards a reddish colored form that is as yet unnamed. The last-mentioned race we propose to eall: Pipilo fuscus petulans, new subspecies San Francisco Brown Towhee Type—Male; no. 36439, Mus. Vert. Zool.; Palo Alto, Santa Clara County, California; January 26, 1901; collected by J. Grinnell; original no. 4576. Distinguishing characters.—The reddish tone of general coloration is the most noticeable feature of this subspecies; it exhibits the extreme of ruddiness seen in all the brown towhees. In size it is intermediate between the interior race carolae and the southern coast form crissalis. (See accompanying table of measurements, and Swarth, 1918, p. 121.) Range.—The north-central coast region of California from the vicinity of Santa Cruz north to Humboldt Bay; in typical form, in the immediate vicinity of San Francisco Bay. The few specimens available from Humboldt County, while referable, we think, to the present form, are appreciably less ruddy and slightly larger, thus showing a tendency toward the condition in carolae. South of the San Francisco Bay region, there is intergradation with crissalis in northeastern Monterey County. Specimens from the Salinas Valley, at the same latitude as Monterey but some twenty miles inland, show a decided approach toward the reddish coloration of petwlans. It further develops from our renewed study of the brown towhees that the seemingly rather isolated colonies of this species resident in the interior valleys of southern Oregon differ sufficiently from the subspecies carolae as now restricted to warrant naming. This we do as follows: 1926] Grinnell-Swarth: Pacific Coast Brown Towhees 431 Pipilo fuscus bullatus, new subspecies Oregon Brown Towhee Type.—Male; no. 46578, Mus. Vert. Zool.; Eagle Point, Jackson County, Oregon; January 9, 1926; collected by William E. Sherwood. _ Distinguishing characters—At the maximum size of the races of Pipilo fuscus; the bill, in particular, long and heavy. Coloration, tending to slate gray, a shade that becomes accentuated in worn spring plumage ; sides and flanks more heavily and extensively saturated with brownish slate than in carolae. Range.—Known only from the Upper Sonoran valleys of Josephine and Jackson counties, southwestern Oregon. The habitat of bullatus is apparently entirely separated by mountainous territory from the northern boundary of carolae, at the upper margin of the Sacramento Valley. Remarks.—Specimens from Kerby, much nearer the coast than the other localities represented, are slightly more tinged with brown, though at that they are more grayish than carolae. (For comment upon the Kerby specimens, see Grinnell, 1912, p. 199.) Birds from Ashland, Eagle Point, and Trail exhibit the extreme of slatiness seen in the brown towhees of the Pacific coast. In nomenclatural treatment of the Pacific coast forms of the brown towhee we are following the procedure advocated by Oberholser (1919, p. 210), who asserts the existence of intergradation between the fuscus and crissalis groups, usually heretofore regarded as specifically dis- tinct. His claim of intergradation between senicula of northern Lower California, and albigula’ of the Cape San Lucas region, through the subspecies aripolius (Oberholser, loc. cit.) of middle Lower California, is corroborated in statements by Thayer and Bangs (1907, p. 140). The Pacific coast races of the brown towhee now stand as follows, listing them from north to south. Reference should be made to the accompanying map for approximate geographic allocation. 1. Pipilo fuscus bullatus Grinnell and Swarth. Oregon Brown Towhee. 2. Pipilo fuscus carolae McGregor. Sacramento Brown Towhee. 3. Pipilo fuscus petulans Grinnell and Swarth. San Francisco Brown Towhee. 4. Pipilo fuscus crissalis (Vigors). California Brown Towhee. 5. Pipilo fuscus senicula Anthony. Anthony Brown Towhee. 6. Pipilo fuscus aripolius Oberholser. San Pablo Brown Towhee. 7. Pipilo fuscus albigula Baird. San Lucas Brown Towhee. 1 According to Oberholser (loc. cit., footnote) this name should take the form albigulus. However, Baird named the bird Pipilo albigula; even though albigulus be correct, the fact is that albigula is what the original describer called it, and that is also correct! [ Von. 21 Zoology Publications in ua ty of Californ Wersr Un 432 (¢ 61-061) 361 (0'1Z-0'61) 2°61 (002-0 ST) 2 61 (0 02-¢ 9T) 2ST (0'SI-S 91) O'LT ABO JNOGIIM 90} B[PPIy (Z 66-0 LZ) 2 '8Z (S 62-0 '8Z) 9°S8Z (0 62-¢ LZ) I’ 8% (0'8Z-G EZ) 8°9z (G°L-0'G2) F'9% snsie J, (Z°91-0'FT) TST (0° 91-0'ST) FST (G’FI-G ST) I FI (0°SI-@ FI) 2°F1 (O0'FI-G ZI) 2 SI (¢ OLI-0 801) 2°601 (0 OLI-¢ 86 ) Z°SOI (0 III-O0 ZOT) 8 SOT (0 60I- ‘T0T) 9° 90T (0 OII-0 001) F 0T “UOSII() ULOJSOMYYNOS ULOI} SoTBUI g snjpjyyng snosnf ondig tee ene e ene e ween ene e esas teecreens, BIULOJI[BO ‘sory -unod ULIBI, puw BUuIOUOg ‘Bpou -B[ Vy ‘eie[Q BURG UIOLJ] Sa[BUl (CT supjnjad snasnf opndig FEES HE CAGOD IE MOPS OACOSIOT BIULOJI[BO ‘Aqunog Aogioyuoyy ‘VpIsveg UlOIJ SeTVUl 6 ssypssila snosnf opdid Die Rare Nee nae eieaneungess BIUIOPTEO ‘Aqunoy OsaIq UBg WIOIJ So[VUl OT 2 isypssiia snosnf opdig Sisco sah coe Seana BIUIOpTRD IIMO'T [B1JU90-Y}LOU UOLJ SOTBUL OT ‘pynavuas snasnf opidig usul[ND Te (2°66 -0'¥6) $°26 (¢ OOI-E 16) &°S6 (0°66 -0°16) 6 #6 (0° 6 -¢ 06) 6 26 (0°6 -0°28) $06 Bul snosnf opdrg 0 SHINTdSANG LSVOH olMIOVd NIVLUGD dO ‘SUMLAWITII NI ‘(WOAWIXVJT GNV WOWINIJY “ADVaEAY) SLNAWAYOSVaIN 1926] Grinnell-Swarth: Pacific Coast Brown Towhees 433 LITERATURE CITED AMERICAN ORNITHOLOGISTS’ UNION COMMITTEE 1910. Check-list of North American birds. Ed. 3 revised (New York, Am. Orn. Union), 430 pp., 2 maps. ANTHONY, A. W. 1895. Description of a new Pipilo from southern and Lower California. Auk, 12, 109-112. GRINNELL, J. 1912. The northern brown towhee. Condor, 14, 199. OBERHOLSER, H. C. 1919. Description of a new subspecies of Pipilo fuscus. Condor, 21, 210-211. SHARPE, R. B. 1888. Catalogue of the Passeriformes or perching birds, in the collection of the British Museum. Fringilliformes: Part III. Cat. Birds British Mus. (London), 12, xvi + 872, 16 pls. SwartH, H. S. 1918. The distribution of the subspecies of the brown towhee (Pipilo crissalis). Condor, 20, 117-121, 2 figs. in text. THAYER, J. E., AND BANGs, O. 1907. Catalog of birds collected by W. W. Brown, Jr., in middle Lower California. Condor, 9, 135-140. Vicors, N. A. 1839. Ornithology, in Zoology of Captain Beechey’s voyage (London, Henry G. Bohn), pp. 13-40, pls. 3-14. Transmitted February 15, 1926. bs A Be ee ti hae \ i e uh an ” “i WV INDEX Univ. Calif. Publ. Zool., vol. 21. Titles of papers and new systematic names are in boldface; synonyms are in italics. Acknowledgments, 2, 54, 81, 225, 276, 311, 356, 405, 417. Arvicola montana, 145, 262. longirostris, 263. Atsatt, Sarah R., 303. Behavior of the Leaf-nosed Snake Phyllorhynchus decurtatus, 303. Boving, Dr. A., acknowledgment to, 250. Brown Towhee, list of races on Pacific Coast, 431. Oregon, 431. San Francisco, 430. Brown Towhees, of the Pacific Coast, Systematic Review of the, 427. Bunting, Unalashka, 85. ' California, A New Race of the White- breasted Nuthatch from Lower, 405. A Synopsis of the Microtus mordax group of Meadow Mice, in, 275. Five New Five-toed Rats from, 43. Jumping Mice of the Genus Zapus, A Study of the, 225. Notes on the Natural History of the Bushy-tailed Wood Rats from, 49, Revision of the Avian Genus Passe- rella with Special Reference to the Distribution and Migration of the Races in, 75. Revision of the genus Lynx in, 339. Systematic List of the Mammals of, 313. Systematic Status of the Mountain Lion of, 325. . The Trout of the Sierra San Pedro Martir, Lower, 419. Two New Rodents (Genera Thomo- mys and Marmota) from the Eastern Border of, 239. Changes During Growth in the Skull of the Rodent Otospermophilus grammurus beecheyi, 356. Dixon, J., 49, 325, 339, 411. Emberiza rufina, 87. unalaschcensis, 85. Hees Cr S:, cited; 327. Felis oregonensis californica, 325. Fox Sparrow. See Sparrow, Fox. Fringilla crissalis, 428. ferruginea, 87. iliaca, 84. meruloides, 85. 256, [435] TUSA, Sls townsendi, 85. Gopher, Pocket, Armagosa, 239. Grinnell, J., 43, 239, 313, 326, 339, 405, 411, 427. Ground Squirrel. See Squirrel, Ground. Hall, E. R., 356. Howell, A. B., 225. Jumping Mouse. See Mouse, Jumping. Kangaroo Rat. See Rat, Kangaroo. Kangaroo Rats, from California, Five New Five-toed, 43. Kellogg, R., 1, 245, 275. Lamb, C. C., cited, 419. Leaf-nosed Snake, Phyllorhynchus decurtatus, Behavior of the, 303. Lower California, A New Race of the White-breasted Nuthatch from, 405. The Trout of the Sierra San Pedro Martir, 419. Lynx, California, 339; external varia- tions, 341; distribution, 345; measurements, 351. Lynx ruffus baileyi, 349. ealifornicus, 346. fasciatus, 348. pallescens, 350. Lynx, in California, Revision of the Genus, 339. J Mammals of California, A Systematic List of the, 313. Marmot, White Mountains, 242. Marmota flaviventris fortirostris, 242; measurements, 242. Martes caurina humboldtensis, 411; measurements of skulls, 416. vancouverensis, 414; measure- ments of skulls, 416. Meadow Mouse. See Mouse, Meadow. Merriam, C. H., cited, 325. Microtus californicus Group of Meadow Mice, a Revision of the, 1 Microtus californicus aestuarinus, 3, 15; measurements, 39. ealifornicus, 2, 5. constrictus, 2, 9. eximius, 2, 12; measurements, 39. huperuthrus, 3, 34; measurements, 39. kernensis, 3, 26; measurements, 41. mariposae, 3, 19; measurements, 40. w (0CT28 1926 * ah TONAL muse> Index mohavensis, 3, 29; measurements, 41. neglectus, 3, 42. scirpensis, 3, 40, vallicola, 3, 40. edaxz, 15. montanus, diagnostic criteria, 246; variation, 246; dentition, 249; enamel patterns, 251, 252, 254, 255; key to subspecies, 257; distribution in California, 258. montanus, 259; type locality, 257. arizonensis, type locality, 257. earyi, type locality, 257. dutcheri, 268; type locality, 257. montanus, 259; type locality, 257. rivularis, type locality, 257. yosemite, 264; type locality, 257. Microtus montanus Group of Meadow Mice, A Study of the Californian forms of the, 245. Microtus mordax, individual variation, 26; enamel patterns, 277, 278, 279; age and sex variation, 281; key to 91. vo ) measurements, 24; measurements, 21; ; measurements, subspecies, 282; distribution in California, 283. angusticeps, 293; type locality, 282. bernardinus, 296; type locality, 282. mordax, 284, 288; type locality, 282. sierrae, 288; type locality, 282. Microtus mordax Group of Meadow Mice in California, A Synopsis of the, 275. Miller, G. S., acknowledgment to, 249; cited, 327. Mountain Lion, California, 325. Mountain Lion of California, The Systematic Status of the, 325. Mouse, Jumping, Allen, 232. Humboldt, 229. Point Reyes, 231. Warner Mountain, 235. Mouse, Meadow, Armagosa, 24. Cantankerous, 284. Cape Mendocino, 9. Kern River, 26. Lower California, 34. Mariposa, 19. Mohave River, 29. Mount Whitney, 268. Northwest Coast, 293. Owens Valley, 21. Peale, 259. San Bernardino, 296. Sanhedrin, 12. Sierra Nevada, 288. Southern California, 31. [436] Tule, 15. West Central California, 5. Yosemite, 264. Neotoma cinerea, in California, fossil forms, distribution, behavior, popu- lation, ete., 57-67. cinerea, 50; measurements, 55. occidentalis, 56; measurements, 55. New Race of the White-breasted Nuthatch from Lower Califor- nia, 405. New Subspecies of Birds from Pata- gonia, 333. Notes on the Natural History of the Bushy-tailed Wood Rats of Cali- fornia, 49. : Nuthatch, from Lower California, A New Race of the, 405. Nuthatch, San Pedro Martir White- breasted, 405. Otospermophilus grammurus beecheyi, morphological discussion of, 356— 390; criteria for determining age groups, 403. Otospermophilus grammurus beecheyi, Changes during Growth in the Skull of the Rodent, 356. Passerella, history of the genus, 84; variation in, 87; distribution and migration, 101. Passerella, Revision of the Avian Genus, with Special Reference to the Distribution and Migra- tion of the Races in California, 75. Passerella iliaca altivagans, 87; de- scribed, 119; listed, 185; measure- ments, 182. annectens, 86; described, 140; listed, 194; measurements, 182. brevicauda, 87; described, 165; listed, 204; measurements, 183. canescens, 87; described, 169; listed, 205; measurements, 183. fuliginosa, 86; described, 149; listed, 199; measurements, 182. fulva, 87; described, 158; listed, 201; measurements, 183. iliaca, deseribed, 114; listed, 184; measurements, 182. insularis, 86; described, 132; listed, 188; measurements, 182. mariposae, 87; described, 176; listed, 209; measurements, 183. megarhynchus, 86; described, 161; listed, 202; measurements, 182. monoensis, 87; described, 170; listed, 206; measurements, 183. obscura, 87. schistacea, 86; described, 152; listed, 200; measurements, 182. Index sinuosa, 87; deseribed, 135; listed, 189; measurements, 182. stephensi, 87; described, 176; listed, 209; measurements, 193. townsendi, described, 144; listed, 197; measurements, 182. unalaschcensis, deseribed, 127; listed, 186; measurements, 182. Pemberton, J. R., collection of Pata- gonian birds, 334. Perodipus elephantinus, 43. swarthi, 44. dixoni, 45. leucogenys, 46. monoensis, 46. Phyllorhynchus decurtatus, 303; motor impulses of, 3804-307; feeding, 310. Pine Marten, distribution in Califor- nia, 415. Humboldt, 411. Vancouver Island, 414. Pine Marten, from the Pacific Coast of North America, Two New Races of the, 411. Pipilo fuscus, distribution of sub- species on the Pacific Coast, 429. Pipilo fuscus bullatus, 431. petulans, 430. senicula, 427. Plectrophanes townsendi, 85. Pocket Gopher, Amargosa, 239. Pteroptochos rubecula nemorivaga, 333. Rat, Kangaroo, Carrizo Plains, 44. Hlephant-eared, 43. Mereed, 45. Mono, 46. Pale-faced, 46. Rat, Wood, Gray Bushy-tailed, 50. Western Bushy-tailed, 56. Rats, from California, Five New Five-toed Kangaroo, 43. Rats, Bushy-tailed Wood, of Califor- nia, Notes on the Natural His- tory of, 49. Revision of the Avian Genus Passe- rella, with Special Reference to the Distribution and Migration of the Races in California, 75. Revision of the Genus Lynx in Cali- fornia, 339. Revision of the Microtus californi- cus Group of Meadow Mice, 1. Salmo nelsoni, 419; measurements, 420. irideus, 424; measurements, 420. Sierra San Pedro Martir, Lower Cali- fornia, Trout of the, 419. Sitta carolinensis alexandrae, 405. Skull of the Rodent Otospermophilus grammurus beecheyi, Changes during Growth in the, 356. [437] Snyder, J. O., 419. Sparrow, Fox, Hastern, 114. Kadiak, 132. Mono, 170. Shumagin, 127. Sooty, 149. Thick-billed, 161. Townsend, 144. Valdez, 135. White Mountains, 169. Yakutat, 140. Yolla Bolly, 165. Yosemite, 173. Spizitornis parulus lippus, 336. Squirrel, California Ground, 356. Study of the Californian Forms of the Microtus montanus Group of Meadow Mice, 245. Study of the California Jumping Mice of the Genus Zapus, 225. Swarth, H.S., 75, 427. Synopsis of the Microtus mordax Group of Meadow Mice in Cali- fornia, 275. Systematic List of the Mammals of California, A, 313. Systematic Review of the Pacific Coast Brown Towhees, 427. Systematic Status of the Mountain Lion of California, 325. Taenioptera pyrope ignea, 334. Thomomys perpallidus amargosae, 239; measurements, 240. Towhee. See Brown Towhee. Towhees, Brown, Systematic Review of the Pacific Coast, 427. Trout of the Sierra San Pedro Mar- tir, Lower California, 419. Turdus magellanicus pembertoni, 335. Two New Races of the Pine Marten from the Pacific Coast of North America, 411. Two New Rodents (Genera Thomo- mys and Marmota) from the Eastern Border of California, 239. Wetmore, A., 333. Wildcat, California, 346. Desert, 349. Northwestern, 348. Pallid, 350. Wood Rat. See Rat, Wood. Zapus, A Study of the California Jumping Mice of the Genus, 225. Zapus, cranial differences, 226; molt and coloration, 226; distribution in California, 227. major, 235; measurements, 237. orarius, 231; measurements, 237. pacificus alleni, 232; measurements, 937 trinotatus eureka, ments, 237. 229; measure- Oy eh PO iL ie Ay ea mi | Ny i PA ie , OF aa i cn : HM oun ibe mnie Ann ay hal Ue at ' me “ F; y ti AY y a nae ! ier en ae ¥ eve ‘ i i) mat } i xt \ * feat sites) J Ae 7 ig ek _ UNIVERETTY OF CALIFORNIA PUBLICATIONS (Continued) A ghee ee es Agee tT My RN RY ua SAR PRICE DR LE SD Ae San eel Ne ‘Dixon. | Pp, 339-354, plate 11, 1 figure in ‘text. January, 1924 0 eus RS 3 mares beecheyi, by E, Raymond Hall. Pp. 856-404, 43 figures: in ‘text. Sana pe March, 1926.2... Opals PRE ep Bont lara Bs as Si ATG HD ISLE PET Ap Day RENO Gd ME Vg A RAR 15, A New Race, ‘of: the White-breasted Nuthatch from Lower California, 16. Two New Races of the Pine Marten from the Pacific Coasti of ' North ‘ America, by Joseph Grinnell and Joseph ranean Pp, 411-417, 9 figures ; in text. March, go, py ae sneer Oey SRD Sk tea Bis rhea cine Sate wD oe he ANON ap eetiea wae 17. The Trout of the Sietra San Pedro Martir, Lower Catitomia,, by John > Otterbein: Snyder. © Pp, 419-426, March, 1926 00000 1-470, Plates 1-50, 165 figures in text. . May, 1921 Pac Wee ag AT Mee Wars Ae Ni Grinnell.” Pp. 1-124, plates 1-7, 24 figures in text. June, 1922 2000. 2, Birds and “Mammals of the Stikine River: Region of. Northern British Columbia and Southeastern Alaska, by H..8, Swarth. - Pp, 125-314, plate: 8; S4-cures in: text.; ine, 10e2 se oe ee ee “3. Birds and Mammals of the Skeena River Region of Northern British Columbia, by Bey 8. Swarth.. Pp. 315-394, plates 9-11; 1 figure in text. PARUATY, TIA ea ca ct eeons chara tgranpeensewinctbemsesinseenlintantnegnees 4, Report on a Collection of Birds Made by’ J. BR. Pemberton in Patagonia, hy Alexander Wet rire: Pp. 395-474, plates 12-14, 11 figures in text. “Wol 25. A Bibliography of Eugenics, by Samuel z Holmes. Pp, 1-514: ee nash 1024 plates 1-18, 42 figures.in text. June, 1926 oceans ectnee by David Causey. Pp. 1-18, plates 1-3, sie 4, Nos. 1 and 2 in one cover. N lovemiber, RODE Ss ene ct nS oS Oe eee hole Richard P, Hall. Pp. 29-64, plates 5-9, 5 figures in text. November, 1925. ess 4. ‘The Cultivation of Endamoeba gingivalis (Gros), by Beatrice Fay Howitt, Pp. 65-126, plates 10-12, 4 figures in text. ‘November, 1925 000.00. 6, The Behavior of Endamoeba dysenteriae in Mixed, Cultures. with Bacteria, ASRS vi a A ‘Kofoid aut Edna panto aiecncmar ee Pp. 127-154, plates i Nos. 6 and 6 in one cover. Novemiber, 1926 Wolume 26, 1923-1925, 453 pages, 84 plates . PE BIDE NS MAD Ein 8 SEN Sy TEAM E Oa PR Me BSBA aD LNG * Be Vol, 27. A Synopsis of the Amphibia of California, by Tracy I. ‘Aisles Pp. 1-949, 2s Vol. 28. 1. Mitochondria and Golgi Bodies in Hndamoeba gingivalis (Gros) Brunpt, | & Mitochonaria in Leishmania brasiliensis, by David. Causey. Bp, 19-28, . er ‘ ee Mitosis in Ceratium hirundinella.O. F. M., with Notes on Nuclear Pheriom- : : ena in Encysted Forms.and the Question of Sexual Reproduction, by — oh ae : a ae Some New and Some Previously Unreported Hydroids, Mainly from the SF AE ; Californian Coast, by C. ‘McLean Fraser. Pp. 167-172, 7 figures in text. 9%. Experiments with Endamoeba. gingivalis (Gros) in Mixed’ Bacterial Cul- _ hF : “10. On Oxyphysis oxytoxoides gen nov., sp. nov., A Dinophysoid Dinoflageliate - A ae ORR ase 203-216, plate 19, February, UCR Eset REN ABD A ts a 1 B 60 Vol 24. LA Geographical ‘Study of the Kangaroo. Rats. of ‘California, by. Joseph ganed I@c Shedies OF sib. Eiecie OF Wertaln “Diuen tigon. idemoela. Queenterias nis vitro, by Charles A, Kofoid and Edna Hannibal: Wagener. Pp. 165-166. 40 and with Digestive Setretions, by. Beatrice Gish Howitt... Pp. 183-202, ey Wr td he ¥ gh) > eae) Tht F ai ‘ ih : ig Se ek Te Te NYY i a ' Peek ; are ae F Og) 4) UNIVERSITY OF CALIFORNTA PUBLICATIONS (Continued) ‘U1. Mitochondria in Euglena gracilis ‘Blebs, ‘by David Causey. Pp. a plates 19-20. °°.» » 12. Mitochondria in Noctiluca sointillang (Macartney, 1810), by David Causey. a Pp. 225-230, plate 21. 3. Mitochondria’ in’ Ciliates. with. Especial: Reference to ‘Paramecium Ebr, _ by David'Causey. Pp. 231-250, plates 22-24, 7 Nos. 11, 12, and 18 in one cover. February, 1926 0a 14. Studies on the Ingestion of Leucocytes, and..on Mitosis, - in Endamoeba — - gingivalis, by Horace J. Child. Pp. 251-284, plates 25-29, 9 figs. ia texte 4 40. 1b. On Oxymonas, a Flagellate ‘with an Extensile and Retractila. ‘Proboseis, ok February, 1926. LS oS Rs Te ee eee ea oe eas from Kalotermes from British Guiana, by Charlies A. Kofoid: ‘and Olive j coh ad PRS, marth ie Loy x bey Nay CER A Tee Bh, Aly? ‘Swezy. Pp. 285-300, plate 30, 6 figures in text. /.:..0202-224.2.. wesabondneegtatbentopete see 16. On Proboscidiella. multinucleata gen, nov., sp. Nov., from Planocryptotermes _nocens From the Philippine’ Islands, by Charles A. ‘Kofoid and Olive ~ i ‘Swezy.. Pp. 301-316, plates 81, $2, 4 figures in text 20.20.0001. sep ot SS igo edi Nos, 15:and 16 in one cover, February, 1926 <2. inc ctt ec cteeseepene Z 17. On the Cestode Genus Dipylidiwm from Cats and Dogs,. by. ‘a scone Marie — Millzner. Pp. 317-356, plates . 83-39. February, 19260. soa eek 18. A Useful Modification of 8 Clearing Fluid Formulated by Spalteholz, by A | The Earliest Blood Vessels of the Mammalian Embryo, Studied by Means er oe a * < of the Injection Method, : by Franklin P. Reagan. Pp. 361-364. ~’ Nos, 18 and 19 in one cover. ‘February, BORG Ay Ce Snes ey 20; The Biological Relationships: ‘of ‘Leishmania’ and Certain ‘Herpetomonads, | by Edna. Hannibal Wagener and Dorothy Ann Koch, Pp. 365-388; ae Caine a suit Be rs plates. 40-43, Avisreh LRG ch iii ai less oe eS Ur le apap 21. Notes on Termites from Arizona with Descriptions of Two New. ‘Species, “by Thomas E. Parner Pp. 980-897, 6 AEE in . text, Mataed TOL iiik es a EO eee ~ CR ra bie Wiehe ipa re Ph its v , Wh Uta ten) nad ee Ma sy Pai hata in ny) Ys) ret a tal SMITHSONIAN INSTITUTION LIBRARIES AIR AO nt aca st { ht t hac} Mh} we " +r ‘: PUNY 5 Af deel} hae pat oe igen Sak Aa vt Pion ie Mae! we Wye Hn OR | math DAROR DLN neh ema SA , Veter at ark a ae eles oleae Rah Re % put rt aesa Sh a fa Sta th Regnier ey ye ory * P 4 rt! ‘oh heal Heres HN Oe 1) ti Coab . 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